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Resources of South-East is a multivolume handbook that aims to summarize knowledge about useful for workers in education, research, extension and industry. The following institutions are responsible for the coor­ dination ofth e Prosea Programme and the handbook: - Forest Research Institute of (FRIM), Karung Berkunci 201, Jalan FRI Kepong, 52109 Kuala Lumpur, Malaysia - Indonesian Institute of Sciences (LIPI), Widya Graha, Jalan Gatot Subroto 10,Jakart a 12710, - Institute of Ecology and Biological Resources (IEBR), Nghia Do, Tu Liem, Hanoi, - University ofTechnolog y (UNITECH), Private Mail Bag, Lae, Papua New Guinea - Philippine Council for Agriculture, Forestry and Natural Resources Research & Development (PCARRD), Los Banos, Laguna, the - Institute of Scientific and Technological Research (TISTR), 196 Phahonyothin Road, Bang Khen, 10900, Thailand - Wageningen Agricultural University (WAU), Costerweg 50, 6701 BH Wage­ ningen, the Netherlands 25 /y«C M Y^ CK a \ ù UG. ?? 'O ^ J Plant Resources of South-

No 4

Forages

L. 't Mannetje and R.M. Jones (Editors)

"\ r Pudoc Scientific Publishers, Wageningen 1992

ItfVi^T^Coâ» Professor L. 'T MANNETJE obtained hisfirs t degree in crop and science from Wageningen Agricultural University in 1958an d his PhD for athesi s about taxonomie studies in the Stylosanthes from University, Bris­ bane, in 1967. From 1958 to 1982 he was a research scientist with the CSIRO Division of Tropical Crops and , Brisbane, Australia. During that period he was seconded to the Malaysian Agricultural Research and Devel­ opment Institute (MARDI) at Serdang, Selangor for a period of one and a half years. In 1982h e was appointed Professor of Grassland Science at Wageningen Agricultural University and in 1983h e became head of the Department of Field Crops and Grassland Science. His publications cover species evaluation, metho­ dology of grassland research and primary and secondary production of tropical . He has visited many tropical countries for consultations, to lecture and to supervise research. R.M. JONES is a agronomist who graduated in Agricultural Science from the University of Sydney in 1958. From 1959 to 1965 he was employed by the Conservation Service, . During that period he spent a year at the University of the Witwatersrand, South , during which time he gained his MSc degree in Plant Ecology. Since 1965 he has been employed by the CSIRO Division ofTropica l Crops and Pastures at the Cunningham Labo­ ratory, Brisbane. He has written extensively on the agronomy of tropical for­ ages, with particular emphasis on the persistence of .

Cip-Data Koninklijke Bibliotheek, Den Haag

Plant

Plant resources of South-East Asia. -Wageningen : Pudoc-Ill., map, drawings. No. 4 : Forages / L. 't Mannetje and R.M. Jones (ed.).Wit h index, réf. ISBN 90-220-1032-5 bound NUGI835 Subject headings: forages / stock breeding ;South-Eas t Asia.

ISBN 90-220-1032-5 NUGI 835

Design: Frits Stoepman GVN.

© Prosea Foundation, Bogor, Indonesia, 1992. © Pudoc-DLO, Wageningen, the Netherlands, 1992.

No part of this publication, apart from bibliographic data and brief quotations embodied in critical reviews, may be reproduced, re-recorded or published in any form including print, photocopy, microfilm, electric or electromagnetic record without written permission from the copyright holders, c/o Pudoc Scien­ tific Publishers, P.O. Box 4,670 0A A Wageningen, the Netherlands.

Printed in the Netherlands. Contents

Editors and contributors 9

Prosea Board of Trustees and Personnel 13

Foreword 15

1 Introduction 17

1.1 Forages and 17 1.2 Sources of 18 1.2.1 18 1.2.2 Annual and perennial crops 20 1.2.3 Crop residues and agricultural by-products 20 1.3 Forage and livestock production systems 20 1.3.1 Current inputs toforage systems 20 1.3.2 Input offorage into different livestock production systems 21 1.4 Main limitations to forage production 22 1.4.1 Climate 22 1.4.2 Soil conditions 24 1.4.3 Species 24 1.4.4 Management25 1.4.5 Socio-economic constraints 25 1.5 Overcoming limitations to improving forage resources 26 1.5.1 Climate 26 1.5.2 Soil fertility 26 1.5.3 Improved species 27 1.5.4 Management and socio-economic constraints 31 1.6 Selection of species to be included in this volume 31 1.7 Species primarily in cultivation as a food crop 32 1.8 Prospects 34

2 Alphabetical treatment ofspecie s 35

Aeschynomene americana : American jointvetch 37 Aeschynomene falcata :jointvetc h 39 Albizia lebbeck : siris 40 vaginalis : alyce clover 42 gayanus : gamba grass 44 : rhizoma peanut 46 Arachispintoi : pinto peanut 48 Arundinaria pusilla :bambo o grass 50 Asy stasia gangetica :rumpu t bunga putih 51 :broadlea f carpet grass 53 : n couch grass 54 Brachiaria brizantha :palisad e grass 56 Brachiaria decumbens :signa l grass 58 Brachiaria dictyoneura :y a signal luey 60 Brachiaria distachya : green summer grass 61 Brachiaria humidicola :koronivi a grass 62 :par a grass 64 :ruz i grass 65 Brachiaria subquadripara : cori grass 67 Calliandra calothyrsus : calliandra 68 caeruleum :caeruleu m calopo 71 : calopo 72 Canavalia ensiformis :jac k bean 74 ciliaris :buffelgras s 77 Centotheca latifolia :juku t kidang 79 Centrosema acutifolium :-80 Centrosema macrocarpum :-82 Centrosema pascuorum :centurio n 84 Centrosema pubescens : centro 86 Chamaecrista rotundifolia :roundlea f cassia 88 Chloris gay ana :Rhode s grass 90 aciculatus :lov e grass 92 Chrysopogon orientalis :tinlo y 93 Clitoria ternatea :butterfl y pea 94 Codariocalyx gyroides :codarri o 97 Crotalaria juncea :sun n hemp 98 dactylon : Bermuda grass 100 Cynodon nlemfuensis :stargras s 102 Dactyloctenium aegyptium : crowfoot grass 104 Desmanthus virgatus Desmodium heterocarpon :fals e 105 Desmodium heterocarpon : carpon desmodium 106 ssp. ovalifolium :-108 Desmodium heterophyllum :heter o 110 Desmodium incanum :kaim i 112 Desmodium intortum :greenlea f desmodium 114 Desmodium triflorum :three-flowere d beggarweed 116 Desmodium uncinatum :silverlea f desmodium 116 Dichanthium annulatum :hind i grass 118 ciliaris :tropica l finger grass 119 :commo n finger grass 121 Digitaria milanjiana :digi t grass 123 colona :jungl e rice 125 Echinochloa crus-galli :barnyar d 126 Eragrostis tenella :bug' s egg grass 127 Eragrostis unioloides :padang togu128 Ficus subcordata bunut lengis 129 Flemingia macrophylla apa apa 131 Gliricidia sepium gliricidia 133 Heteropogon contortus speargrass137 Hymenachne acutigluma wick grass 139 Imper ata cylindrica blady grass 140 ciliare batiki blue grass 142 Ischaemum magnum rumput melayu 144 Ischaemum muticum seashore centipede grass 145 Ischaemum rugosum wrinkle duck-beak 146 Ischaemum timorense centipede grass 148 chinensis timunan 149 Leucaena leucocephala leucaena 150 Lotononis bainesii lotononis 154 Macroptilium atropurpureum siratro 155 Macroptilium lathyroides phasey bean 158 Macroptilium longepe- dunculatum :llano s macro 159 Macrotyloma axillare :axillari s 161 Medicago sauva :lucern e 163 Microstegium ciliatum :banyona n tembaga 165 Mikania cordata :mile-a-minut e 166 Mimosa pudica :sensitiv e plant 167 Neonotonia wightii :glycin e 169 Ottochloa nodosa :slende r panic grass 171 maximum :guine a grass 172 Panicum maximum var. trichoglume green panic 174 Panicum repens torpedo grass 176 conjugatum buffalo grass 177 Paspalum dilatatum Dallis grass 178 Paspalum distichum ginger grass 180 Bahia grass 181 Paspa lum plicatulum plicatulum 183 Paspalum scrobiculatum kodo millet 185 clandestinum kikuyu grass 187 Pennisetum polystachion feather pennisetum 189 Pennisetum purpureum elephant grass 191 Pueraria phaseoloides tropical kudzu 192 Saccharum spontaneum glagah 195 Sesbania grandiflora agati sesbania 196 Sesbania sesban sesban 198 sphacelata setaria 201 x almum Columbus grass 203 Sorghum, artificial perennial hybrids -205 Sorghum x drummondii Sudan grass 206 Stenotaphrum secundatum St. Augustine grass 208 Stylosanthes capitata -209 Stylosanthes guianensis : stylo 211 Stylosanthes hamata :Caribbea n stylo 213 Stylosanthes humilis : Townsville stylo 216 Stylosanthes macrocephala : - 218 Stylosanthes scabra : shrubby stylo 219 Themeda triandra : red oat grass 222 Thysanolaena latifolia : tiger grass 224 Trifolium repens : white clover 225 Trifolium semipilosum : white clover 226 Tripsacum andersonii : Guatemala grass 228 mosambicensis : Sabi grass 230 Vignaparkeri : creeping vigna 232

Zoysia matrella : Manila grass 234

3 Minor forages 236

4 Forages with other primary use 245

Literature 254

Acknowledgments 258

Acronyms of organizations 259

Glossary 261

Sources of illustrations 271

Index of scientific plant names 277

Index of vernacular plant names 292

The Prosea Foundation 297 Editors and contributors

General editors of the Prosea Handbook

P.C.M. Jansen, E. Westphal and J. Kartasubrata

Editorial staff of this volume

- Editors: L. 't Mannetje and R.M. Jones - Associate editors: C.P. Chen, P.C.M. Jansen (botanical aspects), C. Manidool and I.M. Nitis - Illustrators: P. VerheijJ£aye s (drawings) and R. Boekelman (map) - Publishing consultant and in-house editor: R.J.P. Aalpol - Language corrector: J. BurroughJ3oenisc h

Contributors '

- N.O. Aguilar, Museum of Natural History, University of the Philippines at Los Bafios, College, Laguna 4031, the Philippines (Imperata cylindrica, Mimosa pudica) - A. Aminah, Livestock Research Division, Malaysian Agricultural Research and Development Institute (MARDI), G.P.O. Box 12301, 50774 Kuala Lumpur, Malaysia (Calopogonium mucunoides) - B.B. Baki, Malaysian Agricultural Research & Development Institute (MARDI), 3C Rubber Road, 93400 Kuching, Sarawak, Malaysia (Digitaria ciliaris, Ischaemum ciliare, I. muticum, I. rugosum, I. timorense, Paspalum notatum, P. scrobiculatum, Pennisetumpolystachion) - A.K. Benjamin, Department of Primary Industry, P.O. Box 1075, Goroka, Papua New Guinea (Neonotonia wightii, Trifolium semipilosum, Vigna par­ ken) - H.G. Bishop, Department of Primary Industries, P.O. Box 668, Mackay, Q 4740,Australi a (Aeschynomene americana) - J.L. Brewbaker, Department ofHorticulture , University ofHawai i at Manoa, 3190 Maile Way, Honululu, 96822, United States (Leucaena leucoce- phala) - A. Budelman, Royal Institute for the , Mauritskade 63, 1092 AD Amsterdam, the Netherlands {Flemingia macrophylla) - A.G. Cameron, Department of Primary Industries and Fisheries, P.O. Box 79, Berrimah, NT 0828,Australi a (Macroptilium longepedunculatum) - Y.K. Chee, Rubber Research Institute of Malaysia, G.P.O. Box 10150, 50908 Kuala Lumpur, Malaysia (Calopogonium caeruleum, Canavalia ensiformis, Crotalaria juncea, Mikania cordata) 10 FORAGES

- C.P. Chen, Malaysian Agricultural Research and Development Institute (MARDI),G.P.O . Box 12301,50774KualaLumpur , Malaysia (Asy stasia gange- tica, Calopogonium caeruleum, C. mucunoides, Canavalia ensiformis, Centro- sema pubescens, Crotalaria juncea, Digitaria ciliaris, Ischaemum timorense, Mikania cordata, Panicum maximum, Paspalum notatum, Pennisetum poly- stachion, Stenotaphrum secundatum, Tripsacum andersonii) - R.J. Clements, CSIRO Division of Tropical Crops and Pastures, 306 Carmody Road, St. Lucia, Q4067 ,Australi a (Centrosema pascuorum, Medicago sativa) - B.G. Cook, Department of Primary Industries, P.O. Box 395, Gympie, Q4570 , Australia (Aeschynomene falcata, Arachis glabrata, A.pintoi, Vignaparkeri) - L.A. Edye, CSIRO Davies Laboratory, Private Mail Bag, P.O. Aitkenvale, Q 4814,Australi a (Stylosanthes hamata. S. humilis, S. scabra) - R.C. Gutteridge, Department of Agriculture, University of Queensland, St. Lucia, Q4067 ,Australi a (Sesbania grandiflora, S. sesban) - J.B. Hacker, CSIRO Division of Tropical Crops and Pastures, 306 Carmody Road, St. Lucia, Q 4067, Australia (Canavalia ensiformis, Desmodium hetero- carpon, D. heterophyHum, D. incanum, D. intortum, D. uncinatum, Digitaria eriantha, D. milanjiana, Setaria sphacelata, Sorghum x almum, Sorghum ar­ tificial perennial hybrids, Sorghum x drummondii) - R.A. Halim, Department of Agronomy & Horticulture, Universiti Pertanian Malaysia, 43400U.P.M . Serdang, Selangor, Malaysia (Alysicarpus vaginalis, Pueraria phaseoloides) - W.W. Hanna, Department ofAgriculture , Coastal Plains Experiment Station, Tifton, Georgia 31793,Unite d States (Cynodon dactylon, C. nlemfuensis) - J.H. Heering, ILCA, P.O. Box 5689,Addi s Ababa, (Sesbania grandi­ flora, S. sesban) - S.M. Howden, Bureau of Rural Resources, Department of Primary Industries and Energy, P.O. Box Ell, Queen Terrace, ACT 2600, Australia (Bothriochloa pertusa) - E.M. Hutton, 11Norwoo d Place, Station Road, Indooroopilly, Q4068 , Austra­ lia (Panicum maximum) - LB. Ipor, Universiti Pertanian Malaysia, P.O. Box 396, 97008 Bintulu, Sa­ rawak, Malaysia (Digitaria ciliaris, Ischaemum ciliare, I. muticum, I. ­ ense, Paspalum notatum, P. scrobiculatum) - R.J. Jones, CSIRO Division of Tropical Crops and Pastures, Private Bag, Ait­ kenvale, Q4814 ,Australi a (Leucaena leucocephala) - R.M. Jones, CSIRO Division of Tropical Crops and Pastures, 306 Carmody Road, St. Lucia, Q 4067, Australia (Chamaecrista rotundifolia, Lotononis bai- nesii, Macroptilium atropurpureum, M. lathyroides, Mimosapudica, Trifolium repens, T. semipilosum) - S.M.M. Kersten, Department of Agriculture, University of Sydney, Sydney, NSW 2001,Australi a (Cenchrus ciliaris, Chloris gayana, Panicum maximum var. trichoglume, Paspalum dilatatum, P.plicatulum, Trifolium repens) - A.E. Kretschmer, University of Florida, Institute of Food and Agricultural Sciences, P.O. Box 248,For t Pierce, Florida 34954-0248,Unite d States (Desmo­ dium heterocarpon) - S.A. Lee, Malaysian Agricultural Research and Development Institute (MARDI),G.P.O . Box 12301,50774Kual a Lumpur, Malaysia (Asystasia gauge- tica) EDITORS AND CONTRIBUTORS 11

J.B. Lowry, CSIRO Davies Laboratory, University Road, Townsville, Q4814 , Australia (Albizia lebbeck) J.G. Mclvor, CSIRO Davies Laboratory, Private Mail Bag, P.O. Aitkenvale, Q 4814,Australi a (Bothriochloapertusa, Urochloa mosambicensis) C. Manidool, AUSTREX, 66/2-3 Sukumuiv Soi 2, Bangkok 10110, Thailand (Arundinariapusilla, Axonopus compressus, Brachiaria distachya, Centothe- ca latifolia, Chrysopogon aciculatus, C.orientalis, Dactyloctenium aegyptium, Dichanthium annulatum, , E. crus-galli, Eragrostis tenella, E. unioloides, Hymenachne acutigluma, Ischaemum ciliare, I. rugosum, Lep- tochloa chinensis, Microstegium ciliatum, Ottochloa nodosa, Panicum repens, Paspalum conjugatum, P. distichum, Pennisetum polystachion, Saccharum spontaneum, Thysanolaena latifolia, ) P.T. Mears, Department of Agriculture and Fisheries, Agricultural Research Station, Grafton, NSW 2460,Australi a (Pennisetum clandestinum) T.T. Ng, Department ofAgriculture , Jalan Simpang Tiga, 93632 Kuching, Sa­ rawak, Malaysia (Ischaemum magnum) I.M. Nitis, Department of Nutrition and Tropical Forage Science, Udayana University, Jalan Jendral Sudirman, Denpasar, Indonesia (Ficus subcordata, Gliricidia sepium) B.C. Pengelly, CSIRO Division of Tropical Crops and Pastures, 306 Carmody Road, St. Lucia, Q4067 , Australia (Alysicarpus vaginalis, Desmanthus virga- tus, Desmodium triflorum, Neonotonia wightii) L. Ramirez, CSIRO, 306 Carmody Road, St. Lucia, Q4067 ,Australi a (Canava- lia ensiformis) I.K.Rika , Department of Extension Services, Udayana University, Jalan Jen­ dral Sudirman, Denpasar, Indonesia (Calliandra calothyrsus) R. Schultze-Kraft, Universität Hohenheim, 380 Institut für Pflanzenproduk­ tion in den Tropen und Subtropen, Kirchnerstr. 5, Postfach 700562, D-7000 Stuttgart 70, Germany (Andropogon gayanus, Brachiaria brizantha, B. decumbens, B. dictyoneura, B. humidicola, B. mutica, B. ruziziensis, B. sub- quadripara, Centrosema acutifolium, C. macrocarpum, Desmodium heterocar- pon ssp. ovalifolium, Stylosanthes capitata, S. macrocephala) M.E. Siregar, Balai Penelitian Ternak, Ciawi, P.O. Box 123800, Bogor, Indo­ nesia (Flemingia macrophylla) R. Soedomo, Faculty of Husbandry, Gadjah Mada University, Yogyakarta 55281, Indonesia (Codariocalyx gyroides) C.T. Sorensson, Department of Horticulture, University of Hawaii at Manoa, 3190 Maile Way, Honululu, Hawaii 96822, United States (Leucaena leucoce- phala) LB. Staples, Department of Primary Industries, P.O. Box 1054, Mareeba, Q 4880,Australi a (Clitoria ternatea, Macrotyloma axillare) J.K. Teitzel, Department ofPrimar y Industries, P.O.Bo x20 ,Sout h Johnstone, Q 4859, Australia (Brachiaria decumbens, B. humidicola, B. mutica, B. ruzi­ ziensis, Centrosema pubescens, Desmodium heterophyHum) L. 't Mannetje, Department of Field Crops and Grassland Science, Wage­ ningen Agricultural University, Haarweg 333, 6709 RZ Wageningen, the Netherlands (Cenchrus ciliaris, Chloris gayana, Macroptilium atropurpu- reum, M. lathyroides, Panicum maximum var. trichoglume, Paspalum dilata- tum, P.plicatulum, Pennisetum purpureum, Stylosanthes guianensis) 12 FORAGES

A. Topark-Ngarm, Department of Agronomy, Faculty of Agriculture, Khon Kaen University, Khon Kaen 40002, Thailand (Desmanthus virgatus, Stylo- santhes hamata, S. humilis, S. scabra) J.C. Tothill, CSIRO Division of Tropical Crops and Pastures, 306 Carmody Road, St. Lucia, Q4067 ,Australi a (Heteropogon contortus, Themeda triandra) K.F. Wiersum, Department of Forestry, Wageningen Agricultural Universi­ ty, P.O.Bo x 342,6700AH ,Wageningen , the Netherlands (Calliandra calothyr- sus, Gliricidia sepium) C.C. Wong, Malaysian Agricultural Research and Development Institute (MARDI), G.P.O. Box 12301, 50774Kual a Lumpur, Malaysia (Digitaria milan- jiana) Prosea Board of Trustees and Personnel

(May 1992)

Board of Trustees

Aprilani Soegiarto (LIPI, Indonesia), chairman H.C. van der Plas (WAU, the Netherlands), vice-chairman Salleh Mohd. Nor (FRIM, Malaysia) Misty Baloiloi (UNITECH, Papua New Guinea) B.P. del Rosario (PCARRD, the Philippines) Santhad Rojanasoonthon (TISTR, Thailand) Dang Huy Huynh (IEBR, Vietnam)

A. De Villepin (European Communities) Soekiman Atmosoedaryo (Yayasan Sarana Wanajaya)

J.M. Schippers (Pudoc-DLO, à titre personnel) Sampurno Kadarsan (à titre personnel)

Personnel

Indonesia

Sampurno Kadarsan, Programme Leader Hadi Sutarno, Country Officer Sarkat Danimihardja, Assistant Country Officer

Malaysia

Salleh Mohd. Nor, Programme Leader Lillian Chua, Country Officer

Papua New Guinea

A.M. Gurnah, Programme Leader R. Matu, Country Officer

The Philippines

P.S. Faylon, Programme Leader N. Altoveros, Country Officer 14 FORAGES

A. del Mundo, Assistant Country Officer N.B. Pena, Assistant Country Officer

Thailand

Prapandh Boonklinkajorn, Programme Leader Soonthorn Duriyaprapan, Country Officer Sayan Tanpanich, Assistant Country Officer

Vietnam

Nguyen Tien Ban, Programme Leader Zuong Due Huyen, Country Officer La Dinh Moi, Assistant Country Officer Nguyen Van Du, Assistant Country Officer

Network Office, Bogor, Indonesia

L.S. Anema, Head Ch. Aipassa, Secretary Jajang, Office Assistant J. Kartasubrata, General Editor A.J.G.H. Kostermans, Scientific Adviser C.L. Nurcahyanti, Financial Officer & Secretary N.W. Soetjipto, Coordinator Regional Data Bank

Publication Office, Wageningen, the Netherlands

J.S. Siemonsma, Head C.L. Crow, Secretary H.C.D. de Wit, Scientific Adviser J.M. Fundter, Forestry Officer J.W. Hildebrand, Forestry Officer P.CM. Jansen, General Editor J. Jukema, Tropical Agronomy Officer R.H.M.J . Lemmens, Plant Officer L.P.A. Oyen, Documentation Officer M.S.M. Sosef, Plant Taxonomy Officer E. Westphal, General Editor W.P.M. Wolters, Programme Secretary Foreword

According to FAO estimates, the livestock population of South-East Asia com­ prises approximately 55millio n equivalents. Of the 435millio n ha of land in the , only 16millio n ha are classified as permanent grasslands, which is the typical form of land use devoted to livestock. Since the productivity of these grasslands is poor, they would be unable to carry these numbers of ani­ mals. Furthermore, the greatest livestock densities occur in the most densely populated , where grasslands are not common. These simple facts indi­ cate that in South-East Asia livestock feed is not produced solely on permanent grasslands. Most ofth e livestock is kept on small mixed farms. Cattle and buffa­ loes provide draft power and they produce small amounts of milk, meat and manure. The cultivation of food crops has the greatest priority on these farms, so that there is little land available to grow forage. Instead, the are tethered on fallow land and along roadsides. Animals are also stabled, fed on harvested forage, whilst the manure is used to fertilize food crops, so that the soil fertility of forage-growing areas is gradually depleted. Additional animal feed is obtained from crop residues, such as straw, but these are generally of poor quality. However, by-products of industrially processed agricultural prod­ ucts provide better quality animal feed such as rice bran and molasses. The quality ofmos t tropical grasses ispoor , leading to poor animal performance. Both their yield and crude protein concentration could be improved by the use of nitrogen fertilizer, as long as the forage is harvested at a young stage of growth. However, small farmers, who do not own the land from which they har­ vest forage, are not in a position to use fertilizers for forage production. For this reason the smallholder is better off using legumes which fix atmospheric nitrogen and which maintain a reasonable feeding value while ageing. Grass- mixtures which are used on large scale animal farms, however, are not feasible in the more densely populated areas of South-East Asia. Therefore, legumes in sole cropping, which can be used in rotation with crops, thus contri­ buting to soil fertility, and woody legumes which can be grown as hedges or in alley cropping are a more appropriate way of improving the forage supply on small farms. The improvement of native grasslands requires an inventory and evaluation of available grasses and legumes. Generally speaking, native forage species do not give an economic return to improved soil fertility, whilst most improved grasses and legumes are not productive and persistent on poor . Therefore, large-scale grassland improvement for extensive animal production has to rely on species which can thrive on poor soils. Introduced grasses require a source of nitrogen, which can be supplied by legumes that are adapted to poor soils (e.g. Stylosanthes spp.). However, native pastures, even Imperata grasslands, can also be improved by the addition of such legumes. Much knowledge and 16 FORAGES experience in these matters has been accumulated by the CSIRO Division of Tropical Crops and Pastures in Queensland, Australia, with which the editors of this volume have been closely associated. In this Prosea volume, much useful information is provided about the , ecology and agronomy of a large number of native and introduced grasses, legumes, and other forage species that can be gainfully used in South-East Asia. I hope that this information will be an inspiration to all those who are involved in the improvement of the forage supply in the region.

Wageningen, June 1992

J.G.P. Dirven Emeritus Professor of Tropical Grassland Science, Wageningen Agricultural University 1 Introduction

1.1 Forages and livestock

Grassland and fodder plants serve as feed for domestic and wild herbivores. They mostly belong to the families Gramineae and Leguminosae, although other herbaceous and woody plants are also eaten by animals. Grassland and fodder plants are collectively named forages. Forage can be defined as feed for herbivores, usually with low nutrient concentration and low digestibility, thus contrasting with concentrates such as grain. Forage plants can occur in the wild, but are more commonly found in grasslands, in tree crop plantations or in open spaces in and near forests and along roads, bunds between rice paddies and canals. In South-East Asia such areas are sometimes termed 'waste' areas - this does not mean they do not provide useful forage, but that they are not used for food or forage crops. However, forages can be cultivated and sown for the specific purpose ofproducin g feed for livestock. Residues or by-products from food crops, and in cropping lands, are also very important sources of forage in South-East Asia. Forage is either grazed in situ by tethered or free ranging animals or is cut and carried to penned or tethered animals. Even within one region the proportion of forage obtained from as compared with hand feeding can change appreciably between different sites and seasons, as is the case for West Java (Thahar &Petheram , 1983). Although South-East Asian agriculture is mostly geared to the production of rice and plantation crops, livestock play an important role in providing draft power and for the production of meat, milk and dung. The estimated livestock numbers in South-East Asia comprise about 33millio n cattle, 19 million buffa­ loes, 6millio n and 15millio n goats (Table 1). Most animals are kept in the densely populated areas and about 90% of rumi­ nants occur on mixed farms run by smallholders. Such farms are typically of 1-2 ha with 1-4 large ruminants and a few sheep and goats (Ranjhan, 1986). Between 1969-1971 and 1988 the estimated number of cattle, sheep and goats increased, but the number of buffaloes did not change. Yet there are differences in the trends of large and small ruminants within and between countries. For example, in Indonesia and the Philippines the number of goats increased sub­ stantially, whereas there was no such increase in Malaysia. The estimated 20% increase in the numbers of large ruminants and 68% increase in the numbers of small ruminants between 1970an d 1988clearl y point to an increased demand for fodder. Moreover, many of these animals suffer malnutrition and also the demand for meat and milk is rising as a result of increased buying power of the population, so that the increase in animal numbers may underestimate the need for increased quantity and quality of forages. It has been estimated that the demand for forage would double by the year 2000(Remeny i &McWilliam , 1986). 18 FORAGES

Table 1.Ruminan t livestock (x 106)i n South-East Asia; means for 1969- -1971an d 1988(FAO , 1989).

Country1 Cattle Buffaloes Sheep Goats

1969-1971 1988 1969-1971 1988 1969-1971 1988 1969-1971 1988

Burma 7.0 10.0 1.6 2.2 0.2 0.3 0.6 1.1 Cambodia 2.2 2.0 0.9 0.7 Indonesia 6.3 10.1 3.2 3.3 3.2 5.5 6.9 10.6 Laos 0.4 0.8 0.9 1.1 Malaysia 0.3 0.6 0.3 0.2 0.1 0.4 0.3 Philippines 1.7 1.5 4.4 2.8 * 0.8 2.2 Thailand 4.5 5.3 5.6 5.4 0.1 * 0.1 Vietnam 1.8 3.0 2.3 2.9 * 0.2 0.4

Total 24.2 33.3 19.2 18.6 3.4 6.0 8.9 14.7

1. Bruneian d Singaporehav ebee nomitte dbecaus e ofsmal lnumber s ofanimals . *Les stha n 100000animals .

Although there have been considerable forage research programmes in the region (Blair et al., 1986; Halim, 1989), comparatively little use is made of improved forage plants. However, the use of these species is increasing and there is a large potential for increased forage production in the region.

1.2 Sources of forage

1.2.1 Grasslands

In tropical areas with high rainfall, grasslands are usually sub-climax vegeta­ tion, because the natural climax is almost always closed forest. These sub-cli­ max grasslands have been formed by man clearing forest for shifting cultivation and are then maintained as a sub-climax by grazing and burning of abandoned cropland. These grasslands are almost always found on areas that are not suited to food crops by virtue of factors such as slope and soil type. Natural climax grasslands in South-East Asia are rare and usually restricted to frequently flooded lands, as has been reported for parts of Vietnam (Whyte, 1974). Grasslands are sometimes referred to as '', a term that denotes a contin­ uous graminoid stratum, more or less interrupted by trees or (Johnson & Tothill, 1985). One of the characteristics of savanna is that the climate is seasonal with wet, warm to hot periods alternating with more or less dry, warm to cool ones (Johnson & Tothill, 1985). But the term savanna is still subject to confusion and here the term 'grassland' is used, defined as vegetation types in which the tree cover is less than 40% (UNESCO, 1979). Grasslands can be divided into pure (i.e.treeless ) and wooded grasslands. It has been estimated by FAO (1989) that there are about 14millio n ha of perma­ nent grassland in South-East Asia and the western Pacific Islands, whereas Soer- jani (1970) estimated that there were 16millio n ha in Indonesia alone. The domi­ nant species in these grasslands is usually Imperata cylindrica (L.) Raeuschel. INTRODUCTION 19

Major areas of grassland include: - the large central plains ofThailan d andth e Korat , which extend into Laos andth e northern parts of Cambodia; - parts of north and northwestern Thailand, extending into Burma and north­ ern Laos; - moderately high areas which are almost totally converted to wooded grass­ lands by livestock husbandry, especially on the plateaux of northern Laos and Vietnam. The grazing lands of Indonesia are wooded grasslands resulting from degrada­ tion of climax forest. These mayb eth e result ofdrie r conditions combined with annual fires, as on the eastern islands (Nusa Tenggara), or they may be aban­ doned crop lands used for grazing. Very often these lands are impoverished as a result of frequent exploitative cropping. On Java and Madura grazing lands occupy less than 5% of the land. Nevertheless these two islands support 65% of the livestock of Indonesia, but their feed comes from roadsides, banks of ca­ nals, bunds between rice fields, open areas within and alongside forests and from crop residues. In the Nusa Tenggara region of Indonesia pure or wooded grasslands cover large areas, e.g. 35% on Lombok and 65% on Sumba and Timor. About 12% of the Philippines is grazing lands which are maintained by frequent fires. There are approximately 15millio n ha of plantation crops in South-East Asia (FAO, 1989). The area under many of these plantation crops is used for food crops, but often itha sa herbaceou s understorey ofnativ e species, or sown cover crops which can be grazed. There is a great opportunity for forage production from pastures consisting of improved forages under , and to a lesser extent under young stands ofrubbe r andoi lpalm . Grazing canenhanc e nutrient cycling, control waste herbage andreduc e useo fweedicides , enable easier man­ agement of the plantation crop and may improve crop yields and income (Rey­ nolds, 1988). The species occurring in these unimproved grasslands arenativ e or naturalized and, except for fire, no cultural ormanagemen t practices are imposed. Dry mat­ ter yields and the nutritive quality of the species are frequently low.Th e grass genera most commonly encountered are: Andropogon L., Arundinella Raddi, Arundo L., Axonopus Beauv., Capillipedium Stapf, Chloris Swartz, Chrysopo- gon Trin., Coix L., Cyrtococcum Stapf, Cymbopogon Sprengel, Dactyloctenium Willd., Digitaria Haller, Echinochloa Beauv., Eleusine Gaertner, Eragrostis v. Wolf., Heteropogon Pers., Hyparrhenia Andersson, Imperata Cyr., Ottochloa Dandy, Paspalidium Stapf, Paspalum L., Pennisetum Rich., Phragmites Adan- son, Saccharum L. and Themeda Forssk. None ofth e species concerned, except Pennisetum purpureum Schum., is purposely established as grassland or forage crop. There arefe w legumes in these grasslands, some ofth e more common spe­ cies being Desmodium heterophyllum (Willd.) DC, Desmodium triflorum (L.) DC., Mimosa pudica L., Mimosa invisa Martius ex Colla and Indigofera spp. The species composition of grazing lands changes with the climatic regime and management imposed (Manidool &Chantkan , 1986).

Grass species of temperate origin (C3-species) occur at high altitudes in the region, such as on Mt Kinabalu in Sabah and on mountains in Indonesia and in the highlands in Malaysia and NewGuine a (Whyte, 1974). 20 FORAGES

1.2.2 Annual and perennial fodder crops

Fodder crops in present use fall into three categories, viz. herbaceous crops, root crops, and trees and shrubs. The herbaceous crops are mostly grasses and legumes. The main grass crops are: Pennisetumpurpureum Schum., P. america- num (L.) K. Schum. ex Leeke, P. purpureum x P. americanum, Panicum maxi­ mum Jacq., Tripsacum andersonii J.R. Gray, Saccharum officinarum L. and Zea mays L. The second and the last two species listed are primarily used as food crops. The main herbaceous legume fodder crops are Cajanus cajan (L.) Millsp., Lablab purpureus (L.) Sweet and Vigna unguiculata (L.) Walp. and these are used primarily as food crops, and Stylosanthes guianensis (Aublet) Swartz. The root crops used for human and animal nutrition are Manihot esculenta Crantz and Ipomoea batatas (L.) Lamk. For further information from the viewpoint of forage, see Section 1.7. Ofth e trees and shrubs Leucaena leucocephala (Lamk) deWi t isth e most extensi­ vely used, but other species such as Calliandra calothyrsus Meissn., Flemingia macrophylla (Willd.) Merrill, Gliricidia sepium (Jacq.) Kunth ex Walp., Sesba- nia grandiflora (L.)Poire t and S. sesban (L.)Merril l are also important (Topark- Ngarm, 1990). Trees and shrubs can be used in many ways, as living fences, vegetation in uncropped areas, hedgerows in alley cropping and as component species in intercropping (Topark-Ngarm, 1990).Ther e is great interest in fodder trees and shrubs in South-East Asia (Devendra, 1990). Fodder crops are used by smallholder farmers as well as by large-scale animal production units, mostly dairy farms near the larger cities. However, in most cases the cultivation of forage crops is subservient to that of food crops. Nevertheless, there is scope for forage crops in combination with food crops, either as alley cropping or undersown to provide better quantity and quality of forage after the food crop is harvested.

1.2.3 Crop residues and agricultural by-products

Nearly all food crops leave residues that can be used as forage. They are a very important forage resource in South-East Asia. Major sources include: rice hulls and straw, pseudostems, stover, sorghum stover, cassava leaves, sugar cane bagasse and tops, and pulse straws (Ranjhan, 1986). Residues from cereal crops are of poorer quality than residues from pulse crops and consider­ able research has been carried out on ways of improving their digestibility and intake (Wanapat &Devendra , 1985). The importance of different crop residues in animal feeding systems varies very widely between sites and between seasons at one site (Moog, 1986; Soedomo et al., 1986). In addition, by-products from crop processing, such as rubber seed meal and oil palm meal (Tinnimit, 1985), are used as concentrate feedstuffs.

1.3 Forage and livestock production systems

1.3.1 Current inputs toforage systems

Permanent native grasslands in this region can be classed as extensive forage production systems, as there are no managerial inputs such as irrigation, fertil- INTRODUCTION 21

izer or controlled grazing. Even though the pastures may be over-utilized to the point of overgrazing, the only inputs of fire and grazing are often uncon­ trolled. Semi-intensive forage production systems have inputs such as fertilizer, weeding and irrigation primarily applied to the main cash crop,whic h also bene­ fit associated forage. Intensive forage production systems have inputs applied for the sole purpose of forage production. An example of this would be large- scale dairy farms near urban centres, where fertilizers could be used on planted forage crops. Based on the classification of Perkins et al. (1986), five forage production sys­ tems, each with its typical level of input, can be distinguished in South-East Asia: - extensive permanent grassland: (a) privately owned land, (b) communally grazed hill-land and (c) communally grazed and cut roadsides; - semi-intensive permanent forages: (a) understorey of tree crop plantations, (b) forage from shade trees in plantations, (c) paddy field bunds and edges of crop fields, and (d)perennia l forage in alley cropping; - semi-intensive annual forages: (a) forage crops sown after harvest of food crops and (b) crop residues; - intensive permanent forages: (a) improved grasslands, and (b)protei n banks; - intensive short-term forages: (a) fodder crops, replacing food crops and (b) fodder crops on special areas. or silage is not widely used in South-East Asia and is not likely to be so in the future (Ranjhan, 1986). However, conservation is successfully used in some situations. For example, in Indonesia some commercial dairy farmers are using silage of elephant grass (Pennisetum purpureum) and some smallholders are using dried grass for feed in the dry season.

1.3.2 Input offorage into different livestock production systems

All livestock production systems can conveniently be grouped into three cate­ gories: pastoralism, livestock-crop, and crop-livestock. In pastoral systems people are dependent for livestock on all their needs and there is no interaction with crops for human food. There are three main subgroups: nomadism, ranch­ ing and intensive dairying. Nomadism is widely practised in semi-arid and arid areas of Africa and is not applicable to South-East Asia. Ranching takes place in northern Australia and parts of , but is of very little impor­ tance in South-East Asia. Commercial dairying in South-East Asia is still re­ stricted to areas with good access to large urban centres, but is likely to become more widespread in the future. The second or 'livestock-crop' production sys­ tem, where there is some dependence on crop residues for feed, but livestock are more important in the farm system than cropping, is typified by many areas in Africa, but to a lesser extent in South-East Asia. The third or 'crop-livestock' system is typical of South-East Asia, where the crop can be an annual such as rice or a long-lived tree such as . The food crops are the major and essential part of the farm system. The conceptual potential for incorporating the different forage sources pre­ viously mentioned in Section 1.2 into these different systems of livestock pro­ duction is summarized in Table 2. This table illustrates that there are poten­ tially more ways ofincorporatin g forages into crop-livestock systems than into 22 FORAGES

Table 2.Forag e production systemsuse d indifferen t livestock production systems.

Forageproductio n system Livestock production system

nomadism ranching dairying livestock- crop- crop livestock Extensivepermanen t grassland privately used — communally used +

Semi-intensive permanent forages understorey tree crops — — + — + forage from shade trees — - + - + edgeso fcro pfield s + - + + + forage in alley cropping — — + — + Semi-intensive annual forages after harvest ofmai n crop — _ + + cropresidue s + — + + + Intensivepermanen t forages improved grasslands — + + _ protein banks — + + + + Intensive short-term forages fodder crops —

systems such as nomadism or ranching. But this potential is restricted by sever­ al important limitations to forage production.

1.4 Main limitations to forage production

Forage production constraints in South-East Asia can be attributed to climate, soil conditions, species, management and socio-economic conditions.

1.4.1 Climate

In humid regions without a dry season of any significance, climatic conditions are generally conducive to good forage growth. However, even in such areas cloud cover will frequently reduce incoming radiation, resulting in lower photo­ synthesis. Daylength in equatorial regions can also limit growth in comparison to higher latitudes. The combination of longer days and less cloud cover in northern Australia, for example, results in more total incoming radiation than at the equator (Cooper, 1970). Forage growth under trees is also restricted through the interception of light by the tree canopy, as well as by competition for nutrients and moisture. Temperature is not a limiting factor in equatorial regions except at higher alti- INTRODUCTION 23

Table 3. The climatic zones oftropica l andsubtropica l South-East Asia and adjacent regions as defined by Troll (1966).

Troll Climatic Number of humid Observations indice zone months per year

V! tropical rainy climate 12-9.5 occurrence: wetter areas of Indonesia, Papua New Guinea, Malaysia, the Philippines, south­ ern Thailand

V2 tropical humid-summer 9.5-7 occurrence: eastern Java, Lesser Sunda Is­ climate lands, southern New Guinea, western Philip­ pines,Indo-Chin a

V3 wet-and-dry tropical 7-4.5 occurrence:centra l Thailand climate

IV4 dry-winter climate with 9-6 occurrence: northern Indo- extending long summer-humidity into southern China; basically this climatic zonei sa subtropica l extension ofV 2an dV 3.

IV6+y permanently humid 10-12 occurrence: northern Vietnam with hot sum­ climate mer extending into eastern China; basically this climatic zone is a subtropical extension of

Vj and V2.

tudes. However, as one moves away from the equator, temperature will be limit­ ing during the time of year with the shortest daylengths. Both at high altitudes and latitudes, frosts can destroy leaves and stems or even kill the plants of trop­ ical species, such as legumes and C4-grasses. At high altitudes, where growing conditions are temperate, the adaptation of the vegetation is reflected in the increased presence of C3-species.Bu t at higher latitudes within and immediately north and south of the tropics of Cancer and Capricorn, the growing conditions are primarily tropical. This is because the period ofadequat e moisture coincides with that of high temperatures and longest daylength. The grasses occurring in these regions are mainly of the C4-type, which cannot grow at temperatures below about 15°C (McWilliam, 1978). Rainfall is not often limiting in equatorial regions, although periods without rain of up to six weeks have been encountered (Niewolt, 1982). However, total rainfall and the period of the year in which it is received generally decrease with increasing distance from the equator. Using the classification of Troll (1966), the following tropical climatic zones can be distinguished in South-East Asia (Table 3):tropica l rainy climates (V^), tropical humid-summer climates (V2), and wet-and-dry tropical climates (V3). To adequately describe South-East Asia, when the region is defined in broader terms, the following subtropical climatic zones should be included: dry-winter climates with long summer-humidity and cool dry winter (IV4),an d permanently humid climates with hot summer and maximum rainfall in the summer (IV6+7). Mountainous areas in equatorial regions pose a problem in climatic classifica- 24 FORAGES tion. From the point of view of forage plant adaptation, such areas can be regarded as equivalent to the subtropical climatic zones IV6+7 and IV4, depend­ ing on rainfall distribution. Soil moisture will be limiting in the climatic zones V2> V3 and IV4 during the period of shortest daylength, and in the latter zone this will coincide with tem­ perature constraints. In climatic zones V3 and IV4 the unreliability of rainfall during the rainy period also imposes strong limitations on forage production.

1.4.2 Soil conditions

Soil fertility varies widely over the region. The more fertile soils are derived from basic volcanic rocks, coralline material, or alluvium. These soils are inten­ sively cropped. Some 60% of the soils in South-East Asia have severe nutrient deficiencies or toxicities (Kerridge et al., 1986), even though they have medium to excellent soil physical conditions. These soils are leached and have low orga­ nic matter levels and low cation exchange capacities. They are acid, with conse­ quently high availability of soluble Al and Mn and low availability of P, Ca and some trace elements. Many tropical forage species, including legumes, are adapted to acid conditions and can tolerate high levels of available Al. Soils in Troll's V3 and IV4 climatic zones tend to be less acid than those in V1; V2 and IV6+7. Regional studies of soil fertility, such as described by Kerati-Kasi- korn (1984),ca n help document the severity ofnutrien t deficiencies on different soil types.

1.4.3 Species

The native and naturalized forage species are adapted to the prevailing climatic and soil conditions. Rainfall in climatic zones VltV 2 and IV6+7 is sufficiently regular that drought tolerance is not a requirement for the plants. Adaptation to dry conditions as a result of irregular and/or seasonally low rainfall is, how­ ever, required for plants in zones V3 and IV4. In all climatic zones soil fertility is a major constraint to growth although species differ in their adaptation to low fertility (Kerridge et al., 1986). Plant species that are adapted to the unfa­ vourable rainfall and soil fertility have developed strong survival mechanisms. These are based on vegetative propagation and competitive ability to exclude other species (e.g. Axonopus compressus (Schwartz) P. Beauv. and Imperata cylindrica) or on the ability to produce copious seed (e.g. Mimosapudica). High dry matter production and nutritive value are not necessary attributes for spe­ cies to survive, and sosom e native species have low productivity and poor nutri­ tive value. Some would not persist unless soil fertility is amended, or ifthe y do,the y are often no more productive than native or naturalized species. Conversely, some native and naturalized species may yield as much as intro­ duced species under improved conditions of soil fertility. Introduced legumes should be checked to ensure that they are nodulated. Preferably, introduced legumes should be able to nodulate with indigenous strains of . Species differ in their adaptation to defoliation. Some species, such as Themeda triandra Forssk., are unable to persist even under moderate grazing pressure. Other species, such as Imperata cylindrica and Mimosa pudica, avoid high graz- INTRODUCTION 25 ing pressures because of low palatability while others, such as Axonopus com- pressus, are resistant to grazing through their prostrate growth habit. Recent research has also shown that there is considerable variation between species in their ability to grow under the reduced light conditions in plantation crops (Wong et al., 1985a, 1985b;Ahme d Tajuddin, 1991; Shelton &Stiir , 1991).

1.4.4 Management

The production of forage from a given area of land depends not only on climatic and edaphic conditions. Within these given limitations the farmer has a large influence on both quantity and quality offorag e harvested. The options are: - to return removed nutrients or not; - to fertilize or not; - to alter frequency and height of cutting; - to alter numbers and movement of animals which are freely grazing or are tethered on grazing land; - to alter the integration offodde r and food crops. The first two options are seldom applied to native forage species. In fact the usual system is to remove forage, feed it to stalled or tethered animals and to use the excrement asmanur e on food crops.Thi s amounts to mining of nutrients, leading to impoverishment of the soil under forages and consequently lower forage yields. In the case of grazing most of the nutrients are returned to the field, but the distribution is very uneven and losses of inorganic nitrogen will occur through volatilization of ammonia and leaching of nitrate. A high frequency of harvesting can accelerate exploitation of nutrients and disappearance of species not morphologically adapted to frequent defoliation. On the other hand, forage harvested in a young stage of growth is of higher protein and mineral concentration and ofhighe r digestibility than material har­ vested at an older stage of growth. A stocking rate exceeding the carrying capacity of a grazed area will lead to overgrazing, often resulting in the disappearance of palatable species and the dominance of species that are not readily eaten, such as Imperata cylindrica and Mimosa pudica. Even Imperata cylindrica can be eliminated by very heavy grazing and be replaced by unpalatable weeds (Falvey & Hengmichai, 1979). Overgrazing thus leads to reduced productivity of the land. The unpalatable species at least protect the land against severe soil erosion which would other­ wise occur. A detailed discussion of how grazing pressure effects pasture pro­ ductivity, botanical composition of pastures and animal production is given by Humphreys (1991). Food crops are the key to agriculture in South-East Asia and any attempt to improve fodder production must work within this limitation. However, there are possibilities for combining quality requirements for human food as well as for animal feed in one crop. For example, rice can be selected for grain as well as straw quality, and cassava for root quality as well as for low HCN content ofth e leaves.

1.4.5 Socio-economic constraints

The improvement of forage resources requires investment of money and labour 26 FORAGES and therefore depends on factors such as: - the economic value and marketability of the animal products; - the availability of land and its lease conditions; - accessibility to finances, knowledge and continued support; - motivation of the farmer and his attitude to risk; - cultural barriers; - economic alternatives within the farm systems. Much animal production in South-East Asia has no direct market value, as in the case of draft power. Many farmers have no land available for direct forage production, and if they do,the y may not have access to finance or may not know howt o improvethei r production offorage . Inparticular , there arema jo r problems in improving communally grazed lands where any improvements are not for indi­ vidual, but common use (Raghavan, 1990).Socio-economi c barriers are, in gener­ al, the main limitation to improving animal production in South-East Asia. This is further illustrated by Ghani & Wong (1990), who list factors that constrain the adoption ofimprove d agroforestry systems.Thes e arefactor s of socio-econom­ ic, socio-cultural and institutional nature which are prevalent in South-East Asia, as in most developing countries. The most important of these are lack of motivation, lack of finance, lack of knowledge, lack of entrepreneurship, lack of community leadership, limitations caused by customs and tradition, lack of research and extension facilities and personnel, and problems ofpropert y rights.

1.5 Overcoming limitations to improving forage resources

1.5.1 Climate

Nothing can be done to alter the macro-climate for forage production, although it is possible to make the most of the potential for forage production in areas with a changed micro-climate as, for example, under coconut plantations. Where irrigation is available, it is usually used to grow food crops in preference to forage.

1.5.2 Soil fertility

It is highly unlikely that inorganic or organic fertilizers will be used on a wide­ spread basis on extensive grasslands in South-East Asia in the near future. There are more prospects offertilizer s being applied to vegetables, grain or tree crops, thus having an indirect impact on forages in semi-intensive systems and to intensive forage production areas. Phosphorus will be the main element required for legumes though S, K, Cu, Mo, B and Ca may also be needed. Nitro­ gen is the primary limitation for grasses. In grass/legume systems the legume supplies some nitrogen to the grass, but rarely enough for the grass to reach maximum yield. In some localities there is potential for increasing animal pro­ duction by supplying Co and Na directly to animals, as these elements can be deficient for animals but not for plants. Many tropical forages are low in Na (Little et al., 1989). Some forage species such as Stylosanthes spp. can grow at low levels of available soil P and so can have low P concentrations in their forage. In these cases, P can be a limiting factor for animal growth when it is not limiting for plant growth. INTRODUCTION 27

1.5.3 Improved species

The need for new forage species is obvious, because native and naturalized spe­ cies frequently lack production capacity and/or nutritive value. Improved for­ age species have been selected for yield potential and nutritive value, but will frequently require amendments to soil fertility and some control of defoliation to express this potential. Since the main emphasis of this book is on species and not on management, some of the very large number of native or naturalized species and potentially useful introduced species are described in detail in Chapter 2. They are also listed in Table 4, with an indication of the climatic zones to which they are best adapted. The importance of adequate nutrition and correct management in optimizing the use ofthes e improved species isdiscusse d in a series of reviews edited by Blair et al. (1986). There is considerable variability within many of the promising native and introduced pasture species and this variability needs to be evaluated and any weaknesses clearly defined before commencing breed­ ing programmes. Where species are sown by seed, improving seed production can be important in reducing seed costs (Hopkinson, 1986) and in enhancing adoption by farmers. This is well illustrated by experience in Thailand where a government backed guaranteed price for seed of forage species has enabled smallholders to treat seed production as a cash crop.I n 1990,abou t 3000 farmers participated in this programme and produced about 500 t of seed of grass and Stylosanthes hamata (L.)Taub . (Manidool, 1990). ,

Table 4. Important or potentially useful forage species for South-East Asia described in Chapter 2 and an indication of their adaptation to the climatic zones defined by Troll (1966) (x = adapted, - = not adapted).

Forage species Climatic zones

1 Vj&Va V3 IVe+v IV, a. Native/naturalized grasses

Arundinaria pusilla X Axonopus compressus x X Brachiaria distachya x X Centotheca latifolia x Chrysopogonaciculatus x X Chrysopogonorientalis x X Dactyloctenium aegyptium x X Dichanthium annulatum x X Echinochloacolona x X Echinochloacrus-galli x X Eragrostis tenella x X Eragrostis unioloides x Heteropogoncontortus x X Hymenachne acutigluma X X Imperata cylindrica X X Ischaemum ciliare X 28 FORAGES

Table 4. Continued.

Forage species Climatic zones

1 Vi&V2 V. IVe.v IV,

Ischaemum magnum Ischaemum muticum Ischaemum rugosum Ischaemum timorense Leptochloa chinensis Microstegium ciliatum Ottochloa nodosa Panicum repens Paspalum conjugatum Pennisetum polystachion x Themeda triandra x Thysanolaena latifolia x Zoysia matrella x b. Selected grasses

Andropogon gayanus X X - - Bothriochloa pertusa - X - X Brachiaria brizantha X X - - Brachiaria decumbens X X - - Brachiaria dictyoneura X X - - Brachiaria humidicola X X - - Brachiaria mutica X X - - Brachiaria ruziziensis X X - - Brachiaria subquadripara X X - - Cenchrus ciliaris - X - X Chloris gayana - X X X Cynodon dactylon X X X X Cynodon nlemfuensis X X - - Digitaria ciliaris X X X - Digitaria eriantha X X X - Digitaria milanjiana X X X - 2 Oryza sativa X X - - Panicum maximum X X - - Panicum maximum var. trichoglume - - X X Paspalum dilatatum - - X - Paspalum distichum X X - - Paspalum notatum X X X X Paspalum plicatu lum X X X - Paspalum scrobiculatum X X X - 2 Pennisetum americanum X X - - Pennisetum clandestinum - - X - Pennisetum purpureum X X - - 2 Saccharum officinarum X X X - Saccharum spontaneum X X X - Setaria sphacelata X X X - Sorghum x almum - - - X Sorghum (art. perennial hybrids) - - - X INTRODUCTION 29

Table 4. Continued.

Forage species Climatic zones

1 VX&V2 V, IVe+v IV.

Sorghum x drummondii Stenotaphrum secundatum Tripsacum andersonii Urochloa mosambicensis Zea mays2

Shrub legumes

Albizia lebbeck Calliandra calothyrsus Codariocalyx gyroides Flemingia macrophylla Gliricidia sepium Leucaena leucocephala Sesbania grandiflora Sesbania sesban d. Herbaceous legumes

Aeschynomene americana Aeschynomene falcata Alysicarpus vaginalis x Arachis glabrata x Arachis pintoi 2 x Cajanus cajan x Calopogonium caeruleum x Calopogonium mucunoides x Canavalia ensiformis x Centrosema acutifolium x Centrosema macrocarpum x Centrosema pascuorum x Centrosema pubescens x Chamaecrista rotundifolia x Clitoria ternatea x Crotalaria juncea x Desmanthus virgatus x Desmodium heterocarpon x Desmodium heterocarpon ssp. ovalifolium x Desmodium heterophyllum x Desmodium incanum x x Desmodium intortum x Desmodium triflorum X Desmodium uncinatum X Lablab purpureus2 X Lotononis bainesii X Macroptilium atropurpureum X Macroptilium lathyroides X 30 FORAGES

Table 4. Continued.

Forage species Climatic zones

1 Vi&Va v3 IVe+7 IV4

Macroptilium longepedunculatum _ X - - Macrotyloma axillare - X X X 2 Macrotyloma uniflorum - X — - Medicago sativa - - X X Mimosa pudica X - - - Neonotonia wightii - - X - Pueraria phaseoloides X X - - Stylosanthes capitata X X - - Stylosanthes guianensis X X X X Stylosanthes hamata - X - X Stylosanthes humilis - X - X Stylosanthes macrocephala X - - - Stylosanthes scabra - X - X Trifolium repens - - X - Trifolium semipilosum - - X - Vignaparkeri - - X - 2 Vigna unguiculata X X - X e. Other species

Asystasia gangetica Ficussubcordata Manihot esculenta2 Mikania cordata

1. Species adapted to climatic zoneIV 6+7 are alsousuall y adapted to high altitudes inth e wettropics . 2.Specie sprimaril y in cultivation asa foo d crop,bu t frequently used asa forag e (seeSectio n 1.7).

The grasses and herbaceous legumes can be used in mixtures in grasslands and under plantation crops, or separately as forage crops. The tree and legumes can be used in protein banks, in alley cropping, in hedges, or as shade trees or single trees.I n selecting species for improving forage production, consi­ deration should be given to particular management requirements. For example, in cut-and-carry systems, erect grasses such as Panicum maximum and Pennise- tumpurpureum have advantages over more prostrate grasses such as Brachiaria spp. which are better suited to grazing. Ability to tolerate shading is a very important selection criterion for species under plantation crops. Ease of estab­ lishment under realistic conditions is also important and experimental studies can help to define the most appropriate establishment techniques (Wilaipon & Pongskul, 1984).I t is very advantageous if legumes can nodulate readily and consistently with native rhizobia. INTRODUCTION 31

1.5.4 Management and socio-economic constraints

In general terms, there is enough knowledge to predict the reaction of many introduced forage species to defoliation by cutting or grazing. This is shown by the many descriptions of the reaction of species to defoliation that are in the second chapter ofthi s book, and, in the case ofshrub s and trees, as indicated by Devendra (1990). However, much less is known about the potential benefits of controlled management or the introduction of herbaceous or tree legumes into natural grassland. Although our agronomic or biological understanding is still incomplete, it should be pointed out that socio-economic conditions are usually a greater constraint to introducing and managing forage species than our present understanding ofthei r biology and agronomy.

1.6 Selection of species to be included in this volume

As indicated in the previous sections, a very wide range ofplan t species contrib­ utes to the forage resources ofSouth-Eas t Asia. Some ofthes e species are native or naturalized and others have been deliberately introduced. Species evaluation throughout the region has also shown that there is considerable potential for using cultivars or accessions offorag e species that have been developed in other countries. In most instances more is known about the ecology and agronomy of these introduced species than there is of the important native or naturalized species, though useful information on the agronomy of some native grasses is given by Mehra &Fachruroz i (1985) and Skerman &Rivero s (1990). Forage legumes are often also cover, or pulse crops, particularly in humid climates. In many cases legumes serve more than one purpose simulta­ neously. The cover crops in plantation agriculture are grazed by or cut for domestic animals and the green manure crop can also be grazed or cut before being allowed to grow before being ploughed in. It would be impossible to give details about every forage species used in South- East Asia in this book, so the species included in Chapter 2 were selected after consultation with specialists throughout the region. Important native and naturalized forage species were included, provided there was sufficient agrono­ mic and ecological information. Section 1.7 gives some information about the forage properties of selected species that are primarily used for food crops, but where some parts of the crop are used for forage, (e.g., sugar cane, cassava and rice). In these cases, botanical and other relevant information will be included in the appropriate Prosea volumes. Forage species that are commercially available from Australia and South and , which are as yet little used in South-East Asia, have also been included. Many of these species are showing considerable promise in South- East Asia and are being used or are recommended for use on farms. They are also frequently mentioned in current documents such as journals or workshop proceedings. It was therefore appropriate to include them in this book so that their evaluation and subsequent use in farms may be made faster and more effi­ cient by making readers more aware of their strengths and weaknesses. Some temperate and subtropical legumes have been included as they show potential in highland areas (e.g. Nurjaya et al., 1983). An enumeration of species which were not included in Chapter 2,bu t are known 32 FORAGES

to be used as forages, is presented in Chapter 3. Although it does not include every species which is eaten by animals, it includes the species noted in the historical handbooks on plants of economic value in South-East Asia as being used for forage. Forages with another primary use are listed in Chapter 4. Information given under the headings such as 'Properties' and 'Yield' which refers to nutrient concentrations or dry matter yield is usually only in the form of broad ranges. These properties are much more variable than they are in the case of commodity groups such as pulse crops. They vary with the age of the material sampled, the grazing or cutting regime practised, soil fertility, and climatic conditions during the growth period prior to sampling. Consequently, it is usually misleading to quote specific figures or even narrow ranges of nutrient concentration and yield. Where ranges are quoted, they merely serve to give some idea of the values that are likely to be achieved under the usual conditions of field variability. Furthermore, if pastures are grazed, the quality ofth e material ingested by animals will depend upon the opportunities for selec­ tive grazing.

1.7 Species primarily in cultivation as a food crop

Practically all grass and legume species which are palatable to livestock and ofadequat e yield and quality are used as forage. The primary use ofsom e species depends on local conditions, tradition and on the availability of certain types of food. For example, Cajanus cajan, Lablab purpureus, Macrotyloma uniflorum and Vigna unguiculata are important pulse crops in many parts of Asia, but used only as forage in Australia. Oryza sativa, Pennisetum americanum, Sac- charum officinarum and Zea mays are cereal and sugar-producing crops and Manihot esculenta is a starch crop in much of the tropics, but in many places they are solely or partially used as forage. In addition, when these crops are used for food, they leave residues that can be used as forage. Some information relating to the forage qualities ofth e species just mentioned follows below. The species are treated in detail in other Prosea volumes. Cajanus cajan (L.) Millsp. (pigeon pea). A short-lived perennial shrub which can be grown as a protein bank or in alley cropping systems mixed with other food crops or grass as forage. The shrubs must be well established before they can be grazed intermittently, cut for forage, hay or silage. Defoliation should belenien t to achieve maximum persistence ofu p to about 5years .I t can produce DM up to 25 t/ha per year. Pigeon pea hay, made when a large percentage of pods are mature, serves as a substitute for industrial concentrate feed. The N concentration varies between 2.5% and 4.5%, and the plant is a good source of vitamin A. At the young stage, pigeon pea is not highly palatable, but it isreadil y eaten in the green pod stage (Skerman et al., 1988). Lablab purpureus (L.) Sweet (lablab). An annual or short-lived perennial crop with potential to provide early dry season forage when sown at the end of the rainy season. It can be leniently grazed 10 weeks after sowing, 2 or 3 times in a season, but is not tolerant ofheav y grazing. Livestock may require a few days before readily accepting the green material. Lablab can be cut for hay or silage. Dry matter yields of 20-30 t/ha can be expected during a 4-6 month growing period, with an N concentration of 2.5-4.2% and a DM digestibility between 50-65 % (Skerman et al., 1988). INTRODUCTION 33

Macrotyloma uniflorum (Lamk) Verde, ( gram). An annual herb grown in the rainy season, with reasonable drought tolerance, which can be used as a dry season feed reserve. Dry matter yield of 6.6 t/ha, consisting of over 2 t of seed, has been recorded in Queensland (Australia). N concentration ranges be­ tween 2.5-4.0% (Staples et al., 1983). Manihot esculenta Crantz (cassava). A perennial shrub with storage roots con­ sisting mainly of starch. The roots and leaves are used as livestock feed, but both may contain high levels of cyanogenic glucosides (HCN), which can be poisonous if eaten in large quantities. Heat treatment of roots and wilting of green leaves detoxify the material sufficiently for safe use. The roots have an N concentration of about 0.5%, but leaves may contain up to 5% N. Cassava leaves are an excellent supplement to poor quality forage. Oryza sauva L. (rice). An annual grass grown in either upland (dry) or lowland (flooded) conditions. Floating rice has a long vegetative growth period, and in deepwater rice areas in , India, Thailand and the Philippines the green vegetative material is harvested for forage for livestock, without reduc­ tion of the grain yield. N concentration of rice herbage ranges between 1.5-4.0% . The in vitro DM digestibility 40day s after transplanting lies between 70-84%. Deepwater rice cultivars have a greater nutritive value than lowland rice cultivars. In addition, throughout South-East Asia rice straw and bran are important livestock feeds. N concentration of rice straw ranges from 0.5-1.6% depending on maturity and and DM digestibility is about 50%. Straw quality can be marginally improved by treatment with urea. Rice bran is high in energy, but the N concentration varies greatly and ranges between 0.5-3.8 % (Lopez &Vergara , 1988). Pennisetum americanum (L.) K. Schum. exLeek e (). Arobus t annual grass used as a cereal crop in Africa and India, but elsewhere mostly as forage. Pearl millet should be grazed frequently but leniently, or cut at about 6-week intervals. DM yield up to 20t/h a can be obtained with N concentration of young regrowth ranging from 2.4-3.8% and DM digestibility from 60-75%. Young pearl millet is very palatable (Skerman &Riveros , 1990). Saccharum officinarum L. (sugar cane). A tall perennial grass used for the pro­ duction of sugar and as a dry season forage for livestock. Sugar cane has a high sugar content, but low N concentration (1.0%). DM digestibility at stage of harvest is about 50%. The tops of sugar cane grown for sugar production or the whole crop can be cut for hand feeding or chopped for silage (Skerman &Riveros , 1990). Vigna unguiculata (L.) Walp. (). An annual pulse crop which is also grown as a forage crop for grazing or cutting for hay or silage, giving DM yields of about 5t/h a with a N concentration ranging from 2.5-4.5 %. Cowpea is quite palatable to livestock after they have become accustomed to it in a few days. It can be grown together with annual grass forage crops (Skerman et al., 1988). Zea mays L. (maize). A tall annual grass grown for grain or forage for fresh feeding or silage. Maize can yield up to 8t/h a of DM, with a high energy content and N concentration ranging from 2.4% in young leaves to 1.4% in 10-week-old material (Skerman &Riveros , 1990). 34 FORAGES

1.8 Prospects

In most parts of South-East Asia animal production based on forages is one way of increasing the incomes and security of smallholder farmers. Although all the arable land in densely populated areas is usually already in use, there are still opportunities to either collect or cultivate forage to feed animals. The main restriction will often be lack of surplus labour. Despite the importance of socio-economic limitations to forage production, there is ample evidence that improved practices will be adopted. This is illus­ trated by the use of Stylosanthes spp. in the dry tropics of Thailand, the use of Leucaena leucocephala in many countries and the promising early adoption of the 'three strata farming system' in the drier areas of Indonesia (Nitis et al., 1990).Th e agronomic areas where there is likely to be the greatest potential for improvement, within existing socio-economic conditions, are in the use of shrub or tree legumes, backyard or home garden forage production, making better use ofbund s between rice paddies, forage under plantation crops - partic­ ularly coconuts, and use of annual legumes with or after food crops. Recent progress in improving animal production by shrub legumes or by forages under plantation crops has been reported by Devendra (1990) and Shelton & Stiir (1991), respectively. A small but increasingly important system with the poten­ tial for improvement is that involved in the supply ofmil k to urban areas. If improved forage systems can be effectively incorporated into individual smallholdings, as in the case of the 'three strata farming system', there is more likely to beread y acceptance ofimprovement s than when they affect a commun­ ity, as in the case of common grazing land. Forage improvement schemes will be accepted by a community much more easily, if the community 'owns' the scheme rather than if the scheme is imposed on the community. Nevertheless, governments can also encourage improvements, as in the case of leucaena and other shrub/tree legumes in Indonesia (Rangkuti et al., 1990) and the Philip­ pines (Trung, 1990) and through the development of 'mini-farms' in Malaysia (Chin, 1989).I t ishighl y desirable to have an effective and sustained educational and 'follow-up' scheme to support farmers adopting any improved practice (Nitis et al., 1990). The key point is that although the constraints imposed by socio-economic factors must be recognized, they are not insurmountable (Per­ kins et al., 1986). There are many forage species to choose from and proven species are available for all climates in South-East Asia. This diversity should be utilized as much as possible, because it is undesirable to rely on one or a few species or cultivars. Reliance on a narrow germplasm base has the danger that a pest or disease will destroy all or a large part of the feed supply and thus endanger the income of farmers. This was well illustrated by the hardship experienced by farmers who were relying heavily on Leucaena leucocephala prior to the advent of the leucaena psyllid in South-East Asia (Trung, 1990). 2 Alphabetical treatment of species AESCHYNOMENE 37

Aeschynomene americana L.

Sp. PL: 713(1753) . LEGUMINOSAE In = 20 Synonyms A.javanica Miquel (1855). Vernacular names American jointvetch (Am). Indonesia: kacang meongan, asem-aseman, anjang. Philippines: makahiyang-lalaki (Taga- log), karaparak (Mar.). Thailand: sano-don, sano- bok (central), sano khon (north-east). Vietnam: pötuk, rokdönao, rui köjing. Origin and geographic distribution American jointvetch is native to the and adjacent areas of the between latitudes 30° N and 30°S . It has been introduced into several countries in South-East Asia. Uses American jointvetch is used as a cut-and- carry forage for animals, and as a hay crop. It is used as a pasture legume in Florida (United States), the tropical east coast of Australia, and Vanuatu. It is also applied as a green manure crop in rice and other cropping systems. Properties Nitrogen concentrations of green leaf range from 2.5-4.0% and of stems from 0.3-1.0%. In vitro digestibility of leaves is 60-70 %. Although tolerant oflo w soil P,yiel d and P concentrations respond greatly to applied P. Ripepod s divide into single-seeded segments (seed- Aeschynomene americana L. - 1, branch with in-pod) with 150-220 segments/g. Dehulled or leaves; 2,flowering and fruiting branch. naked seed varies in size with 350-400 seeds/g. Description Annual or perennial herb or sub- seeding, self regenerating annuals or short-lived shrub, growing 1-2 m tall and 1.5-2 m wide, glan­ perennials with crowns surviving up to 2-4 years. dular-hispid to subglabrous, with prostrate to The seedling phase is relatively slow, but subse­ erect, hard but pithy main stem 5-10 cm thick at quent growth is rapid in hot and moist conditions. base. Branches form roots where they touch the Nodulation isprolifi c and, under waterlogged con­ ground. Habit changes dramatically under graz­ ditions, nodules form on lower parts of the main ing where plants branch close to the ground, form­ stem and branches. It has a short-day flowering ing a leafy sward. Leaf 2-7 cmlong ,pinnatel y com­ response. Flowering commences 60-200 days after pound, 20-60-foliolate; leaflet more or less sickle- germination, with another 30-60 days to ripe seed. shaped, 3-15 mm x 1-3 mm, glabrous but ciliate Away from the equator, perennial plants will also at least along one margin, 2-several-costate, fold­ flower and seed in spring. Although most of the ing together at night or when touched. Inflores­ pollen is shed before flowers open, recent studies cence axillary, racemose, few-flowered, about the indicate that up to 30% outcrossing occurs. Yield length ofth e subtending leaf;pedice l ca. 1 cm long; of DM is higher in the late maturing types. Up to flower papilionaceous, 3-10 mm long, yellow, 90% of freshly harvested seed is 'hard' and or mauve, usually with red or purple requires scarification before sowing. Natural stripes. Fruit a straight or slightly curved pod, 1-3 breakdown ofhardseedednes s is sufficient to allow cm long, 3-9-articulate; articles (segments) 2.5-5 re-establishment in the field. mm x 3-6 mm, glabrous to villous-hispid, light- Other botanical information Sometimes A. brown, more or less muricate, margins thickened. americana and A. villosa Poiret (syn. A. javanica) Seed kidney-shaped, 2-3 mm x 1.5-2 mm, dark- are considered as different species. In that view, A. brown to blackish. americana originates from the Caribbean and adja­ Growth and development Plants are heavy cent areas, usually in wet or moist places, up to 1400 38 FORAGES m altitude, and A. villosa originates from southern tions neither disease isa problem. Powdery mildew Arizona (United States) to northern South Amer­ (Oidium sp.) turns mature leaves white towards ica and the Antilles, usually in drier areas up to the end of the growing season, but does not affect 2250 m altitude. Discriminative morphological quality or acceptability. In South America an- characteristics of A. americana and A. villosa thracnose {Colletotrichum gloeosporioides) has would be respectively: flowers 6-8 mm versus 3-5 been found on American jointvetc h but not in Aus­ mm long;leaflet s more than, versus less than, 5m m tralia or Florida. long; mature fruit-centre muricate versus non- Harvesting A. americana should be cut for hay muricate. Young material of both species is impos­ or ploughed in for green manure at full flower. For sible to distinguish. Unification ofbot h taxa seems cut-and-carry, the forage should be cut above 50 tobe justified. cm prior to flowering to encourage regrowth. Ripe Cultivar 'Glenn' was registered for use in tropical seed remains on the plant so plants should be coastal Queensland, Australia, in 1983 and 'Lee', allowed to mature if commercial seed production a more perennial form, in 1991. Common American isth e aim. Ripe, dry seed will store and remain via­ jointvetch has been used in Florida, United States, ble for many years. since the early 1970s. Descriptions of collections Yield Dry matter yields of 10-15 t/ha for a full grown in Florida and Australia indicate wide season's growth have been recorded in Queens­ diversity ofmateria l differing in range of maturity land. In Florida, cutting for hay, initially at 30 cm and perenniality, plant habit and size. with a second cut at 90cm ,yielde d 4.5t/h a of quali­ Ecology American jointvetch grows in poorly ty hay per season. Five cuts between March and drained and waterlogged conditions. Best growth November produced 7.5t/h a of hay in Puerto Rico. of American jointvetch occurs in tropical areas For seed production, direct header harvesting with hot, moist climates, normally with an annual yielded up to 1000 kg/ha seed-in-pod in northern average rainfall of over 1000 mm and most com­ Queensland. monly in poorly drained or run-on situations. How­ Genetic resources Collections are held at ever, its natural distribution covers a wide range CIAT (Columbia); CENARGEN/EMBRAPA (Bra­ of latitude and altitude (up to 2250 m). It should silia, ); ATFRGRC (CSIRO, Australia); Uni­ be possible to select for cold tolerance and differ­ versity of Florida, Fort Pierce, Florida; IPB, Uni­ ent rainfall requirements. 'Glenn' has a summer versity of the Philippines (the Philippines). minimum rainfall requirement of about 1000 mm. Breeding Most effort is directed at evaluating It prefers wet, poorly drained or waterlogged areas existing collections, but limited breeding work is on sandy to clay soils. being undertaken at the University of Florida to Propagation and planting Suggested seeding achieve nematode resistance and increased forage rate is4- 8 kg/ha seed-in-pod or 2-4 kg/ha dehulled yield. seed. Half these rates can be used in a mixture. Prospects American jointvetch is a high yield­ Dehulling is the best method of scarification and ing, high N fixing legume which tolerates water­ will raise germination to about 30%. When sown logging and is palatable to stock. Its role in South- with a companion grass or grown as a pure stand East Asian areas will include green forage in pure for forage or hay, it is best sown into a well pre­ stands or in mixed pastures, hay production, green pared seed-bed. Satisfactory establishment can be manure, and ley pastures for rice paddies.I t should achieved by spreading seed onto the surface of be possible to select cultivars to suit climatic and well-grazed pasture. growth habit requirements. Husbandry In grazed pastures, American joint­ Literature |lj Bishop, H.G.,Ludke , D.H. &Ruth ­ vetch should be kept 50-60 cm tall to promote leaf erford, M.T., 1985.Glen n jointvetch: anew pasture production. Grazing pressure should be reduced legume for Queensland coastal areas. Queensland after flowering to promote maximum seed produc­ Agricultural Journal 111: 241-245. |2j Bishop, tion, particularly in the year of planting. Seed is H.G., Pengelly, B.C. &Ludke , D.H., 1988. Classifi­ spread in dung following ingestion by animals. cation and description ofa collection ofth e legume Diseases and pests Commercial seed crops may genus Aeschynomene. Tropical Grasslands 22: require control of Heliothis larvae which eat the 160-175. |3| Hodges, E.M., Kretschmer, A.E. Jr, flowers and green pods. Where cool, showery Mislevy, P., Roush, R.D., Ruelke, O.C. & Synder, weather occurs at flowering, spraying with ben- G.H., 1982. Production and utilisation of the trop­ late may be required to control the fungal disease ical legume Aeschynomene. Circular S-290. Flor­ botrytis stem rot (Botrytis cinerea). In grazed situa­ ida Agricultural Experimental Stations Institute AESCHYNOMENE 39 of Foods and Agricultural Services. University of Florida, Gainesville. 11 pp. |4| Paul, W.K.C.,1951 . Notes on legumes. 1.Tropica l Agriculturalist 107: 15-20. |5SRudd , V.E., 1955.Th e American species of Aeschynomene. Contributions from the United States National Herbarium 32: 23-30. |6| Rudd, V.E., 1959. The genus Aeschynomene in Malaysia (Leguminosae-Papilionatae). Reinwardtia 5: 23-36. 17] Singh, R.G., 1971.Prospect s of utilising wild legumes as green manure in a paddy - rotation. The Allahabad Farmer 55: 489-495. |8j Skerman, P.J., Cameron, D.G. &Riveros , F., 1988. Tropical forage legumes. FAO, Rome. pp. 205-211. H.G. Bishop

Aeschynomene falcata (Poiret)DC .

Prodr. 2:32 2(1825) . LEGUMINOSAE In = unknown Synonyms Hedysarum falcatum Poiret (1804), H. diffusum Vellozo (1825, 1835), Aeschynomene apoloana Rusby (1910). Vernacular names Jointvetch (used for all members of the genus) (En).Indonesia : turi rawa (Javanese). Philippines: torog-torog (Bikol). Origin andgeographi c distribution A. falcata isnativ e toth ehighe r altitude tropical or subtropi­ Aeschynomene falcata (Poiret) DC. - 1, habit; 2, cal parts of , Bolivia, Brazil, Colombia, flowering and fruiting branch. and Paraguay. It has been introduced into many South-East Asian countries and Australia. curved, stipe 6-14 mm long; articles 3-4 mm x Uses A. falcata is used in grazed pastures and 2.5-3.5 mm, the body of the articles tending to is not appropriate to cut-and-carry systems. It has break away from the margins. Seed yellowish- been tested as a low-growing cover crop. brown toblack , 2-2.5 mm x 1.4-1.7mm . Properties Nitrogen concentrations range from 'Bargoo' plant s develop slowly andma y take sever­ 1-2%, and in vitro DM digestibilities from al years to reach maximum size. They flower 42-65 %. Phosphorus concentrations are typically throughout thegrowin g season provided moisture 0.1-0.2%. A. falcata is readily eaten by livestock is adequate. Pod segments open when seed is ripe. although older stems mayb e rejected in favour of Spread of seed is assisted by passage through the young grass. In the Australian cultivar 'Bargoo' grazing animal. Persistence is ensured by build-up there are37 5 seeds/g. of soil seed levels from 1000 to 8000/m2, depending Botany A prostrate, herbaceous perennial with on management history and environment. a short tough taproot. Stems up to 1 m long and Ecology A. falcata occurs naturally on rocky 1-3 mm diameter, puberulent, decumbent, show­ hillsides, in , and in cultivated fields, at ing little tendency to produce roots at the nodes; elevations to about 1800 m. 'Bargoo' originates small nodules to about 2 mm diameter occur on from 25°30'S in central Paraguay where it was both the taproot and secondary roots; older stem growing on a rocky sandstone ridge in forest bases woody to 3mm thick. Leaf 5-7(-9) foliolate; country. In eastern Australia, 'Bargoo' is adapted leaflet obovate-elliptical, 6-12 mm x 2.5-4mm , to a variety of poor to moderately fertile soils der­ pubescent. with usually only 1o r 2 ived from sandstone, shale and granite between flowers developing, longer than the subtending 20-30°S. Most areas where 'Bargoo' has proven leaf; flower 7-10m mlong , yellow, with orbiculate successful have an annual rainfall between standard. Pod4-8-articulate , puberulent, slightly 700-1200 mm. It can tolerate temporary waterlog- 40 FORAGES ging and periods of drought, but grows best on 276-277. |4j Rudd, V.E., 1955. The American spe­ moist, well-drained soils. It does not tolerate sus­ cies of Aeschynomene. Contributions from the tained wet conditions. On suitable soils it is a very United States National Herbarium 32: 88-90. |5| persistent legume. Skerman, P.J., Cameron, D.G. &Riveros , F., 1988. Agronomy 'Bargoo' is readily established from Tropical forage legumes. FAO, Rome. pp. 211-212. seed. Because it is often extremely hard, seed may |6| Wilson, G.P.M., 1980. Bargoo jointvetch: tough need scarification if a high level of immediate ger­ legume for tough country. Agricultural Gazette of mination is desired. Seed should be inoculated New South Wales 91: 51-53. ,7! Wilson, G.P.M., with an appropriate Bradyrhizobium strain, al­ Jones, R.M. & Cook, B.G., 1982. Persistence of though effective nodulation may be achieved from jointvetch (Aeschynomene falcata) in experimen­ native strains in some soils. Best establishment is tal sowings in the Australian subtropics. Tropical obtained from shallow or surface sowing on a Grasslands 16:155-156. clean, well prepared seed-bed. Although tolerant B.G. Cook of low fertility, it can respond to applications of P. The only disease of any consequence is caused by Albizia lebbeck (L.) Benth. Colletotrichum sp. which attacks the stem near the growing tip, ultimately resulting in tip death. This Lond. Journ. Bot. 3:8 7(1844) . is a problem in humid environments, particularly LEGUMINOSAE in seed crops. 'Bargoo' is not affected by amnemus 2n = 26 weevil {Amnemus quadrituberculatus) and is resis­ Synonyms Mimosa lebbeck L. (1753), Mimosa tant to rootknot nematodes (Meloidogyne spp.) sirissa Roxb. (1832),Albizia latifolia Boivin (1838). It is extremely tolerant of, and can still produce Vernacular names Siris, koko, East Indian seed under heavy grazing. 'Bargoo' is too low- walnut (En). Indonesia: tekik (Javanese), kitoke, growing for ease of hand harvest, and is therefore tarisi (Sundanese). Philippines: aninapla, langil grazed. Yields are generally low and may not (Tagalog). Cambodia: chreh. Thailand: kampu exceed 1-2 t/ha of DM per year. 'Bargoo' is not (general), chamchuri (central). Vietnam: bô kêt suitable for hay as leaves shed readily. tây, lim xanh, trat. Genetic resources and breeding A small col­ Origin and geographic distribution Siris is in­ lection, including 'Bargoo', is held at ATFGRC digenous to the and to those (CSIRO, Australia). No breeding programmes are areas of South-East Asia with a marked dry season anticipated. It was tested with moderate short- (e.g. north-east Thailand, the eastern islands of term success on Sumba Island (10°S, 770m m rain­ Indonesia) and to the areas of northern fall with a 4 month wet season) in eastern Indone­ Australia. It has been introduced widely through­ sia. out the tropics and has become naturalized in Prospects Since the only commercial cultivar many places. ('Bargoo') originates from near the southern ex­ Uses The green leaf is a valued forage for rumi­ tremity of its natural distribution, which has only nants and it should be useful in extensive grazing been evaluated to a limited extent globally, it is systems because of the feed value of the natural difficult to speculate on the prospects of the spe­ drop of leaves, flowers and pods. Free-standing cies. 'Bargoo' should have a role in those parts of trees appear to enhance pasture production and the sub-humid tropics and subtropics with heavily quality beneath the canopy due to increased N sta­ grazed pastures on light, infertile and well drained tus of the soil. In India plantation-grown siris soils. yields a high quality hardwood traded in Literature |1| Bishop, H.G., Pengelly, B.C. & as 'Indian walnut' or 'koko'. More generally it is Ludke, D.H., 1988. Classification and description useful for fuelwoo d because of its high productivi­ ofa collection ofth e legume genus Aeschynomene. ty. It is a valued honey tree because of its produc­ Tropical Grasslands 22: 160-175. |2| Hennessy, tion of both nectar and pollen. It is a popular ame­ D.W. & Wilson, G.P.M., 1974. Nutritive value of nity tree throughout the dry tropics because of its Bargoo jointvetch (Aeschynomene falcata) and shady spreading habit, although the copious litter companion grasses when fed to sheep. Journal of is often regarded as a disadvantage. the Australian Institute of Agricultural Science Properties In feeding experiments with sheep, 38: 82-84. |3|Oram , R.N., 1990.Registe r of Austra­ DM digestibility and N concentration, respective­ lianherbag e plant cultivars. CSIRO,Australia , pp. ly, were as follows: green leaf fed fresh: 64% and ALBIZIA 41

3.5%; mature green leaf fed dry: 48% and 3.0 %; what asymmetrical with midrib nearer upper mar­ fallen leaf: 42% and 1.6%; fallen flower: 57%> and gin, subglabrous, initially bright green and folding 3.8%; fallen pods: 44% and 3.0%. Leaf is notable at night, maturing to a duller glaucous green with for the absence of antinutritional factors such as position fixed on rachis. Inflorescence a terminal phenolics, but pods contain saponins. There are and axillary globular cluster of 15-40 pedicellate 7-8 seeds/g. flowers, often 2o r more together per axil; peduncle The wood is dense (specific gravity 0.5-0.6), easily up to 10 cm long, pedicel 1.5-5(-7.5) mm long; worked, yellow-brown, with a very distinct bound­ calyx tubular, 2-5 mm long, ending in 5 triangular ary between heartwood and pale sapwood. A range teeth; corolla tubular, 5-11 mm long, ending in 5 of uses such as cabinet timber have been listed, triangular lobes which are hairy at the apex; sta­ including the doors of Chinese temples.Th e massic mens at base united in a tube, free filaments energy of air-dried wood is 21840kJ/k g (5200 kcal/ numerous, 1.5-3 cm long. Pod flat-oblongoid, kg). 12-35 cm x 3-6 cm, much swollen on the seeds, Description Deciduous tree, in plantations subglabrous, glossy, veined, pale yellowish, dehis­ 3-15 m tall, bole length 3 m with diameter 45 cm; cent. Seeds 3-12 per pod, flattened ellipsoidal, when grown in the open, often much larger, up to 7-11 mm x 6-9 mm x 1-1.5 mm, brown. 25 m tall, often multi-stemmed and widely spread­ Growth and development Siris has a strongly ing (up to 30 m diameter); bark rough, grey, some­ seasonally-dependent growth pattern with a leaf­ what flaky; inner bark reddish; branches terete, less period of 1-2 months in the middle of the dry puberulous or pubescent when young. Leaves season. Production of new season growth com­ bipinnate with 1-5 pairs of pinnae on a rachis of mences in the dry season, shortly followed by 8-9 cm length; leaflets in 3-11 pairs, oblong to abundant flowering which produces very little elliptical-oblong, 1.5-6.5 cm x 0.5-3.5 cm, some- seed. Further growth and fruiting occur as the wet season develops. Flowers are pollinated. Pods mature in the early dry season but;remai n on the tree for 3-4 months. Seed dispersal seems to occur mainly due to strong wind, when intact pods can be carried hundreds of metres. Some seed passes through the digestive tract of cattle but not of smaller ruminants. Light transmission through the canopy of free-standing trees is 40-50%). In an open woodland environ­ ment it has been repeatedly observed that there is modification of the ground cover with enhance­ ment of grass production and quality beneath the canopy. Growth stops in the early dry season, 2-3 months before leaf drop. New growth may be induced by fire damage, grazing or lopping. Other botanical information Albizia Durazz. is a large genus and in parts ofit s range non-fertile material of siris may be quite similar to other local species. There is little problem with identification when both flowers and pods can be examined. Use of 'albizia' as a vernacular name should be avoided assom e workers apply it also toParaserianthes fal- cataria (L.) I. Nielsen, a very important species in South-East Asia and quite different from siris. Albizia is often misspelt as Albizzia and lebbeck as lebbek. Ecology Although siris will grow in the humid tropics, its natural range is in semi-arid to sub- Albizia lebbeck (L.) Benth, - 1,flowering branch; humid areas with marked dry and reliable wet sea­ 2,flower; 3. fruit. sons. However, it may be established under condi- 42 FORAGES

tions of low (400 mm/year) and irregular rainfall. accessions. The value of siris is critically depen­ Seedlings will not tolerate frost or waterlogging. dent upon resistance to insect attack and this Reserves in the root system enable young plants should be the first objective of development work to survive total defoliation from fire or grazing, on the species. but with an obvious setback in growth. Siris is tol­ Prospects Siris offers excellent prospects for de­ erant of salinity and can be established on most veloping more productive agroforestry systems in soils except cracking clays. areas in the semi-arid tropics. It would appear to Propagation and planting The species is not have excellent multipurpose aspects, being poten­ particularly hard-seeded and requires only mild tially useful for cabinet timber, fuelwood, animal treatment (e.g. in water at 50°C for 3 minutes) to feed, and for enhancing crop production. Perhaps germinate successfully; a proportion of seeds ger­ most remarkable is the prospect of obtaining minate immediately without any treatment. There enhanced pasture production in addition to the is nothing published on preferred rhizobial strains tree products. Despite its great economic impor­ but it appears to nodulate readily without inocula­ tance on the Indian subcontinent and its wide­ tion. Plants can be direct-sown, container grown, spread distribution in South-East Asia, its quali­ or raised in a massed seed-bed and planted out as ties are little known there. This may reflect a lack bare-rooted stems. of research and development in the drier ('outer') Husbandry Siris is probably not productive areas in the region. Other species of Albizia are under repeated cutting (more than 2cut s per year). worth considering for the same purposes, in partic­ It does not develop a shrubby habit and is thus not ular A.procera (Roxb.) Benth. suitable for direct browsing. However, larger trees Literature |1! Lowry, J.B., 1989.Agronom y and can be lopped annually with removal of the entire forage quality of Albizia lebbeck in the semi-arid green crown without loss of vigour. tropics. Tropical Grasslands 33: 84-91. i2i Lowry, Diseases and pests Establishment can be J.B., 1991. Integrated production from Albizia leb­ adversely affected by grazing of young plants by beck trees and tropical pastures. Journal of the mice,rabbit s and other wildlife. Leaves are largely Australian Institute of Agricultural Science 4: unaffected by ,bu t young leaves may be sub- 36-38. |3|Lowry , J.B., Lowry, J.B.C., Jones, R.J., ject to heavy prédation by larvae of the grass yel­ 1988. Enchanced grass growth below canopy of low butterfly {Eurema hecoba); this appears to be Albizia lebbeck. Nitrogen fixing Tree Research a short-lived effect. The most serious pests are Reports 6: 45-46. |4| Murugan, M. & Kathaperu- bark-feeding larvae of longicorn beetles. These do mal, V., 1987. Nutritive evaluation of vagai (Albi­ not affect small stems and have little effect on zia lebbeck) leaves for goats. Indian Journal of An­ large stems, but through complete girdling can imal Nutrition 4: 61-62. |5i Nielsen, I., 1981. cause dieback in stems with diameters ranging Légumineuses-Mimosoïdées. Flore du Cambodge, from 4-10 cm. There is considerable variation in du Laos et du Vietnam. Vol. 19.pp . 82-84. |6I Prin­ susceptibility of individual trees. Trees may be sen, J.H., 1987. Potential of Albizia lebbeck as a more susceptible under prolonged drought stress. tropical fodder tree - a review of literature. Trop­ Harvesting Branches may be lopped for fuel and ical Grasslands 29: 78-83. |7|Schlink , A.C., Lowry, forage. In extensive grazing systems, the free-stand­ J.B. & Gibson, D.S., 1990. Products from the tree ing tree itself provides feed. Production of sawn legume Albizia lebbeck as supplements for sheep timber involves felling the wholetree ,wit h concom- in dry tropics. Proceedings Australian Society for mittant production offuelwoo d and forage. Animal Production 18:546 . Yield Comprehensive yield data have not been J.B. Lowry published. Under best conditions plants can grow 5 m/year. Fuelwood plantations produce 5 m3/ha per year. Isolated mature trees produce edible DM Alysicarpus vaginalis (L.) DC. at the rate of 100-120 kg/year. Roadside trees in the dry tropics show a crown diameter expansion Prodr. 2:35 3(1825) . of 2-2.2 m/year until mature. LEGUMINOSAE Genetic resources The wide natural range sug­ In = 16,20 gests a broad genetic base, but there are no com­ Synonyms Hedysarum vaginale L. (1753), Alysi­ prehensive germplasm collections. carpus nummularifolius (Willd.) DC. (1825). Breeding There are no reported programmes in Vernacular names Alyce clover, buffalo breeding, collection, or evaluation of existing clover, one-leaf clover (En, Am). Indonesia: bro- ALYSICARPUS 43 bos, gudé oyod, tebalan (Java). Malaysia: akar Botany Annual or short-lived perennial, erect seleguri. Philippines: banig-usa, mani-manian to prostrate herb with many stems 10-100 cm long (Tagalog). Thailand: thua lisongna. Vietnam: cây emanating from the rootstock. Stem glabrous to me dât, cây the the. pubescent, rooting at the nodes under sustained Originan dgeographi c distribution A. vaginalis moist conditions. Leaf unifoliolate; leaflet lanceo­ is native to and widespread throughout East Africa late to ovate, 5-65 mm x 3-25 mm, glabrous to including , the Indian sub-, puberulous; petiole 4-15 mm long; stipules lanceo­ South-East Asia and the Pacific, and is naturalized late. Flowers about 6m m long in terminal and leaf- in northern Australia (especially in the Northern opposed up to 13 cm long; corolla Territory), in South America and the United States. orange to purple and very variable. Fruit 12-25 Uses A. vaginalis is a useful component of native mm long, well exserted from the calyx, puberulent, pastures in and northern Australia, especially reticulate, not or slightly constricted between the where fertilizer has been applied, and isa useful fod­ 4-7 articles which are subcylindrical, 2.5-3 mm der legume in the Philippines, Indonesia and south­ long with raised ridges. Seeds dark red, 1-1.5 mm ern China. It has been used as a cover cropi n Papua long, copiously produced. New Guinea and in rubber plantations in Java, and The species is extremely variable in habit, leaf also as a hay crop in the United States.I n Indonesia shape and flower colour. Annual specimens with and Malaysia a decoction ofroot s isapplie d against lax inflorescences have been distinguished as A. coughs. In Vietnam an infusion of powdered seeds ovalifolius (Schum.) Leonard in West Africa. Dis­ isuse d against dysentery and colics. tinction below the species level seems more appro­ Properties The few recorded analyses of priate. nutrient concentration and nutritional value of A. Ecology A. vaginalis grows on a wide range of vaginalis are within the usual ranges reported for soil types, from coraline sands to clays, but prefers tropical legumes. There are about 650 seeds/g. lighter soils. It has been collected from very acid (pH(H20) 4.5) to neutral soils. The species usually occurs in seasonally dry climates with total annu­ al rainfall of between 700 and 1700 mm. It does not tolerate waterlogging and good drainage is essen­ tial. The species is a common of throughout the Asian region and so appears adapted to frequent defoliation and grazing. It is in these situations that the species perennates, whereas in the seasonally dry climates it usually behaves as an annual. Stands in northern Austra­ lia are variable between seasons and this can be attributed in part to the species behaving as an an­ nual in this environment and to the high propor­ tion of hard seed. Agronomy Propagation is by seed, which usually has a high percentage ofhar d seed. Scarifi­ cation isthe n required for immediate germination. Sowing rates of 2-4 kg/ha should be adequate, al­ though in the United States, where it is used as hay crop,rate s of up to 16kg/h a are recommended. Rhizobiarequirement s are non-specific so inocula­ tion prior to sowing is unnecessary. It is susceptible to root-knot nematodes (Meloido- gyne spp.) so may sometimes be restricted to hea­ vier soils where nematodes are less of a problem. On soils with high populations of nematodes, yields decline after the first year. Germplasm has not been evaluated for resistance to nematodes. Alysicarpus vaginalis (L.) DC. - 1, flowering and A. vaginalis ispalatabl e to livestock and is usually fruiting branch; 2,flower; 3,fruits; 4, seed. grazed directly, but it can be made into hay. Al- 44 FORAGES

though it will persist on soils of low fertility, pro­ Andropogon gayanus Kunth ductivity in northern Australia has been markedly improved by the addition ofP .Whe n grown in pure Enum. PL 1:49 1(1833) . swards, DM yields are typically 4-6 t/ha. GRAMINEAE Genetic resources and breeding Germplasm 2re = 20 (var. tridentatus (Höchst.) Hack.), 40 collections are maintained by ATFGRC (CSIRO, (other varieties) Australia) and CIAT (Colombia). Synonyms Andropogon bisquamulatus Höchst. No breeding programmes are in progress or have (1842), A. squamulatus Höchst. (1842), A. tridenta­ been attempted. The extreme variation in plant tus Höchst. (1842). morphology and in climatic and edaphic origin has Vernacular names Gamba grass (En). Philip­ enabled the selection of accessions for advanced pines: batag (Tagalog), batad (Bikol), bukakau evaluation from wild collections. (Ilokano). Thailand: ya kumba. Vietnam: hung Prospects A. vaginalis is a useful forage legume th'ao. in parts of the tropics and subtropics both for hay Origin and geographic distribution Gamba and as a component in permanent pastures. The grass originates in Africa where it extends widerang e ofgeneti c material available should en­ throughout the tropical region from 15°N to able further development of the species within almost 25°S. Var. bisquamulatus (Höchst.) Hack., South-East Asia. which is of particular interest as a forage, only Literature |1| Duke, J.A., 1981. Handbook of occurs as an important component of fire-climax legumes of world economic importance. Plenum savanna vegetation north of the equator from Press, New York & London, pp. 15-17. |2| Gillett, Senegal to Sudan in regions with annual rainfall J.B., Polhill, R.M. &Verdcourt , B., 1971. Alysicar- of between 400-1500 mm. Commercial or semi- pus. In: Milne-Redhead, E. & Polhill, R.M. (Edi­ commercial gamba grass cultivars or lines have tors): Flora of tropical East Africa. Leguminosae spread to most tropical regions ofth e world, partic­ 4 - Papilionoideae 2. Crown Agents for Oversea ularly in tropical America. Governments and Administrations, London, pp. Uses The main use of gamba grass is as forage 491-501. |3[Gramshaw , D., Pengelly, B.C., Muller, in permanent pastures grazed by ruminants. The F.W., Harding, W.A.T .& Williams , R.J. , 1987.Clas ­ stems are used for thatching, and when flattened, sification ofa collection ofth e legume Alysicarpus for coarse matting. using morphological and preliminary agronomic Properties Gamba grass provides a palatable attributes. Australian Journal of Agricultural forage when young but feeding value declines rap­ Research 38: 355-372. |4| Martin, T.J. & Torssell, idly with forage age and a decreasing leaf/stem B.W.R., 1974. Buffalo clover (Alysicarpus vagina­ ratio. Nitrogen concentrations range from lis (L.) DC): a pasture legume in Northern Austra­ 0.5-2%, and in vitro DM digestibilities from lia. Journal of the Australian Institute of Agricul­ 40-55%. Mineral contents are low (0.08-0.14% P tural Science 40: 232-234. I5| Nguyen van Thuân, and 0.27-0.39% Ca). Seed of commercial quality Dy Phon, P.,Niyomdham , C.& Vidal , Y., 1987.Aly ­ contains 60-120 viable seeds/g. sicarpus Desv. In: Morat, Ph. (Director): Flore du Botany An erect, coarse, and -forming, Cambodge, du Laos et du Vietnam. Vol. 23. Légu- perennial bunch grass with short and mineuses-Papilionoïdées. Muséum National stems 1-3 m high. Root system consists of fibrous, D'Histoire Naturelle, Laboratoire de Phanéroga- horizontally growing roots, vertical roots pene­ mie, Paris, pp. 128-133. |6| Skerman, P.J., trating as deep as 2-3 m and strong cord roots con­ Cameron, D.G. &Riveros , F., 1988.Tropica l forage taining starch granules. Leaf-sheath up to 20 cm legumes. FAO, Rome. pp. 246-247. |7| van long, densely hairy at base; leaf-blade linear-lan­ Meeuwen, M.S., van Steenis, C.G.G.J. & Stem­ ceolate, up to 100 cm x 4-30 mm, acute, usually merik, J., 1961. Preliminary revisions of some narrowed to the prominent midrib at the base to genera of Malaysian Papilionaceae 2. Alysicarpus form a pseudopetiole; pubescent on both sides, par­ Desv. Reinwardtia 6:86-89 . |8| Verdcourt, B.,1979 . ticularly when young. Inflorescence consists of A manual of New Guinea legumes. Botany Bulle­ paired , 4-9 cm long, bearing about 17 tin No 11. Office of Forests, Division of Botany, spikelet pairs; sessile spikelet up to 8 mm long, Lae, Papua New Guinea, pp. 420-422. with a long (10-30 mm) conspicuous awn; pedi- R.A. Halim &B.C . Pengelly celled spikelet 5-8 mm long, hairy (var. bisquamu­ latus), with a short (1-10 mm) awn; each spikelet with 2florets , but only upper floret ofsessil e spike- ANDROPOGON 45

humid tropics with an annual rainfall between 800-1500 mm. It still grows well in environments with annual rainfall up to 2500 mm with a marked dry season. Due to its deep rooting, it is remarka­ bly drought-tolerant and survives up to 6 dry months. It is also tolerant of fire. It is adapted to a wide range of soils, but grows best on those that are medium-textured. It has low nutrient require­ ments and exhibits excellent growth on acid, highly Al-saturated oxisols and ultisols of low fer­ tility, including low available P. Agronomy Gamba grass is established by drill­ ing or broadcasting viable pure seed at a rate of 0.75-1.25 kg/ha (equivalent to 10-15 kg/ha of aver­ age-quality commercial seed). Seed should not be sown immediately after it is harvested, as dor­ mancy will result in poor establishment. If the seed-bed is not too fine, adequate plant popula­ tions can be obtained even by surface-sowing. Fer­ tilization with P and K is generally necessary for establishment on low-fertility soils. Newly sown gamba grass pastures are quite susceptible to weed infestation because of slow initial growth. Gamba grass can be successfully associated with legumes of creeping, climbing, semi-efec t to erect growth habit such as Centrosema spp., Pueraria phaseoloides (Roxb.) Benth. or Stylosanthes spp. The grass requires a grazing management which prevents it from becoming mature and stemmy Andropogon gayanus Kunth - 1, plant part with (reduced quality), and, in mixtures with legumes, stem and leaf; 2,flowering stem. from becoming too vigorous and too tall (increased competition affecting legume persistence). Rota­ let isbisexua l and fertile. Caryopsis oblong, plano- tional grazing at appropriately high stocking rates convex, 3 mm x 0.75 mm, purple or hyaline (var. is recommended. Although its soil-fertility requir­ bisquamulatus). Germination of fresh seed is often ements are low, gamba grass responds to fertiliza­ reduced by dormancy, but under appropriate stor­ tion with N, P, and K. age conditions, improves within nine months after The major pest problem of gamba grass, which is harvest. Initial growth is slow but once estab­ restricted to tropical America, is leaf-cutting ants lished, the grass is very competitive and, despite of the genera Atta and Acromyrmex. Not only can its erect growth habit, it may consequently affect they completely destroy a pasture in the seedling persistence of associated legumes. It has a short- stage, but they can also severely affect the persis­ day flowering response with a critical photoperiod tence of an established pasture. of 12-14 hours. Gamba grass is usually harvested by grazing ani­ In A. gayanus four botanical varieties are distin­ mals but can be used to make hay. Depending on guished by morphological and ecological charac­ soil fertility, moisture and cutting regimes, it pro­ teristics. Of these, the most important one as a for­ duces DM yields ranging from 1-18 t/ha per year. age is var. bisquamulatus. Most A. gayanus Under favourable conditions, however, 12-week literature refers to this variety, with the possible DM yields ashig h as 5-15 t/ha are possible. Gamba exception of cultivar 'Kent' in northern Australia. grass is a prolific seeder with potential pure seed All commercial cultivars released in tropical yields as high as 350 kg/ha. Animal production America belong to var. bisquamulatus; they are from pure gamba grass pastures is low in savanna derived from accession CIAT 621, an introduction climates (90-120 kg liveweight gain per head per from Shika, Nigeria. year) because of liveweight losses during the dry Ecology Gamba grass is best adapted to the sub- season; by introducing a legume, annual weight 46 FORAGES gains of about 150k g per head can be achieved. has proven useful for hay production in Florida, Genetic resources and breeding There is con­ and is showing promise under coconuts in Indone­ siderable variation within the gamba grass culti- sia. vars. As the grass reproduces sexually by cross- Properties In the United States,i n vitro organic pollination, there is a continuous recombination matter digestibility (IVOMD)value s ranging from ofgenes .Additiona l variability within var. bisqua- 45-68%, and N concentrations from 1.6-2.9%, mulatus seems to be adequately represented in the have been measured in DM from stands cut twice major germplasm collections of tropical forage a year. Up to 74% IVOMD and 3.5% N have been grasses at CIAT (Colombia) and ATFGRC (CSIRO, measured from stands cut every two weeks.I n Aus­ Australia). Several exploratory breeding efforts tralia, IVOMD of 6-week-old regrowth of seven aimed at improving gamba grass cultivars are accessions varied from 70-77% with N varying under way in Colombia and Brazil. from 2.5-3.5%o. Phosphorous levels in the DM Prospects Because of its adaptation to acid, largely reflect soil P levels, and have been mea­ low-fertility soils, to burning and drought, and sured from 0.15%o in A. glabrata growing in extre­ also its high productivity, gamba grass is likely to melyinfertil e soilst o 0.52% in well-fertilized soils. continue playing an important role in pasture de­ Description Herbaceous perennial with erect to velopment in low-input systems of the subhumid decumbent unbranched, hollow above-ground tropics characterized by acid soils and savanna cli­ stems 5-35 cm long, 2-3 mm thick, arising from mate. In more humid environments, other grasses a mat of rhizomes which range in thickness from (such as Brachiaria spp.) seem to have a compara­ 3-5(-10) mm. Rhizomes which form a dense mat tive advantage. in the top 5 cm of soil arise from the deep, woody Literature |1| Bogdan, A.V., 1977. Tropical pas­ taproot; roots covered with a multitude of small, ture and fodder plants. Longman, London, pp.. oblate nodules. Leaf glabrous to sparsely pubes- 34-38. |2| Bowden, B.N., 1963, 1964. Studies on Andropogon gayanus Kunth. 1:Empir e Journal of Experimental Agriculture 31: 267-273 (1963); 2: Journal of the Linnean Society (Botany) 58: 509-519 (1964). |3| Skerman, P.J. & Riveros, F., 1990. Tropical grasses. FAO, Rome. pp. 185-190. |4| Toledo, J.M., Vera, R., Lascano, C. & Lenné, J.M. (Editors), 1990.Andropogo n gayanus Kunth: a grass for tropical acid soils. CIAT, Cali, Colom­ bia. 381 pp. R. Schultze-Kraft

Arachis glabrata Benth.

Linn. Soc. London, Trans. Bot. 18:159(1841) . LEGUMINOSAB In = 40 (tetraploid) Synonyms Arachisprostrata Benth. (1841). Vernacular names Rhizoma peanut (Am). Origin and geographic distribution A. gla­ brata and its near relatives are native to Brazil, Argentina and Paraguay in a large triangular area between 13°S and 28°S. Over the last 50years , var­ ious sets of the group have been sent to Australia and the United States, and more recently to India, Thailand, Malaysia and Indonesia. Uses Although A. glabrata is primarily used as a high quality forage legume for intensively grazed pastures on infertile, acid soils, it also has poten­ Arachis glabrata Benth. habit tial for soil conservation and as an ornamental. It with rhizomes. ARACHIS 47 cent, tetrafoliolate; leaflet ranging from linear- (probably the Australian accession CPI12121) has lanceolate to oblanceolate, obovate or cuneate up persisted well in Indonesia. to 4 cm x 2 cm; apex acute to mucronate, base Ecology A. glabrata is productive from about mostly obtuse; petiole grooved, up to 7.5 cm long, latitudes 30° N and °S to near the equator. It has 1-2 mm diameter with pulvinus 10-15 mm above persisted in areas receiving rainfall as low as 750 axil; stipules linear-lanceolate, falcate, up to 3 cm mm per year but is probably best suited to areas long, adnate to the petiole and membranous below receiving 1000-2000 mm. It grows successfully on the pulvinus; petiolule about 1 mm and rachis soils with textures ranging from sands to clays pro­ 10-15 mm long. Flowers sessile, axillary; hypan- vided they are well-drained. While apparently pre­ thium filiform, tubular, up to 10 cm long, pilose, ferring acid soils, it has produced good yields on containing the ovary at its base; standard more or neutral to slightly alkaline soils. It grows well on less orbicular, 15-25 mm wide, yellow, soft orange soils low in P. Unlike many tropical legumes, A. to brilliant orange without red veins on back. glabrata can compete successfully with sward- Fruit set geocarpic, but usually scarce; fruit ovoid forming grasses such as bahia grass (Paspalum ca. 10m m x 5-6 mm. Seeds ovoid, whitish. notatum Flueggé), narrowleaf carpet grass (Axo- Growth and development Because seed set is nopus affinis A. Chase), pangola grass (Digitaria rare, A. glabrata is usually propagated from rhi­ eriantha Steudel) and bermuda grass (Cynodon zomes. With adequate temperature and moisture, dactylon (L.) Pers.). shoots usually emerge 2-3 weeks after planting; Moderate to heavy grazing pressures are neces­ dense swards develop as rhizomes form and extend sary for best performance. It grows best when the margins of the stand at up to 2 m per year in mean monthly temperatures are above about 20°C. the absence of competition, or 5-30 cm per year Although tops are cut by frost, plants regrow with grass competition. Accessions differ in rate vigorously with the onset of warmth and moisture. of spread. Despite often dense flowering, few seeds During very dry conditions, top growth may die are set, except in some accessions prior to dense off, but rhizomes mostly survive, providing a nu­ sward development. Seedlings are usually quite cleus for recovery. large before they form rhizomes. Propagation and planting A. glabrata is best Other botanical information Although not propagated from rhizomes. While pieces of rhi­ validly published according to the International zome as short as 5 cm may strike, it is generally Code of Botanical Nomenclature, there is a system considered best to plant 30c m square pieces of rhi­ of classification in common use which divides the zome mat about 1.8 m apart. Alternatively, the rhi­ genus Arachis L. into a number of sub-generic sec­ zomes can be teased out, broadcast over the soil tions and series, based largely on the diversity of surface and disked in. In either case, it is best to growth habits within the genus. Section Rhizoma- plant about 3.5 m3 of rhizomes/ha at a depth of 3 tosae Krap. & Greg. (nom. nud.) which comprises cm in clay soils to 6.5 cm in coarse sands. It is gen­ therhizomatou s types is divided into series Prorhi- erally not necessary to inoculate rhizomes. A zomatosae Krap. &Greg . (nom. nud.) and Eurhizo- clean seed-bed is preferable, to minimize competi­ matosae Krap. & Greg. (nom. nud.). The former is tion for the developing plants. Ideally, rhizomes a group of delicate diploids, the latter is a group should be produced in sandy soils for ease of dig­ of more robust tetraploids of which A. glabrata is ging. Planting appears to be most successful when a member. In the absence of current description, rhizomes are dormant. it is not clear which of the 100 extant representa­ Husbandry Weeds should be controlled during tives of the Eurhizomatosae actually belong to A. establishment. This can be achieved through man­ glabrata. The foregoing description of A. glabrata ual weeding or through use of pre- and post-emer­ was drawn from a diverse set of Eurhizomatosae. gence herbicides such as trifluralin or vernolate, Four cultivars of A. glabrata have been released post-emergence applications of alachlor and dino- in Florida: 'Arb', 'Arblick', 'Florigraze' and seb, and routine applications ofbentazo n or 2,4-DB 'Arbrook'. 'Arb' and 'Arblick' were selected initial­ for broadleaf weed control, and sethoxydim and ly, based on productivity and spreading ability re­ fluazifopbutyl for grass control, as required. Mow­ spectively; 'Florigraze' was selected because it ing reduces shading from taller weeds and pro­ combined both characteristics, and 'Arbrook', the motes spread of A. glabrata. Adressin g ofP fertiliz­ most recent release, because it ismor e drought-tol­ er may be advisable if soils are extremely low in erant than 'Florigraze' but is of similar productivi­ P. However, liming is rarely necessary. ty. One accession known as the Maiwa peanut Diseases and pests A. glabrata is rarely trou- 48 FORAGES bled by insects or disease. It is immune to the com­ American Peanut Research & Education Associa­ mon groundnut leaf-spots caused by Cercospora tion, Inc.,Stillwater , Oklahoma, United States, pp. arachidicola and Cercosporidium personatum, and 47-133. |3| Prine, G.M., Dunavin, L.S., Glennon, most accessions are immune to groundnut R.J. & Rouch, R.D., 1986. 'Arbrook' rhizoma pea­ (Puccinia arachidis). Although susceptible to nut, a perennial forage legume. University of Flor­ other leaf-spots caused by Phyllosticta and Stem- ida, Gainsville. Agricultural Experiment Station phylium, and to white mould {Sclerotium rolfsü), Circular S-332. 16 pp. j4| Prine, G.M., Dunavin, these diseases have not caused serious long-term L.S., Moore, J.E. &Rousch , R.D., 1981. 'Florigraze' damage. Resistance to root-knot nematodes rhizoma peanut, a perennial forage legume. Uni­ (Meloidogyne spp.) has been recognized in a versity ofFlorida , Gainsville.Agricultura l Experi­ number of studies. ment Station Circular S-275. 22pp . Harvesting Low-growing accessions are best B.G. Cook used in grazed pastures. Taller-growing accessions also lend themselves to hay-making or cut-and- carry systems. Arachis pintoi Krap. & Greg., nom. nud. Yield Dry matter yields ofu p to 16t/h a have been measured in Florida and up to 12t/h a in Australia. In: Wilson, C.T. (Editor): Peanuts - culture and Yields of only 5t/h a have been obtained in Malay­ uses: 121(1973) . sia and Indonesia, but these may have been LEGUMINOSAE influenced by shading from rubber and coconuts. 2n = 20 Genetic resources Large collections are held Vernacular names Pinto peanut (En). Thai­ by various germplasm banks at Texas A & M Uni­ land: thua lisong tao. versity (Stephenville, United States) and North Origin and geographic distribution Pinto pea­ Carolina State University (Raleigh, United nut originates from the valleys of the Jequitin- States), at CENARGEN/EMBRAPA (Brasilia, honha, Sao Francisco and Tocantins rivers in cen­ Brazil) and at ICRISAT (Patancheru, India). A tral Brazil. Since its collection in 1954, it has been more limited set is held at ATFGRC (CSIRO, Aus­ distributed to Argentina, Australia, Colombia and tralia) and QDPI (Gympie, Queensland, Austra­ the United States, and more recently to many lia). countries in South-East Asia, Central America Breeding 'Florigraze' is the result of a chance and the Pacific. intra-specific cross in Florida. While A. glabrata Uses Because of its high degree of shade toler­ is compatible with species from sections Arachis ance, pinto peanut is finding application as a pas­ and Erectoides Krap. & Greg. (nom. nud.), the ture legume in tree plantations, but also as a resultant hybrids are infertile. An intersectional ground cover in plantations from which grazing rhizomatous hybrid, more productive than either animals must be excluded. It is showing potential parent, has been developed in Tamil Nadu, India. in intensively managed grass/legume pastures on Prospects A. glabrata is a persistent, very pro­ cleared land. ductive, high-quality forage, well-adapted to the Properties In vitro digestibility of pinto peanut often infertile, acid soils and heavy grazing pres­ varies from 60-76%, N concentrations from sures encountered in the humid and sub-humid 2.5-3.0% and P concentrations from 0.18-0.37%. tropics and subtropics. The main constraint to It is well accepted by cattle at all stages of growth. evaluation and adoption is the need to propagate There are 6-8 seeds in pod/g. from rhizomes. With the development of mechani­ Description A stoloniferous, perennial herb de­ cal methods or with the availability of manual veloping a strong taproot on the older crowns and labour for propagation, A. glabrata may prove to large numbers of small oblate nodules, mostly to be one of the more useful forage legumes in South- 2 mm diameter, on both taproot and subsidiary East Asia. roots. Stems initially prostrate, becoming ascen­ Literature |1[ Gregory, M.P. & Gregory, W.C., dant to 20c mheigh t in dense swards. Leaves tetra- 1979. Exotic germ plasm ofArachi s L. interspecific foliolate, margins entire, ciliate; distal leaflets hybrids. The Journal of Heredity 70: 185-193. j2| obovate and proximal leaflets oblong-obovate, Gregory, W.C., Gregory, M.P., Krapovickas, A., obtuse at the apex and slightly cordate at the base; Smith, B.W. & Yarbrough, J.A., 1973. Structure leaflets may attain 4.5 cm x 3.5cm ,bu t are smaller and genetic resources of peanuts. In: Wilson, C.T. in regularly defoliated stands; the upper surface (Editor):Peanut s - culture and uses.A symposium. of leaflets is glabrous and a darker green than the ARACHIS 49

ground for more than one season. Other botanical information The genus Ara­ chis L. has been divided into a number of invalidly published sections based on growth form and gross morphology. A. pintoi is one of two species in the section Caulorhizae Krap. &Greg. , (nom. nud.), a section defined by the possession of stolons. The original line collected by G.C.P. Pinto near the mouth of the Jequitinhonha River in 1954 and var­ iously catalogued as GK 12787 (Collection Number), PI 338314 (United States), CPI 58113 (Australia) and CIAT 17434 (Colombia), has been released as cultivar 'Amarillo' in Australia. More recent collections of the species differ slightly from the original in leaf shape and size, and inter- node length. Ecology The climate in central Brazil is humid tropical, with some 1800-2000 mm of rain falling in the October-May wet season, and an additional 200 mm in the June-September dry season. Pinto peanut has been mostly collected in red, sandy- loam alluviums, apparently growing best in low areas which are wet to flooded during the wet sea­ son. Soils are generally of low fertility with high Al content. Native vegetation is low forest with a fairly dense canopy. In cultivation, pinto peanut has proven adaptable to soils ranging from sand to clay texture, with low to neutral pH and low to high fertility. It failed to persist on seasonally Arachis pintoi Krap. & Greg., nom. nud. - 1, habit waterlogged, poorly structured clays.I n pot exper­ flowering plant; 2, fruits. iments, it has shown a tolerance of soil Mn at levels detrimental to many other legumes, a fair pubescent lower surface. Flowers emerge indivi­ tolerance ofsoi lAl , and a low tolerance of salinity. dually from short axillary racemes and are similar It is capable of growing under shaded conditions, in shape to commercial groundnut (A. hypogaea often appearing more vigorous in shade than in L.), but smaller (standard 12-17 mm wide) and yel­ full sunlight. low. The terminal pod on the peg usually contains Propagation and planting Fresh seed of pinto a single seed but may contain 2,whil e pods formed peanut has a high level of dormancy which may along the peg contain only one. Pod moderately be reduced by drying the seed at 35-40°C for 10 reticulated, 10-14 mm x 6-8 mm, containing days. Seed should be inoculated with a specific mostly a single seed. Seed light-brown, 8-11 mm strain of Bradyrhizobium, which is different from x 4-6 mm, weighing 0.11-0.20 g. that used on commercial groundnuts. A well-pre­ Growth and development Seedlings of pinto pared seed-bed is desirable but not essential. Seed peanut develop quickly following epigeal germina­ should be sown 2-6 cm deep at 10-15 kg seed in tion, and with good growing conditions and sever­ pod/ha, followed by rolling. If seed isno t available, al plants per square metre, complete ground cover pinto peanut is readily propagated from cuttings. can be achieved by a network of stolons in less Husbandry Pinto peanut is tolerant of and than six months. Flowering commences three to increases under heavy grazing, but, under light four weeks after emergence and continues through grazing, is not shaded out by taller grasses. It does the growing season, appearing to intensify follow­ not require highly fertile soil, but on an oxisol in ing rain or irrigation. The ovary is borne on a Colombia it responded to additions of P, K, Ca and gynophore or peg, which elongates to up to 27 cm Mg. after pollination and pushes the ovary up to 7 cm Diseases and pests The only information avail­ depth into the soil. Seed remains viable in the able relates to 'Amarillo', which is resistant to the 50 FORAGES major groundnut diseases, rust (Puccinia arachi- 47-133. |3|Grof , B., 1985. Forage attributes of the dis) and leaf-spot (Mycosphaerella spp.). Other perennial groundnut Arachis pintoi in a tropical fungi (Phomopsis sp., Cylindrocladium sp.an d Col­ savanna environment in Columbia. In: Proceed­ letotrichum gloeosporioides) have been isolated ings ofth e XVInternationa l Grasslands Congress, from leaf-spots, the latter also being associated August 24-31, Kyoto, . Science Council of with black stem lesions in Colombia. None of these Japan and Japanese Society ofGrasslan d Science, diseases causes long-term or serious damage. Nishinasuno, Japan, pp. 168-170. |4iLascano , CE. 'Amarillo' has moderate to high resistance to the & Thomas, D., 1988. Forage quality and animal various root-knot nematodes (Meloidogyne spp.) selection of Arachis pintoi in association with but issusceptibl e to the root-lesion nematode (Pra- tropical grasses in the eastern plains of Colombia. tylenchus brachyurus). Leaves ofsom e plants have Grass and Forage Science 43: 433-439. |5j Oram, an apparently non-pathogenic variegation. Rats R.N., 1990.B .Legume s 21. Arachis (a) Arachis pin­ and mice are attracted to the nuts and can be a toi Krap et Greg, nom-nud. (Pinto peanut) cv. problem in stands of pinto peanut. Amarillo. Australian Journal of Experimental Harvesting Being a low-growing, ground cover Agriculture 30: 445-446. j6| Vails, J.F.M., 1983. species, pinto peanut is more readily grazed than Collection of Arachis germplasm in Brazil. Plant hand-harvested. Seed should be dried in the sun or Genetic Resources Newsletter 53: 9-14. in an artificial drier and stored under cool condi­ B.G. Cook tions with low humidity. Yield In Colombia, pinto peanut had an annual DM production ranging from 5 t/ha growing with Arundinaria pusilla A. Chevalier & A. Brachiaria dictyoneura (Fig. & De Not.) Stapf, Camus which produced 20 t/ha, to 10 t/ha when grown with B. ruziziensis Germain & Evrard, which pro­ Bull. Mus. Hist. Nat. Paris 27:45 0 (1921). duced 11 t/ha. It has yielded 5 t/ha of DM in pure GRAMINEAE stands under 30% shade in Indonesia and 3 t/ha 2n = unknown in full sunlight in Malaysia. Yields of seed in pod Vernacular names Bamboo grass (En). Philip­ from 1-2 t/ha have been recorded. pines: utod (Igorot). Cambodia: rüssèi préïch. Genetic resources A number of accessions Laos: ph'ek. Thailand: ya-phek, ya-phet. Vietnam: have been collected, most of which are held by le, truc th'ao. CENARGEN/EMBRAPA in Brazil. A more Origin and geographic distribution A. pusilla limited range is held at Texas A & M University occurs in South-East Asia, particularly between (Stephenville, United States), CIAT (Colombia), latitudes 13-17°N in Thailand and Indo-China. and ATFGRC (CSIRO, Australia). Uses A. pusilla is primarily used for traditional Breeding Breeding mechanisms are well under­ grazing of village cattle and buffaloes, particular­ stood, since Arachis wild types have largely been ly on new growth at the onset of the rainy season. collected as a source ofgeneti c material in ground­ Properties Leaves are palatable but stems rap­ nut improvement programmes, but there are no idly become woody and unacceptable to livestock. breeding programmes. Leaves collected at the beginning of the rainy sea­ Prospects Pinto peanut has only been assessed son contain up to 1.7 % N with 52% organic matter as a forage over the last 10 years. At this stage it digestibility. The average dry matter intake by appears to have great potential in the wet tropics, sheep is 620 g/head per day. When mixed with Sty- especially in more shaded situations under planta­ losanthes hamata (L.) Taub, cultivar 'Verano' and tion crops. Arundinaria ciliata A. Camus, it can produce en­ Literature |1| Cook, B.G. &Franklin , T.G., 1988. ough feed to support cattle at 1.2-1.8 animals per Crop management and seed harvesting of Arachis ha throughout the dry season of north-eastern pintoi Krap. et Greg. nom. nud. Journal of Applied Thailand. Seed Production 6: 26-30. |2| Gregory, W.C., Gre­ Botany Ashallo w rooting, erect, perennial bam­ gory, M.P., Krapovickas, A., Smith, B.W. & Yar- boo with short, strong woody rhizomes; leafy stems brough, J.A., 1973. Structure and genetic 0.5(-1.5)m tal l and upt o 8m mdiameter , branching resources of peanuts. In: Wilson, C.T. (Editor): at the nodes. Leaf-blade 8-14 cm x 1 cm, con­ Peanuts - culture and uses. A symposium. Ameri­ tracted into a short stalk at the junction with the can Peanut Research and Education Association sheath, pubescent, acuminate at the top. Flower­ Inc., Stillwater, Oklahoma, United States, pp. ing stems 15-75 cm long, often fascicled on the rhi- ARUNDINARIA 51

Genetic resources and breeding It is unlikely that substantial germplasm collections are being maintained. Prospects A. pusilla is usually not cultivated and its use is limited to grazing. Its value increases when grown with suitable legumes. Literature il! Camus, E.G. & A., 1923. Grami­ nées. In: Lecomte, H. & Gagnepain, F. (Editors): Flore générale de L'Indo-Chine. Tome 7(5). pp. 585-587. ,2,Gutteridge , R.C., Shelton, H.M., Wilai- pon, B. &Humphreys , L.R., 1979.A comparison of cattle liveweight gains on native and improved pastures in northeast Thailand. Annual Report Faculty of Agriculture. Khonkaen University, Thailand. 13; Manidool, C, 1989.Natura l grassland and native grasses ofThailan d [inThai] . Technical Bulletin No 1301-26-32. Division of Animal Nu­ trition, Department of Livestock Development, Bangkok. 45 pp. |4 Manidool, C. &To r Sakool, P., 1971. Digestibility of pek grass (Arundinaria pusilla). Thai Journal of Agricultural Science 4: 119-124. j5j Robertson, A.D. & Humphreys, L.R., 1979. Effects of frequency of heavy grazing and P supply on Arundinaria ciliata association over­ sown with Stylosanthes humilis. Thai Journal of Agricultural Science 9:131-138. C. Manidool

Arundinariapusilla A. Chevalier & A. Camus - 1, habit of plant with ; 2, leafy stem; 3, part Asystasia gangetica (L.) T. Anderson offlowering stem. Thwaites, Enum. pi. zeyl. 3:23 5(1860) . zome clumps of burned plants, usually without ACANTHACEAE leaves. Inflorescence narrow, consisting of 2-8 In = unknown groups of 1-5 spikelets in the axil of a glume-like Synonyms Justicia gangetica L. (1759), Asysta­ bract or sometimes on a short stalk. Spikelets 3-4 sia coromandeliana Nees (1832). cm long with 7-9 florets, the lowest 3-5 bisexual, Vernacular names Malaysia: rumput bunga the upper ones male, the terminal one sterile; putih, rumput hantu, rumput nyonya. Philippines: styles 3, s 6.Littl e or no seed is produced. asistasiya (Tagalog), bulak-bulak (Subanun). In Thailand, it flowers in February and March. Thailand: baya, yaya. Ecology A. pusilla is adapted to warm environ­ Origin and geographic distribution A. gange­ ments with a short wet and a long dry season at alti­ tica is indigenous in and Asia, but tudes from sea-level to 1500m .I t tolerates moderate has been distributed pantropically. In South-East shade but isintoleran t ofwaterlogging . It is usually Asia it is recorded from Malaysia, Thailand, Indo­ found in Dipterocarpus forests, growing as a lower nesia and the Philippines. canopy, on moderately acid upland soils. Uses A.gangetica provides excellent feed for cat­ Agronomy A. pusilla spreads naturally by short tle, goats and sheep. In cocoa, oil palm, rubber, rhizomes, but can be propagated by dividing vegetables and field crops it is not regarded as use­ clumps of roots and stubble, or by stem cuttings. ful, though its ground cover checks erosion and Light grazing is recommended as it is intolerant prevents the infestation by other weeds such as of heavy grazing. It is not used in cut-and-carry Mimosa pigra L. and Imperata cylindrica (L.) systems and is not suitable for making hay or Raeuschel which are considered to be more nox­ silage. Edible DM yields range between 0.8 and 2.5 ious. It is liked as a naturally occurring plantation t/ha per year. cover in some orchards because the bees which pol- 52 FORAGES linate the flowers of fruit trees such as starfruit crops as illumination declined from 54-7 % full or durian, are attracted to the orchard by the sunlight. flowers of A. gangetica. Botany An erect, ascending or clambering herb, A. gangetica is also used as a village medicine. Chi­ 0.3-1.25 m high, with quadrangular pilose stem. nese medicine shops sell it as 'kaw kua chai'. In Leaf ovate to deltoid, 3-7.5 cm x 1.5-5 cm, base the Moluccas, its juice, together with and obtuse to truncate, apex acuminate, 'glabrate to onion juice, is recommended for dry coughs with sparingly pilose, especially on the veins, numerous an irritated throat and discomfort in the chest. bar-like cystoliths visible on the upper surface; Young shoots are used as a food ('ulam') by farmers petiole 1-3 cm long. Inflorescences in terminal in south-western Johore. racemes, up to 16 cm long, with flowers directed Properties A. gangetica has high concentra­ to one side only; flowers on short pedicels, white tions of N, P, K, Mg and Cu in its leaves and excep­ to yellow, white with purplish throat, or violet; tionally high concentrations of K in its stems and calyx 5-lobed, lobes lanceolate 5-7 mm long; corol­ roots. The nutrient concentrations in leaves and la funnelform, up to 4 cm long, with 5 semiorbicu- young stems, expressed on a dry matter basis, typi­ lar lobes ca. 1 cm wide; 4; stigma 2-lobed. cally range from 0.30-0.42% P, 4.56-5.83% K, Fruit an oblongoid capsule, up to 13 mm x 2 mm. 0.98-1.08% Mg. Large accumulations of Cu Seeds 2-4, ovoid, flat, ca. 1m m x 0.75 mm. (12-19 mg/kg) occur. Potassium concentrations of Seeds germinate readily, with a germination per­ 3.6%) and 1.6% have been measured in stems and centage of up to 90%). The period from seedling roots respectively. In vitro DM digestibilities of 3, emergence to seed dispersal can be as early as 8 6, 9, 12 and 16 weeks old top growth were 71%, weeks in open areas but, in partially shaded areas, 71%),69%o ,61% o and 61%) respectively. Leaf diges­ this can take 2week s longer. tibility increased from 69-74% under plantation It takes one month from floral initiation to seed dispersal. The seeds are thrown as far as 6 m by an explosive mechanism on hot afternoons. Ecology A. gangetica is found along roadsides and riverbanks, in semi-waterlogged areas as well as well drained cultivated areas. In 1976 it became very widespread in oil palm plantations in Penin­ sular Malaysia and by 1983 it had infested large tracts of land on peat soils in the south­ eastern region ofJohore . It is a shade-loving plant and optimum photosynthesis occurs between £ to \ full sunlight. With no weeding, its proportion in the undergrowth of young oil palm plantation increased in a period of 2year s from 25% to 84% . It grows, even if slowly, under a closed canopy of oil palm with less than 10% full sunlight. Howev­ er, in areas with a dry season of 4month s or more, it may not survive. It thrives on sedentary soils, coastal alluvium, peat soils with 85% organic mat­ ter and pH(H20) 3.5-4.5, sandy loams and clays. Agronomy A. gangetica can bepropagate d from seeds or by stem cuttings with 1, 2 or 3 nodes. Sin­ gle node cuttings buried in soil produce flowers and fruits within 6weeks . It isdifficul t to establish in unshaded open areas. The high palatability and digestibility of A. gange­ tica make it attractive to grazing animals. Its agressiveness, high uptake of soil nutrients and ability to smother other species have character­ Asystasia gangetica (L.) T. Anderson -1, flowering ized it as a weed in plantation management. How­ and fruiting branch; 2, flower, cut lengthwise; 3, ever, when grazing animals are introduced, it is a ripe fruit. useful plant. Villagers hand harvest its forage to AXONOPUS 53

supplement the forage which they feed to their ani­ Axonopus compressus (Swartz) mals at night. Good cattle production under oil P. Beauv. palm from native forages mixed with A. gangetica can be achieved within the range of 110-135 kg/ha Ess. Agrost.: 12,154,167 (1812). per year, equivalent to 270-310 g/head per day. GRAMINEAE Care is needed when grazing with sheep, since if 2n = 40 they graze for extended periods on swards contain­ Synonyms compressum Swartz (1788), ing more than 60% A. gangetica they can accumu­ Paspalum platicaulon Poiret (1804), Anastrophus late toxic levels of over 1000 mg/kg Cu in their compressus Schlechtendal ex Doell. (1877). liver. Excessive consumption by sheep can also Vernacular names Broadleaf carpet grass result in bloat. (En). Indonesia: rumput pahit (Indonesian), papa- No major diseases or pests have been observed on hitan, jukut pahit (Sundanese). Malaysia: rumput A. gangetica. parit. Philippines: kulape (Tagalog). Thailand: ya A. gangetica is harvested by grazing animals or by baimaln, ya-malaysia. cutting for stall feeding. Frequent cutting induces Origin and geographic distribution It is early dieback because the stems have long inter- native to the , and from nodes and growing points high up in the canopy. to Brazil, and has been introduced into Low grazing pressures or long intervals between many warm countries, including South-East Asia. grazing allow the plant to flower and set seed. It Uses Broadleaf carpet grass is used for grazing, is usually consumed fresh by animals but it can be especially in plantation crops, particularly coco­ conserved asha y ifproperl y dried. Pure swards are nut. It is also a good grass and can be used rarely found in full sunlight. Under heavy shade for controlling soil erosion. It can be a troublesome (6-16% full sunlight) DM yields under cutting weed. Under dense shade in established oil palm from 2-5 t/ha have been recorded, but under a and rubber plantations, it is often allowed to grow more open canopy of Leucaena leucocephala as a soil cover. (Lamk) deWi t at 2m x 1m spacing , yields between Properties Nitrogen concentrations of broad­ 3.5-8 t/ha were obtained. leaf carpet grass range between 1-2%. It is Genetic resources and breeding There are no reported to have an ability to fix atmospheric N known germplasm collections of A. gangetica and through associated micro-organisms. no breeding programmes. Botany Aperennial , stoloniferous, short spread­ Prospects Due to its ability to grow under shade ing grass; stolons often long, branched, rooting at and its high nutritive value, A. gangetica is an the nodes; culms geniculately ascendent, 20-50 cm important forage species in South-East Asia, tall, solid, laterally compressed. Leaf-sheath either when grazed or cut for stall feeding. It could strongly compressed, finely hairy along the outer have potential as a substitute for legumes in the margin, the nodes densely pubescent; ligule very production of leaf meal. short, fringed with short hairs; leaf-blade lanceo­ Literature [1| Backer, C.A. & Bakhuizen van late, flat, 2.5-38 cm x 2.5-16 mm, base broadly den Brink, R.C., 1965.Flor a of Java. Vol. 2. Noord- rounded, margin ciliate, apex obtuse. Inflores­ hoff, Groningen, the Netherlands, p. 576. |2| Kang, cence compound, l-2(-8) peduncles exserting L.C., 1981. Asystasia. Nature Malaysiana 6(2): from the final sheath, each composed apically of 14-17. ;3| Lee, S.A., 1984. Morphology, reproduc­ 2 conjugate, one-sided spikes 3-11 cm long, lower tive capacity and seed germination of Asystasia down often with a third, rarely a fourth; spikelets intrusa. Proceedings seminar on control of Asysta­ oblong, rather acute, 2-3.5 mm x 1-1.25 mm, pale sia. Pejabat Pengarah IADP Johor Barat, Johor, green or purplish tinged, 1-flowered, solitary on Malaysia, pp. 215-238. !4jRajaratnam , J.S., Chan, alternate sides of rachis and forming two rows, K.W. &Ong , H.T., 1976.Asystasi a in oil palm plan­ ciliate on the margins; lower glume absent, upper tations. In: Earp, D.A. & Newell, W. (Editors): glume and lemmas equalling the spikelet in length. International developments in oil palm. Proceed­ Caryopsis elliptical, 1.25 mm long, dorsally com­ ings of the Malaysian International Agricultural pressed, yellow-brown. Oil Palm Conference, Kuala Lumpur, 14-17 June Broadleaf carpet grass flowers all year round, al­ 1976. Incorporated Society of Planters, Kuala though little seed is produced in some environ­ Lumpur, pp. 454-470. ments. Young plants start growth in a circular S.A. Lee &C.P . Chen patch. If it has no serious competition, the patch may reach a size of up to 1 m in diameter in one 54 FORAGES

sia, yields up to 1 t/ha have been measured. In Bra­ zil, Zebu steers grazing over 672 days achieved an average daily gain of 0.18 kg. A. compressus is an alternate host of Rhizoctonia solani. In Malaysia the grass as weed is controlled by spraying with 1.1 kg MSMA + 0.6 kg sodium chlorate in 273 1 water. Genetic resources and breeding It is unlikely that any substantial germplasm collections are being maintained. Prospects Broadleaf carpet grass is a useful for­ age source for livestock raised under plantation crops, particularly under heavy shade, and it is adapted to extremes in soil types. Studies are war­ ranted to see if its productivity can be improved. Literature |1|Boonklinkjorn , P., 1979. Herbage production of grasses under coconuts in Southern Thailand. Thai Journal ofAgricultura l Science 12: 43-50. |2| Bor, N.L. 1960. The grasses of Burma, Ceylon, India and Pakistan. Pergamon Press, Oxford, p.278 . ;3| Chen, C.P. &Bon g Julita, I.,1983 . Performance of tropical forages under closed canopy of oil palm. I. Grasses. MARDI Research Bulletin 11: 248-263. |4; Holm, J., 1971. Feeding tables. Nutrition Laboratory of Thai-German Dairy Project, Livestock Breeding Station, Chiangmai, Thailand, p. 19. |5j Holm, L.G., Pluck- Axonopus compressus (Swartz) P. Beauv. - habit. nett, D.L., Pancho, J.V. & Herberger, J.P., 1977. The world's worst weeds:distributio n and biology. season. It often crowds out all other weeds and The East-West Center, University ofHawai i Press, grasses and forms a dense mat-like cover. Honolulu, Hawaii, United States, pp. 180-184. j6! Ecology Broadleaf carpet grass is best adapted Mehra, K.L. & Fachrurozi, Z., 1985. Indonesian to moist warm environments. It is moderately economic plant resources: forage crops. Lembaga shade tolerant but grows well in full sunlight. It Biologi Nasional-LIPI, Bogor, No 31.p .2 . |7j Sker- occurs in Indonesia up to 2300 m altitude. It is fre­ man, P.J. & Riveros F., 1990. Tropical grasses. quently noted as being one of the most persistent FAO, Rome. pp. 211-214. |8| Soerjani, M., Koster- and productive native grasses in plantations and mans, A.J.G.H., Tjitrosoepomo, G. (Editors), 1987. will persist under heavy shade where introduced Weeds of rice in Indonesia. Balai Pustaka, Ja­ grasses may not survive. It will not stand water­ karta, pp. 388-389. logged conditions although it grows on a range of C. Manidool soil types, particularly sandy soils. It is dormant during cool periods. Agronomy Broadleaf carpet grass is usually Bothriochloa pertusa (L.) A. Camus vegetatively propagated by planting stolons. When seeded, a seeding rate of 6 kg/ha is recom­ Ann. Soc. Linn. Lyon 76:164(1931) . mended. Frequent grazing is preferred, to main­ GRAMINEAE tain its palatability and quality. It is encouraged In = 40 (tetraploid), 50 (pentaploid), 60 (hexa- by close grazing, and will oust other grasses. ploid) Broadleaf carpet grass is usually grazed by teth­ Synonyms pertusus L. (1771), Andropo- ered or freely grazing animals; it is rarely used in gon pertusus (L.) Willd. (1806), Amphilophis per­ a cut-and-carry system.Whe n used as a lawn grass, tusa (L.) Nash ex Stapf (1916). it should be frequently mown. With mixed fertiliz­ Vernacular names Indian couch grass, Indian er application at 300 kg/ha it has yielded up to 5 bluegrass, sweet pitted grass (En). Indonesia: t/ha of DM. Under oil palm plantations in Malay­ suket putihan (Javanese), rebha las-alasan BOTHRIOCHLOA 55

(Madura). Malaysia: rumput embun. Philippines: salay-parang (Tagalog), salay (Pangasinan). Thai­ land: ya-hom (central), ya-hangma (northeast). Vietnam: huyêt tha'o lô. Origin and geographic distribution Indian couch grass is widely distributed in Asia from Ara­ bia to South-East Asia, and is particularly promi­ nent on the Indian subcontinent. It has been intro­ duced to other tropical areas and is now naturalized in tropical America, Australia and on several island groups (e.g. Hawaii). Uses Indian couch grass is used mainly as a for­ age for ruminants, but also for soil conservation, revegetating mine sites, lining floodways, lawns, recreation areas and playing fields. Properties Chemical composition and nutritive value depend upon age and soil fertility. Young green leaves may contain more than 2% N and 0.2% P, and be 70% digestible. These values de­ crease with age and at the end of the wet season will be considerably lower. Indian couch grass de­ velops a high stem to leaf ratio with age and this reduces both the quality and palatability of her­ bage. Standing forage in the dry season may con­ tain only 0.5% N, 0.05% P and be 45% digestible. Indian couch grass is moderately palatable as cat­ tle sometimes avoid eating it whilst other grasses are available. It has a strong odour when crushed. There are 650-750 seeds/g. Botany A low growing, stoloniferous perennial of variable size and growth habit. Stolons root Bothriochloa pertusa (L.) A. Camus - 1, flowering readily at the nodes and produce small tufts of plant; 2, ligule; 3, inflorescence; 4, sessile spikelet; leaves. Culms are smooth, erect or geniculately as­ 5,pedicelled spikelet. cending, up to 60 cm tall and 3 mm in diameter; nodes are prominent and may be glabrous or cover­ growth ceases. Seed matures in 3-4 weeks. The ed with hairs up to 3 mm long. Leaf linear-acumi­ onset of flowering can be delayed by poor growing nate, 5-10(-30) cm x 4-6 mm, bright to dark conditions and flowering is checked by frost. green, usually glabrous except for a few hairs up B. pertusa is morphologically indistinguishable to 3m m long especially in the throat; ligule a shal­ from the mainly African B. insculpta (A. Rich.) A. low membrane surrounded by long ciliate hairs. Camus, but they do not interbreed. B. pertusa is Inflorescence 3-6 cm long, subdigitate, bearing less robust, more stoloniferous, and bears fewer 3-13 racemes each 2-7 cm long; spikelets in alter­ and shorter racemes. It is also closely related to nate pairs, one sessile, one pedicelled, on a slender B. decipiens (Hack.) C.E. Hubbard, B. longifolia rachis; sessile spikelet with a short blunt ciliate- (Hack.) Bor, B. panormitana (Pari.) Pilger and B. hairy callus, a lower glume with a circular depres­ radicans (Lehm.) A. Camus. sion (pit) in upper half and upper lemma with a Seed of naturalized strains (particularly Bowen kneed and twisted scaberulous awn 1.5 cm long; strain) is available in northern Australia. Culti- pedicelled spikelet usually neuter, pitless. Seed vars 'Medway' and 'Dawson' were released in awned, with weak antrorse hairs. Queensland in 1991.The y are later flowering and Seeds germinate early in the wet season and vege­ leafier than the Bowen strain. 'Dawson' is particu­ tative growth continues until soil water is larly useful as a lawn grass. exhausted. Time of flowering varies widely be­ Ecology Indian couch grass grows in tropical tween strains but may commence 3-4 weeks after areas with warm season dominated rainfall of the start of the wet season and continue until 500-1400 mm per year. It survives in areas which 56 FORAGES

are subject to frost, although its vigour is reduced, onizing grass which provides good ground cover, as frost kills the plant tops leaving only the basal reasonable quality herbage, is tolerant of heavy parts alive. Indian couch grass hays off rapidly grazing and suited to the drier regions of South- when soil moisture becomes depleted but can with­ East Asia. It has no rhizomes and is easily killed stand moderate drought. Prolonged drought or fire by cultivation so is unlikely to become a weed in will substantially reduce the basal area of the cropping areas. stand, although regrowth is rapid when growing Literature Ü! Bisset, W.J., 1980. Indian blue- conditions become favourable. Indian couch grass grass has special uses. Queensland Agricultural is found on a wide range of soil types. It is most Journal 106:507-517 . |2| de Wet, J.M.J. &Higgins , competitive on infertile to moderately fertile soils. M.L., 1963. Species relationships within the Bo- Prolonged waterlogging will kill or reduce its thriochloa pertusa complex. Fyton 20:205-211 . |3i basal area. Mclvor, J.G., 1984. Phosphorus requirements and Agronomy Indian couch grass can be estab­ responses of tropical pasture species: native and lished vegetatively from runners, but is normally introduced grasses and introduced legumes. Aus­ established from seed. Freshly harvested seed is tralian Journal of Experimental Agriculture and dormant, but dormancy breaks down after 4-8 Animal Husbandry 24: 571-578. ,'4| Skerman, P.J. months storage. It can be established under a vari­ & Riveros, F., 1990. Tropical grasses. FAO, Rome, ety of conditions, but best results are obtained by pp. 230-233. |5! Truong, P.N. & McDowell, M., sowing on or near the soil surface of a well culti­ 1985. Indian bluegrass for soil conservation and vated seed-bed during the early wet season. Some land stabilization in Queensland. Journal of Soil seed-harvesting ants remove the seed, so early sow­ Conservation Service of New South Wales 41: ing should be avoided where these are present. 38-44. Seeding rate is 1-3 kg/ha. J.G. Mclvor &S.M . Howden Indian couch grass can be used by continuous or rotational grazing, or cutting. It will stand close defoliation and is most prominent in heavily Brachiaria brizantha (A. Rich.) Stapf grazed or closely mown areas. It is used in associa­ tion with legumes, although some problems have Flora of Trop. Africa 9:53 1(1919) . been experienced as it may exclude Stylosanthes GRAMINEAE hamata (L.) Taub, and other low-growing legumes. In = 36,5 4 Indian couch grass isusuall y grown without fertil­ Synonyms Panicum brizanthum Höchst, ex A. izers. However, yields can be increased substan­ Rich. (1851). tially by fertilization as it responds well to both Vernacular names Palisade grass,palisad e sig­ N and P. There are no important diseases or pests nal grass (En). Thailand: ya siknaentontang. although it is attacked by rust. Indian couch grass Origin and geographic distribution The ori­ is harvested by grazing animals but can be made gin of B. brizantha is Africa where the species has into hay. It produces 1-5 t/ha ofdr y matter depend­ a very wide distribution throughout the wetter ing on seasonal conditions, soil fertility and asso­ tropics. In recent decades, experimental lines and ciated species. cultivars have been introduced to other parts of Genetic resources and breeding There is a the world, including tropical Asia and the Pacific wide variation in Indian couch grass in morpholo­ region. gical, flowering and agronomic characteristics. Uses The main use of palisade grass is as forage Collections, mainly originating from India, are in permanent pastures for grazing or for cut-and- held by ATFGEC (CSIEO, Australia) and the carry systems. It has also proved useful as grazed Queensland Department of Primary Industries. ground cover in tree plantations. There are no breeding programmes and any im­ Properties Palisade grass provides a more pal­ provement will depend on selection from naturally atable forage than signal grass (B. decumbens occurring populations. Important objectives for Stapf) but of similar N concentration (2.5%) and any selection process will be to increase herbage in vitro DM digestibility (75%) in 2-week-old yield, quality and palatability whilst maintaining regrowth, falling to 1.0% N and 55% digestibility the adaptability to a range of environments, abili­ in 12-week-old regrowth. Like signal grass, B. bri­ ty to provide effective soil cover, persistence under zantha can cause photosensitization in sheep and heavy grazing and colonizing ability. goats, but in Brazil, where the grass is widely used Prospects Indian couch grass is an effective col­ in pastures, no photosensitization effects on cattle r BRACHIARIA 57

have been observed. There are 130-180 seeds/g. whereas 'La Libertad' is a recent Colombian Botany Atufted , prostrate or semi-erect to erect release. perennial with short rhizomes and stems 30-200 Ecology Like signal grass, palisade grass is well cm tall. Leaf linear to broadly linear, 10-100 cm adapted to the humid and sub-humid tropics where x 3-20 mm, glabrous or hairy. Inflorescence con­ it can withstand dry seasons of up to 5 months. It sisting of 2-16 relatively long racemes (4-20 cm), grows well on a range of soils including sandy and with spikelets generally in one row; rachis 1 mm acid soils, but it requires more fertile soil than sig­ wide, usually purple; spikelets elliptic, 4-6 mm nal grass. It does not tolerate poorly drained soils. long, glabrous or with a few hairs at the tip; lower Like signal grass, it tolerates light to moderate glume £th e spikelet length, clasping; upper glume shade. cartilaginous, dully shining; upper lemma granu­ Agronomy Although palisade grass can be es­ löse. tablished by spreading out stolon cuttings, it is B. brizantha intergrades with B. decumbens and mainly sown at rates of 1.5-12 kg/ha, depending the species are difficult to distinguish. It is, there­ on seed quality. Seed is broadcast or drilled 2-4 fore, likely that in the literature the name B. bri­ cmdee p into conventionally prepared seed-bed. As zantha sometimes actually refers to B. decumbens, germination offres h palisade grass seed is affected and vice versa. The main difference between the by dormancy (hard-seededness), seed should be cultivars or lines ofB. brizantha and B. decumbens scarified with concentrated sulphuric acid, or seed is in the growth habit: B. brizantha is tufted with that has been stored for 6-8 months should be a rather erect growth whereas B. decumbens is used. low-growing and forms a dense cover. Two culti­ Palisade grass grows quickly, and 3-5 months vars have been released in South America: the best after sowing, it can be ready for a first, light graz­ known and most widespread is 'Marandu' (Brazil) ing. It responds well to fertilization with N, but also with P and K. Although experience regarding persistence of legumes in association with palisade grass is limited, the less competitive growth habit of this grass, as compared with signal grass, suggests that it can be grown with a range of legumes represent­ ing distinct growth habits, such as Desmodium heterocarpon (L.) DC. ssp. ovalifolium (Prain) Ohashi (prostrate-stoloniferous), Centrosema pubescens Benth. or Puerariaphaseoloides (Roxb.) Benth. (trailing-climbing), Stylosanthes guianen- sis (Aublet) Swartz (semi-erect herb/subshrub) and Leucaena leucocephala (Lamk) de Wit (shrub/tree). Like signal grass, palisade grass pastures can be severely affected in their productivity and persis­ tence by spittlebug (mainly the genera Aeneola- mia, Deois and Zulia in the family). This pest, however, seems to be restricted to trop­ icalAmerica . The most wide-spread cultivar, 'Mar­ andu', is spittlebug-resistant. Palisade grass can be continuously or rotationally grazed down to aheigh t of20-3 0 cm.I t is harvested by grazing animals or is mown and fed fresh to ani­ mals in cut-and-carry systems. Dry matter yields of palisade grass vary, according to growth condi­ tions, from 8-20 t/ha per year. At stocking rates of 1.5 steers/ha during the dry season and 2.5 steers/ha during the rainy season, annual livew- eight gains can range between 400-500 kg/ha. It flowers and sets seed abundantly; a first seed crop Brachiaria brizantha (A. Rich.) Stapf-habit. is possible 6-8 months after sowing. Depending on 58 FORAGES environmental conditions, up to 3 seed crops per proved to be useful as grazed ground cover in tree year can be obtained with a total seed yield of plantations, and is suitable for erosion control. 100-500 kg/ha. Properties Signal grass provides a forage that Genetic resources and breeding Palisade is palatable to ruminants (but not to ); grass, particularly the material from East Africa, depending on regrowth age, it is of moderate to is well represented in the major germplasm collec­ high quality. Nitrogen concentrations decrease tions of tropical forage grasses such as CIAT (Col­ from 2.7-0.7% and in vitro DM digestibility from ombia). At CIAT, parallel to evaluation and selec­ 75-50% as age of regrowth increases. When con­ tion within the available large B. brizanthajB. sumed as a pure diet, signal grass can occasionally decumbens gene pool, a breeding programme is in cause skin-photosensitization combined with progress aiming at cultivars that combine spittle- hepatic disorders, particularly in sheep and goats, bug resistance, low soil-fertility requirements, and and also in young cattle. There are 200-300 seeds/ high nutritive value. g- Prospects Palisade grass,mainl y the spittlebug- Botany A low-growing, decumbent, rhizoma- resistant cultivar 'Marandu', will continue to play tous and stoloniferous, apomictic (in cultivation) an important role in the development of tropical perennial forming a dense soil cover. Stem pros­ pastures in regions where spittlebug is a major trate to ascending, 30-150 cm high. Leaf broadly constraint until new Brachiaria cultivars with linear to narrowly lanceolate, 5-25 cm x 7-20 spittlebug resistance as well as lower soil-fertility mm, sparsely to densely pubescent, bright green. requirements become available. Inflorescence composed of 2-7 racemes on an axis Literature |1|Bogdan , A.V., 1977. Tropical pas­ 2-10 cm long; racemes 1-5 cm long, often borne ture and fodder plants. Longman, London, p. 54. at almost right angles to the axis (hence the name |2| Cuesta, P.A. &Pérez , R.A., 1987.Past o La Liber- 'signal' grass), bearing spikelets usually in 2 rows tad. Instituto Colombiano Agropecuario (ICA), on a broad, flattened rachis; spikelets elliptic, 4-5 Bogota, Colombia. Boletin Técnico No 150. 16 pp. mm long, pubescent at the tip; lower glume -j—j the |3| Empresa Brasileira de Pesquisa Agropecuâria, spikelet length, clasping; upper glume membra­ Centro Nacional de Pesquisa de Gado de Corte, nous; upper lemma granulöse. 1984. Brachiaria brizantha cv. Marandû. B. decumbens intergrades with B. brizantha (A. EMBRAPA-CNPGC, Campo Grande-MS, Brazil. Rich.) Stapf, and the species are difficult to dis- Documento No 21. 31pp . |4|Skerman , P.J. & Rive- ros, F., 1990. Tropical grasses. FAO, Rome. pp. 234-237. R. Schultze-Kraft

Brachiaria decumbens Stapf

Flora of Trop. Africa 9:52 8(1919) . GRAMINEAB 2n = 36 Synonyms Brachiaria bequaertii Robyns (1932). Vernacular names Signal grass (En). Malay­ sia: rumput signal. Thailand: ya-siknaentonnon, ya-surinam. Origin and geographic distribution The ori­ gin of signal grass is in East Africa (Uganda, Kenya, Tanzania, Rwanda, Burundi, Zaire). Ex­ perimental lines and cultivars have spread very widely to all tropical regions of the world. Thus, signal grass is becoming increasingly naturalized in South-East Asian countries and the Pacific region. Uses The main use of signal grass is as forage Brachiaria decumbens Stapf- 1,habit leafy stems; in permanent pastures that are grazed. It has also 2, inflorescence. BRACHIARIA 59 tinguish. Therefore, in the literature the name B. mown in cut-and-carry systems and fed asfres h ma­ decumbens probably sometimes actually refers to terial. It withstands heavy grazing and trampling B. brizantha, and vice versa. The main difference and thus can be grazed continuously or rotatio- between the cultivars or lines of B. brizantha and nally at stocking rates that should be high enough B. decumbens is the growth habit: B. decumbens to prevent the grass from becoming old and of poor is low-growing and forms a dense cover whereas quality. Depending on soil fertility and moisture B. brizantha is tufted with a rather erect growth. regime, DM yields of signal grass range between The best-known and most widely used cultivar of 6-36 t/ha per year. Animal production varies ac­ signal grass is 'Basilisk', released in Australia. cordingly (liveweight gain of 250-800 g/steer per Ecology Signal grass is well adapted to the day and up to 1200kg/h a per year). humid and sub-humid tropics where it withstands Genetic resources and breeding There is no dry seasons of up to 5month s but does not tolerate variability within cultivars or lines ofsigna l grass. flooding for more than a couple of days. It grows As their reproduction is apomictic, progenies are well on a range of soils, provided they are well essentially clones of the mother plant. However, drained, including acid, highly Al-saturated soils the major germplasm banks of tropical forage of medium to low fertility. It is moderately shade grasses at ATFGRC (CSIRO, Australia) and CIAT tolerant. (Colombia) hold a range of quite variable, distinct Agronomy Signal grass is easily established by B. decumbens accessions. At CIAT, besides selec­ seed, at rates that vary according to seed quality tion within a recently assembled, large B. decum- from 2-10 kg/ha. Germination of fresh seed may bensjB. brizantha gene pool, a breeding pro­ be poor because of dormancy (hard-seededness), gramme is in progress aimed at B. decumbensjB. but improves with time (up to one year of storage) brizantha cultivars that combine spittlebug resis­ or through scarification with sulphuric acid tance, low soil fertility requirements, and high (10-15 minutes). Signal grass can also be propa­ nutritive value. gated vegetatively by stem cuttings that are Prospects Until new lines from breeding and planted 60-100 cm apart in moist soil, or are broad­ selection programmes become available, signal cast and then disked in. The highly competitive grass cultivars or lines will continue playing a vigour exhibited by signal grass under appropriate very important role in tropical pasture develop­ conditions helps in suppressing growth of weeds ment in regions where spittlebug is no major con­ but creates problems with legume persistence. In straint, and where other Brachiaria species such spite of the competitive vigour of signal grass, per­ as B. brizantha, B. dictyoneura (Fig. & De Not.) sistent associations with legumes are feasible in Stapf or B. humidicola (Rendle) Schweick. have no certain environments. Suitable legumes are comparative advantage. mainly stoloniferous species such as Arachis pin- Literature |1|Bogdan , A.V., 1977. Tropical pas­ toi Krap. & Greg., nom. nud. (Pinto peanut), Des- ture and fodder plants. Longman, London, pp. modium heterophyllum (Willd.)DC .(hetero ) and D. 54-57. |2j Loch, D.S., 1977. Brachiaria decumbens heterocarpon (L.) DC. ssp. ovalifolium (Prain) (signal grass) - a review with particular reference Ohashi, but also the trailing-climbing Centrosema to Australia. Tropical Grasslands 11: 144-157. |3i pubescens Benth. (centro) and Pueraria phaseo- Oram, R.N., 1990. Register of Australian herbage loides (Roxb.) Benth. (tropical kudzu), the herba­ plant cultivars. CSIRO, Australia, pp. 89-90. |4j ceous to shrubby Stylosanthes guianensis (Aublet) Skerman, P.J. & Riveros, F., 1990. Tropical Swartz (stylo), and the shrub/tree Leucaena leuco- grasses. FAO, Rome. pp. 238-242. |5I Toutain, B., cephala (Lamk) de Wit. 1985. Graminées fourragères des genres Bra­ Although it is tolerant of low soil-fertility, it res­ chiaria et Urochloa pour lePacifique . Revue d'Ele­ ponds well to fertilization with N,P and K. Flower­ vage et de Médecine Vétérinaire de Nouvelle Calé- ing and seed-setting can occur year-round and donie 7:47-56 . high seed yields can be obtained. R. Schultze-Kraft &J.K . Teitzel The major pest problem of signal grass (which, however, seems to be restricted to tropical Amer­ ica) is spittlebug (mainly the genera Anaeolamia, Deois and Zulia in the Cercopidae family). This pest can very severely affect the productivity and persistence of signal grass. Signal grass is harvested by grazing animals or is 60 FORAGES

Brachiaria dictyoneura (Fig. &D e Not.) Stapf

Flora of Trop. Africa 9:51 2(1919) . GRAMINEAE 2n = 42 Synonyms Panicum dictyoneurum Fig. & De Not. (1854), Brachiaria obvoluta Stapf (1919), B. keniensis Henr. (1940). Vernacular names Thailand: ya signal luey. Origin and geographic distribution B. dictyo­ neura has its origin and natural distribution in eastern and southern Africa. It is increasingly being introduced to South-East Asia and the Pacif­ ic region and is spreading quite quickly in tropical America. Uses B. dictyoneura is used as a forage in perma­ nent pastures that are grazed. It has potential for erosion control and also as ground cover in tree plantations. Properties The nutritive value ofB. dictyoneura is regarded as moderate, yet higher than that of koronivia grass (B. humidicola (Rendle) Schweick.). Nitrogen concentrations range mostly between 1-2% but can be as low as 0.5% during Brachiaria dictyoneura (Fig. & De Not.) Stapf- 1, the dry season. The range for in vitro DM digestibi­ habit leafy stem; 2, inflorescence. lity is 55-70%. The grass has a high seed-produc­ tion potential; there are about 170 seeds/g. Ecology B. dictyoneura is well adapted to the Botany A densely tufted, semi-erect, stolonifer- humid and sub-humid tropics where it withstands ous perennial with short rhizomes and with stems dry periods of up to 4-5 months. It grows well on 40-120 cm high; stolons slender but strong and of a wide range of soils, provided they are well reddish colour. Leaf linear to lanceolate, 4-40 cm drained, including acid and highly Al-saturated x 3-18 mm, glabrous and with strongly denticu­ soils of low fertility. late margins. Inflorescence consisting of 3-8 Agronomy B. dictyoneura is preferably propa­ racemes on an axis 5-25 cm long; racemes 1-8 cm gated by seed, but can be propagated vegetatively long, bearing spikelets in two rows;spikelet s ellip­ by rooted stem segments or stolons. Recommended tic, 4-7 mm long, pubescent; lower glume seeding rate is 2-12 kg/ha, depending on seed qual­ 75-100%o of spikelet length, 11-nerved; upper ity. Germination of fresh seed is very poor because glume 7-9-nerved; lower lemma 5-nerved; upper of dormancy, mainly physiological. Scarification lemma slightly papillose. with sulphuric acid improves germination only Flowering seems to be long-day induced, and seed- slightly; therefore, it is recommended to use seed setting improves at higher latitudes. In cultivars, that is 6-8 months old. reproduction is apomictic. Its slow intial growth and weakly stoloniferous B. dictyoneura can easily be confused with B. growth habit enable association with a range of humidicola (koronivia grass). The latter is, howev­ legumes of varying growth habits, such as Desmo- er, much more stoloniferous than B. dictyoneura. dium heterocarpon (L.)DC . ssp.ovalifolium (Prain) Further important differences, at least regarding Ohashi (prostrate-stoloniferous), Centrosema the cultivars or commercial lines, are that B. dic­ pubescens Benth. or Puerariaphaseoloides (Roxb.) tyoneura has a noticeable, undulating protuber­ Benth. (trailing-climbing), Stylosanthes guianen- ance along the leaf-collar (outer surface of the sis (Aublet) Swartz (semi-erect herb/subshrub), junction of leaf-sheath and blade), and that the and Leucaena leucocephala (Lamk) de Wit (shrub/ -rachis of B. dictyoneura is remarkably tree). long-ciliate at the margins. The only cultivar Although its soil-fertility requirements are low, B. released so far is 'Llanero' in Colombia. dictyoneura responds well to fertilization, particu- BRACHIARIA 61 larly with N. B. dictyoneura should be leniently Brachiaria distachya (L.) Stapf grazed until it isfull y established. Then, using con­ tinuous or rotational grazing, stocking rates can Flora ofTrop . Africa 9:56 5(1919) . be relatively high (2-4 steers/ha), with seasonal GRAMINEAE adjustments as necessary. The grass should be In = 36 grazed down to 15-20 cm height. Synonyms Panicum distachyon L. (1771), Digi- Like most other species of Brachiaria, B. dictyo­ taria distachya (L.) Pers. (1805), Urochloa dista­ neura is affected by spittlebug (species of the chya (L.) Nguyen (1966). genera Aeneolamia, Deois, and Zulia in the Cerco- Vernacular names Green summer grass (En). pidae family) in tropical America. However, Indonesia: blembem, kadalan, blabakan (Java). unlike signal grass (B. decumbens Stapf), it recov­ Malaysia: rumput minyak, rumput meiera minyak. ers quickly. Philippines: gome-gome, tanageb. Thailand: B. dictyoneura isusuall y harvested as fresh materi­ yateenka. Vietnam: co' mât. al by grazing animals. Depending on growth condi­ Origin and geographic distribution B. dista­ tions, including the severity of the dry season, the chya originates and occurs from the Indian sub­ grass produces DM yields ranging from 4-11 t/ha continent throughout South-East Asia, Australia per year on acid, low-fertility soils.Anima l produc­ and the Pacific Islands. tion on a hectare basis varies accordingly. Daily Uses B. distachya is used primarily as forage. In liveweight gains of 250-500 g/steer can be coastal sand dunes it acts as a soil-binder. expected. Properties Results from Thailand have shown Genetic resources and breeding There is no that 8-week-old growth of B. distachya had 65% variation within 'Llanero'; as its reproduction is digestible DM and low N concentrations of 1.5% apomictic, the progenies are essentially clones of at 4week s and 0.9% at 12weeks . the mother plant. The species is not very well Botany Creeping annual; stems up to 50 cm tall, represented in the major germplasm banks of trop­ ical forage grasses at ATFGRC (CSIRO, Australia) and CIAT (Colombia), and only a few but distinct accessions are so far available. There are no breed­ ing programmes involving B. dictyoneura. Prospects Compared with other Brachiaria spe­ cies adapted to acid, low-fertility soils, B. dictyo­ neura offers the advantage of being less affected by spittlebug than signal grass and the advantage of better forage quality and compatibility with legumes than koronivia grass. For these reasons, it has excellent potential for pasture development in acid-soil regions. Literature |1|Centr o Internacional de Agricul­ ture Tropical, 1983-1990. Annual reports. Trop­ ical Pastures Program. CIAT, Cali, Colombia. ,2\ Instituto Colombiano Agropecuario, 1987. Pasto 'Llanero'. ICA, Bogota, Colombia. Boletin Técnico No 151. 12 pp. |3| Keiler-Grein, G. (Editor), 1990. Red Internacional de Evaluation de Pastos Tropi­ cales RIEPT-Amazonia. I Reunión-Lima, Peru, 6-9 Noviembre 1990.Document o deTrabaj o No75 . Centro Internacional de Agricultura Tropical, Cali, Colombia. 2Vols . 1119pp . R. Schultze-Kraft

Brachiaria distachya (L.) Stapf - 1, habit leafy stem; 2, inflorescence; 3, spikelet. 62 FORAGES

ascending from aprostrat e base, onth e soil surface Brachiaria humidicola (Rendle) rooting at the nodes. Leaf broadly linear to Schweick. narrowly lanceolate, 2-8 cm x 3-7 mm; sheath keeled, up to 3 cm long, hairy at the margin when Kew Bulletin 1936:29 7(1936) . young. Inflorescence consisting of 2-3 racemes on GRAMINEAE an axis 0.5-2 cm long; racemes 1-3 cm long with In = 72 spikelets attached singly on either side ofth e medi­ Synonyms Panicum humidicola Rendle (1899). an ridge of the rachis; spikelets narrowly elliptic, Vernacular names Koronivia grass (En). Thai­ 2.5-3 mm long, glabrous, acute; lower glume £-•£ land: ya humidicola. as long as the spikelet, clasping; upper glume sepa­ Origin and geographic distribution The ori­ rated from the lower by a short internode; upper gin and natural distribution of B. humidicola lemmarugulose . Caryopsis flattened ovoid, 1.6mm extends throughout eastern and southern Africa. long, light yellow. During the past 2-3 decades, it has been intro­ B. distachya commences flowering early in the duced to other tropical regions and has become a growing season and then flowers continuously fairly widespread species in South-East Asia and throughout the year, but little seed is set. B. dista­ the Pacific region. chya much resembles B. subquadripara (Trin.) Uses Koronivia grass is mainly used as a forage Hitchc, with which it is sometimes united. The in permanent pastures that are grazed, and for ero­ range of spikelet length seems to be disjunct (2.5-3 sion control. It has also proved to be useful as mm and 3.3-3.7 mm respectively), thus also justify­ grazed ground cover in tree plantations. ing the maintenance of two species. Properties Koronivia grass has a reputation as Ecology B. distachya is adapted to humid trop­ a palatable forage with very low N concentrations, ical lowlands. It can grow under light shady condi­ even during the growing season (0.6-1.0 %), which tions such as occur in old coconut plantations or can severely limit forage intake by ruminants. In orchards. It is often found in small patches along vitro DM digestibility values range from 50-70%. forest edges and waste places where it is freely The grass tolerates shade and poor drainage and grazed by village cattle. It does not tolerate water­ is an effective seed producer. There are 200-260 logging but is well adapted to sandy loam soils. seeds/g. Agronomy B. distachya is vegetatively propa­ Botany A prostrate, strongly stoloniferous and gated through division ofroo t stocks. It is very pal­ rhizomatous perennial, forming a dense ground atable and light grazing ispreferred . Yields of3 t/ha cover. Stolons slender but strong and of reddish ofD M without fertilizer and 9t/h a with mixed fertil­ colour; flowering stems ascend up to 100 cm. Leaf izer have been obtained in southern Thailand. linear to narrowly lanceolate, 4-30 cm x 3-10 Genetic resources and breeding It is unlikely mm, glabrous. Inflorescence consisting of 2-5 that any substantial germplasm collections are racemes on an axis 2-13 cm long; racemes 2-7 cm being maintained. long, bearing the spikelets in two rows; spikelets Prospects B. distachya is an interesting natural broadly elliptic, 4-6 mm long, slightly pubescent; forage grass in South-East Asia, deserving more lower glume 75-100% of spikelet length; attention because of its high palatability and its 11-nerved; upper glume 5-9-nerved; lower lemma tolerance of light shade and poor soils. 5-nerved; upper lemma slightly papillose. Literature |1| Gilliland, H.B., Holttum, U.E. & Koronivia grass can easily be confused with B. dic- Bor, N.L., 1971. Grasses of Malaya. In: Burkill, tyoneura (Fig. & De Not.) Stapf. The main differ­ H.M. (Editor): Flora of Malaya. Vol. 3. Govern­ ence is that koronivia grass is very strongly stolo­ ment Printing Office, Singapore, pp. 176-178. |2! niferous while B. dictyoneura is distinctly tufted Holm, J., 1971. Feeding tables. Nutrition Labora­ and considerably less stoloniferous. There are sev­ tory of Thai-German Dairy Project, Livestock eral cultivars throughout the tropics; however, Breeding Station, Chiangmai, Thailand, pp. 9-10. they probably represent the same genotype. Offi­ |3| Kotepat, W. &Sookase m P., 1989. Effects of cow cially released cultivars are 'Tully' in Australia dung and chemical fertilizer on the yields and and 'INIAP-701' in Ecuador. The reproduction in nutritive values of Brachiaria distachya on Ban- cultivars is apomictic. thon soils [in Thai]. 1987-89 Kesearch Report. Ecology Koronivia grass is well adapted to the Lampang Animal Nutrition Research Center, humid tropics but also thrives under sub-humid Department of Livestock, Bangkok. 42pp . tropical conditions where it withstands 3-4 dry C. Manidool months. It grows well on a range ofsoils , including BRACHIARIA 63

Because ofit s strongly stoloniferous growth habit, koronivia grass can withstand very heavy grazing. Overstocking, however, will result in very low ani­ mal growth rates. Prostrate-stoloniferous legumes such as Arachis pintoi Krap. & Greg., nom. nud. (Pinto peanut), Desmodium heterophyllum (Willd.) DC. (hetero), and D. heterocarpon (L.) DC. ssp. ova- lifolium (Prain) Ohashi have the potential to per­ sist readily in associations with koronivia grass. However, when all soil nutrient deficiencies have been corrected, Centrosema pubescens Benth. and Pueraria phaseoloides (Roxb.) Benth. have also persisted with it. Although its requirements for soil fertility are low, it responds to fertilization. As with most other Brachiaria species, the produc­ tivity of koronivia grass can be affected by spittle- bug (Aeneolamia spp., Deois spp., and Zulia spp. in the Cercopidae family). Unlike most other Bra­ chiaria species, however, it recovers quickly from attacks by this insect pest. Rust (Uromyces setaria- italicae) has recently been identified as a potential major disease in tropical America. Koronivia grass is regarded as a high-yielding grass. Depending on climate and soil fertility, DM production ranges from 7-33 t/ha/ per year. Because of its high carrying capacity, liveweight gain per ha can be high, but with rather low growth rates (150-500 g/steer per day). Although Brachiaria humidicola (Rendle) Schweick. 1, the grass isusuall y consumed fresh by grazing ani­ habit leafy sterns; 2, inflorescence. mals, hay making is occasionally practised in sub- humid environments in Brazil and Venezuela. moist, poorly drained sites. It is particularly well Genetic resources and breeding There is no adapted to acid, highly Al-saturated soils of low variation in cultivars of koronivia grass as their fertility. reproduction is apomictic, resulting in progenies Agronomy Koronivia grass can be propagated being clones of the mother plant. The major germ- and established by seed at a rate of 2-8 kg/ha plasm banks of tropical forage grasses at CIAT depending on seed quality, or vegetatively by root­ (Colombia) and ATFGRC (CSIRO, Australia) hold ing stolon cuttings that are spread and disked in, a range of quite variable B. humidicola accessions. or planted by hand, at 1m x 1m distances or less. At present, there are no B. humidicola breeding The latter method is particularly convenient in programmes. Research activities to improve the smallholder systems. species are limited to evaluation of the available Under favourable conditions, koronivia grass can genepool. grow very quickly and form a dense ground cover Prospects Because of its aggressive, weed-sup­ ready for grazing 3-5 months after sowing. pressing growth, tolerance of overgrazing, adapta­ Because ofit s growth habit and highly competitive tion to poorly drained sites, and low requirements vigour, the growth of weeds is easily suppressed, regarding soil fertility, the present Koronivia but associations with legumes are difficult to grass cultivars will continue to play an important maintain. role in tropical pasture development. Their impor­ In contrast to lower altitudes, koronivia grass tance will only decline when new cultivars or grown at higher altitudes has the potential to other species become available which combine the flower profusely and to produce seed yields of up above-mentioned features with better forage quali­ to 150kg/h a during the first year. Once the pasture ty. isfull y established, flowering and seed production Literature jl| Bogdan, A.V., 1977. Tropical pas­ are scarce. ture and fodder plants. Longman, London, pp. 64 FORAGES

57-58. |2I Centro Internacional de Agricultura Tropical, 1983-1990. Annual reports, Tropical Pastures Program. CIAT, Cali, Colombia. j3| Oram, R.N., 1990. Register of Australian herbage plant cultivars. CSIRO, Australia, pp. 91-92. |4| Sker- man, P.J. & Riveros, F., 1990. Tropical grasses. FAO, Rome.pp . 245-248. |5| Toutain, B.,1985 . Gra­ minées fourragères des genres Brachiaria et Urochloa pour le Pacifique [Grass forages of the genera Brachiaria and Urochloa for the Pacific]. Revue d'Elevage et de Médecine Vétérinaire de Nouvelle Calédonie 7:47-56 . R. Schultze-Kraft &J.K . Teitzel

Brachiaria mutica (Forssk.) Stapf

Flora of Trop. Africa 9:52 6(1919) . GRAMINEAE 2n = 36 Synonyms Panicum muticum Forssk. (1775), P. purpurascens Raddi (1823), Brachiaria purpuras- cens (Raddi) Henr. (1940). Vernacular names Para grass, Mauritius grass (En). Herbe de para (Fr). Indonesia: rumput malela, sukut kolonjono, jukut inggris. Malaysia: rumput melela, rumput para. Philippines: babaka- nalabaga, mara-kawayan (Ilokano). Cambodia: smau kôô. Thailand: ya khon. Vietnam: co' long Brachiaria mutica (Forssk.) Stapf- 1, habit culm tây. with leaves; 2, ligule; 3, inflorescence; 4, part of Origin and geographic distribution The ori­ raceme; 5, spikelet. gin of para grass is probably tropical Africa. Today, the grass occurs naturally all over the trop­ 2-15 cm long, basal ones generally branched; spik- ical world, including South-East Asia and the Pa­ elets paired (often single in upper part), elliptic, cific region. 2.5-5 mm long, glabrous, in several untidy rows; Uses It is mostly grazed but can also be cut and fertile (upper) floret 3 mm long, pale yellow when fed to tethered animals or animals confined in ripe. Reproduction is mainly apomictic. stalls. It is suitable for erosion control at moist Under appropriate conditions, para grass is very sites, e.g. on river banks. As a cover crop in planta­ competitive and vigorous, thus suppressing growth tions it is often considered too agressive. of weeds and making persistent associations with Properties Para grass is a palatable forage of legumesdifficult . Flowering and seed-setting occur, reputedly high quality. Depending on plant age, particularly at low latitudes and in humid environ­ N concentrations can range from 1.6-2.2% for ments. There is no formally released cultivar of leaves and from 0.5-1.0% for stems, and in vitro para grass. Para grass closely resembles tanner DM digestibility from 40-65 %. grass {Brachiaria arrecta (Th. Dur. &Schinz ) Stent Botany A robust, coarse, creeping perennial from tropical and southern Africa, but the latter with short rhizomes and long stolons. Stems species always has single spikelets. decumbent to ascending, rooting at nodes, densely Ecology Para grass is best adapted to poorly covered with long, white hairs. Leaf-sheath hairy drained (swampy or periodically waterlogged) and or glabrous on upper portion, with densely hairy wet areas of the warm tropics with soils of medium collar; leaf-blade linear to lanceolate, 6-30 cm x to high fertility; it tolerates salinity. As long as 5-20 mm, flat, glabrous or slightly hairy. Inflores­ soil moisture is appropriate, it also thrives in well- cence a panicle 6-30 cm long with 5-20 densely drained areas. Growth is poor on dry sites. It can flowered racemes somewhat separated; racemes become a weed in irrigation ditches and drains. BRACHIARIA 65

Agronomy Although establishment by seed is 'ponded pastures' (shallow earth dams) in drier en­ feasible (using 4 kg/ha), para grass is usually and vironments. However, because of insect problems, easily established from stem cuttings with at least it is gradually being replaced by B. humidicola 3-4 nodes each, two of which should be buried (Rendle) Schweick. in the humid tropics of Austra­ 10-15 cm deep. Planting density depends on weed lia. potential of the site and desired speed of ground Literature 11, Blair, G.J., Ivory, D.A. & Evans, cover, and should not be at spacings wider than T.R., 1986. Forages in Southeast Asian and South 1 m x 1m . If machinery is available, spread stem Pacific agriculture. ACIAR Proceedings Series No cuttings can be disked in. It responds to fertiliza­ 12. Canberra, Australia. 202 pp. !2i Bogdan, A.V., tion, mainly Nbu t also P and K,whic h become par­ 1977. Tropical pasture and fodder plants. Long­ ticularly important if the grass is cut regularly. man, London, pp. 59-62. |3! Skerman, P.J. & Rive- Legumes compatible with para grass at poorly ros, F., 1990. Tropical grasses. FAO, Rome. pp. drained sites are the annuals Macroptilium lathy- 249-253. ;4j Toutain, B., 1985. Graminées fourra­ roides (L.)Urba n (phasey bean) and Aeschynomene gères des genres Brachiaria et Urochloa pour le americana L. (American jointvetch), and at well- Pacifique [Grass forages of the genera Brachiaria drained sites the perennials Puerariaphaseoloides and Urochloa for the Pacific]. Revue d'Elevage et (Roxb.) Benth. (tropical kudzu), Centrosemapubes- de Médecine Vétérinaire de Nouvelle Calédonie 7: cens Benth. (centro), and the shrub Leucaena leuco- 47-56. cephala (Lamk) de Wit. R. Schultze-Kraft &J.K . Teitzel Para grass can be grazed continuously, or rotatio- nally at intervals of 5-8 weeks, but a residual height of 15-20 cm should be maintained. Brachiaria ruziziensis Germain & InAustralia , para grass isattacke d by a leafhopper Evrard which sucks the sap and severely reduces pasture productivity. Bull. Jard. Bot. Etat Brux. 23: 373(1953) . Para grass is harvested by grazing animals or cut GRAMINEAE for stall-feeding. It can be used for hay-making if 2n = 18, 36 environmental conditions allow but is difficult to Vernacular names Ruzi grass, Congo grass, dry. Depending on growth conditions, DM yields Congo signal grass (En).Thailand : ya ruzi. can range between 3-39 t/ha per year, mostly 5-12 Origin and geographic distribution Ruzi t/ha per year. Animal production varies according­ grass has its origin in the Ruzizi valley in eastern ly and annual liveweight gains of 300-800 kg/ha Zaire and Burundi. It is widely distributed on can be expected from para grass. Depending on N farms and experiment stations in most tropical fertilization, seed yields of up to 60 kg/ha are pos­ countries including South-East Asia and the Pacif­ sible. However, except in Australia, commercial ic region. seed production of para grass is practically non­ Uses Ruzi grass is used as a forage for direct existent because of the ease of establishment from grazing of permanent pastures, in the open or stem cuttings. under coconuts, and for feeding tethered or stalled Genetic resources and breeding Para grass is ruminants. not well represented in the major germplasm col­ Properties Ruzi grass is palatable and of excel­ lections of tropical forage grasses held by CIAT lent quality, with N concentrations ranging from (Colombia) and ATFGRC (CSIRO, Australia). The 1.5-2.5% and in vitro DM digestibility from variability in the available gene pool is limited 50-75% (4-16 weeks old regrowth). It tolerates because ofth e species' apomictic reproduction and shade moderately, and drought fairly well. It has clonal propagation. However, important varietal a high seed production potential. There are differences have been observed in Thailand and 230-250 seeds/g. Burma. Because of lack of a gene pool, there has Botany A tufted, creeping perennial, forming a been no description of the variation within para dense and leafy cover. Stems leafy and hairy, aris­ grass and no evaluation or breeding programmes ing from many-noded, creeping shoots and short to select improved lines. rhizomes up to 1.5 m high when flowering. Leaves Prospects Para grass will continue to be one of hairy, broad-lanceolate, 10-25 cm x 10-15 mm. the most widely used good-quality forages for low- Inflorescence consisting of 3-9 relatively long lying, seasonally waterlogged areas of the tropics. (4-10 cm) racemes, bearing spikelets in 1o r 2 rows In Australia it has also become popular for use in on one side of a broad, flattened and winged rachis 66 FORAGES

nation of fresh seed is poor because of dormancy, but increases following storage or treatment with concentrated sulphuric acid. Because ofit s high soil-fertility requirements, ade­ quate N, either through a legume or fertilizer, P and K must be supplied in order to maintain high productivity of ruzi grass, particularly under cut­ ting. Ruzi grass can be sown with a range of legumes such as stylo (Stylosanthes guianensis (Aublet) Swartz), greenleaf desmodium (Desmodium intor- tum (Miller) Urban), centro (Centrosemapubescens Benth.), and the shrub legume Leucaena leucoce- phala (Lamk) de Wit. No diseases or pests of economic importance are known to affect ruzi grass in South-East Asia. In tropical America it can be severely damaged by spittlebug (Aeneolamia spp., Deois spp., and Zulia spp.i n the Cercopidae family). In Zaire the inflores­ cences are parasitized by the fungus Sphacelia sp. Ruzi grass is harvested by grazing animals or cut for stall-feeding, and is also suitable for making hay. Depending on growth conditions (mainly soil fertility), annual DM yields of 10-20 t/ha can be expected from ruzi grass. Annual liveweight gains ofbee f cattle can behighe r than 500kg/ha , depend­ ing on soil N status. Brachiaria ruziziensis Germain &Evrard-1, habit Genetic resources and breeding The gene flowering culm; 2, inflorescence. pool ofruz i grass is limited to a few accessions held in the collections of ATFGRC (CSIRO, Australia) of generally purplish colour; spikelets hairy, 5 mm and CIA T (Colombia). Because of its sexual repro­ long; lower glume 3m m long, 0.5-1 mm distant (= duction and a high degree of cross-pollination, below) from rest of spikelet. It produces seed abun­ these accessions are quite variable. There are no dantly. ruzi grass breeding programmes, but a tetraploid Brachiaria ruziziensis is very closely related to B. form of this species is used at CIAT as a source decumbens Stapf, but differs morphologically (in of sexuality in a breeding project involving the B. ruziziensis therachis is subfoliaceous and 2-3.5 tetraploid, apomictic B. decumbensjB. brizantha mm wide and the lower glume is at a considerable complex. distance from rest of spikelet; in B. decumbens the Prospects The main value of ruzi grass lies in rachis is flat, 1-1.7 mm wide and internode be­ its reasonably high nutritive value and high seed tween lower and upper glume is shorter) and in re­ production potential, but it is markedly less pro­ production (sexual in B. ruziziensis, apomictic in ductive than B. decumbens. Its importance will de­ B. decumbens). The best known cultivar is 'Ken­ crease rapidly as current Brachiaria selection pro­ nedy', released in Australia. grammes result in alternatives which combine the Ecology Ruzi grass is well adapted to the warm above mentioned attributes with adaptation to tropics with annual rainfall in excess of 1000 mm; low-fertility soils. it can withstand up to 5 dry months. It tolerates Literature |1| Bogdan, A.V., 1977.Tropica l pas­ a range of soils but requires very fertile soil. It ture and fodder plants. Longman, London, pp. thrives best on well-drained sites. 64-65. |2|Reynolds , S.G., 1988.Pasture s and cattle Agronomy Although it can also be propagated under coconuts. FAO plant production and protec­ vegetatively by stem cuttings with rooting nodes, tion paper 91. FAO, Rome. pp. 44-46. |3| Skerman, ruzi grass is easily established by seed at rates of P.J. & Riveros, F., 1990. Tropical grasses. FAO, 2.5-10 kg/ha, depending on seed quality. Seed Rome.pp .255-259 . |4|Toutain , B., 1985. Graminées should not be sown deeper than 1.5-2.5 cm. Germi­ fourragères des genres Brachiaria et Urochloa BRACHIARIA 67 pour le Pacifique [Grass forages ofth e genera Bra­ chiaria and Urochloa for the Pacific]. Revue d'Ele­ vage et de Médecine Vétérinaire de Nouvelle Calé- donie 7:47-56 . R. Schultze-Kraft &J.K . Teitzel

Brachiaria subquadripara (Trin.) Hitchc.

Lingnan Sei. Journ. 7:21 4(1931) . GRAMINEAE 2n = 54-56, 72 Synonyms Panicum subquadriparum Trin. (1828),P. miliiforme J. &C .Pres l (1830), Brachiaria miliiformis (J. &C .Presl ) Chase (1920). Vernacular names Cori grass (En). Origin and geographic distribution Of trop­ ical Asian origin (India, Sri Lanka, Burma, Malay­ sia, Pacific Islands), this grass has been introduced to other regions such as tropical Africa. Uses The main use of cori grass is as a forage for ruminants in coconut plantations where it is grazed by animals, and as a shade-tolerant ground cover. Properties Cori grass is very palatable, and young regrowth is of acceptable forage quality Brachiaria subquadripara (Trin.) Hitchc. - habit. (1-2% Nat4-8 weeks). Botany A creeping stoloniferous perennial Agronomy Cori grass is easily and quickly pro­ (reported as annual by some authors), mat-forming pagated and established by stolon cuttings par­ under close grazing, with slender stems up to 1 m tially buried 10-15 cm deep. Planting density long, ascending from a prostrate base up to 50 cm depends on weed potential and desired speed of high, rooting at hairless nodes.Lea f broadly linear ground cover; establishment of a minimum of 1 to narrowly lanceolate, 2-20 cm x 5-10 mm, gla­ plant/m2 is recommended. brous, with rounded base and sharp tip. Inflores­ Once well established, Cori grass can withstand cence consisting of 2-5 widely spaced racemes on heavy grazing and competes well with weeds, par­ an axis 3-10 cm long; racemes 1-6 cm long, bear­ ticularly under some shade. Under coconuts, an­ ing singly attached spikelets in two rows; spikelets nual DM yields vary from 4-9 t/ha. Suggested stoc­ narrowly elliptic, 3.3-3.7 mm long, glabrous, pale king rates are 1.5-2.0 steers/ha, and annual green; lower glume shorter, upper glume as long liveweight gain up to 400 kg/ha can be expected as spikelet; upper lemma rugulose; stigmas purple. without affecting yields of coconuts. The taxonomie situation of B. subquadripara and The grass responds well to N fertilization and is B. miliiformis is not clear. Some authors consider considered to be moderately drought-tolerant. them to be synonymous whereas others claim these Legumes suitable for association with Cori grass species are distinct but closely related, with B. sub­ are centro (Centrosema pubescens Benth.) and quadripara being a perennial with 2ra = 72 and B. tropical kudzu (Pueraria phaseoloides (Roxb.) miliiformis an annual with 2n = 54 chromosomes. Benth.). Ecology Cori grass is widely adapted to the Genetic resources and breeding In the major warm lowland tropics, but particularly suited to germplasm collections of tropical forages, CIAT monsoon environments. It prefers medium- to (Colombia) and ATFGRC (CSIRO, Australia), B. light-textured soils of medium to high fertility. It subquadripara seemst o berepresente d by only one is one of the most shade-tolerant tropical grasses, distinct genotype. and this makes it particularly valuable for pas­ Prospects Further development of the existing tures in tree plantations. genotype to improve adaptation, production or 68 FORAGES quality is unlikely. However, the genetic base of promise as an understorey in coconut plantations Cori grass should be broadened through plant col­ with about 60% light transmission. lection in view of its potential for pastures in tree Properties Calliandra leaves contain (DM plantations, particularly under coconuts. based): N 3-3.5%, fibre 30-75%, ash 4-5% and Literature |1|Bogdan , A.V., 1977. Tropical pas­ fat 2- 3% . No toxic substances have been reported, ture and fodder plants. Longman, London, p. 58. but high concentrations of condensed tannins (up |2i Plucknett, D.L., 1979. Managing pastures and to 11% ) are responsible for the rather low in vitro cattle under coconuts. Westview Tropical Agricul­ digestibility of 35-40%. The wide range of ture Series, No 2. Westview Press, Boulder, Color­ recorded tannin contents may reflect differences ado, United States, pp. 211-213. |3| Skerman, P.J. in analytical procedures as much as variation due & Riveros, F., 1990. Tropical grasses. FAO, Eome. to growth stage and environment. The digestibi­ pp. 260-262. lity data should also be treated with caution, as R. Schultze-Kraft when dried, material with a high tannin content can have a marked effect on digestibility and intake by animals. There is increasing evidence Calliandra calothyrsus Meissn. that wilted or dried material is less acceptable to animals than fresh material. There are 14-19 Linnaea 21: 251(1848) . seeds/g. LEGUMINOSAE Description Shrub or small tree, (1.5-)4-6(-12) 2n = 22 m tall, with trunk diameter up to 30 cm, blackish- Synonyms Calliandra confusa Sprague & Riley brown bark and a dense canopy. Leaves bipinnate, (1923). alternate, rachis 10-17 cm long, without glands, Vernacular names Calliandra (En). pinnae 4-7 cm long in 15-20 pairs with 25-60 pairs Origin and geographic distribution Although ofdar k green leaflets each; leaflets linear, 5-8 mm originally described from Surinam, C. calothyrsus x 1mm . Inflorescences composed of few to many is native in humid and sub-humid Central America flowered heads, in terminal raceme-like clusters of from southern Mexico to north-western , between 8-16°N. From Guatemala it was intro­ duced into Java in 1936.Durin g the 1970si t became well established in Indonesia, with over 30000 ha ofplantations . In view ofit s excellent performance in Indonesian plantations and its multiple uses, it is now being planted in many other countries of South-East Asia and to a lesser extent in other tropical areas. Uses Calliandra isuse d as a high quality protein leaf crop to supplement low quality forages and crop residues. Only the leaves and young stems are used for fodder. In Indonesia calliandra is primar­ ily used for fuelwood for households and small industries. The wood can also be used for pulp and paper production but its small dimensions make handling and chipping difficult. It is widely planted to control erosion on sloping lands and ravines. It is used to improve soils by nitrogen fixa­ tion and good litter production by cultivating it in rotation with arable crops. It is also incorpor­ ated in alley-cropping systems as a source of green manure, or used as a nurse tree for partially shade- tolerant timber species. Calliandra is also planted in firebreaks. Its beautiful red flower clusters make it a popular ornamental, and the flowers are a good nectar source for bees. It is a suitable host Calliandra calothyrsus Meissn. - 1, flowering for shellac insects (Kerria lacca). It has shown branch; 2, branch with fruits. CALLIANDRA 69

10-30 cm length. Flower showy, purplish-red, 4-6 opment occurs at moderate elevations below 1300 cm long; calyx 2 mm long; corolla 5-6 mm long, m. On Java the species is planted up to 1500m alti­ pale green; stamens numerous, 4-6 cm long, pur­ tude, but it grows best between 250-800 m in areas plish-red. Fruit broadly linear, flattened, 8-11 cm with 2000-4000 mm annual rainfall and a 3-6 x 1cm , slightly tapering from top to base, margins month dry period. The plants require amea n annu­ thickened and raised, finely pubescent or gla­ al temperature of (20-)22-28°C, with mean maxi­ brous, brownish, dehiscent, 3-15-seeded. Seed mum temperature in the hottest month between ellipsoidal, flattened, 5-7 mm long, dark brown. 24-30°C and mean minimum temperature in the Wood characteristics The wood of calliandra coldest month between 18-24°C. The species ismoderatel y heavy, volumic mass 510-780 kg/m3, occurs in secondary vegetations, often in thickets. strong and easy to saw. It provides good fuelwood It isa n aggressive colonizer ondisturbe d sites such (massic energy 18900-19950 kj/kg) and is suitable as recent landslides and roadsides. It grows on a for charcoal production. Cellulose content varies large variety of soil types ranging from deep, vol­ from 44-56%, fibre length from 0.66-0.84 mm, canic, sandy loams to alluvial soils and shallow or extractive material 3% DM. eroded metamorphic sandy clays.I t is well adapted Growth and development Mature seeds germi­ to acid soils ofpoo r fertility but can respond to fer­ nate readily. Seedlings grow quickly up to 2.5-3.5 tilizer application on such soils. m in 6 months and up to 3-5 m in the first year. It prefers light soil textures and slightly acid con­ Trees mature rapidly; flowering may start in the ditions. Best growth isobtaine d on acid soils ofvol ­ first year, but good fruit setting normally starts canic origin. Growth decreases on compacted soils the second year. Normally large quantities of seed and trees die after 2week s of oxygen depletion due are produced each year. In its native region, flow­ to waterlogging. ering is concentrated at the end ofth e rainy season Propagation and planting Calliandra is nor­ and the beginning of the dry season, and fruits are mally propagated by seeds, directly in the field or set in the dry season. On Java, calliandra flowers in the nursery. Seeds germinate without pre-treat- and fruits throughout the year, but most seeds are ment, but best germination is obtained if seeds are produced in the dry season. Flowers are insect-pol­ briefly treated with almost boiling water and linated and seeds mature 2 months after pollina­ soaked for 24hours . Seeds retain their viability for tion. Fruits are mature when they turn brown. The at least 2-3 years if stored in a refrigerator but stem turns brittle around the age of 12 years, but viability decreases within one year at room tem­ the rootstock remains vigorous and new sprouts perature. Direct sowing in the field can be done are easily formed. After cutting, trees coppice in planting holes or bybroadcastin g on previously vigorously. With annual coppicing of stems of 3-5 ploughed or burned lands. Seedlings usually nodu- cm in diameter, plants can survive for many years. late with native rhizobia and inoculation is not Roots develop quickly and may reach 1.5-2 m in required. Potted plants and transplants can be plants of 4-5 months old. Both surface and deeply raised with standard nursery techniques. Potted penetrating roots are formed. plants are transplanted when they are 20-50 cm In humid climates the tree is evergreen, in areas tall and have a root collar of 0.5-1 cm. Plants ap­ with a long dry season it is semi-deciduous. In proximately 1m tall can be transplanted after cut­ severe dry conditions, trees die back but recover ting the stem back to 30 cm and the roots to 20 cm. at the beginning of the rainy season. Spacing within plantations varies according to Other botanical information Calliandra purpose. For firewood, planting distances are 1m Benth. is a genus of about 100 species of shrubs x 1 m to 1 m x 2 m; in alley-cropping, spacing of and small trees of tropical and warm temperate 25-50 cm in the contour lines is used. Areas to be regions, some of which are widely cultivated as planted are cleared completely. To provide fodder ornamentals. Together with the introduction of C. calliandra is often planted in fences, rows or in calothyrsus in Java, the white-flowered C. tetra- small blocks, usually under a cut-and-carry gona Benth. was also introduced from Guatemala. regime. For optimal leaf production dense stands Because of its slower growth, this species became of up to 40000trees/h a can be used. less popular for plantations. Husbandry Because seedlings grow quickly, no Ecology In its native habitat C. calothyrsus special plantation management is needed, except grows at altitudes of 400-1800 m in areas with an for weeding in the first year. As yet there is not average annual precipitation from 700 mm to 3000 much specific knowledge on management prac­ mm and with 1-7 dry months per year. Best devel­ tices for optimal fodder and wood production from 70 FORAGES

C. calothyrsus. Neither is much known about the exceptional. Further provenance trials are needed feasibility of combining calliandra and cultivated to assess species variability, forage quality, and forage grasses. adaptability to different environmental condi­ Diseases and pests There are no serious dis­ tions. easeso rpest srecorde d inIndonesia , buti nth ePhi ­ Prospects C. calothyrsus has become popular lippines trees are damaged by a stem-borer. In because it can readily produce high-quality fuel- Kenya, damage from Pachnoda ephippiata (a rose wood in annual coppice rotations, can be used in flower beetle) hasrestricte d seed production toth e different farm systems as an auxiliary crop, and extent that it has been suggested that the insect can grow under a wide range of soil fertility. Spe­ could limit theus eo fC. calothyrsus. cial attention hasbee n given to theus eo fthi s spe­ Harvesting Harvesting of calliandra can nor­ cies as an alternative to Leucaena leucocephala on mally start after the first year. Forage is usually acid lands or areas infested with Leucaena psyllid. cut by hand. In Indonesia a cutting height of 35 Itshig h potential production and high protein con­ cm is commonly used, although forage yields may tent make it a promising fodder species to supple­ be slightly larger with a cutting height of10 0cm . ment low-quality forages. However, due to high Yield In plantations, forage DM yields are typi­ tannin content, the palatability of calliandra is cally in the range of 7-10 t/ha per year, although less than that of L. leucocephala or Gliricidia yields willb elowe r inarea s of low rainfall or infer­ sepium. Further studies areneede d to ascertain its tile soil. In one experiment, leaf yields were full potential as a fodder crop, including its true slightly higher when harvested at intervals of 12 nutritive value andit svalu e asa plan t for grazing. weeks instead of 6 weeks. When grown as fences, Care must also be taken that this hardy species fodder DMyield s of1.8-3. 2t/k mi n 10month s have does notbecom e a weed. been obtained. In one trial, yields of Panicuni- Literature 111 Ahn, J.H., Robertson, B.M., maximum Jacq. cultivar 'Eiversdale' were similar Elliott, R., Gutteridge, R.C. & Ford, C.W.,1989 . to those of C. calothyrsus, Leucaena leucocephala Quality assessment of tropical browse legumes: (Lamk) deWi tan d Gliricidia sepium (Jacq.) Kunth tannin content and protein degradation. Animal exWalp . Yields ofth e three legumes were also sim­ Feed Science and Technology 27:147-156. &Bag - ilar. When plots were harvested every 6 weeks, gio, A. &Heuveldop , J., 1984.Initia l performance grass yields were higher at denser tree spacings, of Calliandra calothyrsus Meissn. inliv efence s for but when harvested every 12 weeks, grass yields the production of biomass. Agroforestry Systems were higher at the wider tree spacings. On reason­ 2: 19-29. |3|Ella , A. & Blair, G.J., 1989. Effect of able soils on Java first wood harvests produced tree density and cutting frequency on the produc­ 5-20 m3/ha per year of fuelwood . On favourable tion of four tree legumes and understorey grass. sites annual coppice harvests continued for 10-20 Nitrogen Fixing Tree Research Reports 7: 14-16. years with yields of35-6 5 m3/ha per year. |4j Ibrahim, T.M., Palmer, B.,Boer , M. & Santhes, Genetic resources A collection of C. calothyr­ M., 1988. Shrub legume potential for integrated sus germplasm has been made by CATIE (Tur- farming systems innorther n Sumatra-nutritional rialba, Costa Rica) and the Oxford Forestry Insti­ constraints and palatability. In: Maximizing live­ tute (United Kingdom). stock productivity. Proceedings ofth e 11th Annu­ Breeding C.calothyrsus wasintroduce d in Java al Conference, Malaysian Society of Animal Pro­ as part of a trial of green manure crops for timber duction, pp. 128-132. |5j National Academy of plantations. Its introduction was based on only Sciences, 1983.Calliandra , a versatile small tree two seed samples from Guatemala. All existing from the humid tropics. National Academy Press, plantations inIndonesi a arederive d from this first Washington, D.C., United States. 52 pp. |6| Ver­ introduction. In Central America the species did hoef, L., 1941.Voorlopig e resultaten met enige uit not receive anyattentio n until itssucces s asa fire­ tropisch Amerika ingevoerde Leguminosae [Preli­ wood crop inIndonesi a became known inth e early minary results with someLeguminosa e introduced 1980s. The species in Central America has a wide from tropical America].Tecton a 25(10): 711-719. geographic distribution and is quite variable. It K.F. Wiersum &I.K . Rika seems that the seeds introduced from Guatemala to Indonesia were derived from a fast-growing, les­ ser branched andtalle r ecotype. Insom e trials spe­ cific calliandra accessions have been observed which produce root suckers, but this seems to be CALOPOGONIUM 71

Calopogonium caeruleum (Benth.) Botany A sturdy, somewhat woody, twining, Sauv. short-lived perennial with pubescent or glabres- cent stems up to several m long, rooting at nodes Ann. Acad. Habana 5: 337(1869) . when in contact with moist soil. Leaves trifolio- LBGUMINOSAE late, petiole up to 16 cm long; leaflets elliptical, 2n = unknown ovate or rhomboid-ovate, (1.5-)6-9(-20) cm x Synonyms Stenolobium caeruleum Benth. (1837). (l-)4-6(-15) cm, the lateral ones oblique, pubes­ Vernacular names Caeruleum calopo (En). cent above, velvety pubescent below. Inflores­ Thailand: thua sealulium. cences elongated many-flowered racemes, up to 50 Origin and geographic distribution Native to cm long; peduncle l-5(-17) cm long; rachis sul- Mexico, Central America and the and cate, tomentose; calyx campanulate, 5-lobed; co­ eastern tropical South America, C. caeruleum is rolla blue or violaceous, ca. 1c m long. Pod linear- now widespread throughout the humid tropics. It oblong, 4-8 cm x 0.8 cm, straight or curved, was introduced to South-East Asia in 1940. impressed between the seeds,pubescent . Seeds 4-8 Uses C.caeruleum is a forage of low palatability, per pod, orbiculate, 4-5 mm across, shiny brown, but it gives good soil cover, builds up soil fertility compressed. as a green manure crop, and can beuse d to smother In the literature, the species name is also written weeds such as Imperata cylindrica (L.) Raeuschel. 'coeruleum', but Bentham published the name as Properties Mineral concentrations in top 'caeruleum' (= blue). growth usually range from 2.1-3.6% N, 0.17-0.29% Although it grows vigorously once established, P, 2.4-2.6% K and 0.91-1.05%) Ca. In vitro DM growing as high as 1m in a pure sward, its growth digestibility was not affected by shading. It is much is not compact and it allows free aeration of the less palatable to animals than C.mucunoides Desv. soil. When grown as a cover crop it will smother young oil palms and rubber trees unless weeded out from around the base of the trees, it is a poor seeder in the first and second years after establish­ ment. Ecology C. caeruleum is adapted to the humid tropics but it is more drought-tolerant than C. mucunoides and Pueraria phaseoloides (Roxb.) Benth. It grows better in cooler conditions than centro (Centrosema pubescens Benth.) and in the elevated tropics can grow up to an altitude of about 800m .I t isver y tolerant ofshading . C. caeru­ leum is adapted to a wide range of soil textures and soils with a pH(H20) as low as 4.0. It grows best on well-drained soils. Agronomy C. caeruleum is usually established from seed, sowing at the start of the wet season into a seed-bed prepared mechanically or by hand. Fertilizing and weeding aids establishment and de­ velopment of the sward. However, the use of C. caeruleum seed is restricted by its high cost, which is a reflection of its poor seed production. It can bevegetativel y established from stem cuttings, but only about 5% of the cuttings may establish. The best success is obtained from using older stem ma­ terial, 50c m or more away from the terminal grow­ ing point. Establishment from stem cuttings can be improved by hormone treatments to induce root development. It is slower to establish than tropical kudzu (Pueraria phaseoloides) and may take 20 Calopogonium caeruleum (Benth.) Sauv.-l, flower­ months to establish a complete ground cover. It ing branch; 2,flower; 3,fruit; 4,opened fruit. persists longer under the increasing shade of 72 FORAGES young oil palm or rubber than most other legumes. Performance of tropical forages under the closed In Malaysia it still produced DM 1 t/ha 5 years canopy of oil palm. Legumes II. MAEDI Research after planting. Bulletin 12: 21-37. |3|d'Arcy , W.G., 1980. Calopo­ Although C. caeruleum is a productive species, gonium. In: Dwyer, J.D. et al. (Editors): Flora of especially under shade, it has very limited value Panama. Annals ofth e Missouri Botanical Garden as a forage because of its low palatability. Experi­ 67: 557-561. !4| Han, K.J. & Chew, P.S., 1981. ence in Malaysia and northern Australia has Growth and nutrient contents of leguminous shown that it can become dominant in grazed pas­ covers in oil palm plantation in Malaysia. Interna­ tures because it is virtually ungrazed, whereas the tional Conference on the Oil Palm in Agriculture companion species are selectively grazed. It is in the Eighties. Organized by the PORIM and the probably eaten slightly more during the dry sea­ Incorporated Society of Planters held on 17-20 son, when there is less opportunity for animals to June 1981a t Kuala Lumpur, Malaysia. Article 30. graze selectively. The herbage can be harvested pp. 182-199. |5! Pillai, K.R., Thiagarajan, S. & and then spread onto the soil and ploughed in as Samuel, C, 1985. Weed control by sheep grazing a green manure crop, but it is better to leave the under plantation tree crop. In: Sivarajasingam, S. sward untouched and to allow leaf fall and decom­ et al. (Editors):Proceeding s ofth e 9th Annual Con­ position. Leaf fall can be as much as 7 t/ha of DM ference of the Malaysian Society of Animal Pro­ per year. duction, March 11-12, 1985. Serdang, Selangor, No major diseases or pests have been reported on Malaysia, pp. 43-52. j6! Skerman, P.J., Cameron, this species in South-East Asia. D.G. &Riveros , F., 1988.Tropica l forage legumes. It is one of the most productive herbaceous FAO, Rome. p. 223. j7| Wong, C.C., 1990. Mineral legumes, with DM yields of 10 t/ha in the year of composition and nutritive value of tropical for­ establishment and up to 15t/h a in following years. ages as affected by shade. MARDI Research Jour­ Under low light intensities of 6-16% of full sun­ nal 18:135-143. light under an oil palm canopy, C. caeruleum C.P. Chen &Y.K.Chee outyielded other herbaceous legumes such as Sty- losanthes guianensis (Aublet) Swartz, Macropti- lium atropurpureum (DC.) Urban, Centrosema Calopogonium mucunoides Desv. pubescens Benth., Desmodium heterocarpon (L.) DC. ssp. ovalifolium (Prain) Ohashi, Desmodium Ann. Sei. Nat. Sér. 1,9 :42 3(1826) . heterophyllum (Willd.) DC. and Calopogonium LEGUMINOSAE mucunoides with DM yields of 1-1.5 t/ha. It also 2n = 36 had the best survival rate. No information is avail­ Vernacular names Calopo (En). Indonesia: able on its use as hay or silage, although it is pos­ kacang asu (Javanese), kalopogonium (Indone­ sible that such treatment could increase its pala- sian). Philippines: santing (Sulu), karaparapak tablity. sara naw (Mar.).Thailand : thua karopo. Genetic resources and breeding Limited col­ Origin and geographic distribution Calopo is lections of C. caeruleum are held by CIAT (Colom­ indigenous to tropical America and the West bia) and ATFGRC (CSIRO, Australia). There are Indies. It was introduced in the early 1900s to trop­ no breeding programmes on this species. ical Africa and Asia. Calopo was used as a green Prospects The potential of C. caeruleum as a for­ manure and cover crop in the central and eastern age is limited unless genotypes of better palatabi­ parts of Java from 1922 onwards. It was brought lity can be identified. The potential benefits of to Malaysia soon after that as a cover crop for rub­ using this species to improve soil fertility in farm ber. Itbecam e naturalized in Malaysia and Indone­ systems involving forage production could be sia, and has spread to most humid tropical areas explored further. It is likely to become a standard ofth e world. It was introduced to Australia in 1930, component of mixed cover crops in plantation but has not been widely used there. crops. Uses Calopo is used as forage for animals during Literature |1| Allen, O.N. & Allen, E.K., 1981. the latter part of the dry season, and is an impor­ The Leguminosae: a source book of characteris­ tant cover crop for plantations and as green ma­ tics, uses and nodulation. Macmillan Publishers nure for soil improvement. It is well recognized as Ltd., London, United Kingdom, and The Universi­ being a valuable pioneer legume to protect the soil ty of Wisconsin Press, Madison, Wisconsin, Uni­ surface, reduce soil temperature, fix nitrogen and ted States, p. 127.|2 |Chen , C.P. &Othman , O.,1984 . improve soil fertility. CALOPOGONIUM 73

Properties Considering that calopo is widely emarginate standard, about 1c m long. Pod linear- used as a green manure crop, there are few pub­ oblongoid, 2-4 cm x 3.5-5 mm, straight or curved, lished data on its chemical composition. Nitrogen softly pilose with coarse reddish-brown hairs, percentages of 2.6-3.8% have been recorded, but impressed between the seeds. Seeds 3-8, compres­ lower values can be anticipated in older, stemmy sed squarish, 2-3 mm long, yellowish or reddish- material. Calopo forage is not very palatable to brown. cattle, but animals are forced to eat it during the Growth and development Calopo grows rapid­ dry season when little green feed is available. Its ly, able to form a dense entangled sward in 4-5 low palatibility is usually ascribed to the abun­ months after sowing, but the plants are short-lived dance of hairs on the stems and leaves. There are and may only persist for 1-2 years. When grown 70-75 seeds/g. in a mixture with puero (Pueraria phaseoloides Description A vigorous, creeping, twining or (Roxb.) Benth.) and centro (Centrosema pubescens trailing herb, up to several m long, forming a tan­ Benth.), calopo is the first one to become estab­ gled mass of foliage 30-50 cm deep, with densely lished but also the first one to be shaded out. Long- pilose stems with long spreading ferruginous term persistence is through recruitment of new hairs. Leaves trifoliolate, petiole up to 16 cm long, plants from seedlings. Flowering in calopo is pilose; leaflets elliptic, ovate or rhomboid-ovate, initiated by short days. It is self-pollinated and (1.5-)4-10(-15)cm x (l-)2-5(-9) cm, the laterals seeds freely. oblique, adpressed pilose or pubescent onbot h sur­ Other botanical information The name 'Tor­ faces. Inflorescence a slender raceme, up to 20 cm tilla' is used to indicate seed of calopo sometimes long, peduncle 0-17 cm long, ferruginous pilose; harvested from naturalized stands in the Adelaide flowers in fascicles of 2-6, blue or purple; calyx River area of the Northern Territory (Australia). campanulate, unequally 5-lobed; corolla with It was at one time thought to have been a long-term locally adapted ecotype, but it is now believed to have come to the area as a contaminant in puero seed from Queensland which had been sown in the late 1960s at the Tortilla Flats Research Farm. 'Tortilla' is likely to be similar to Queensland com­ mercial material, which is rarely harvested and has never been allotted a cultivar name. Ecology Calopo is suited to the hot humid trop­ ics with an annual rainfall exceeding 1250 mm. It grows at altitudes from sea-level to 2000 m but is more suited to altitudes from 300-1500 m. It is moderately drought tolerant but may die out if the dry season is prolonged. Vigorous growth occurs on soils of all textures, even with a low pH(H20) range of4.5-5 . As a result ofit s self-seeding nature and twining growth habit, calopo is well adapted to a range of ecological conditions. It can be used in a mixture of species provided it does not become too dominant. Calopo is poorly adapted to shade, with top growth, root growth and nodulation declining markedly with decreasing light intensities. This may be attributed to the 'non-plasticity' of leaves under shade as compared with other shade-toler­ ant plants such as Desmodium heterocarpon (L.) DC. ssp. heterocarpon var. ovalifolium (Wallich ex Prain) Rugayah, Centrosema pubescens Benth., and Calopogonium caeruleum (Benth.) Sauv. Under low light intensitites (< 20%), calopo Calopogonium mucunoides Desv. - 1, flowering leaves are reduced in size by 70% compared with branch; 2,fruiting branch. leaves in full sunlight. In contrast, centro and C. 74 FORAGES caeruleum leaves are reduced by only 10-25 % and Harvesting Whether grazed or cut and fed, calopo leaves of var. ovalifolium are 20% larger under is often refused by cattle although they eat it less such a low light intensity. reluctantly during the dry season. It is usually cut Propagation and planting Calopo is normally byhan d and is seldom conserved as hay or silage. propagated by seed. Seed is sown at 1-3 kg/ha, Yield When pods are mature, peak DM yields of usually drilled inrow swhe n sown into new planta­ up to 14t/h a can be obtained in a single cut. Lower tions or broadcast in stands to be used for forage yields of 4-6 t/ha per year are obtained when production. After seed is broadcast, the seed-bed calopo is cut every 9-12 weeks. Seed yields of should be rolled to improve establishment. Newly 200-300 kg/ha have been recorded. harvested seed usually has a high percentage of Genetic resources Collections of calopo are hard seed (> 75%). Consequently, mechanical held at CIAT (Colombia) and ATFGRC (CSIRO, scarification, soaking in concentrated sulphuric Australia). acid for 30minutes , or soaking in hot water (75°C ) Breeding There are no breeding programmes on for 3 minutes is recommended. Although calopo calopo. stems root at the nodes when in contact with moist Prospects Low palatability is perhaps the main soil, there is usually poor establishment of stem reason why interest in this species as a forage cuttings placed directly into soil. Seeds are usually plant has faded during the last decade. However, not inoculated, as this species nodulates promis­ this low acceptability may open up opportunities cuously with native rhizobia. If inoculum is for incorporating calopo into forage systems as a applied then cowpea strains such asth e Australian way of improving soil fertility and the growth rate CB 756 are used. When planted as a cover crop in and quality of pastures. plantations, it is usually sown in a mixture with Literature |1| Bogdan, A.V., 1977. Tropical pas­ other legumes such as Centrosemapubescens, Calo- ture and fodder plants. Longman, London, pp. pogonium caeruleum and Pueraria phaseoloides 328-329. !2| Bunting, B. & Milsum, J.N., 1928. with 1-3 kg/ha of calopo in a total mixture of Cover crops and green manures. The Malayan Ag­ 12-15 kg/ha of legume seed. When sown for forage ricultural Journal 16:256-280 . |3| Duke, J.A.,1981 . production, calopo has been successfully used in Handbook of legumes of world economic impor­ mixtures with stoloniferous grasses, such as mo­ tance. Plenum Press, New York. pp. 37-39. |4| lasses grass ( minutiflora Beauv.) and Humphreys, L.R., 1980. A guide to better pastures Rhodes grass (Chloris gayana Kunth), and with for the tropics and sub-tropics. 4th ed. Wright, Ste­ tussock grasses such as setaria (Setaria sphacelata phenson & Co., Silverwater, Australia, p. 52. |5| (Schumacher) Stapf & Hubbard ex M.B. Moss). Skerman, P.J., Cameron, D.G. &Riveros , F., 1988. Good results have been obtained from oversowing Tropical forage legumes. FAO, Rome. pp. 224-228. it into existing stands of pangola grass (Digitaria |6| Wong, C.C., 1990. Mineral composition and eriantha Steud.) which have been harrowed. nutritive value of tropical forage legumes as Husbandry Because of the vigorous growth of affected by shade. MARDI Research Journal 18: calopo, a large amount of leaf litter falls onto the 135-143. |7|Yate s Seeds, 1987. Better pastures for soil. Fertilization with ground dolomite and Mo in the tropics. 2nd ed. Yates Seeds, Toowoomba, Aus­ acidic soils results in higher yields. Use of pre- and tralia, pp. 40-41. |8| Yost, R. & Evans, D., 1986. post-emergence herbicides reduces weed infesta­ Green manure and legume covers in the tropics. tion and results in faster establishment of calopo. Research Series 055. College of Tropical Agricul­ The effect of calopo and associated legumes in ture and Human Resources, University of Hawaii, improving soil fertility may last for 14-16 years. United States, p.14 . If calopo is grazed, it is advisable to use rotational C.P. Chen&A.Aminah grazing with rest periods of from 8-12 weeks when calopo growth is erect rather than prostrate. With calopo as cover in young oil palm and rubber plan­ Canavalia ensiformis (L.) DC. tations, regular slashing is needed to prevent the cover from overgrowing the trees. Prodr. 2:40 4(1825) . Diseases and pests Calopo is susceptible to vir­ LEGUMINOSAE uses in Guatemala, Costa Rica and Panama. Al­ 2re = 22 though leaf-eating caterpillars and beetles have Synonyms Dolichos ensiformis L. (1753), Cana­ been observed on calopo in Malaysia, they have valia gladiata (Jacq.) DC. var. ensiformis (L.) not been a serious problem. Benth. (1859-1862). CANAVALIA 75

"Vernacular names Jack bean, horse bean (En). Fève Jacques (Fr). Indonesia: kacang parang (Malay), kara bedog, kacang mekah (Sundanese). Malaysia: kacang parang putih. Philippines: habas (Tagalog), lagaylay (Visaya), badang- badang (Ilokano). Cambodia: tiehs. Laos: thwâx fak ph'aaz. Thailand: thua khaek. Vietnam: (cây) dâu ra, dâu tây, dâu ngua. Origin and geographic distribution Jack bean originates from South and Central America. It is widely cultivated in the southern United States since prehistoric times, and has been discovered in archaeological sites in Mexico dated 3000 BC. Now it is commonly cultivated throughout the tropics. Uses Jack bean is a forage for ruminants and is grown extensively as a cover crop or green manure crop in rotation with a wide range of crops. As a forage it is usually sun-dried before feeding to cat­ tle, but may also be made into silage. Heat-treated ripe seed, when ground, is used as a concentrate in cattle and poultry rations. Half-ripe seeds are mixed with sorghum for feeding cattle in Hawaii. Young seeds and pods may be used in human nutri­ tion as a vegetable but only when cooked. Soaked and roasted ripe seeds are eaten as a delicacy in Indonesia, but outside Japan and tropical Asia, the Canavalia ensiformis (L.) DC. - 1, branch with species is now little grown as a food crop for leaves; 2,flowering and fruiting branch; 3,fruit; 4, humans. The seed issometime s roasted and ground seeds. to make a coffee substitute. In Indonesia flowers and young leaves are steamed and used as a fla­ and some secondary branching also occurs. Leaves vouring. alternate, 3-foliolate; petiole 11-17 cmlong , rachis Urease extracted from the seed is used in analyti­ 3-4.5 cm; leaflets ovate-elliptic, 5-20 cm x 3-12 cal laboratories. In Indonesia and China, heat- cm, acute or rounded and mucronate at top, spar­ treated seeds and pods are used as medicine, and sely covered with short hairs on both surfaces, it has been suggested that some of the chemical venation raised and reticulate. Flowers mauve to constituents ofjac k bean might be used for medici­ purple, or sometimes white, borne on an axillary nal purposes or for control of pests. raceme with swollen nodes each bearing 1-3 Properties The forage is not especially palat­ flowers on pedicels 2-5 mm long; peduncle 10-35 able,bu t cattle can acquire a taste for it;dr y forage cm, raceme up to 20 cm long; calyx campanulate, is more palatable than fresh. Nitrogen concentra­ 5-lobed, sparsely pubescent; corolla with rounded tions of the forage and seed are 2.2-2.6% and standard, 2.7 cm long, rose to purplish. Pod oblon- 3.8-5.7% respectively. Non-ruminants fed on the goid, laterally compressed, 15-35 cm x 3-3.5 cm, seed show reduced weight gains, due mainly to the containing 8-20 seeds, each valve with a suturai toxic amino-acid canavanine, which is an antime­ rib and an additional rib just below it. Seed oblon- tabolite of arginine. Lectins (concanavalin A and goid, laterally compressed, ca. 21 mm x 15 mm x B) are present in the seed and can inhibit the ab­ 10 mm, ivory or white, the hilum brown, 6-9 mm sorption of nutrients. Heat treatment of the seed long. overcomes the toxic effects. Seed weight ranges Growth and development Germination is epi- from 0.85-1.65 g/seed. geal; the juvenile leaves are fully expanded about Description Annual or short-lived perennial a week after sowing. Hard-seededness has not been climber with a deep root system, bushy, twining recorded. Early growth is slow, but after establish­ or prostrate, up to 2-3 m long. Stems become some­ ment the species is capable of rapid growth. Jack what woody with age; branching is at lower nodes bean flowers 50-110 days after sowing, depending 76 FORAGES on the accession and on climate and soil condi­ 50 kg/ha in rows 0.6-1 m apart. With accurate tions. The time from sowing to seed harvest is nor­ placement, sowing rates of 10 kg/ha will suffice. mally about 170 days. In temperate regions it Sowing at the start of the wet season is preferred flowers in summer or autumn; in the tropics it in regions with about 1000m m annual rainfall, but flowers throughout the year. In temperate areas in more humid environments sowing should be beans remain unaffected when light frost kills the delayed until the end of the wet season. ' foliage. As a pulse, jack bean may be grown with other Other botanical information In the literature crops such as maize or cassava. Although the grain on Canavalia Adans., confusion exists on the iden­ yield of jack bean is then likely to be reduced, tity of 3closel y related species: C.ensiformis (jack sometimes by asmuc h as 50% ,th e total productivi­ bean), C.gladiata (Jacq.) DC. (sword bean) and C. ty of the system is improved. virosa (Roxb.) Wight & Arnott. C. ensiformis and Husbandry Inoculation is normally unneces­ C. gladiata are cultivated taxa, possibly derived sary as jack bean nodulates promiscuously and from the wild (occasionally cutivated) C. virosa. effectively with rhizobia present in most soils. These 3tax a are considered as one species by some, Twining forms grown for grain are sometimes and by others they are kept separate. A thorough grown on trellises since the improvement in light taxonomie revision is needed to bring clarifica­ utilization improves pod yield and the pods are tion. Most recent floras separate the 3tax a as fol­ kept off the ground. lows: Diseases and pests Jack bean is little affected - C. ensiformis: standard ca. 2.5 cm long, rose to by diseases and pests. A fungal root disease and purple; pod up to 35 cm x 3.5 cm; seed ivory or a stem-borer sometimes cause serious losses. Plati- white, hilum less than half as long as the seed. prosopus acutangulus, a leaf-eating chrysomelid - C.gladiata: standard ca. 3.5 cm long, white; pod beetle, may cause serious problems during the up to 40c m x 5cm ; seed red or red-brown, rarely early growth ofth e crop. Stored seed is quite resis­ white, hilum more than half as long as the seed. tant to pests and diseases, but can become infested - C. virosa: standard ca. 3 cm long, whitish veined with Tricorinus tabaci. mauve; pod up to 17 cm x 3 cm; seed brown or Harvesting Forage and seeds are mostly har­ red-brown, marbled with black, hilum ca. half vested by hand. Although flowering is fairly well the length of the seed. synchronized, some green pods may still be present There are several cultivars and accessions of jack at harvest time, especially if water is still avail­ bean which differ in their growth habit. They are able. being tested and produced in Venezuela (Universi- Yield Forage DM yields of up to 23 t/ha have dad de Venezuela, Facultad de Agronomia, Mara- been obtained in Hawaii and green fodder yields cay). The bushy types are early-flowering, better may exceed 50t/ha . Grain yields average 800-1000 adapted to low-rainfall areas and have low capaci­ kg/ha (range: 400-1500 kg/ha, depending on rain­ ty for forage production. The viny or twining types fall distribution) but in experimental plots yields are late-flowering but are better adapted to more as high as 6000kg/h a have been recorded in highly humid areas and produce greater amounts of for­ intensive agriculture. age. Fresh forage is not palatable to ruminants and is Ecology Jack bean is well adapted to the humid eaten only in small amounts, although cattle grad­ tropics, yet is hardy and can withstand periods of ually become more accustomed to it. Conse­ drought. Consequently, it grows in regions with quently, it is usually fed after drying as this annual rainfall ranging from 700-4000 mm. Al­ increases intake. Ruminants can consume small though jack bean is a lowland plant, it is also quantities of untreated grain or meal without ill grown at altitudes up to 1800 m. It grows best in effects, but larger quantities can be toxic. Affected full sunlight but has moderate shade tolerance. It animals have a clear nasal discharge, become lame istoleran t ofa widerang e ofsoi l conditions includ­ and cannot rise. Mucous membranes become ing acid soils and highly-leached infertile tropical muddy in appearance and clear urine is passed soils. It is less affected by waterlogging or salinity more frequently than usual. Heat-treatment over­ than other pulses. comes this toxicity. Meal prepared from jack bean Propagation and planting Jack bean is propa­ seed ismor e palatable to cattle ifmolasse s is added gated by seed in a range of densities from 30 cm to it. For non-ruminants, extensive boiling with x 30 cm to 100 cm x 150 cm at shallow depth. For one or two changes of water and peeling off the green manure crops, it is sown at a rate of about seed-coat is required before the mature seeds are CBNCHRUS 77 edible.Th e beans may also beprocesse d to produce itan (Tagalog), sagisi (Ilokano). Thailand: ya-bup- a protein concentrate for use in formulated foods. fen. Genetic resources There are very limited gene­ Origin and geographic distribution Bufful- tic resources available of this species. Germplasm grass naturally occurs throughout Africa and from collections are available at the Faculty de Agrono- Arabia and the to India. It has been mia of the Universidad de Venezuela (Maracay, widely introduced and is naturalized throughout Venezuela) and the USDA (Georgia, United the semi-arid and sub-humid tropics and subtrop- States). ics Breeding Although largely self-pollinating, Uses The main use of buffelgrass is as a pasture some 20% of flowers may be cross-pollinated by grass for ruminants and horses. It is usually insects. Some selection for lines with low concen­ grazed, but can be used as silage or hay. trations of canavanine has been carried out in Properties The N concentration of green buffel­ Venezuela. Also, some cultivars have been grass ranges from 1.0% to 3.0% of the dry matter, selected with climbing or dwarf attributes. depending on growth stage, soil fertility, fertiliza­ Prospects As a forage crop, prospects for jack tion rate and cultivar. Phosphorus concentration bean are limited since it is ofonl y mediocre accept­ is usually adequate for animal requirements and ability to ruminants, and has toxic properties if higher than that of most other tropical grasses, eaten when fresh. Furthermore, other annual legu­ with published values up to 0.65% of the DM. The minous forages are equally or more productive. As DMdigestibilit y ofgree n buffelgrass leaves ranges a leguminous crop with the residue providing use­ from 65-70% . Buffelgrass ishighl y palatable to all ful forage, the comparative freedom of jack bean kinds of grazing animals. from disease and pest problems and its high yield Horses eating this grass can develop a calcium in low input systems justifies the development of deficiency syndrome ('bighead disease') caused by jack bean as a sole crop or in intercropping sys­ soluble oxalate (concentrations of 1-2% of the tems as a vegetable or a pulse crop. DM have been recorded). Variation in the diet or Literature 111 Allen, O.N. & Allen, E.K., 1981. supplementation with limestone or dolomite will The Leguminosae: a source book of characteris­ control it. Ruminants are able to break down the tics, uses, and nodulation. Macmillan Publishers oxalate in the rumen, although occasionally toxi­ Ltd., London, United Kingdom, and The Universi­ city symptoms have been recorded with sheep and ty of Wisconsin Press, Madison, Wisconsin, Uni­ cows. ted States, p. 132. |2j Bogdan A.V., 1977. Tropical There are 90-1200 fascicles/g. A fascicle is a clus­ pasture and fodder plants. Longman, London, p. ter of spikelets which may contain one to five 330. |3! National Academy of Sciences, 1979. Trop­ caryopses, making up 25-30% of the fascicle ical legumes: resources for the future. Wash­ weight. ington, D.C.pp . 11,54-59, 305.|4 |Perry , L.M., 1980. Description A tufted or rarely spreading peren­ Medicinal plants of east and : attri­ nial, often with short rhizomes. Culm 10-150 cm buted properties and uses. MIT Press, Cambridge, tall, erect or ascending, branching and rooting at Massachusetts, United States, p. 209. |5|Sauer , J., the nodes. Leaf-sheath compressed; ligule a hairy 1964.Revisio n of Canavalia. Brittonia 16:142-144. ring, up to 1.5 mm tall; leaf-blade linear, 3-30 cm i6| Skerman, P.J., Cameron, D.G. & Riveros, F., x 2-13 mm, glabrous, but hairy at the mouth. 1988. Tropical forage legumes. FAO, Rome. pp. Inflorescence a cylindrical to ovoid panicle, 2-14 229-230. cm x 1-2.6 cm, grey, purple or straw-coloured, Y.K. Chee,J.B . Hacker, L. Ramirez &C.P . Chen bearing numerous clusters of spikelets (fascicles), each fascicle surrounded by an elongated invo­ lucre of bristles 6-16 mm long; rachis angular and Cenchrus ciliaris L. puberulous; inner bristles much exceeding the spikelets, somewhat flattened and connate at the Mant.pl. alt.: 302(1771) . base, ciliate below, one of them longer and stouter GRAMINEAE than the rest; outer bristles shorter, filiform; spike­ 2ra = 32,34 ,3 6 (tetraploid), 40,52 ,5 4 lets 1-4 per cluster, acutely lanceolate, 2-5.5 mm Synonyms Pennisetum cenchroides Rich. (1805), long, sessile; glumes distinct, acute, upper ones up P. ciliare (L.) Link (1827). to as long as the spikelet; lemmas subequal, 2.5-5 Vernacular names Buffelgrass (Australia) mm long, minutely awned. Caryopsis ovoid, 1.5-2 (En).Africa n foxtail (Am).Philippines : kawit-kaw- mm x 1 mm. 78 FORAGES

'Biloela', but with green leaves), 'Molopo' (tall growing with a lower tiller density, more rhizoma­ tous and distinctly glaucous leaves, more greyish), 'Lawes' (similar to 'Molopo'), 'Gayndah' (short, semi-prostrate to ascending, non-rhizomatous), 'American' (short, similar to 'Gayndah'), 'West Australian' (short, non-rhizomatous, least produc­ tive,highl y palatable, most drought tolerant). Cul­ tivars are very homogeneous, because buffelgrass is an apomict, although there isa small percentage of sexual seed production. 'Higgings grass' was de­ veloped from a single sexual plant in Texas. There are also some African cultivars. Ecology Buffelgrass grows best in climates with 300-750 mm annual summer-dominant rainfall. Nevertheless it also grows well in the Philippines on 1600-2900 mm annual rainfall. Optimum tem­ perature is about 35° C and minimum temperature between 5-16°C. Buffelgrass prefers free-drain­ ing, light-textured neutral to alkaline soils (pH(H20) > 6.0). 'American' is more tolerant to acid soils than the other cultivars. The more rhizo­ matous cultivars also grow well on many clay soils. Buffelgrass is sensitive to soils containing high levels of Al, has a moderate tolerance to salinity and is intolerant offlooding . It tolerates fire. Propagation and planting Seed dormancy can be broken by high temperatures (60°C) for 4-12 Cenchrus ciliaris L. - 1,habit; 2, ligule; 3, inflores­ weeks, by removal of the hulls of the fascicle by cence; 4,spikelet with involucre. hammermilling, or by storing the seed for up to 18 months. Treatment of the seed with sulphuric acid Growth and development Buffelgrass seed also improves germination. Seeds are not easy to germination rate can be as low as 15% soon after clean; hammermilling to pulverize stems and bris­ harvest, when the caryopses remain within the fas­ tles improves seed handling and sowing. The seed cicles, but over 90% germination has been mea­ can be sown in soil without or with light tillage, sured. Naked caryopses have a higher germination onth e surface followed byrolling , but germination rate, but seedlings are then more sensitive to is best when sown at a depth of 1-2 cm. Suggested drought. Glumes contain water-soluble germina­ seed rates vary from 0.5 kg/ha of caryopses to 12 tion inhibitors, which is the main cause ofth e seed kg/ha offascicles ; good results have been obtained dormancy. with 4kg/h a of fascicles. Germination rate improves with storage of 6-18 Sowing whole fascicles is preferred to sowing months. Seedling growth rate is relatively slow. naked caryopses. In India vegetative propagation Buffelgrass has a deep root system with coarse from tuft splits or replanting ofyoun g plants sown roots. Maximum rate of tiller formation occurs in a nursery has also been advocated. just after first inflorescence appearance; tillering Buffelgrass can be grown in association with Sty- is sustained at a reduced rate during seed matura­ losanthes hamata (L.) Taub., Macroptilium atro- tion. Buffelgrass is an early flowering short-day purpureum (DC.) Urban, Chloris gayana Kunth plant with an optimal photoperiod of 12 hours. and Panicum maximum Jacq. var. trichoglume Other botanical information Most cultivars Robijns. are developed in Australia: 'Biloela' (tall growing, Husbandry Buffelgrass grown as a sole crop is rhizomatous, greyish leaves), 'Nunbank' (similar usually not fertilized, although it responds well to to 'Biloela'), 'Boorarra' (similar to 'Biloela', but P and N fertilizer. less rhizomatous), 'Tarewinnebar' (similar to When grown in association with a legume, fertil- CENTOTHECA 79

ization with superphosphate is beneficial on poor Indonesia. Journal of Agricultural Science (Uni­ soils. Irrigation is not usual or necessary as the ted Kingdom) 111: 11-17. |6|Skerman , P.J. &Rive - grass is drought tolerant. ros, F., 1990. Tropical grasses. FAO, Rome. pp. Diseases and pests Buffelgrass is quite free of 266-274. \T\ 't Mannetje, L. & Jones, R.M., 1990. diseases or pests. In wetter areas the seed can be Pasture and animal productivity of buffelgrass destroyed by ergot or smut. with siratro, lucerne or nitrogen fertilizer. Trop­ Harvesting First harvest of buffelgrass can be ical Grasslands 24: 269-281. 4-6 months after sowing by cutting or grazing. L. 't Mannetje &S.M.M . Kersten Suggested cutting height is about 7 cm above ground level, with 6-8 week intervals. For hay, buffelgrass is usually cut in the early flowering Centotheca latifolia (Osbeck) Trinius stage. Buffelgrass can be grazed continuously or rotationally. Fund. Agrost.: 141(1820) . Yield Dry matter yields range from 2-9 t/ha per GRAMINEAE year without fertilizer and up to 24 t/ha per year 2n = 24 with complete fertilizer. Seed production depends Synonyms Holcus latifolius Osbeck (1757), on cultivar and growing conditions and recorded Cenchrus lappaceus L. (1763), Centotheca lappacea (fascicle) yields range from 150-500 kg/ha. Mean (L.) Desv. (1810). liveweighf gains of steers of 160kg/annu m grazing Vernacular names Indonesia: jukut kidang buffelgrass pastures with Macroptilium purpu­ (Sundanese), suket lorodan (Javanese), karetet reum at 1 steer/ha have been recorded in a sub- lempad (Alfuru). Malaysia: rumput lilit kain, rum- humid subtropical climate in south-east Queens­ put darah, rumput temaga. Papua New Guinea: land. kuang. Philippines: baylu-patong (Tagalog), andu- Genetic resources A large collection of buffel­ dukot aridekdiket (Bikol). Thailand: ya khon moi grass germplasm is held at ATFGRC (CSIRO, Aus­ maemai, ya-enieo. Vietnam: co' móc. tralia). Origin and geographic distribution C. latifo­ Breeding The apomixis of buffelgrass is linked lia is widely distributed in the tropics of the Old with pseudogamy. Hybridization is possible World, from Africa to and Australia. It because of the discovery of a rare fertile female. is abundant in southern Thailand and in Peninsu­ Male pollen from the apomictic lines are used to lar Malaysia. fertilize the rare fertile female with consequent Uses C. latifolia is grazed by village cattle and release of much variation. A selection programme regarded as a good fodder. is in progress at CSIRO, Brisbane, Australia with Botany A perennial rosette grass with erect the objectives of developing cultivars of better culms up to 1.25 m tall. Leaf-sheath closely striate, quality forage and better spring vigour. 4-6 cm long; leaf-blade elliptical or ovate-oblong, Prospects Buffelgrass has little prospect in 4-30 cm x 0.8-3.5 cm, hairy to glabrescent; ligule humid climates, but can play a major role in forage membraneous, 2-3 mm tall. Inflorescence a production in areas with a dry season in South- branching terminal panicle, up to 40c m long; spik- East Asia. elets 2-3-flowered, about 8 mm long; lemma of Literature |1|Bogdan , A.V., 1977. Tropical pas­ upper floret(s) ca. 3m m long, 5-nerved with 2prom ­ ture and fodder plants (grasses and legumes). inent rows of bulbous-based retrorse spiny hairs Longman, London and New York. pp. 66-74. |2| on either side of the back. Caryopsis about 1 mm Clayton, W.D. &Renvoize , S.A., 1982. Gramineae long, dark brown, glossy. (Part 3).In :Polhill , R.M. (Editor): Flora of tropical Adjacent branches from inflorescences become East Africa. A.A. Balkema, Rotterdam, pp. entangled at maturity because ofth e reflexed hairs 691-692. |3| Hacker, J.B., 1989. The potential for on the lemmas. These hairs also adhere to hair or buffelgrass renewal from seed in 16-year-old buffel- fur ofpassin g animals,thu s providing effective dis­ grass-siratro pastures in south-east Queensland. persal ofth e seeds. Journal of Applied Ecology 26: 213-222. |4| Some spikelets seem to be viviparous; the upper Humphreys, R.L., 1967. Buffelgrass in Australia. lemmas each grow into a small leaf-blade and when Tropical Grasslands 1: 123-134. |5|Rudolf , I.G.W., the whole spikelet eventually falls, it has the abili­ Blair, G., Orchard, P.W., Till, A.R. & Hunt, M., ty to grow into a new plant. 1988. The performance of Ongole heifers grazing Flowering occurs throughout the year. Often seed­ native and introduced pasture species at Sumba, lings can befoun d around older plants. 80 FORAGES

Malaya. In: Burkill, H.M. (Editor): Flora of Malaya. Vol. 3. Government Printing Office, Singapore, pp. 53-54. |3i Mehra, K.L. & Fachru- rozi, Z., 1985. Indonesian economic plant resources: forage crops. Lembaga Biologi Nasio- nal-LIPI, Bogor. No 31. p. 5. |4|Mono d de Froide- ville, Ch., 1971.A synopsis of Centotheca and re­ duction ofRamosia . Blumea 19: 57-60. C. Manidool

Centrosema acutifolium Benth.

Comm. legum. gen.: 54(1837) . LEGUMINOSAB 2n = 20? Origin and geographic distribution The ori­ gin of C. acutifolium is in tropical America, re­ stricted to 4-6°N in Colombia and Venezuela, and in central-west and south-east Brazil. A cultivar and several experimental lines have been spread for testing from South America to other tropical regions, including South-East Asia. Uses C. acutifolium is used as forage for rumi­ nants in grazed pastures or in cut-and-carry sys­ tems. Properties C.acutifolium is similar to centro (C. pubescens Benth.) in several regards, including nutritive value. Depending on plant age, N con­ Centhotheca latifolia (Osbeck) Trinius - 1, flower­ centrations in leaves range from 3.5-5.0%, and in ing plant; 2, spikelet. vitro DM digestibility from 45-75%. Phosphorus concentrations in leaves range from 0.14-0.27%), Ecology C. latifolia grows from sea-level to 1500 and Ca from 0.38-1.13 %. There are 15-20 seeds/g. m altitude in humid shady places receiving over Description A perennial, trailing-twining 2000m m ofrai n per year. The species is often found herb; stems slender, pubescent, with tendency to on disturbed sites along shaded roads, fields and root at nodes. Leaves trifoliolate, young leaflets plantations but is natural in swamps and open distinctly purplish; stipules deltoid-acuminate, places in rain forests and thickets. pubescent; petioles and petiolules pubescent, Agronomy C. latifolia is propagated vegetati- reddish at their base; leaflets ovate to ovate-lan­ vely or by seed. It is moderately palatable and is ceolate, apically acuminate, membranaceous, usually eaten by grazing stock but can be cut and puberulous to subglabrous on both surfaces; cen­ fed to animals. Animals avoid eating the seed tral leaflet symmetrical, 5-8.5 cm x 3-3.5 cm; lat­ heads. Kegrowth is very slow so frequent defolia­ eral leaflets asymmetrical, 4-7.5 cm x 2.5-4 cm. tion is not recommended. Inflorescence an axillary raceme with up to 24 Genetic resources and breeding It is unlikely flowers inserted by pairs along the rachis; pedun­ that substantial germplasm collections of C. latifo­ cle conspicuously long, up to 24 cm, pubescent; lia are being maintained. flower papilionaceous, subtended by a pair of Prospects C. latifolia is shade tolerant and short, ovate-acuminate bracteoles; calyx campa- should be evaluated for its suitability in agrofor- nulate, 5-toothed, pilose, carinal and lateral teeth estry systems. short; petals light violet; standard orbicular Literature |1| Bor, N.L., 1960. The grasses of emarginate, 28-35 mm x 32-40 mm, pubescent Burma, Ceylon, India and Pakistan. Pergamon outside. Pod linear, straight to slightly bent, up Press, London, pp. 457-459. |2| Gilliland, H.B., to 20 cm long, beaked, scabrid, containing 10-15 Holttum, K.E. & Bor, N.L., 1971. Grasses of seeds, dehiscent. Seed cylindrical, 5-7 mm x ca. CENTEOSEMA 81

Ecology C. acutifolium is best adapted to sub- humid, tropical environments with 1000-2500 mm rainfall per year and a distinct dry season. It toler­ ates up to 5 dry months. Var. orinocense requires well-drained, light-textured soils, whereas var. matogrossense grows particularly well on heavier soils, including those with seasonal drainage prob­ lems. C.acutifolium tolerates soil acidity and toxic levels of Al and Mn very well; its nutrient require­ ments are low; its shade tolerance is moderate. Propagation and planting C. acutifolium seed is drilled in rows or broadcast, alone or simultane­ ously with a grass, or strip-sown into an existing grass-sward at a rate of 3-4 kg/ha. Mechanical or acid-scarification of seed is necessary in order to overcome hard-seededness. Although it can nodu- late from native cowpea rhizobia, inoculation with a Bradyrhizobium strain of known effectiveness is recommended. Grasses suitable for association with C. acutifolium are bunch grasses such as gui­ nea grass (Panicum maximum Jacq.) on fertile soils, or gamba grass (Andropogon gayanus Kunth) on infertile soils. However, successful as­ sociations have also been obtained with the stolo- Centrosema acutifolium Benth. - flowering and niferous Brachiaria dictyoneura (Fig. $c De Not.) fruiting branch. Stapf. Husbandry Fertilization with P and K enhances 3mm , greenish-yellow with dark, fine mottles. establishment of C. acutifolium on most tropical Growth and development C. acutifolium soils. Despite the legume's adaptation to soils of flowers late in the season, and this can reduce seed medium to low fertility, it responds well to mainte­ production in years with an early onset of the dry nance fertilization with these nutrients. Besides season. Seed yields are, therefore, often low, but maintenance fertilization, appropriate grazing under adequate soil-moisture conditions, seed pro­ management is needed to maintain an adequate duction potential is high. legume proportion in a C.acutifolium] grass associ­ Other botanical information C.acutifolium has ation. Grazing must allow sufficient seedling re­ only recently been rediscovered. Germplasm now cruitment and/or rooting from stolons to maintain known to be C.acutifolium was, until recently, fre­ plant density. quently referred to as Centrosema sp., Centrosema Diseases and pests C. acutifolium shows good sp. nov., or Centrosema new sp. No 2.Ther e are two tolerance, mainly in sub-humid environments, to morphologically and physiologically distinct forms the major Centrosema diseases. It is, however, of C. acutifolium related to their geographical ori­ more susceptible to Pseudomonas bacterial blight gin:(1 )var . orinocense nom. nud. from a small distri­ than other Centrosema species.Leaf-eatin g insects bution niche in Colombia and Venezuela, and (2) can be a problem during dry periods. var. matogrossense nom. nud. from central-west Harvesting The legume is usually harvested by Brazil. The species description given above refers grazing but can be cut and fed to animals. Cutting to var. orinocense; it is represented by cultivar intervals will depend on soilmoistur e and fertility; 'Vichada' recently released in Colombia. a 10-14 week cutting interval and a 10-15 cm cut­ C. acutifolium is closely related to C. pubescens ting height are suggested. Benth. and the species are morphologically very Yield Wet-season DM yields on acid soils of low similar. The main characteristics that distinguish fertility range mostly between 1-3 t/ha per 12 C.acutifolium are: (1)th e purplish colour of young weeks; on soils of somewhat better fertility they leaflets, (2) the short bracteoles and calyx teeth, can be as high as 5t/ha . Dry-season DM yields sel­ (3) the scabrid pod indumentum, and (4) the cylin­ dom reach 1t/h a per 12weeks . Year-round animal drical seeds. production can be improved considerably if it is 82 FORAGES

present in a pasture. For example, in association grazed in pure legume stands ('protein bank') or with Andropogon gayanus, 180 kg liveweight gain in a mixture with grass. It is suitable for cut-and- per steer per year has been measured as compared carry systems and can also beuse d as ground cover with 110 kg/steer per year from A. gayanus alone. in plantation agriculture. The seed production potential is high; up to 700 Properties C. macrocarpum provides a palat­ kg/ha can be obtained. able, high-quality forage. Depending on plant age Genetic resources A limited number of acces­ and soil fertility, N concentration in leaves ranges sions of both C. acutifolium forms is available at from 3.5-5.0% and in vitro DM digestibility from CIAT (Colombia). They appear to adequately rep­ 45-70 %;P concentrations are about 0.20% . There resent its natural variability. are 15-25 seeds/g. Breeding Some exploratory cross-compatibility Description Arobus t perennial, taprooted vine, studies have been conducted with C. acutifolium prostrate in absence of a support; trailing stems and closely related species such as C. pubescens with variable tendency to root at nodes. Stem and C.macrocarpum Benth. However, there is cur­ pilose with greyish hairs when young, glabrescent, rently no programme to improve C. acutifolium lignified at base.Leave s trifoliolate; stipules trian­ through breeding. gular, petioles and petiolules pubescent; leaflets Prospects The main value of C.acutifolium lies broadly to narrowly ovate, apically acute to acu­ inth e fact that it combines the high nutritive value minate, rounded or slightly wedge-shaped at the and drought tolerance of the closely related C. base; central leaflet larger and longer petiolated pubescens with adaptation to acid, low-fertility than laterals, mostly 8-13 cm x 3-8 cm, papyra­ soils and good disease tolerance. For cut-and-carry ceous to subcoriaceous, almost glabrous to pubes­ systems, however, it is inferior to C. macrocarpum cent on lower or both surfaces; frequently with a because of lower DM production. light-green marking along midrib. Inflorescence Literature |1| Instituto Colombiano Agrope- an axillary raceme with up to 30 flowers inserted cuario, 1987. Centrosema 'Vichada'. ICA, Bogota, Colombia. Boletin Técnico No 152. 13 pp. |2| Schultze-Kraft, R.,Benavides , G. &Arias , A., 1987. Recolección de germoplasmay evaluación prelimi- nar de Centrosema acutifolium [Recollection of germplasm and preliminary evaluation of Centro­ sema acutifolium]. Pasturas Tropicales - Boletin 9: 12-20. |3| Schultze-Kraft, R. & Clements, R.J. (Editors), 1990. Centrosema: biology, agronomy, and utilization. CIAT, Cali, Colombia. 667pp . R. Schultze-Kraft

Centrosema macrocarpum Benth.

Ann. Nat. Hist. (Paris) 3:43 6(1839) . LEGUMINOSAE In = 22,20 Synonyms Centrosema lisboae Ducke (1922). Origin and geographic distribution C. macro­ carpum is of tropical American origin, and occurs naturally between latitudes 20CS and 20°N; it is particularly frequent north of the equator where it extends from the extreme north of Brazil to Venezuela, Colombia and all Central American countries north to Mexico. Experimental lines have spread to many tropical countries for testing, including in South-East Asia. Uses The main use of C. macrocarpum is as a for­ Centrosema macrocarpum Benth. ousering and age for ruminants. In pasture systems it can be fruiting branch. CBNTROSEMA 83 in pairs along rachis; flower papilionaceous, sub­ grass (Andropogon gayanus Kunth) and guinea tended by a pair of ovate-lanceolate-falcate brae- grass (Panicum maximum Jacq.); however, suc­ teoles; calyx campanulate, 5-teethed with carinal cessful associations have also been obtained with tooth considerably longer than others; petals the stoloniferous Brachiaria dictyoneura (Fig. & showy and cream-coloured with purple centre; De Not.) Stapf. standard orbicular-emarginate, 3-6 cm in diame­ Husbandry Fertilization with P and K enhances ter, pubescent outside; wings and keel much establishment of C. macrocarpum on most tropical smaller than standard, directed upwards. Pod soils. Despite the legume's adaptation to modera­ linear, compressed, up to 30 cm x 1 cm, straight tely fertile to infertile soils, it responds well to to slightly bent and beaked, subglabrous, contain­ maintenance fertilization with these nutrients. ing up to 25 seeds, dehiscent. Seeds transversely When used in grazed pastures, legume persistence oblong to rectangular, on average 5 mm x 3 mm, will be enhanced if C. macrocarpum is allowed to yellowish-brownish unicoloured, mottled or mar­ establish well before it is first grazed and if stoc­ bled. king rates are not excessive. Growth and development C. macrocarpum is Diseases and pests C. macrocarpum is one of extremely photoperiod-sensitive. Even as close as the Centrosema species most tolerant of diseases 3° to the equator, flowering is induced by shorten­ and pests. None ofth e economically important dis­ ing of daylength. Further conditions which pro­ eases of the genus (Rhizoctonia foliar blight, ant- mote flowering are the removal of accumulated hracnose, Cercospora leaf-spot, and bacterial biomass and the provision of support for plants to blight) has been observed to affect C. macrocarpum climb up. seriously. Leaf-eating insects can be a problem Tripping of flowers, usually by large insects such during dry periods. as bumblebees, is required for seed-setting. The Harvesting The legume is usually grazed or is legume has specific Bradyrhizobium strain requir­ consumed as fresh material after mowing. When ements. mown, cutting intervals will depend on soil mois­ Other botanical information There are sever­ ture and fertility; 10-14 weeks is suggested as an al groups ofdistinc t C.macrocarpum forms related adequate interval with a cutting height of 10-15 to their geographical origin. Of these, the low-alti­ cm. tude ecotypes from northern South America (Col­ Yield Depending on soil-moisture conditions, ombia, Venezuela) are particularly promising as DM yields on acid, moderately fertile to infertile forage plants. As yet, no cultivar has been soils range from less than 1 t to almost 5 t/ha per released. C. macrocarpum has occasionally been 12 weeks; annual DM yields in Colombia, at a site confounded with C. grandiflorum Benth. with bimodal rainfall distribution and a highly Al- Ecology C. macrocarpum is best adapted to the saturated ultisol, averaged 15 t/ha. Under opti­ humid and sub-humid tropics with annual rainfall mum conditions seed production reaches 800 kg/ above 1000 mm. Once established, it is very ha. drought-tolerant and can remain green during dry Genetic resources and breeding Evidence of seasons as long as 3-4 months. It tolerates moder­ considerable outcrossing has been observed in C. ate shade. The legume grows well on a range of macrocarpum. This is a consequence of the trip­ soils, provided they are well drained, but prefera­ ping of flowers by bumblebees which also transfer bly onmedium-texture d soils.I t has good tolerance pollen. Because of the large degree of outcrossing, of soil acidity including Al and Mn toxicity, and plant populations within a given ecotype can show oflo w available P. considerable variation. The species is well repre­ Propagation and planting C. macrocarpum sented in the collection held by CIAT (Colombia) seed is drilled in rows or broadcast, alone or simul­ where a large number of quite variable accessions taneously with a grass, or strip-sown into an exist­ are available. A breeding project aimed at intro­ ing grass sward, at a rate of 3-5 kg/ha. In order ducing the acid-soil tolerance of C. macrocarpum to break hardseededness, mechanical or acid-scar­ into C. pubescens Benth. is presently being con­ ification ofsee d isnecessary . Asi t has specific Bra­ cluded in Brazil. dyrhizobium requirements, seed must be inocu­ Prospects Because of its adaptation to acid, lated with an appropriate strain if the legume is infertile soils and drought, and its high productivi­ to be established where it has not been sown ty and nutritive value and good disease tolerance, before. Grasses suitable for association with C. C. macrocarpum isa valuable legume for the humid macrocarpum are bunch grasses such as gamba and sub-humid tropics. Its potential seems to be 84 FORAGES

greater in cut-and-carry systems and for protein banks than as a component in a grass-legume pas­ ture, where it is sensitive to grazing mismanage­ ment. Literature |1| Keiler-Grein, G. (Editor), 1990. Red Internacional de Evaluación de Pastos Tropi­ cales, RIEPT-Amazonia. I Reunión-Lima, Peru, 6-9 Noviembre 1990.Document o deTrabaj o No75 . CIAT, Cali, Columbia. 2Vols . 1119pp . |2| Schultze- Kraft, R., 1986. Natural distribution and germ- plasm collection of the tropical pasture legume Centrosema macrocarpum Benth. Angewandte Botanik 60:407-419 . |3| Schultze-Kraft, R. & Clem­ ents, R.J. (Editors), 1990. Centrosema: biology, agronomy, and utilization. CIAT, Cali, Colombia. 667pp . R. Schultze-Kraft

Centrosema pascuorum Martius ex Benth.

Comm. legum. gen.: 56(1837) . LBGUMINOSAB 2n = 22 Centrosema pascuorum Martius ex Benth. -1, flow­ Vernacular names Centurion (Australia) (En). ering branch; 2,calyx; 3,flower; 4, fruits. Origin and geographic distribution The ori­ gin and natural distribution of this species is in cal, glabrous to scarcely pilose, branched at the tropical South and Central America, mainly in nodes, up to 2 m long. Leaf trifoliolate and often semi-arid regions in north-eastern Brazil, Vene­ held erect; stipules narrowly triangular, 4-9 mm zuela, Guyana, Ecuador, Panama, Costa Rica, long; petiole 1.7-5 cm long including the upper , Guatemala and southern Mexico. C. rachis; leaflets long and narrow, 2-15 cm x pascuorum also occurs naturally in the Brazilian 0.3-1.7 cm, glabrous to scarcely pilose, with acute Pantanal. It has been introduced recently to Aus­ to acuminate apices. Inflorescence racemose, with tralia (Northern Territory and Queensland) and to 1-2 peduncles per leaf axil; flower resupinate, sin­ most countries in South-East Asia, including Indo­ gly or in pairs at the end of a short (0.5-2 cm) nesia, Malaysia, the Philippines and Papua New peduncle; pedicel 4-10 mm long, subtended by a Guinea. single ovate bract 2-4 mm long with at the distal Uses C. pascuorum is sown as a pasture legume end two conspicuously paired bracteoles which in northern Australia, mainly in regions with a are ovate, 4-6 mm x 2-4 mm, with acuminate long dry, but reliable wet season, or in seasonally apices; calyx tube 3-4 mm long, with 5 narrow flooded parts of the Northern Territory. It is some­ teeth 3-7 mm in length; the lowest tooth (4-7 mm) times cut for hay. It has been planted experimen­ isth e longest; corolla wine red, 15-25 mm long and tally in South-East Asia. wide;th e standard with a spur on the back towards Properties C.pascuorum is regarded as a relati­ the base. Pod linear, 4-8 cm x 3-4 mm, laterally vely palatable, high quality tropical forage. Nitro­ compressed, with a dark longitudinal stripe near gen concentrations range from 2.5-4.3%. In vitro each suture, containing up to 15 seeds; the pod DM digestibility of young plant material has been shatters at maturity. Seed ovoid to cylindrical, ca. measured at 64%, and P concentrations varies 4m m long, slightly compressed laterally, greenish- from 0.15-0.18%. There are 48-58 seeds/g in the yellow to brown, rarely mottled (accessions from two Australian cultivars. Venezuela). Description An annual, herbaceous plant with Minute hooked hairs occur on the outer surfaces a twining or scrambling habit, producing roots on of the calyx, bracts and bracteoles, and more spar­ trailing stems in moist conditions. Stem cylindri- sely on the leaves and stems. CENTKOSEMA 85

Growth and development Photoperiod and to the fungal pathogens Cercospora canescens (leaf- temperature influence flowering and seed produc­ spot), Colletotrichum truncatum (anthracnose), tion. In cultivar 'Cavalcade', flowering occurs in Pseudocercospora bradburyae (leaf-spot), Rhizoc­ short days (12 hour photoperiod or less). In longer tonia solani (foliar blight) and Neocosmospora days (13hou r photoperiod) buds may be produced, vasinfecta. However, these diseases are usually not but they abort at high temperatures (33/28°C day/ serious in areas where it is well-adapted. It is also night) and no seeds are produced. Soil seed susceptible to root-knot nematodes (Meloidogyne reserves of25 0kg/h a are common in good pastures. spp.), and to sucking insects during seed produc­ Hard seed content is high (90-100%) when seeds tion. Control by chemicals usually is not economic ripen, but declines to 10-30 % at the end of the dry except for commercial seed production. season. Harvesting C.pascuorum is usually grazed, or Other botanical information Two botanical is cut for hay. varieties of C. pascuorum have been named, but Yield Annual DM yields of4- 6 t/ha are obtained neither is widely recognized. Ecotypes from Ecua­ from legume-dominant pastures in the semi-arid dor often have larger leaves and seeds, and paler tropics. In small plots, yields of up to 9 t/ha have flowers, than those from elsewhere. Ecotypes from been measured in Thailand. Under favourable con­ Venezuela have smaller, mottled seeds. Accessions ditions, large quantities of seed are produced, ex­ from north-eastern Brazil are more diverse and ceeding 1t/h a in pure stands. usually more vigorous than those from elsewhere. Genetic resources Representative germplasm A few accessions from Brazil and Venezuela have collections (80 accessions) are held by ATFGRC ovate leaflets. Two cultivars, 'Cavalcade' and (CSIRO, Australia) and by CIAT (Colombia). 'Bundey', have been released in Australia. Breeding There are no active plant improve­ Ecology C. pascuorum is adapted to tropical ment programmes at present. A breeding pro­ regions having reliable wet (4-6 months, 700-1500 gramme in Australia from 1976-1981 led to the mm) and dry seasons. It is drought resistant, but release of 'Cavalcade'. The breeding objectives can also survive prolonged waterlogging or flood­ were seed and forage yield, and freedom from symp­ ing (partial immersion). It will not persist in the toms of nematode infestation. Small numbers of subtropics and is not frost-tolerant. It is adapted accessions have been tested in most South-East to soils of near-neutral pH, ranging from sand to Asian countries, notably Thailand. heavy clay. Prospects C.pascuorum has potential for wider Propagation and planting C.pascuorum is pro­ use as apastur e or hay legume in semi-arid tropical pagated by seed, at a seeding rate of 2-5 kg/ha. regions. Its ability to tolerate both drought and Heat treatment to reduce hard-seededness may be prolonged waterlogging is unusual in legumes. necessary for hand-harvested seed. Inoculation of However, its proven area of adaptation in Austra­ seed with an appropriate strain of Bradyrhizobium lia islimited , and its variable performance in trials is advised, but is usually not essential. A culti­ in South-East Asia suggests that its use may be re­ vated seed-bed is required, and the seed should be stricted to a few areas in the drier parts of the covered with soil to a depth of 1-2 cm. Seed should region. be planted at the start of the wet season. Control Literature |1: Clements, R.J., Winter, W.H. & of existing weeds and grasses during establish­ Reid, R., 1984.Evaluatio n ofsom e Centrosema spe­ ment is helpful. Grasses such as Cenchrus ciliaris cies in small plots in northern Australia. Tropical L., Urochloa mosambicensis (Hack.) Dandy and Grasslands 18: 83-91. j2; Nulik, J., Andrews, A. & Andropogon gayanus Kunth can be planted with Jacobsen, C.N., 1986. Evaluation of grass and C. pascuorum. legume species in swards in Nusa Tenggara. In: Husbandry Heavy applications of fertilizer are Annual report of the Forage Research Project. not required, but C. pascuorum will usually res­ Balai Penelitian Ternak (Research Institute for pond well to low rates of P (5 kg/ha). In pastures Animal Production), Bogor, West Java, Indonesia, it is a valuable dry season forage, but it can be pp. 28-31. |3|Oram , R.N., 1990.Registe r of Austra­ grazed throughout the year provided the stocking lian herbage plant cultivars. CSIRO,Australia , pp. rate during the second half ofth e wet season is low 272-274. ,4j Schultze-Kraft, R. & Clements, R.J. enough to allow good seed production. In Austra­ (Editors), 1990. Centrosema: biology, agronomy lia, 2-3 steers/ha can be carried during the dry sea­ and utilization. CIAT, Cali, Colombia. 667 pp. |5| son on good pastures. Skerman, P.J., Cameron, D.G. &Riveros , F., 1988. Diseases and pests C.pascuorum is susceptible Tropical forage legumes. FAO, Rome. pp. 238-242. 86 FORAGES

[6| Stockwell, T.G.H., Clements, E.J., Calder, G.J. & Winter, W.H., 1986. Evaluation of bred lines of Centrosema pascuorum in small plots in north­ west Australia. Tropical Grasslands 20: 65-69. R.J. Clements

Centrosema pubescens Benth.

Comm. legum. gen.: 55(1837) . LEGUMINOSAE 2n = 22 Synonyms Centrosema molle Martius ex Benth. (1837). Vernacular names Centro, butterfly pea (En). Indonesia: sentro. Philippines: dilang-butiki (Tagalog), lesu-kesu (Subanun). Thailand: thua- lai, thua-sentro. Vietnam: day trung châu long. Origin and geographic distribution Originat­ ing in South and Central America, centro is now one of the most widely distributed of all legumes in the humid tropics. Uses Centro has been used as a green manure and plantation ground cover in Java and Peninsu­ lar Malaysia since the 19th century. Centro is widely used as a plantation cover and/or pasture legume in South-East Asia, the Pacific Islands, the wet tropics of Australia and indeed much of the humid tropics worldwide. Properties Centro is an efficient fixer of N with Centrosema pubescens Benth. - 1, flowering N concentrations generally ranging from branch; 2,fruits; 3, seed. 2.4-2.7%. Centro is one ofth e most palatable trop­ ical legumes. There are about 40 seeds/g. on the outside, bright or pale lilac on either side Description A vigorous, climbing, perennial of a median greenish-yellow band with numerous herb; trailing runners have a tendency to root at dark violet stripes or blotches. Pod linear, 4-17 cm the nodes if soil moisture is high, giving it a stolo- x 6-7 mm, flattened, margins prominent, straight niferous appearance; roots deeply penetrating; de­ or slightly twisted, acuminate, dark brown when velopment of taproots and lateral roots is almost ripe, containing up to 20seeds . Seed shortly oblon- equal, although soil type exerts some influence. goid to squarish with rounded corners, 4-5 mm x Stems leafy, arising from the main runners at 3-4 mm x 2 mm, brownish-black, with mottled 0.5-1.5 m intervals, climbing rather than trailing, darker blotches. slightly hairy, possibly becoming woody when Growth and development Centro is notor­ older than 18months . Leaves trifoliolate; leaflets iously slow to establish and requires good condi­ elliptical, ovate-oblong or ovate-lanceolate, 1-7 tions during the establishment period, but when cm x 0.5-4.5 cm, rounded at the base, rounded to grown in a pure sward it forms a dense, compact acuminate at the apex, dark green, slightly hairy cover 35-45 cm deep, 4-8 months after sowing. It especially on the lower surface; petiole up to 5.5 isfull y established and vigorous by at least the 2nd cm long, stipules 2-4 mm long, persistent. Flower year. In ungrazed mixtures with Panicum maxi­ papilionaceous, cleistogamous, large, pale mauve mum Jacq. it forms an impenetrable vine canopy with purple lines in the centre, borne in axillary some 2m high . In a mixture with calopo (Calopogo- racemes, 3-5 per raceme, subtended by 2 striate nium mucunoides Desv.) and puero (Pueraria pha- bracteoles; calyx tube campanulate, teeth un­ seoloides (Roxb.) Benth.), centro persists longest equal, 2 upper ovate triangular 1.5-3 mm long; under the closing canopy of plantation crops. Up standard rounded, up to 3 cm in diameter, hairy to 60% ofth e seedma y behard , requiring scarifica- CENTROSEMA 87 tion before planting. In Australia, common centro and timber regrowth. The tolerance of centro to flowers in April and October with main seed har­ certain herbicides has been determined, which can vesting periods in June-July and November- assist in weed control. In mixtures with grasses, December. Cultivar 'Belalto' flowers in June and notably Panicum maximum, centro tolerates rota­ has a main seed harvesting period in early August. tional or continuous grazing, but is intolerant of Nodulation occurs with a range of rhizobia but grazing in pure stands. Under acidic soil condi­ optimal growth has been achieved with very few tions, centro is more responsive to Mo, Ca, K and strains.Inoculatio n with an effective strain of Bra- P than puero which, in turn, is more responsive dyrhizobium is, therefore, recommended. Esti­ than stylo. Centro is more sensitive than stylo to mates of N fixation range from 120-270 kg/ha per soil P deficiency but is less sensitive to Cu and pos­ year. sibly S deficiencies. Other botanical information Two lines of cen­ Diseases and pests Centro has been relatively tro are in commercial use: common centro and cul­ free of major diseases and pests. Some virus and tivar 'Belalto'. 'Belalto' is now identified as C. bacterial diseases have been noted from time to schiedeanum (comb, ined., syn.: Clitoria schie- time and there are seasonal infestations of Cercos- deana Schlecht.) rather than as C. pubescens as pora leaf-spot and red spider (Tetranychus sp.). In wasliste d originally. It was selected as an improve­ the humid tropics it is noticeable that the infesta­ ment over common centro because of its superior tion offolia r blight (Rhizoctonia solani) may cause cool season growth, greater tolerance of pests and some dieback and that the attack of ladybird bee­ diseases, and its stronger stoloniferous growth. tles may affect plant growth. The damage can be Ecology In the humid tropics, the preferred severe when centro is planted in a pure sward, legumes for fertile and infertile soils have traditio­ especially during the wet season. However, none nally been centro and stylo (Stylosanthes guianen- of these have warranted commercial control mea­ sis (Aublet) Swartz) respectively. However, when sures. / soil mineral deficiencies are corrected and seed is Harvesting Harvesting is by grazing animals, inoculated with an effective Bradyrhizobium, cen­ when it can be selectively overgrazed unless care tro has been more productive than stylo on all land is taken, or it can be cut for stall feeding. Centro classes. The preferred annual rainfall for centro has persisted for decades as plantation cover or in is 1500 mm or more but it has also proved to be well-managed grazed associations with grasses, more tolerant of lower rainfall than once consid­ but it has not been very stable in cut-and-carry sys­ ered, having persisted in pastures receiving an tems. Centro is usually consumed fresh but it can average annual rainfall of 800 mm. Centro toler­ be ensiled or dried for hay or pellets. ates waterlogged conditions when grazing is le­ Yield Pure stands of centro have produced DM nient and it will survive a 3-4 month dry season yields of up to 12t/h a per year. In mixed pastures, but is not adapted to prolonged drought. It is intol­ this is more likely to be about 3 t/ha per year. erant of low temperatures, growing poorly when Standing DM yield of centro in grazed mixed pas­ temperatures fall below 15°C.I t ison e ofth e shade- tures is unlikely to exceed 1 t/ha. Well managed tolerant legumes and can persist under 80% shade. grass/centro pastures have consistently supported Centro combines well with other species in mixed stocking rates of 2.5 steers (375 kg liveweight) per pastures or ground covers under plantation crops. ha in the Australian tropics and 4 steers (250 kg) Propagation and planting Centro is propa­ in the Malaysian wet tropics, producing about 500 gated by seed. Hand harvested seed has a high pro­ kg/ha of liveweight gain per year. Seed production portion ofhard-seedednes s and mechanical scarifi­ can reach more than 200 kg/ha. cation is desirable. Seeding rate is about 5 kg/ha. Genetic resources Seed of centro has been sold Full seed-bed preparation and careful crop plant­ for many decades in South-East Asia and other ing procedures are generally recommended, espe­ humid tropical areas. Large germplasm collec­ cially since centro is somewhat slow to establish. tions are held by ATFGRC (CSIRO, Australia), However, centro has been successfully sod-seeded CENARGEN/EMBRAPA (Brazil) and CIAT (Col­ directly into a run-down grass pasture, following ombia). heavy grazing and low slashing of the residual Breeding Plant breeding programmes have been grass. undertaken by QDPI and CSIRO in Australia and Husbandry Properly fertilized and carefully CIAT in Colombia. However, the most promising grazed grass/centro associations have been persis­ lines are beginning to emerge from plant collec­ tent, productive and competitive against weeds tions in South and Central America. 88 FORAGES

Prospects Except for the humid tropics of Aus­ nutrient levels in cassia are average for tropical tralia and a few areas of Malaysia and the Philip­ legumes, with N concentrations of 1-3%. Al­ pines, the potential of centro as a component of though some species of Cassia are toxic, 6 acces­ grazed pastures has been largely unrealized. Most sions of roundleaf cassia have been fed to rats and pasture developments in South-East Asia have there were no symptoms of toxicity. The live been on infertile soils which are not suitable for weight gains and intake of rats fed with,roundlea f cropping. Similar developments in Australia have cassia have been found to be marginally better only been successful after careful studies of the than with rats fed lucerne. Cultivar 'Wynn' was nutritional status ofth e soil with respect to centro also found to be free of toxic compounds in studies in particular. Very few studies of this type have with pen-feed sheep. There are 200-500 seeds/g. been undertaken in South-East Asia. Centro is a Botany An annual or weak perennial semi-erect recommended legume in the 'Three Strata Forage to prostrate herb, in age basally suffrutescent, System' in Bali, Indonesia. with woody-fibrous blackish taproot up to 1 cm in Literature !1| Clements, R.J., Williams, R.J., diameter; stems up to 1 m long, not rooting at Grof, B. & Hacker, J.B., 1983. Centrosema. In: nodes. Leaves ascending, bifoliolate, petiole up to Burt, R.L, Rotar, P.P.,Walker , J.L. &Silvey , M.W. 1 cm long; leaflets subrotund to broadly obovate, (Editors): The role of Centrosema, Desmodium, 0.5-5 cmlong , by day ascending, face upward from and Stylosanthes in improving tropical pastures. tip ofpetiole , folded face-to-face at night, venation Westview Press, Boulder, Colorado, United slightly prominent on both surfaces; petiolule only States, pp. 69-96. |2[ Nitis, I.M., Lana, K., Suarna, a thickened pulvinule, without a gland. Flowers M., Sukanten, W., Putra, S. & Arga, W., 1989. in racemose axillary clusters of 1-3; pedicel up to Three strata system for cattle feeds and feeding in 6 cm long; sepals 5, greenish or reddish-brown, dryland farming area in Bali.Fina lrepor t to IRDC, 3-13 mm long; petals 5, yellow, as long as or Canada. 252 pp. |3j Schultze-Kraft, R. & Clements, slightly longer than sepals. Pod linear-oblongoid, R.J. (Editors), 1990. Centrosema: biology, agro­ straight or slightly curved, (1.5-)2-5.5(-6) cm x nomy and utilisation. CIAT, Cali, Colombia. 667 pp. 14| Skerman, P.J., Cameron, D.G. &Riveros, F., 1988. Tropical forage legumes. FAO, Rome. pp. 243-255. |S|Teitzel , J.K. &Burt , R.L., 1976. Centro­ sema pubescens in Australia. Tropical Grasslands 10:5-14. J.K. Teitzel & C.P.Chen

Chamaecrista rotundifolia (Persoon) Greene

Pittonia4:31(1899). LEGUMINOSAE 2n = 16 Synonyms Cassia rotundifolia Persoon (1805). Vernacular names Roundleaf cassia, round- leafed cassia (En).Vietnam : muôn'g la trön. Origin and geographic distribution Round- leaf cassia isnativ e to an area extending from Mex­ ico through Brazil and Uruguay, including the Ca­ ribbean region. It has been introduced and naturalized in the southern United States and in parts ofWes t Africa. It has been introduced in Aus­ tralia and is used commercially. Uses Roundleaf cassia is only used for grazing, although good silage has been obtained experi­ mentally. Chamaecrista rotundifolia (Persoon) Greene - flow­ Properties Preliminary data indicate that ering and fruiting branches. CHAMAECRISTA 89

0.3-0.5(-0.6) cm, blackish-brown when ripe, dehis­ grazing, where it has proved to be persistent under cent. Seed rectangular in outline, flattened, 2-3 a wide range of grazing pressure, although persis­ mm long. tence and productivity are poorer with vigorous The species is quite variable in its stature, pubes­ grasses and also at light grazing pressures. In cence and size of flower parts and leaves, allowing mixed cassia/grass pastures, cattle tend to selecti­ distinction of many ecotypes. Provisionally (until vely graze the grass at the beginning and middle better studied), only two varieties have been dis­ ofth e growing season, and the percentage of cassia tinguished by Irwin & Barneby based on size and in the diet increases at the end of the growing sea­ proportion of the flowers alone: son. Seed pods are eaten by cattle, and peak levels - var. rotundifolia: flowers small, the largest petal of 1-3 seeds per g of dry cattle faeces aid in the up to 7 mm long, the longest fertile anther up natural spread ofth e species.Roundlea f cassia can to 4.4mm , the style up to 2m m long. Range: from give DM yields of over 5 t/ha in pure swards, but Florida (United States) and Sinaloa (Mexico) to annual yields are lower in grass/legume pastures. Argentina. Synonyms: Cassia bifoliolata DC. ex Seed yields can range from 200-800 kg/ha. Roun­ Colladon (1816), C. fabaginifolia Kunth (1824), dleaf cassia is suited to either direct heading or C.monophylla Vellozo (1825). hand harvesting of seed. - var. grandiflora (Bentham) Irwin & Barneby: Genetic resources and breeding The largest flowers larger, the longest petal up to 17mm , the germplasm collection is held at ATFGRC (CSIRO, largest fertile anther up to 11.5mm , the style up Australia). The main variation is in the time of to 8.5 mm long. Range: more local within that flowering and growth habit. ofvar . rotundifolia, but never associated with it. Prospects Within its adaptation limits of rain­ Synonym: Cassia bauhiniifolia Kunth (1819). fall and soil type, roundleaf cassia shows consider­ Growth is indeterminate and will continue after able promise. Itma y provide an alternative legume flowering for as long as temperatures and moisture in some of the areas suitable to shrubby stylo (Sty- conditions are suitable. losanthes scabra Vogel). New cultivars'of mid- and One cultivar, 'Wynn' (belonging to the var. rotun­ late-season maturity, as compared to the early- difolia), has been released in Australia. It can seed flowering 'Wynn', will be required to exploit its freely throughout the growing season, but primar­ full potential. ily in autumn. Other lines flower much later and, Literature jlj Ahn, J.H., Elliot, R. & Minson, in areas with a reliable and longer growing season, D.J., 1988.Qualit y assessment ofth e fodder legume are higher yielding. Cassia rotundifolia. Tropical Grasslands 22: Ecology Roundleaf cassia is best suited to areas 63-67. !2|Cook , B.G., 1988.Persisten t new legumes receiving from 700-1400 mm rainfall in the seaso­ for heavy grazing 2. Wynn round-leafed cassia. nally dry tropics and subtropics. It is not suited Queensland AgriculturalJournal 114: 119-121. |3I to poorly drained sites or heavy textured soils, and Irwin, H.S. & Barneby, R.C., 1982. The American is very intolerant of waterlogging. It grows best Cassiinae. Memoirs of the New York Botanical on free-draining, friable sandy soils. Top growth Garden 35(2): 727-733. 4j Lenée, J.M., 1990. Dis­ is readily killed by frost. eases of Cassia species - a review. Tropical Grass­ Agronomy Seed germination should be checked lands 24:311-324 . !5jOram , R.N., 1990.Registe r of before sowing to ensure that hard-seededness is Australian herbage plant cultivars. CSIRO, Aus­ not excessive. Seed can be broadcast into closely tralia, p. 284. 6| Skerman, P.J., Cameron, D.G. & grazed pastures, but establishment ismor e reliable Riveros, F., 1988. Tropical forage legumes. FAO, with some form of soil disturbance. Seeding rates Rome. pp. 232-236. |7| Strickland, R.W., Green­ of2- 4 kg/ha are recommended, but lower rates can field, R.G., Wilson, G.P.M. & Harvey, G.L., 1985. be used especially if sown in a mixture with Morphological and agronomic attributes of Cassia legumes. It nodulates readily with native cowpea rotundifolia Pers., C. pilosa L. and C. trichopoda rhizobia and there are no records of failure to Benth., potential forage legumes for northern Aus­ nodulate. Long-term persistence is assisted by reg­ tralia. Australian Journal of Experimental Agri­ ular recruitment from soil seed reserves, which are culture 25:100-108. frequently 3000/m2 and can be as high as 15000/m2. R.M.Jones No serious disease or pest problems have been recorded although the species is a known host of several viral and fungal diseases. To date, roundleaf cassia has only been used for 90 FORAGES

Chloris gayana Kunth

Révis. gramin. 1(18):29 3(1830) . GRAMINEAE 2n = 20(diploid) , 30, 40 Synonyms Chloris abyssinica A.Richar d (1851), C glabrata Anderss. (1864). Vernacular names Rhodes grass (En, Am). Herbe de Rhode (Fr). Philippines: koro-korosan (Tagalog), banuko (Ilokano). Thailand: ya-rot. Vietnam: tucgiéân. Origin and geographic distribution Rhodes grass is native to East, Central, and the eastern part ofWes t andsouther n Africa. Itwa s first culti­ vated in , and at the beginning ofth e 20th Century it was introduced to a wide range of regions in Africa, South and Central America, the United States, Australia, , Japan, Italy and the southern part ofth e former Soviet Union. The grass is grown in Indonesia (Java, Irian Jaya, northern Sumatra), thePhilippine s and Thailand. Uses Rhodes grass is used in pastures for graz­ ing,ha yan dsilag emaking .I t isa nimportan t grass for sub-humid subtropical regions. Properties Rhodes grass has an N concentra­ tion ranging from 2.7% of the DM in very young, leaves to 0.5% in old leaves. The P concentration ranges between 0.1% and 0.4% depending on age of material and soil fertility. DM digestibility of green material is about 60% (ranging from 40-82% ) depending on growth stage and cultivar. Chloris gayana Kunth - 1, habit; 2,ligule; 3, inflor- Crude protein digestibility canb e ashig h as 70% . escence; 4, spikelet. Palatability is good, but varies with cultivar. Cul­ tivar 'Katambora' is used as nematode suppressor Growth anddevelopmen t Seed germination is in crops oftobacco . There are7000-1000 0 seeds/g. optimal between 8 and 18 months after harvest. Description A perennial, stoloniferous (varies The seed germinates quickly (in1- 7 days, depend­ with cultivar), creeping or occasionally tufted ing ontemperature ) and seedlings establish rapid­ grass, 0.5-2 m tall. Stem fine and leafy, erect or ly.Th e seedlings produce erect tillers, with stolons ascending, occasionally rooting at the lower appearing later and these grow and branch rapid­ nodes. Leaf-sheath glabrous except near theblade ; ly.Widel y scattered seedlings canquickl y produce ligule about 1m mlong , including a fringe of short dense stands. hairs, with long hairs onth eblad e close to it; leaf- Rhodes grass is a short-day plant, flowering blade flat, rarely involute, (15-)25-50 cm x throughout or at the end of the growing season (1.5-)3-9 mm, tapering towards theapex , margins depending on cultivar. The caryopsis is fully ripe rough, glabrous except near thebase , lateral veins 23-25 days after flowering. strongly developed. Inflorescence consisting of Other botanical information Most cultivars are (3-)6-15(-20) digitate ascending to spreading selected from natural stands. Diploid cultivars are spikes 4-15c mlong , all attached at the top ofth e fine-stemmed, fine-leaved and less vigorous than culm, or a few a little below the others; spikelets tetraploid cultivars which arerobus t andvigorous . green orpurplish , about 3.5m mlong ; florets 3(-4), Giant Rhodes grass (tetraploid), released as culti­ with usually one hermaphrodite (lemma with an var 'Callide' in Australia is late flowering and awn of1-1 0 mmlong) , onemal e (awn1.5-5. 5mm ) robust, with thick stems and broad long leaves, and onerudimentary ; sometimesal lfloret s are her­ long awns (6-9mm ) and a long tuft ofhair s at the maphrodite and fertile. awn base. CHLORIS 91

Other Australian cultivars are 'Pioneer' (diploid, range of legumes such as Stylosanthes guianensis with lower nutritive value than other cultivars), (Aubl.) Swartz, Vigna unguiculata (L.) Walp., 'Katambora' (diploid, with many tillers and good Medicago sativa L., Macroptilium lathyroides (L.) seed production), 'Samford' (tetraploid, vigorous Urban, M. atropurpureum (DC.)Urba n and Lotono- stolon development, late flowering and with high nis bainesii Baker and as a companion crop under palatability). Several cultivars have also been wheat and maize. released in Africa. 'Pokot' is a bred cultivar (leafy, Husbandry Rhodes grass responds well to N and vigorous, late flowering and high yielding), P fertilizer. On fertile soils it is usually grown 'Nzoia' (very leafy with high seed yields), 'Masaba' without fertilization. (quite leafy and productive) and 'Karpedo' (good Diseases and pests Most cultivars are little drought tolerance). affected by diseases or pests,bu t majo r damage can Ecology The natural habitats of Rhodes grass be caused by Helminthosporium spp. in 'Nzoia', are grasslands, open or with scattered trees or causing dieback of leaves and shoot bases. shrubs, river banks, lake margins and seasonally 'Masaba' suffers from smut caused by Fusarium waterlogged plains. gramineum giving rise to seed loss in wet years. Rhodes grass has some drought tolerance, but less Harvesting Rhodes grass can be grazed conti­ than Panicum maximum Jacq. var. trichoglume nuously or rotationally from 4-6 months after Robijns. The optimum annual rainfall for Rhodes sowing, but high grazing pressure decreases grass isbetwee n 600-1000 mm but it can withstand vigour. Grazing pressure should be aimed at pre­ a dry season of up to six months. The grass res­ venting flowering, as the nutritive value declines ponds well to irrigation and ismoderatel y tolerant rapidly towards maturity. of flooding (up to 10-15 days with less than 30 cm The best time to cut for hay isjus t before flowering. of water). Six harvests a year are possible with cutting inter­ The optimum temperature for photosynthesis is vals of 25-50 days. The best time for/harvesting 35°C. The temperature range for growth is very seed is at the early stage of seed shedding. broad, from about 0°C to 50°C. Rhodes grass is Yield In North Sumatra 38 t/ha of DM has been more tolerant of low temperatures than most other harvested in 236day s and it was one of the highest subtropical grasses. yielding grasses tested. Seed yields range between Seed germinates best at temperatures between 100-650 kg/ha. Annual liveweight gains of up to 20-35°C . 160 kg/animal and 850 kg/ha have been recorded The optimal daylength for growth is between from well fertilized Rhodes grass pastures. 10-14 hours. Herbage yields are considerably Genetic resources and breeding A large col­ reduced at photoperiods outside this range. lection of Rhodes grass germplasm is held at Rhodes grass is tolerant of fire, but not of shade. ATFGRC (CSIRO, Australia). Rhodes grass is Rhodes grass will grow on a wide range of soils, cross-pollinating with a self compatibility of from heavy clays to sandy loams. Optimum pH lies 1-4%). Breeding work is being done in Kenya between 4.5 and 7. Rhodes grass tolerates a high ('Pokot') and the United States (Texas). Plants are salinity and it can accumulate high concentra­ being selected on leafiness and late flowering char­ tions of Na in the leaves without suffering damage. acteristics. Propagation and planting Rhodes grass can be Prospects Because ofit s wide adaptability, ease propagated from seed and stolons. Propagation of establishment and acceptable nutritive value, from seed gives more rapid establishment. The Rhodes grass will continue to play an important seed is small and should be sown very shallow, or role in pasture improvement in subtropical preferably broadcast on top of a well prepared regions with a pronounced dry season. seed-bed or rather loose rough soil, covered lightly Literature |1| Bogdan, A.V., 1969.Rhode s grass. and/or rolled. It can be sown from the air into the Herbage Abstracts 39:1-13. |2|Bogdan , A.V., 1977. ashes after burning. Seed sown below a depth of Tropical pasture and fodder plants. Longman, 25m m will not germinate. London and New York. pp. 77-86. |3I Jones, R.J. Naked caryopses germinate slightly better than &Jones , R.M., 1989.Liveweigh t gain from rotatio­ those enclosed in glumes. Seed purity is usually nally and continuously grazed pastures of Narok not more than 25-35%, with germination of Setaria and Samford Rhodes grass fertilized with 30-55%. The seeding rate should be 0.5-1 kg/ha nitrogen in southeast Queensland. Tropical Grass­ of pure viable seed. lands 23:135-142. |4|Renvoize , S.A., 1974. Chloris. Rhodes grass grows well in mixtures with a wide In: Polhill, R.M. (Editor): Flora of tropical East 92 FORAGES

Africa. Gramineae (Part 2). Crown Agents for Oversea Governments and Administrations, Lon­ don, p. 346. |5| Skerman, P.J. & Riveros, F., 1990. Tropical grasses. FAO, Rome. pp. 283-288. L. 't Mannetje &S.M.M . Kersten

Chrysopogon aciculatus (Retzius) Trinius

Fund. Agrost.: 188(1820) . GRAMINEAE 2n = 20 Synonyms Andropogon aciculatus Retzius (1789). Vernacular names Love grass (En). Amorseco (Sp). Indonesia: salohot (Batak), jukut domdoman (Sundanese). Malaysia: temuchut, kemuchut, kemunchup. Philippines: marisekos (Tagalog), amorseko (Visaya), pangrot (Bikol). Cambodia: smau kântraëy. Thailand: ya-chaochu, ya-khikh- rok (central), ya-khitroei (south). Vietnam: co'may, co'bông. Origin and geographic distribution C. acicu­ latus originates from tropical Asia, Australia and Polynesia and is very abundant in all South-East Asian countries. It has been introduced to other parts of the tropics, e.g. West and Central Africa. Uses C. aciculatus is used for grazing, as a lawn grass, and for control of soil erosion. However, it can annoy humans and livestock due to its prickly spikelets that stick to and penetrate clothing or Chrysopogon aciculatus (Retzius) Trinius - 1, flow­ skin. The culms are used to make brushes and to ering plant; 2, triad of spikelets. weave small cases. Properties The leaves are highly palatable, but ish-brown hairs on one side and at top an awn 2-8 the abundant flowering culms and spikelets have mm long. Caryopsis yellowish-brown, 2-3 mm low palatability. No data on nutritive value are long. available. At maturity the callus at the base ofa sessile spike- Botany A perennial, spreading, stoloniferous, let separates from the supporting branch near its mat-forming grass. Culms solid, glabrous, erect or base and the sharp free tip easily catches on pass­ creeping, up to 75c mtall , often branching, rooting ing animals or humans. This tip penetrates clo­ at all nodes when creeping. Leaf-sheath glabrous, thing or hairy skins and is difficult to remove often pierced by roots; ligule membranous, trun­ owing to the stiff appressed hairs. The fruit may cate, very short; leaves 2-20 cm x 4-8 mm; leaf- work its way into the flesh, causing extensive ul­ blades on creeping culms ovate-lanceolate, short, ceration. pressed flat against soil; leaf-blades on flowering C.aciculatus spreads and forms a firm mat over the culms more linear. Inflorescence a rigidly erect ground, starting to flower within 6-8 weeks after panicle, 5-12 cm long, composed of several whorls establishment. It flowers throughout the year. of short reddish branches which gradually spread Ecology C. aciculatus can grow from sea-level horizontally when ripe, each branch bearing at its to 1500m altitude. It is adapted to moderately dry end a group of 3spikelets , each group with 1bisex ­ to humid environments and to sandy loamy soils ual sessile and 2 male or neuter pedicelled spike­ of pH 5-6. It is frequently found in overgrazed lets; sessile spikelet with at base a pointed callus, areas, resisting trampling. It cannot withstand 4-6 mm long, which is bearded with short yellow- prolonged dry periods. It is a vigourous colonizer CHRYSOPOGON 93

ofdenude d ground and tends to dominate with reg­ protects sandy coastal areas from wind erosion. ular burning. Properties Nitrogen concentrations in leaves Agronomy C. aciculatus is propagated by seed from flowering plants of C. orientalis range from or rooting tillers. The bristles on the spikelets 0.5-0.7%. The leaf/stem ratio is low and this adhere to the hair oflivestoc k and provide an effec­ stemmy condition persists almost throughout the tive method of seed dispersal. Close grazing is nec­ year.A D Mdigestibilit y of4 6% has been measured essary to keep it in a young growth stage. Cutting for flowering plants. is not practised for forage production. When used Botany A perennial grass with culms up to 80c m as a lawn grass, it should be mown frequently to tall, creeping and branching or sometimes densely keep it green and free of inflorescences. It is a low- tufted, with short stout stolons. Leaf-sheaths and yielding species. internodes distinct below, tight and more overlap­ Genetic resources and breeding It is unlikely ping above; leaf-blades basal and cauline, linear- that any substantial germplasm collections are acuminate to oblong, 6-10(-30) cm x 3-4 mm, flat being maintained. or folded, usually glabrous. Inflorescence a terminal Prospects The main use of C. aciculatus is as a panicle, 10-18 cm long, with whorls of slender naturally occurring grass for livestock, and as a branches, each branch with a terminal tuft of red- control against soil erosion. In improved pastures brown persistent hairs at the base of a group of 3 it is often considered as a noxious weed. spikelets (1 sessile, 2 pedicelled); sessile spikelet Literature |1 Backer, CA. & Bakhuizen van bisexual, with an oblique slender golden-hairy basal den Brink, R.C., 1968. Flora of Java. Vol. 3. callus 2.5m mlon gbelo wth e baseo fth e lower glume, Wolters-Noordhoff, Groningen, the Netherlands, glumes spinulose-hairy,uppe r glume with awn up to pp. 603-604. |2| Gilliland, H.B., Holttum, R.E. & 1-5 cm long, upper lemma with awn to 6 cm long; Bor, N.L., 1971. Grasses of Malaya. In: Burkill, pedicelled spikelets with lower glume awned to 1.5 H.M. (Editor): Flora of Malaya. Vol. 3. Govern­ cm,uppe r glume and upper lemma awnless. ment Printing Office, Singapore, pp. 236-237. 131 Mehra, K.L. & Fachrurozi, Z., 1985. Indonesian economic plant resources: forage crops. Lembaga Biologi Nasional- LIPI, Bogor, No 31.p .7 . !4| Sker- man, P.J. & Riveros, F., 1990. Tropical grasses. FAO, Rome. pp. 296-297. C. Manidool

Chrysopogon orientalis (Desv.) A. Camus

Lecomte, FI. Gén. de 1'Indo-Chine 7:33 2(1922) . GRAMINEAE 2n = unknown Synonyms Rhaphis orientalis Desv. (1831), Andropogon wightianus Nees ex Steudel (1854), Chrysopogon wightianus (Nees ex Steudel) Thwaites (1864). Vernacular names Philippines: tinloy (Taga- log), rukut dukut (Visaya), salsapot (Ilokano). Laos: hnha:z khwa:k. Thailand: ya-phungchu. Vietnam: co' may dông. Origin and geographic distribution C. orien­ talis originates from and is distributed in coastal Malaysia, Thailand, Indo-China and India. Uses C. orientalis is used as green forage. In a traditional grazing system in Thailand, cattle and buffaloes from distant villages are herded oni t dur­ Chrysopogon orientalis (Desv.) A. Camus - 1, culm ing the daytime and return in the evening. It also with leaves; 2, inflorescence; 3,triad of spikelets. 94 FORAGES

When fully mature the panicles become a golden Clitoria ternatea L. colour. Seed production is abundant. First flower­ ing starts about 11week s after germination, and Sp. PL: 753(1753) . seed reaches maturity at 13week s after sowing. LEGUMINOSAE Ecology C. orientalis is adapted to warm, high 2,n= 16 rainfall climates. It is well adapted to sandy soils Synonyms Clitoria zanzibarensis Vatke (1878), but also grows vigorously onheav y soils in central C. tanganicensis Micheli (1897), C. mearnsii De Thailand. It grows best on open fields but can also Wild. (1925). invade oldcoconu t plantations. Vernacular names Butterfly pea, K(C)ordofan Agronomy C.orientalis canb e planted vegetati- pea, blue pea, Asian pigeon-wings (En). Indonesia: vely or sown at a seeding rate of10-1 2 kg/ha. Seed bunga biru (Malay), kembang telang (Javanese, germinates within 5day s after sowing. A spacing Sundanese). Malaysia: bunga biru, kacang telang. of 50 cm x 80 cm is recommended for vegetative Philippines: kolokanting (Tagalog), giting prin- planting, using 3roote d tillers per hill. C. orienta­ cesa (Bikol), balog-balog (Visaya). Cambodia: lis responds well to higher fertility and appro­ rum'choan. Laos: 'ang s'an dam, bang s'an dam. priate rates of complete fertilizer can be used. It Thailand: anchan. Vietnam: dâu biê'c. is palatable and tolerates heavy grazing and fire; Origin and geographic distribution Clitoria frequent grazing and/or burning of old swards is ternatea ispantropica l (20°N-24°S).It stru e origin recommended to keep the plants in a leafy condi­ is obscured by extensive cultivation or natural­ tion. It canb e cut and fed to animals, but grazing ization in the humid lowland tropics of Asia, is more appropriate since the leaves are normally Africa, the Pacific Islands, andth e Americas. It is concentrated at the base of the plant and are too widespread throughout South-East Asia. low to be cut by large mowers. Annual DM yields Uses Butterfly pea has a reputation as a poten­ of 5-8 t/haar eobtaine d inThailand . Itma y be con­ tial fodder plant, hay or cover crop. It has been served as hay but it is then too coarse with little extensively tested assuch , especially inth e subhu- leaf. It isno tsuitabl e for silage making. mid to semi-arid tropics. It has never been used in Genetic resources andbreedin g It is unlikely extensive areas, although it is used by small­ that any substantial germplasm collections are holders. It is used as a cover crop in coconut in being maintained. No variation in growth form southern India and in rubber in Malaysia. It is and adaptation has been observed. Irradiation of widely grown asa n ornamental onfence s and trel­ seeds to obtain leafy mutations has been tried in lises because of its showy blue or white flowers, Thailand butwithou t success. and isgrow n fordy eproductio n andmedicina l pur­ Prospects The species has many advantages, poses in India. In the Philippines young pods are such as its ability to grow on extremely infertile eaten asa vegetable. soils on sandy coastal land, its tolerance of heavy Properties Nitrogen concentrations of whole grazing and light shade and its good seed produc­ tops range from 1.7-4.0% depending onth e season tion. Itha sparticula r potential where cattle graze and stage of growth. For example, in Brazil, hay under oldcoconu t plantations or sandy soils, such cut every 42 days had 3.7% N whereas hay cut as occur in southern Thailand. Accessions with a every 84 days had 3.0%. Hays from Brazil and higher leaf/stem ratio may increase the yield of India contained 24-38% crude fibre, 38-47% acid useful forage. detergent fibre, 11-16 % lignin and21-2 9 % cellu­ Literature |1|Gillilan d H.B., Holttum, R.E.& lose (44%ocrud e cellulose). Bor, N.L., 1971.Grasse s of Malaya. In: Burkill, Seeds contain 38% protein (2.5% lysine), 5% total H.M. (Editor): Flora of Malaya. Vol. 3. Govern­ sugars, 10%o oil and a powerful cathartic com­ ment Printing Office, Singapore, pp. 238-239. |2| pound. Seed size ranges from 15-35 seeds/g. Manidool, C, 1989. Natural grassland and native Description Perennial climbing, scrambling or grasses of Thailand [in Thai]. Technical Bulletin trailing herb with astron g woody rootstock. Stems No 1301-26-32. Division of Animal Nutrition, do not root at the nodes, slender, 0.5-3 m long, Department of Livestock Development, Bangkok, mostly pubescent orglabrescent , sometimes suber- pp. 16-19. |3| Matthew, K.M., 1983. Theflor a of the ect at thebase . Leaves pinnate with 5o r 7 leaflets; Tamilnadu Carnatic. Vol 3(2).Th e Rapinat Her­ leaflets elliptical, oblong, oblong-lanceolate, or barium, Tiruchirapalli, India, pp. 1823-1824. almost round, 1-7 cm x 0.3-4cm ,acute , rounded C. Manidool or emarginate at the apex, acute to rounded at the base, glabrescent above, adpressed pubescent be- CLITORIA 95

geal; the radicle emerges within 48-72 hours and seedlings emerge in 3-6 days, depending on plant­ ing depth. Early growth is rapid in warm moist conditions. Growth of established plants is mostly from the apices ofth e main axis and axillary branches; very few new shoots arise from ground level. Flowers are cleistogamous but a small level of outcrossing occurs.Tim e to flowering in a collection of 58 lines (sown in January at 19°40'S) ranged from 7-11 weeks, with most lines flowering 8-9 weeks after sowing. Subsequent flowering flushes overlap pod maturation from the previous flush, and they con­ tinue throughout the year in frost-free regions. At higher latitudes in the tropics, there is usually a peak at the end of the wet season and again at the end of the cool season, if moisture is available. Pods mature in 8-10 weeks after flowering and shatter readily once fully dry. Growth is more or less continuous in the humid tropics, or with irri­ gation in other hot regions. Individual plants may live for several years and grow into large vines if undisturbed. Other botanical information There is consid­ erable variation in the size of flowers and leaflets. Most 'cultivars' mentioned in the literature to date are ecotypes or agrotypes, not named cultivars in Clitoria ternatea L. 1,flowering branch; 2, flower; the true sense. However, cultivar 'Milgarra' was 3, fruit. released in Australia in 1990. Ecology Butterfly pea is essentially a plant of neath; petiole 1-3 cm long; rachis 1-6 cm long; the humid and subhumid tropical lowlands, but it petiolules 1-3 mm long; stipules persistent, lan­ has a reputation for drought tolerance in the sea­ ceolate, (2-)4-10 mm long, veined. Flowers axilla­ sonally dry tropics (500-900 mm rainfall) and it ry, solitary orpaired ; peduncle 3-10 mm long;pedi ­ has survived moderate frost damage in the cel 6-9 mm long and twisted 180° so that the subtropics (26°S) . It occurs in grassland, open standard is held lowermost; bracteoles ovate or woodland, bush, riverine vegetation, and dis­ round, 4-17mm x 2.5-15mm ,veined ; calyx pubes­ turbed places throughout its natural range. cent, veined, tube 8-12 mm long, lobes oblong-lan­ Butterfly pea grows best in full sun. Its annual ceolate or triangular, 7-10 mm x 2.5-3 mm, acute rainfall requirements for survival may be as low or acuminate, the upper pair joined for less than as 400 mm, but it requires about 1500 mm (or sup­ one third ofthei r length; standard oblong-obovate, plementary irrigation) for best production. Its alti- 25-50 mm x 15-35 mm, white or greenish-white tudinal range is0-1600(-1800 ) man d annual mean often blue margined or entirely blue, basal central temperature range is 19-28°C. area often yellow or greenish, very finely puberu- It has wide soil adaptation (pH 5.5-8.9), but prefers lous, margins sometimes finely ciliate. Pod linear- fertile friable soils and grows poorly on infertile oblong, flattened, 6-12.5 cm x 7-12 mm, mar­ sandy soils if not fertilized. It is one of the few her­ gined, apiculate, glabrous or with a mixture of baceous legumes well adapted to heavy clay soils sparse adpressed long hairs and very short hairs. in the subhumid to semi-arid tropics and the only Seeds 8-10, ellipsoid, oblong or oblong-reniform, one with potential in irrigated pasture mixtures sometimes truncate at one end, 4.5-7 mm x 3-4 on these soils.I t will not tolerate flooding or water­ mm, 2-2.5 mm, olive, pale brown or deep reddish- logging. brown with dark mottling, or almost black, minu­ In the seasonally dry tropics and in cool regions, tely pitted. growth is limited by lack of moisture or low tem­ Growth and development Germination is epi- peratures. Leaves are shed in response to these 96 FORAGES stresses and top growth may be killed by frost or For seed production on a small scale, plants may fire. However, recovery during thefollowin g grow­ be grown on trellises to maximize yield and to ing season is usually good, provided grazing is not make hand harvesting easier. However, for large- heavy and continuous. scale mechanized seed production, butterfly pea is Propagation and planting It is propagated by best grown as an annual row crop suitable for dir­ seed and readily self-propagates and spreads under ect heading. favourable conditions by seed thrown vigorously Diseases and pests Various fungi (e.g. Cercos- from the dehiscing dry pods. Seed is also spread pora, Colletotrichum, Oidium and Rhizoctonia) and in cattle dung. nematodes (Meloidogyne and Pratylenchus) have Normally sown from the beginning until the mid­ been recorded on butterfly pea but damage is rar­ dle of the wet season at rates of (l-)3-5 kg/ha in ely bad and control measures are unpractical or well-prepared seed-beds with seed placement 1.5-4 uneconomic in pastures. Fungicides such as beno- cm deep and lightly covered. Higher rates (5-8 kg/ myl may be useful in seed crops if diseases break ha) may be required when sowing pastures in out. Grass hoppers and leaf-eating caterpillars rough conditions. have caused damage in Australia. Successful stands have been established by broad­ Harvesting Pasture forage is normally har­ casting seed (10 kg/ha) into burnt or heavily vested by direct grazing or as cut-and-carry forage. grazed grassland, but success isver y dependent on Rotational grazing is preferred rather than set good seasonal conditions following planting. stocking whenever the plants are growing acti­ Hand-harvested seed often remains hard-seeded vely. Butterfly pea makes good quality palatable for a long time and requires scarification prior to hay but it isdifficul t to harvest and handle because sowing. Mechanical abrasion, hot water or sul­ of its viny nature. phuric acid can be used to break this dormancy. Yield Yields of butterfly pea vary enormously Mechanically harvested or threshed seed is because ofhighl y variable growing conditions and usually satisfactory for sowing the following wet management. Moreover, levels of productivity season without further treatment. projected from short-term trials are probably Inoculation of butterfly pea seed with rhizobia is unrealistic because ofit s susceptibility to frequent not usually necessary; but, if it is required, broad cutting. For example, when mixed pastures in Col­ spectrum cowpea inoculum should be used. ombia were harvested every 6-8 weeks, the Husbandry Butterfly pea competes fairly well amount ofbutterfl y pea in the sward declined from with weeds once established and can cover the 25-35% to 5% in one year. Worldwide, reported ground in 4-6 weeks when sown at a population DM yields have ranged 0.2-16.5 t/ha in forage of 4plants/m 2. Establishment may be a problem on crops and pastures. Crops grown under irrigation fertile soils if sown with a vigorous companion on good soils in the lowland dry tropics generally grass or oversown into an existing pasture. It com­ yield 7-13 t/ha DM,wherea s dryland crops usually bines better with tussock than stoloniferous yield 3-5 t/ha on good soils and 1-2.5 t/ha on light grasses in mixed pastures. soils or under low rainfall. If a pure stand isrequire d (e.g.fo r seed production), Under upland conditions in the Philippines butter­ cultivation or hand weeding willb erequire d during fly pea produced only 0.1-0.6 t/ha forage DM in early growth. Alternatively, trifluralin (800 g/ha) the first season and 1.3-2.8t/h a inth e second when may be incorporated pre-planting to control grass grown as an intercrop with grain legumes under weeds, and bentazone may be used at rates up to dry conditions. When sown following rice under 1900g/h a to control susceptible broad-leafed weeds. lowland conditions the DM yield of butterfly pea Because butterfly pea is very palatable it must be was 8.8 t/ha (4 cuts) as a forage crop and 3.6 t/ha carefully managed to avoid overgrazing and loss (3 cuts) as an intercrop with mungbean (Vigna of the stand. The location of its growing points at radiata (L.)Wilczek) . Lablabpurpureus (L.) Sweet the ends ofth e main branches makes it susceptible produced twice as much forage as an intercrop but to frequent low cutting as well as to continuous it reduced mungbean grain yield whereas butterfly heavy grazing (for example, it did not persist in pea did not. irrigated pastures on a heavy clay soil in north­ Seed yield may reach 300 g/plant hand harvested west Australia when grazed at a stocking rate of but it varies widely among lines. Yields of up to 2.5 steers/ha). Superphosphate fertilizer applica­ 700 kg/ha have been recorded for direct headed tion will increase DM yield and plant density on seed crops in northeast Australia. infertile soils. Liveweight gain of steers on irrigated pastures in CODARIOCALYX 97

northeast Australia was 0.68 kg/head per day over Codariocalyx gyroides (Roxb. ex Link) 41 weeks for a mixture of butterfly pea and para Hassk. grass {Brachiaria mutica (Forssk.) Stapf) com­ pared with 0.58 kg/head per day for centro (Centro- Flora 25, Beibl. 2:4 9(1842) , pro parte. sema pubescens Benth.) and para grass. In Mexico, LEGUMINOSAE cattle grazing butterfly pea and pangola grass In = 20,22 (Digitaria eriantha Steudel) produced 0.40 kg/head Synonyms Hedysarum gyroides Roxb. ex Link per day over 16week s compared with 0.24 kg/head (1822), Desmodium gyroides (Roxb. ex Link) DC. per day from siratro (Macroptilium atropurpureum (1825),D.papuanum C.T. White (1922). (DC.) Urban) and pangola grass, but both groups Vernacular names Codarrio (En). Indonesia: were also fed 1.5 kg/head per day of supplement sanagori, kadatuwa (Sundanese), julukan (Java­ containing 11.5% crude protein and 64% TDN. nese). Cambodia: kâm'phé:m. Thailand: thua des­ Genetic resources The main collections are modium. held at ATFGRC (CSIRO, Australia) and CIAT Origin and geographic distribution Codarrio (Colombia). is native to the area extending from and Breeding Variation exists among lines in flow­ India through southern China and Indo-China and ering time, pod dehiscence, seed and total dry mat­ throughout . It has been introduced on an ter yield, and plant habit. No breeding pro­ experimental basis into many countries. grammes are known but a major objective would Uses Codarrio is of increasing interest as forage be to improve persistence under grazing and cut­ and at the same time is used as green manure and ting, perhaps by developing forms with more grow­ as a shade tree in coffee and cocoa plantations. ing points at ground level. Properties Preliminary information indicates Prospects Butterfly pea has been considered a that leaves and young edible stems ofcodarri o may promising forage plant wherever it has been eva­ have a slightly lower N but similar P concentra­ luated throughout Australia, Africa, Central and tion than similar regrowth from leucàena (Leu- South America and South-East Asia. It may have caena leucocephala (Lamk) de Wit) cut at the same wider application to smallholder farm systems in height and frequency. Lower digestibility has been South-East Asia. Its drought tolerance and adapta­ reported but this may possibly reflect the presence tion to heavy clay soils, and the palatability and of tannins. If so, the reported values may not be quality of its forage, suggest it could be used to a true indication of the nutritional quality of improve natural grassland in extensive farm sys­ codarrio to ruminants. In feeding experiments tems in the subhumid to semi-arid tropics, given with rats, codarrio gave the same weight gains as appropriate grazing management. Its susceptibili­ equivalent diets with lucerne and there was no evi­ ty to close grazing or cutting is a major limitation dence of anti-nutritional substances. to general use. Botany An erect, usually multi-branched, shrub Literature |1| Crowder, L.V., 1974. Clitoria ter- 1-3 m tall with sparsely to densely pubescent natea (L.) Due. as a forage and cover crop - a stems; older stems can exceed 4 cm diameter. Leaf review. Nigerian Agricultural Journal 11: 61-65. 1-3-foliolate, petiole 1-3 cm long; leaflets 1 or 3, j2jGillett , J.B., Polhill, R.M. &Verdcourt , B., 1971. elliptical to obovate, 2-8 cm x 1-5 cm, obtuse to 44. Clitoria. In: Milne-Redhead, E. &Polhill , R.M. emarginate at the apex, rounded to cordate at the (Editors): Flora of tropical East Africa. Legumino- base, densely adpressed pubescent, sometimes gla- sae 4 - Papilionoideae 2. Crown Agents for Over­ brescent above, usually silvery white beneath; lat­ sea Governments and Administrations, London, eral leaflets usually smaller than terminal leaflet. pp. 515-518. |3j Hall, T.J., 1985. Adaptation and Flower about 1 cm long, usually with 2 together agronomy of Clitoria ternatea L. in northern Aus­ in dense terminal and axillary inflorescences 5-15 tralia. Tropical Grasslands 19:156-163. |4|Reid , R. cm long; corolla initially pinkish turning blue to & Sinclair, D.F., 1980. An evaluation of a collec­ mauve; pedicel ca. 1c m long. Fruit 2.5-5 cm x 4-6 tion of Clitoria ternatea for forage and grain pro­ mm, densely yellowish hairy, with 5-13 segment­ duction. CSIRO, Australia, Division of Tropical like parts but not splitting into articles, the whole Crops & Pastures, St Lucia, Queensland. Genetic fruit opening along the lower margin. Seed about Resources Communication No 1.8 pp . |5| Skerman, 2.5m m x 4m m x 1.5 mm. P.J., Cameron, D.G. & Riveros, F., 1988. Tropical Away from the equator codarrio flowers in the lat­ forage legumes. FAO, Rome. pp. 258-261. ter part of the wet season and early dry season. LB. Staples Late-flowering introductions set very little seed in 98 FORAGES

caena. Reports from Belize and Australia suggest that poor persistence of codarrio may be caused in part by damage from insect larva burrowing in stems, fungal diseases and root-knot nematodes, the latter found particularly on sandy, well- drained soils. Where plants are allowed to seed, recruitment of new plants may be able to compen­ sate for death of older plants. Genetic resources and breeding Collections are held by CIAT (Colombia) and ATFGRC (CSIRO, Australia). Preliminary assessment of lines collected in Sumatra in 1986 is being carried out at CIAT. Prospects Codarrio is showing promise as a for­ age plant, particularly on poorly drained and low fertility sites. In Indonesia further 'on-farm' test­ ing is in progress. Although it may not be persis­ tent in the long term, it could be planted out with slower growing but longer-lived shrub species which may yield less forage initially. Literature |1|Centr o Internacional de Agricul- tura Tropical, 1989.Annua l report, CIAT Tropical Pastures Program. Cali, Colombia, pp. 2-9, 2-13. |2| Jones, R.M., 1984. Yield and persistence of the shrub legumes Codariocalyx gyroides and Leu­ caena leucocephala in south-eastern Queensland. CSIRO, Australia, Division of Tropical Crops and Codariocalyx gyroides (Roxb. ex Link) Hassk. - 1, Pastures, Technical Memorandum No 38. 9 pp. I3| flowering and fruiting branch; 2, fruit. Lazier, J.R., 1981.Effec t of cutting height and fre­ quency on dry matter production of Codariocalyx the subtropics. In Indonesia, codarrio flowers in gyroides (syn. Desmodium gyroides) in Belize, April. Individual plants survive for 2-4 years. Central America. Tropical Grasslands 15: 10-16. Codarrio nodulates with native cowpea rhizobia. |4| National Academy of Sciences, 1979. Tropical C. gyroides is closely related to C. motorius legumes: resources for the future. National Aca­ (Houtt.) Ohashi, which occurs largely in the same demy of Sciences, Washington, United States, pp. area (not in Papua New Guinea). 128-129. |5| Ohashi, EL, 1973. The Asiatic species Ecology Codarrio occurs up to an altitude of of Desmodium and its allied genera (Legumino- 1900 m (e.g. in Papua New Guinea). It grows well sae). Ginkgoana 1: 43-46. |6i Skerman, P.J., on sites with a shallow water table and poor drai­ Cameron, D.G. &Riveros , F., 1988.Tropica l forage nage, and is not drought-resistant. It is tolerant of legumes. FAO, Rome. pp. 555-556. acid and infertile soils with much available Al. It R. Soedomo can survive burning when growing in Imperata cylindrica (L.)Eaeusche l grasslands in Indonesia. Agronomy In Indonesia, it is planted at spac- Crotalaria juncea L. ings of 0.5 m x 1.0 m. Growth may be slow for the first 2months , but ispossibl y faster than leucaena. Sp. PL 2:71 4(1753) . It usually flowers within 12 months. In Indonesia LEGUMINOSAE codarrio is cut and fed to animals, but plants do 2n = 16 not survive low and frequent cutting. A cutting Vernacular names Sunn hemp, Indian hemp, height of 0.5-1.5 m and cutting frequency of 6-10 Madras hemp (En). Chanvre indien (Fr). Indone­ weeks is provisionally suggested. Bushes can de­ sia: orok-orok lembut (Java). Philippines: putok- velop 50o r more basal branches. putukan (Tagalog), karay-kagay (Bikol). Cambo­ Codarrio canb e grazed, but in Australia more dam­ dia: kâk'tung. Laos: po: th'üang, thwax chu:b. age to plants was observed than to grazed leu­ Thailand: po-thuang. Vietnam: sue sat, lue lac. CROTALARIA 99

Origin and geographic distribution Sunn hemp originates from India but is now widely grown throughout the tropics and subtropics. Uses The major significance of sunn hemp lies in its valuable fibre which is extracted from the bark and used to make twine and cord, canvas, fishing nets, etc., and also to make paper and pulp. Moreover, sunn hemp is commonly used as forage, as green manure, and as a cover crop. It is one of the most widely grown green manures throughout the tropics. In Sri Lanka dried leaves, bark and boiled seeds are fed to cattle. With restrictions, seed has been used as fodder in the former Soviet Union and southern Africa. It is showing promise as a forage legume for intercropping with upland rice. In Indo-China it is used medicinally to treat urticaria. Properties Although sunn hemp contains poiso­ nous glucosides, it is widely used as forage. The presence of compounds which cause unpalatabi- lity and/or are poisonous under some conditions is typical of the genus Crotalaria L. Nitrogen con­ centrations of about 3% in hay and 5-10% in seeds have been reported from the former Soviet Union, but normally they are lower. Seeds may contain about 40% starch while stems contain about 40% fibre. There are about 33 seeds/g. Sunn hemp fibre has greater tensile strength and is more durable under exposure than jute. It is not Crotalaria juncea L. - 1,flowering branch; 2, part as strong as hemp (Cannabis saliva L.). ofstem; 3,fruits; 4,opened fruit; 5, seeds. Botany An erect, laxly branched annual, up to 3.5 m tall, with long strong taproot, many lateral growth on such soils is improved by fertilization. roots and numerous irregularly branched and It is not tolerant of salt, nor of sustained waterlog­ lobed nodules. Stem ribbed, pubescent, up to 2 cm ging. in diameter. Leaves simple; blade oblong-lanceo­ For fibre production, light, loamy, well-drained late, 4-13 cm x 0.5-3 cm, pilose; petiole up to 0.5 soils are preferred; on low-lying clay soils it makes cm long. Inflorescence a lax terminal raceme, up vigorous growth, but the fibre is coarser and yields to 25 cm long; flowers yellow, showy; sepals 5, are lower. hairy; standard erect, suborbicular, ca. 2.5 cm in Agronomy Sunn hemp is established by seed. diameter. Pod cylindrical, 3-6 cm x 1-2 cm, Sowing rates of 40-45 kg/ha are used when it is tomentose, light brown, containing ca. 6 seeds. sown as a forage crop or as green manure, but Seed heart-shaped, with narrow end strongly when it is sown for fibre, rates of 100-240 kg/ha incurved, up to 6m m long, dark brown to black. are used. It nodulates readily with native cowpea It is very fast-growing. Normally multibranched, type rhizobia. Seedlings emerge 3 days after sow­ it may have a single stem up to 3.5 m tall in dense ing, and rapidly produce a thick ground cover that stands. It has vigorous lateral roots and a long smothers weeds. taproot that can exploit deeply stored soil mois­ Sunn hemp is attacked by many diseases and pests, ture. Sunn hemp is a short-day plant and long day- including viruses, fungi, insects and nematodes, lengths favour vegetative growth and reduce seed- but they usually cause little economic damage. In set, although daylength neutral selections exist. India, wilt and caterpillar larvae of the Extensive cross-pollination occurs. Utetheisa pulchella can be serious. Pod-boring Ecology Sunn hemp is drought resistant and is insects can reduce seed production. Beetles of the adapted to hot, semi-arid and arid areas, yet can genus Exora can sometimes cause serious defolia­ tolerate light frosts. It grows on poor soils, but tion. Damage from insects is more severe if crops 100 FORAGES are planted in the same area for more than 3 con­ The Leguminosae: a source book of characteris­ secutive years. tics, uses and nodulation. Macmillan Publishers It can be harvested by hand or by machine. Total Ltd., London, United Kingdom, and The Universi­ green matter yields average from 18-27t/h a with ty of Wisconsin Press, Madison, Wisconsin, Uni­ forage yields ranging from 5-19 t/ha. When sown ted States, p. 192. |2i Bogdan, A.V., 1977. Tropical asa green manure crop after ricei nThailand , sunn pasture and fodder plants. Longman, London, p. hemp yielded 2 t/ha of high quality DM in 6-8 338. 3 Carangal, V.R., Tengco, P.L., Miah, N.I. & weeks. When grown for forage it canb e harvested Topark Ngarm, A., 1990. Food and forage crop 4times , starting 6-8 weeks after sowing andsubse ­ intercropping under rainfed lowland conditions. quently every 4 weeks. Sunn hemp should be cut Proceedings Crop-Animal Systems Research for hay or ploughed in for green manure in the Workshop, Serdang, Malaysia, August 15-19, early flowering stage when iti s1.5-2. 5month s old. 1988. MARDI/IDRC/ARFSN. pp. 435-451. ;4 Leaves and stems are dried since animals do not Duke, J.A., 1981.Handboo k of legumes of world eat sunn hemp when it is green. Sheep will not suf­ economic importance. Plenum Press, NewYor k & fer anyadvers e effects ifforce d to eatdrie d forage, London, pp. 63-66. 5 National Academy of but will suffer from toxicity iffe d large quantities Sciences, 1979.Tropica l legumes:resource s for the of seed. It should not be fed to horses, and the future. Washington, D.C. pp. 272-278. 61 White, intake of sunn hemp hay by cattle should be re­ G.A. &Haun , J.R., 1965. Growing Crotalaria jun­ stricted to about 10% of their diet. cea, a multi-purpose legume, for paper pulp. Eco­ When grown for fibre the plants are usually har­ nomic Botany 19: 175-183. 7! Whyte, R.O., Nils- vested at the flowering stage or when the stems son-Leissner, G. &Trumble , M.C., 1953. Legumes turn yellow. Stems are retted either in concrete of agriculture. FAO, Rome. p. 265. 181Yost , R. & tanks or in shallow open water for periods of4-1 4 Evans, D.,1986 . Green manure and legume covers days, depending on the temperature, thickness of in the tropics. Research Series 055. College of stems, and quantity of stems in relation to volume Tropical Agriculture and Human Resources, Uni­ of water. After retting themateria l is dried, decor­ versity ofHawaii , pp.16-17 . ticated by machine and the fibre is then stripped Y.K. Chee& C.P.Chen from the stems by hand, washed, and hung up to dry in the sun for bleaching. For seed production the crop isharveste d when the seeds areripe ; fibre Cynodon dactylon (L.) Pers. is extracted from the stems afterwards. Genetic resources andbreedin g There are no Syn. PL 1: 85(1805) . significant forage breeding programmes with sunn GRAMINEAE hemp, andfe w cultivars with predictable qualities In = 18, 36 are available. Cultivar 'Tropic Sun', of unknown Synonyms Panicum dactylon L. (1753), Cynodon origin, wasselecte d in Hawaii as a green manure. glabratus Steudel (1854), C.polevansii Stent (1927). Seeds andforag e were found tob enontoxi c in feed­ Vernacular names Bermuda grass (Am). ing trials. InIndi a 'Kanpur 14'i sth emos t common Couch grass (En). Chiendent dactyle (Fr). Indone­ cultivar, and 'T6'flower s in 30day s after sowing. sia: jukut kakawatan, gigirinling (Sundanese), Existing lines of sunn hemp are usually variable, sukit grinting (Javanese). Malaysia: rumput and it would appear there is scope for selecting in minyak. Philippines: kawad-kawad (Tagalog), specific regions for specific objectives such as dis­ kapot-kapot (Visaya), bakbaka (Ilokano). Burma: ease and pest resistance and yield of fibre or for­ mye-sa-myet. Cambodia: smao anchien. Laos: age. hnha:z ph'è:d. Thailand: ya-phraek. Vietnam: cô' Prospects Sunn hemp is a fast growing species, chi', co'ôhg. tolerant oflo w fertility and of soil moisture stress. Origin and geographic distribution Bermuda Since the crop is extensively cultivated for fibre grass is thought to originate from Africa and or green manure, there is good potential for using South and South-East Asia but it has been intro­ sunn hemp foliage as a high protein source to sup­ duced to alltropica l andsubtropica l regions of the plement other feeds. It is also showing promise as world andha sbee n found to survive at 50° N inEu ­ a forage intercrop. Eesearch is warranted onselec ­ rope and to 4000 m elevation in the . It tion of improved lines and on clarifying and over­ is also found onisland s inth ePacific , Atlantic and coming problems associated with its toxicity. Indian Oceans. Literature 111 Allen, O.N. & Allen, E.K.,1981 . Uses Bermuda grass is grazed by ruminants, CYNODON 101

used in cut-and-carry systems, to control erosion, whorls; spikelets 2-3 mm long, in two rows tightly and as a turf grass. In the United States improved appressed to one side of the rachis; glumes hybrids are also cut for hay. Worldwide it is also 1-nerved, the upper -j-f as long as the spikelet; a serious weed in maize, cotton, sugar cane, vine­ lemma silky pubescent on the keel;pale a glabrous. yards and plantation crops. Caryopsis ovoid, 1.5 mm long, yellow to reddish. Properties In vitro digestibility of 5-week-old Six botanical varieties have been recognized; var. forage of 500 introductions, uniformly managed, dactylon is distributed worldwide, the other 5 ranged from 40-69%. Nitrogen concentrations of varieties have a limited distribution in Africa and adequately fertilized Bermuda grass range from Afghanistan. Improved hybrid cultivars developed 2-3%, whereas inadequately fertilized grass or in the United States for higher DM yield and old material may only contain 0.5-1.5 % N. It con­ improved DM digestibility include 'Coastal', tains low levels of prussic acid, but toxicity symp­ 'Coastcross-1', 'Tifton 78', 'Tifton 68', 'Hardie', toms in animals are very rare. There are 3000-4000 'Oklan', and 'Brazos'. seeds/g. Ecology Bermuda grass grows best where mean Botany A stoloniferous, prostrate, sward-form­ daily temperatures are above 24°C. Temperatures ing perennial herb with underground rhizomes of -2 to -3°C usually kill leaves and stems back penetrating the soil to a depth of 1 m or more. to the ground but rhizomes survive these tempera­ Culms 8-40 cm tall (rarely taller) and 0.5-1 mm tures and regrowth is rapid in spring. It is deep- in diameter. Leaf-blade linear-lanceolate, 1-16 cm rooted and drought-tolerant. It grows best on well- x 2-5 mm, glaucous, glabrous or hairy on upper drained soils, but will tolerate long periods of surface, midrib prominent with two primary and flooding. It will tolerate a broad soil pH range, but 8-9 secondary nerves on either side; leaf-sheath grows best with a soil pH above 5.5. It also toler­ up to 15m m long, smooth, hairy or glabrous; ligule ates low fertility, but it is intolerant of shade. a conspicuous ring ofwhit e hairs. Flowering stems Agronomy Bermuda grass can bepropagate d by erect, terminated by 3-6 slender radiating 1-sided planting either seeds or stolons. Good-stands can racemes 1.5-10 cm long arranged in 1 (rarely 2) be obtained by planting 5-10 kg/ha hulled seed but higher rates can be used for rapid sward develop­ I * /- ment. Improved hybrid Bermuda grass can only be vegetatively propagated. Plantings of fresh vege­ tative material should be made in moist soil fol­ lowed by a roller to compact the soil. A small nur­ sery for vegetatively planting larger areas can easily be established from one or a few vegetative sprigs or stolons. With adequate fertilization, improved hybrid Ber­ muda grasses will produce twice as much forage as common types. A minimum of 10kg/h a of N per month of growth is needed for moderate to high productivity but Bermuda grass, especially the improved hybrids, will respond to rates of up to 60 kg/ha N per month of growth. 'Coastal' Bermuda grass at Georgia, United States, produced 1.8, 4.7, and 7.2t/h a DM with 1.1,1.2 and 1.3% N when fer­ tilized with 0, 52, and 112 kg/ha N, respectively. Common Bermuda grass could be expected to yield significantly less DM. Low-growing legumes can be grown with Bermuda grass, which improves for­ age quality and provides some of the N require­ ments of the grass. Applications of lime may be needed to bring soil pH to 5.5. Rust and Helminthosporium leaf-spot are the major diseases on common Bermuda grass. The Cynodon dactylon (L.) Pers. - 1, flowering plant improved hybrids are resistant to these diseases. part; 2, ligule; 3, spikelet. Armywor m (Spodoptera frugiperda) and spittlebug 102 FORAGES

(Prosapia bicinata) are the major insects that Cynodon nlemfuensis Vanderyst attack Bermuda grass. Adequate fertilization and defoliation that allows less than 8 cm growth to Bull. Agric. Congo Beige 13: 342 (1922), (11:12 1 accumulate will help control pests. Burning dead (1920),provisiona l name C. lemfuensis). top growth after frosting or after a dormant stage GRAMINEAE will help control spittlebug and the diseases. 2n = 18, 36 Bermuda grass should be cut for hay or silage Vernacular names Stargrass, African star- when it is 30-40 cm tall or after every 4-6 weeks grass (En, Am). Philippines: kolatay (Tagalog), growth. It can be grazed year-round if not dormant rukut-dukut (Visaya), galud-galud (Ilokano). due to frost or drought. A stubble height of 5-10 Thailand: ya-sata. cm should be maintained under grazing or left if Origin and geographic distribution Stargrass cut for hay. Bermuda grass is best utilized by graz­ occurs naturally in East and Central Africa, from ing. It can also be stored as hay after drying, or Ethiopia and Sudan through Zaire to Malawi and green-chopped and fed directly to animals, or made . In other parts of the tropics, including into silage. South-East Asia, it has been introduced as a fodder Genetic resources and breeding Active breed­ grass. ing programmes (each with over 500 introduced Uses Stargrass is grazed by ruminants, and used accessions) to improve Bermuda grass are located in cut-and-carry systems and to control erosion. in Stillwater, Oklahoma and Tifton, Georgia, Uni­ Properties Nitrogen concentration and DM ted States. C. dactylon is highly variable and pre­ digestibility of improved selections range from sent collections do not adequately represent the 1-2% Nand40-70%, respectively. Moststargrass species. Major breeding objectives include has medium to high levels of prussic acid, which increased yield and digestibility, pest resistance, is increased with high nitrogen fertilization. How­ and improved cold tolerance. ever, only few reports of livestock death due to Prospects Much opportunity exists for genetic prussic acid poisoning have been reported. improvement in yield, pest resistance, and quality. Botany A stoloniferous sward-forming perenni­ Proper fertilization and management and use of al without rhizomes; stolons stout, woody, lying current improved cultivars would have major flat on the ground; culms robust to fairly slender, impact on improving animal nutrition and soil 30-60 cm high and 1-3 mm in diameter at the base. conservation. Leaf-blade flat, linear-lanceolate, 5-16 cm x 2-6 Literature |1| Burton, G.W. & Hanna, W.W., mm, thin and green or rather stiff and glaucous, 1985.Bermud a grass.In : Heath, M.E., Barnes, R.F. scaberulous, with or without scattered hairs; & Metcalfe, D.S. (Editors): Forages. Iowa State ligule a scarious rim 0.3 mm long. Inflorescence of University Press. Ames, Iowa, United States, pp. 4-13 digitate 1-sided spikes, usually in 1, some­ 247-254. |2| Clayton, W.D. & Harlan, J.R., 1970. times in 2 whorls; spikes 4-10 cm long; spikelets The genus Cynodon L.C. Rich, in tropical Africa. 2-3 mm long, green or purplish-green, 1-flowered, Kew Bulletin 24:185-189. |3| de Wet, J.M.J. & Har­ strongly laterally compressed, imbricate in 2rows , lan, J.R., 1970. Biosystematics of Cynodon L.C. awnless; glumes 2, 1-3 mm long; lemma silky Rich. (Gramineae). Taxon 19: 565-569. |4j Harlan, pubescent to softly ciliate on the keel; palea gla­ J.R., 1970. Cynodon species and their value for brous. Caryopsis ellipsoid, laterally compressed. grazing and hay. Herbage Abstracts 40: 233-238. Two varieties have been distinguished, but since |5| Harlan, J.R., de Wet, J.M.J., Huffine, W.W. & they intergrade, distinction is often impossible Deakin, J.R., 1970. A guide to the species of Cyno­ and thus questionable: don (Gramineae). Bulletin B-673, Oklahoma State - var. nlemfuensis: culm 1-1.5 mm in diameter; University Agricultural Experiment Station, Still­ leaf-blade 2-5 mmwide ;spike s of 1 inflorescence water, Oklahoma, United States. 37 pp. |6| Sker- 4-9, each 4-7 cm long. man, P.J. & Riveros, F., 1990. Tropical grasses. - var. robustus Clayton & Harlan: culm 2-3 mm FAO, Rome. pp. 310-315. in diameter; leaf-blade 5-6 mm wide; spikes of W.W. Hanna 1 inflorescence 6-13, each 6-10 cm long; a more robust plant than var. nlemfuensis. C. nlemfuensis much resembles C. dactylon (L.) Pers., the main difference being the absence of underground rhizomes and the lack of hardiness. Stargrass grows vigorously and roots at nodes as CYNODON 103

tion, selected stargrasses will produce more than twice as much forage as common types. A minimum of 10 kg/ha N per month of growth is needed for moderate to high productivity, but star- grass (especially the improved selections) will res­ pond to higher rates. Low-growing legumes can be grown with stargrass which improves forage quali­ ty and provides some of the N requirements of the grass. Applications of lime may be needed to bring soil pH to 5.5. Rust and Helminthosporium leaf-spot are the major diseases of common stargrass. Foliar blight (Rhizoctonia solani) has been observed on some of the selected cultivars. Armyworm (Spodoptera fru- giperda) and spittlebug (Prosapia bicinata) are the major insect pests. Adequate fertilization and defoliation that allows less than 15 cm growth to accumulate will help control pests. Burning dead top growth if grass is frosted or after a dormant stage will help control spittlebug and the diseases. Stargrass should be cut for hay or silage when it is 30-40 cm tall or after every 4-6 weeks growth. It can be grazed year-round if temperatures and rainfall are favourable. Overgrazing will decrease stands. A stubble height of 15-25 crrf should be maintained under grazing or cutting. Dry matter yields of improved selections vary with climate Cynodon nlemfuensis Vanderyst - 1, flowering and soil fertility but are at least twice the yield plant part; 2, spikelet. of local or common types. Stargrass is best utilized by grazing. It can also be stored as hay after dry­ it spreads. Some genotypes have a bunch-habit ing. type of growth, even though they spread by sto­ Genetic resources and breeding Individual lons. researchers around the world have a few acces­ Improved cultivars include: 'Florico', 'Florona', sions. C.nlemfuensis is extremely variable and sys­ 'Ona', and 'Costa Rica'. tematic germplasm collection and maintenance is Ecology Productivity and persistence of star- needed. Isolated improvement programmes have grass are limited to lower elevations or where tem­ emphasized selection for higher dry matter yield peratures do not fall below -4°C, as growth rapid­ and digestibility among introduced accessions. ly ceases with low temperatures. In its area of Prospects Much opportunity still exists for origin it occurs up to 2300 m altitude. It does not genetic improvement in yield and quality. Hybrids tolerate long periods of flooding. It grows on many need to be produced and evaluated. Proper fertil­ soil types but does best on moist, well-drained ization and management and wider use of current soils. It will tolerate a broad soil pH range but best improved cultivars could have major impact on growth is made on soils with a pH above 5.5. improving animal nutrition and for soil conserva­ Agronomy Improved selections of stargrass are tion purposes. sterile or almost sterile. It can be propagated by Literature |1| Clayton, W.D. & Harlan, J.R., planting stolons or stem pieces. Plantings of fresh 1970. The genus Cynodon L.C. Rich, in tropical vegetative material should be made in moist soil Africa. Kew Bulletin 24:185-189. |2|d eWet , J.M.J. followed by a roller to compact the soil. A small & Harlan, J.R., 1970. Biosystematics of Cynodon nursery for vegetatively planting larger areas can L.C.Rich . (Gramineae). Taxon 19:565-569 . |3| Har­ easily be established from one or a few vegetative lan, J.R., 1970.Cynodo n species and their value for sprigs or stolons. grazing and hay. Herbage Abstracts 40: 233-238. Fertilizer will need to be added to most soils to |4| Harlan, J.R., de Wet, J.M.J., Huffinae, W.W. & obtain high DM yields. With adequate fertiliza­ Deakin, J.R., 1970. A guide to the species of Cyno- 104 FORAGES

don (Gramineae). Bulletin B-673, Oklahoma State University Agricultural Experiment Station, Still­ water, Oklahoma. 37 pp. |5| Mislevy, P., Brown, W.F., Caro-Costas, E.,Vicente-Chandler , J., Duna- vin, L.S., Hall, D.W., Kalmbacher, R.S., Overman, A.J., Ruelke, O.C., Sonoda, R.M., Sotomayor-Rios, A., Stanley, Jr., R.L. &Williams , M.J., 1989. Flor- ico stargrass . University of Florida Institute of Food and Agricultural Science Circular S-361.15 pp. '6! Skerman, P.J. &Riveros , F., 1990. Tropical grasses. FAO, Rome. pp. 316-318. W.W. Hanna

Dactyloctenium aegyptium (L.) Willd.

Enum. PL Horti Berol.: 1029 (1809) ('aegyptia- cum'). GRAMINEAE 2n = 20, 36, 48 Synonyms aegyptius L. (1753), Eleu- sine aegyptia (L.)Desf . (1798). Vernacular names Crowfoot grass (East Africa), coast button grass (Australia) (En). Indo­ nesia: tapak jalak (Sundanese), suket dringoan (Javanese), rebha kartuut (Madura). Malaysia: rumput miuyak. Philippines: krus-krusan (Taga- log), damong-balang (Visaya), tugot-manok (Ilok- Dactyloctenium aegyptium (L.) Willd. -1, flowering ano). Burma: didok-chi, myet-le-gra, mye-sa myet. plant; 2, ligule; 3, spikelet. Thailand: ya-pakkhwai (central). Vietnam: co' chân gà, co chi. spikes up to 6.5 cm long, ascending or often radiat­ Origin and geographic distribution D. aegyp­ ing horizontally from the top ofth e culm; spikelets tium is widely distributed throughout the tropics, 3-4-flowered, flattened, 3-4 mm long, in 2 alter­ subtropics, and warm temperate regions ofth e Old nate rows at the underside of a flattened rachis; World. It has been introduced to America. glumes subequal, the lower one with a scabrid Uses D. aegyptium is widely used as forage and keel, the upper with a smooth keel extending into is relished by all types of ruminants. Although a a stout divergent scabrid awn up to 2time s as long valuable forage, it can also be a troublesome weed as the glume; lemmas ending in a stout cusp or of cultivation. It makes excellent hay. In times of mucro 1 mm long. Caryopsis broadly obovate to scarcity it is used as a food grain. obtriangular in outline, ca. 1 mm long, transver­ Properties Nitrogen concentrations in D. sely rugose, orange-brown. aegyptium range from 1.2%-2.5%, depending on The inflorescence is similar to that of Eleusine the stage of growth and soil fertility. It contains Gaertn. It can be distinguished by its awned upper high levels of HCN at some stages of its growth. glume, its 2-ranked spikelets, and by its stolonifer- The grain is said to have an unpleasant taste and ous habit. to cause internal disorders in humans. D. aegyptium flowers early in the rainy season and Botany Slender to robust spreading annual, up seeds profusely; it isa n exceedingly variable grass. to 1 m tall, usually geniculately ascending and Ecology D. aegyptium can grow from sea-level rooting at the lower nodes, frequently shortly sto- up to 2100 m altitude and in areas receiving loniferous and mat-forming. Leaf-sheath up to 4c m 400-1500 mm rainfall annually. It is adapted to a long, glabrous, compressed; leaf-blade flat, 3-25 wide range of soil types but is particularly well cm x 0.25-1.2 cm, with bulbous-based hairs, espe­ suited to disturbed areas on light-textured soils. It cially along the margins; ligule a narrow fimbriate will not stand prolonged flooding. membrane. Inflorescence composed of up to 9 Agronomy D. aegyptium is established by sur- DESMANTHUS 105

face sowing at 0.25kg/ha . It grows very vigorously some regions of the tropics including the island of and can be cut or grazed early in the growing sea­ Sulawesi (Indonesia) and in the Pacific islands son. In grazed areas it can be top dressed with N (New Caledonia). fertilizer after its first grazing to stimulate Uses At present D. virgatus is not yet sown on regrowth. It can be cut as a fodder and fed fresh farms in South-East Asia despite recent promising to livestock and it also makes excellent hay. results from species evaluation experiments. In Farmers normally dig up whole plants when weed­ the Americas it makes a contribution to native pas­ ing and feed these fresh to their livestock. A DM tures but is not yet developed to the point of com­ yield of at least 3 t/ha per season can be expected mercial release. In India it is used in hedges to pro­ in unfertilized open fields on loamy sand soils. vide cut forage, and isbein g evaluated in Australia Genetic resources and breeding It is unlikely as a species for permanent pastures. that substantial germplasm collections are being In some areas the species has become a weed. maintained. Properties Nitrogen concentrations of 3.6% in Prospects D. aegyptium is a valuable native for­ the leaves and of 1.1% in the stems have been mea­ age. Studies should be carried out on its agronomy, sured. nutritive value and grazing management. There are 170-340 seeds/g. Literature iljDepartmen t ofLivestoc k Develop­ Botany A perennial or sometimes annual shrub ment, 1986.Repor t on feed analysis [inThai] . Tech­ or herb, prostrate to erect, 0.5-3 m tall, taprooted nical Bulletin No 13-0116-29. Division of Animal (the taproot develops into a storage organ in some Nutrition, Department ofLivestoc k Development, forms); stem unbranched or sparingly branched, Bangkok. 2' Gilliland, H.B., Holttum, R.E. & Bor, hollow, subglabrous, ridged. Leaves bipinnate, N.L., 1971. Grasses of Malaya. In: Burkill, H.M. 2-8 cmlong , with 1-7 pairs ofpinnae , usually with (Editor): Flora of Malaya. Vol. 3. Government a petiolar gland between the lowest pair of pinnae; Printing Office, Singapore, pp. 81-82. |3|Holm , J., petiole up to 5c m long; pinnae up to 7c m long with 1971. Feeding tables. Nutrition Laboratory of Thai-German Dairy Project, Livestock Breeding Station, Chiangmai, Thailand. |4| Holm, L.G., Plucknett, D.L., Pancho, J.V. & Herberger, J.P., 1977.Th e world's worst weeds: distribution and bi­ ology. The East-West Center, University of Haw­ aii Press, Honolulu, Hawaii, United States, pp. 244-248. 5! Skerman, P.J. & Riveros, F., 1990. Tropical grasses. FAO, Rome. pp. 322-324. C. Manidool

Desmanthus virgatus (L.) Willd.

Sp. PI. ed. 4,Vol . 4(2): 1047(1806) . LEGUMINOSAE In = 28 Synonyms Mimosa virgata L. (1753). Vernacular names False tamarind (En). Aca­ cia courant, acacia savane (Caribbean) (Fr). Thai­ land: thua desmanthus. Vietnam: cây diên keo. Origin and geographic distribution D. virga­ tus originated in the Americas where it is wide­ spread, occurring from Arizona, Florida and Texas in the United States, throughout much of Mexico and has been recorded in most countries south to Argentina as well as in the Caribbean and the Galapagos Islands. The main centre of diversity of the genus is Mexico and that is probably true of Desmanthus virgatus (L.) Willd. - 1, flowering D. virgatus as well. The species is naturalized in branch; 2, leaflet; 3,flower; 4, infructescence. 106 FORAGES

10-25 pairs of leaflets; leaflets linear to linear- mainly in their time of flowering in the subtropics oblong, 4-9 mm x 1-2 mm, oblique and eccentric and higher latitude tropics. Cultivar 'Marc' is at the base, obtuse to apiculate at the apex; sti­ early flowering, 'Bayamo' is mid-flowering and pules linear-subulate, up to 6 mm long. Inflores­ 'Yuman ' is late flowering. 'Marc' has a lower yield cence a small globose 6-10-flowered head; pedun­ potential but ismor e likely to persist in grazed pas­ cle up to 7.5 cmlong ; flowers bisexual or with some tures in the long term. The existence of adventive neuter lower flowers; petal white to cream, ca. 4 populations in Sulawesi (Indonesia) and in the Pa­ mm long, sometimes absent; stamens 10. Fruit cific shows that the species has potential as a for­ linear to falcate, 3-10 cm x 3-4 mm, dehiscing age in Asia especially on neutral to alkaline soils. with 2 valves, smooth or subseptate, red-brown, There is also interest in the species as a forage shiny, 20-30-seeded. Seed ovoid to ellipsoid, ca. 2 legume for upland intercropping. The wide range mm long, glossy brown. of material available should enable the develop­ D. virgatus is extremely variable in habit and in ment of commercial material for the region. other characters such as pod shape and size and Literature |1| Isely, D., 1970. Legumes of the leaf morphology. Four varieties are recognized by United States: 2.Desmanthu s and Neptunia. Iowa Isely: var. virgatus, var. glandulosus Turner, var. State Journal of Science 44:495-511 . |2| Skerman, depressus (Humb. & Bonpl. ex Willd.) B. Turner P.J., Cameron, D.G. & Riveros, F., 1988. Tropical and var. acuminatus (Benth.) Isely. These are sepa­ forage legumes. FAO, Rome. pp. 548-550. |3| rated by habit, leaf characteristics, and the Topark-Ngarm, A., Armada, E.C., Tengco, P.L. & number and location of glands on the petiole and Carangal, V.R., 1990. Food and forage intercrop­ leaf. Most accessions belong to var. virgatus. ping under upland condition. In: Proceedings of D. virgatus is a copious seed producer with many the Crop-Animal Systems Research Workshop, forms acting as annuals if not cut or grazed. Serdang, Malaysia, August 15-19, 1988. MARDI/ Ecology D. virgatus is adapted to a wide range IDRC/ARFSN. pp. 453-471. |4| Turner, B.L., 1950. of tropical and subtropical environments. It has Texan species of Desmanthus (Leguminosae). been collected at sites in the Americas with annual Field and Laboratory 18: 54-65. |5| Turner, B.L., rainfall ranging from 250-2000 mm and altitudes 1950. Mexican species of Desmanthus (Legumino­ from sea-level to 2000 m. Most commonly the spe­ sae). Field and Laboratory 18:119-130. cies is associated with neutral to alkaline clay or B.C. Pengelly &A . Topark-Ngarm clay-loam soils. Crowns survive frosting but top growth is frost-sensitive. Desmodium heterocarpon (L.) DC. Agronomy Propagation isb y seed. Seed usually has a high proportion of hard seed and hard-see- Prodr. 2:33 7 (1825) (heterocarpum). dedness can be broken by mechanical scarification LEGUMINOSAE or hot water treatment. Seeding rates of between 2n = 22 2-6 kg/ha should be adequate. D. virgatus should Synonyms Hedysarum heterocarpon L. (1753). preferably be inoculated prior to sowing by a spe­ Vernacular names Carpon desmodium (En). cific strain of Bradyrhizobium. It is adapted to Indonesia: buntut meyong sisir (Sundanese), kaci defoliation and should have cutting or grazing (Javanese), akar entimor (Belitung). Malaysia: applied soon after establishment to induce branch­ rumput kerbau derapah, kacang kayu betina. Phi­ ing and crown development. DM yields of 7.6 t/ha lippines: mangkit-parang (Tagalog), mani-mani have been obtained in Fiji, and fresh material (Visaya), huyo-huyop (Ifugao). Cambodia: baay yields of 23 t/ha (Hawaii) and 70 t/ha (Australia) dâm'nnaëp. Vietnam: trahg qu'a di qu'a. have also been reported. Origin and geographic distribution Carpon Genetic resources and breeding D. virgatus desmodium is distributed naturally from India and germplasm is available from the ATFGRC (CSIEO, Sri Lanka through Burma, Thailand, Indo-China, Australia) and from CIAT (Colombia). No breed­ Malesia, China, Taiwan, Ryukyu, Japan, to the Pa­ ing programmes are in progress. The extreme natu­ cific Islands and Australia. ral variation in climatic origin, plant morphology Uses Although native to South-East Asia, car­ and flowering time of the species has enabled the pon desmodium is grazed as a wild plant, but is not selection of accessions for advanced evaluation sown in the region. Cultivar 'Florida' has been from wild collections. sown to a limited extent in Florida (United States) Prospects Three cultivars have recently been where it is used for grazing in mixtures with peren­ released in Queensland, Australia. They differ nial grasses. DESMODIUM 107

Properties Carpon desmodium is lower in quali­ mostly in pairs, each on a 3-5 mm long pedicel ty than many other forage legumes. A study done within the axil of a 5-8 mm long pointed bract. Pod in the United States found that foliage contained 10-28 mm x 2-3 mm, erect to ascending, straight 1.5-3.2% nitrogen. Tannin percentages of along the upper margin, undulate along the lower 2.1-3.1% have been measured. There are 700-800 margin, the isthmus between the 4-8 articles seeds/g. 2/3-4/5 as wide as the pod which splits along the Description Aperennia l herb or low shrub with lower margin when ripe; articles quadrate, 2.5-3 ascending or creeping stems 0.5-3.0 m long from mm long, glabrous or hairy. Seed broadly ellipti­ a woody rootstock. Stems mostly much-branched cal, ca. 1.5mm x 2mm . at the base, and when young variously hairy, from Growth and development Seedling growth of almost glabrous to densely covered with more or carpon desmodium is slow. Once established, it is less appressed hairs. Leaves with 3 leaflets, or a very long-lived legume which can persist for over often with 1 leaflet on seedlings or at the base of 10 years regardless of grazing management. It older stems; leaflets variable in texture, shape requires a moderate soil P status for maximum and size, mostly papery; the terminal one ellipti­ growth. In Florida, 65% of the growth is during cal, ovate or obovate, (1.5-)2-6(-9) cm x the summer and early autumn growing seasons. (l-)1.5-3(-4) cm, indented or more or less pointed Under heavy grazing it develops a prostrate at the tip, sparsely covered with appressed hairs growth habit. It is self- or cross-pollinating; it selfs on the upper surface, the lower surface more den­ when flowers are tripped. sely covered with appressed silvery hairs and with Other botanical information D. heterocarpon prominent nerves; the lateral ones with the same is very polymorphic and has been variously deli­ proportions but up to 4 cm long. Inflorescence a mited. In the last revision and emendation of Des­ dense axillary or terminal raceme; flowers pink, modium Desv. for Asiatic species, Ohashi defined mauve, purple, violet or white, 4-7 mm long, this species in its widest sense and characterized it as having 1-3-foliolate glabrous leaves, ellipti­ cal, ovate or obovate terminal leaflets, inflores­ cences 3-13 cm long and erect or ascending pods with 4-8 transverse-oblong or tranverse-obovate articles shorter than 4.5 mm long and less than twice as long as broad. He subdivided the species as follows: - ssp. angustifolium Ohashi: synonym: D. reticula- tum Champ, ex Benth.; terminal leaflets narrow­ lyovate , acute or obtuse at apex, 3-6.5 times lon­ ger than broad; inflorescence 10-30 cm long, not branched; Burma, Thailand, Indo-China and China; with 2 formae. - ssp. ovalifolium (Prain) Ohashi: see separate treatment. - ssp. heterocarpon: terminal leaflets obovate, elliptical or oblong,sparsel y to subdensely hairy above; inflorescence elongated; flowers 3-4 mm long,pedicel s 4-7 mm long;pod s sparsely to den­ sely pubescent with white hooked and straight hairs up to 1.5 mm long; with 4 varieties. Ecology Carpon desmodium is adapted to a fairly high rainfall of over 1200 mm a year, can withstand short periods of drought but does not tolerate prolonged flooding. It is tolerant of repeated light frosts, and in the tropics it grows up to 2500m altitude .I t prefers better-drained soils of light texture. It tolerates soil pH values of Desmodium heterocarpon (L.) DC. - 1, fruiting 4.3-5.0 with high Al saturation. branch; 2,fruit; 3, seeds. Agronomy Establishment is by seed, sown at a 108 FORAGES rate of 3-10 kg/ha. A clean, firm seed-bed is desir­ dium heterocarpon (L.) DC. from southeast Asia. able. It nodulates with native cowpea rhizobium. Proceedings soil and crop science society of Flor­ On neutral and alkaline soils, it may become ida 49. pp. 94-99. !5|Lenné , J.M. & Stanton, J.M., micro-nutrient deficient. 1990. Diseases of Desmodium species. Tropical Carpon desmodium isfre e ofmajo r fungal diseases, Grasslands 24:1-14. i6iOhashi , H., 1973.Th e Asiat­ but is sensitive to various leaf-spots (Cercospora ic species of Desmodium and its allied genera sp., Pestalotiopsis versicolor). Anthracnose (Colle­ (Leguminosae). Ginkgoana 1: 210-216. 17!Ohashi , totrichum gloeosporioides, C. truncatum) occurs in H. 1991.Taxonomi e studies in Desmodium hetero­ the south-eastern United States and South Amer­ carpon (L.) DC. (Leguminosae). Journal of Japa­ ica but usually causes only slight to moderate dam­ nese Botany 66:14-25. i8lSkerman , P.J., Cameron, age. Powdery mildew (Oidium sp.) and wilt (Sclero­ D.G. &Riveros , F., 1988.Tropica l forage legumes. tium rolfsii) have also been recorded. Root-knot FAO, Rome. pp. 268-270. nematodes (Meloidogyne arenaria, M, incognita, J.B. Hacker &A.E . Kretschmer, Jr. M.javanica) have been described asth e worst pests of'Florida' in the south-eastern United States, but their significance in South-East Asia has not been Desmodium heterocarpon (L.) DC. ssp. documented. Little leaf, caused by mycoplasma- ovalifolium (Prain) Ohashi like organisms, has caused slight to severe symp­ toms in Central and South America. Web-worms Journ. ofJap . Bot. 66:2 1(1991) . and other insects, including bean leaf roller (Urba- LEGUMINOSAE nus proteus) sometimes attack foliage and pods in 2n = 22 rank stands. They may be controlled by grazing Synonyms Desmodium polycarpum DC. var. and then de-stocking for a period before harvest­ ovalifolium Wallich ex Prain (1897),D. ovalifolium ing the seed crop. Wallich ex Merrill (1910), D. heterocarpon (L.) DC. It is harvested by grazing, and in Florida (United ssp. heterocarpon var. ovalifolium (Wallich ex States), it is reported to be very tolerant of heavy Prain) Rugayah (1987). grazing. Vernacular names General: desmodium Genetic resources and breeding The wide (South-East Asia). Thailand: khonthi din. Viet­ diversity in the species could indicate potential for nam: trahg qu'a xoan. genetic improvement. Studies in Florida, largely Origin and geographic distribution Ssp. ovali­ based on accessions from Thailand, have suggested folium originates in tropical South-East Asia that some accessions are resistant to root-knot where it occurs naturally from northern Thailand nematodes. Germplasm collections are maintained to southern Sumatra between 98°E and 105°E. It at ATFGRC (CSIRO, Australia), CIAT (Colombia) is particularly frequent in southern Thailand and and USDA (Fort Pierce, Florida, United States). is probably also native to the southern parts of Breeding work is being carried out in Florida. Burma, Laos, Vietnam, and Cambodia. The acces­ Prospects There has been little interest in this sion marketed in Singapore has spread, at least for species outside of the south-eastern United States experimental purposes, to most tropical countries since its initial release as a pasture legume in 1979, of the world. and it is not anticipated that future plantings will Uses In South-East Asia, ssp. ovalifolium is be widespread. mainly used as a cover crop in tree-plantation agri­ Literature |1|Bogdan , A.V., 1977.Tropica l pas­ culture where it can be opportunistically grazed. ture and fodder plants. Longman, London, p. 343. Outside South-East Asia, its main use is as forage I2|Imrie , B.C., Jones, R.M. & Kerridge, P.C., 1983. in permanent grass/legume pastures. Desmodium. In: Burt, R.L., Rotar, P.P., Walker, Properties Its nutritive value, including palata- J.L. & Silvey, M.W. (Editors): The role of Centro- bility, is commonly considered moderate to poor, sema, Desmodium and Stylosanthes in improving mainly because of high tannin content (21-43%) tropical pastures. Westview Press, Boulder, Color­ and subsequent low intake and poor digestibility ado, United States, pp. 97-140. I3| Knaap-van (28-51% ) ofDM .Tanni n levels can be reduced and Meeuwen, M.S., 1962. Preliminary revisions of intake of ssp. ovalifolium can be increased by sul­ some genera of Malaysian Papilionaceae 5.A cen­ phur fertilization. Nitrogen concentrations in the sus ofth e genus Desmodium. Reinwardtia 6:93-96, herbage range from 1.5-3.0%. Other mineral con­ 251. |4| Kretschmer, A.E., Bullock, R.C. & Wilson, centrations are low to moderate (0.10-0.24% P C.L., 1990. Evaluation of a collection of Desmo­ and 0.27-0.57 % Ca).Ther e are about 500 seeds/g. DESMODIUM 109

ness. Flowering and seed setting occur in the late wet and early dry seasons. Other botanical information In the literature, this taxon is best known as D. ovalifolium, a name still often used in agricultural publications. In 1973, Ohashi considered it as belonging to the com­ plex species D. heterocarpon, and within this spe­ cies to ssp. heterocarpon. Rugayah gave it a varie­ tal status under Ohashi's ssp. heterocarpon in 1987, justifying this separation by some morphological and anatomical distinguishing characteristics. In 1991 Ohashi agreed that D. ovalifolium deserved a special status, and he gave it the actual subspeci- fic rank. Nevertheless, in the whole D. heterocar­ pon complex, intermediate forms blur the distinc­ tions between the different subspecies recognized by Ohashi. More research isneede d to unravel this complex satisfactorily. All ssp.ovalifolium used in South-East Asian plan­ tation agriculture seems to belong to a single accession marketed in Singapore. The same acces­ sion is also spreading in tropical America for use in pastures. It was recently released in the humid tropics of Bahia, Brazil, as 'Itabela'. Ecology Ssp. ovalifolium is well adapted to the humid and sub-humid tropics with not more than 2-3 consecutive dry months and is very shade-tol­ Desmodium heterocarpon (L.) DC. ssp. ovalifolium erant. It grows well on a range of soils, including (Prain) Ohashi - 1, fruiting branch; 2, rooting light-textured as well as seasonally poorly branch. drained, heavy-textured soils. The species is well adapted to acid, low-fertility oxisols and ultisols Description A creeping, stoloniferous herb, in with high Al saturation and low available P. dense stands or under competition ascending up Propagation and planting Ssp. ovalifolium is to 1m . Stems many-branched, glabrous except for established by seed, at rates of 0.5-2 kg/ha. Seed silky pubescence on young (apical) portions; old samples frequently have a high percentage of hard stems woody at base. Leaves 1- and 3-foliolate, in seed, particularly if newly harvested, and will young plants always 1-foliolate; leaflets variable, require treatment by heat or acid or mechanical mostly ovate or broad-elliptical, sometimes round scarification to break hard-seededness prior to or obovate; terminal leaflet larger than laterals; sowing. 1-foliolate leaves and terminal leaflets on adult To establish pastures, seed is broadcast or drilled plants on average 3-4.5 cm x 1.5-3 cm; leaflets at the same time that the associated grass is sown coriaceous and without any markings, glabrous or planted, or it is strip-sown into an existing grass and glossy on upper surface, whitish pubescent on sward. Because of its stoloniferous growth habit, lower surface, sometimes only along the central ssp. ovalifolium associates well with aggressive, vein. Inflorescence a densely flowered raceme, 2-5 stoloniferous grasses. The species most frequently cm x 1-2 cm; flower papilionaceous, small, with recommended for association with ssp. ovalifolium an obovoid standard about 6 mm x 4 mm, purple is koronivia grass (Brachiaria humidicola (Ren- to dark-pink during anthesis, bluish when wilting. dle) Schweick.). Although the legume nodulates Pod erect or falcate, mostly densely pubescent, freely and effectively with native rhizobia, the in­ dehiscent, comprising 2-8 almost quadrate arti­ oculation of seed with a Bradyrhizobium strain of cles 2.5-3.5 mm long. Seed yellow when ripe. known effectiveness is recommended. Growth and development The initial growth For use as a ground cover in plantation agricul­ of ssp. ovalifolium is slow, but once established, ture, a species mixture with centro (Centrosema plants exhibit excellent vigour and competitive­ pubescens Benth.), tropical kudzu (Pueraria pha- 110 FORAGES seoloides (Roxb.) Benth.), calopo (Calopogonium Australia). Planned plant collection in Burma, mucunoides Desv.) and C. caeruleum (Benth.) Laos, Vietnam and Cambodia should provide Sauv. is recommended. Ssp. ovalifolium will be the important additional variation. most persistent in the mixture because it is the Breeding Ssp. ovalifolium can be crossed with most shade-tolerant component. other subspecies and varieties of D. heterocarpon; Husbandry Fertilization with P and K enhances such crosses sometimes occur even in nature. At establishment, and despite its undemanding re­ present no breeding programme to improve ssp. quirements regarding soil fertility and nutrients, ovalifolium is in progress. ssp. ovalifolium responds to maintenance fertiliza­ Prospects Its excellent adaptation to acid, tion with these nutrients. poorly fertile soils and its aggressive, stolonifer­ Because of its stoloniferous growth habit, ssp. ova­ ous growth habit combined with remarkable shade lifolium tolerates heavy grazing well. Ssp. ovalifo­ tolerance make ssp. ovalifolium valuable for plan­ lium/Brachiaria humidicola pastures can be tation agriculture and pastures. Efforts are justi­ grazed rotationally or continuously, though stoc­ fied to overcome the constraints oflo w nutritional king rate might have to be reduced during drier quality and disease susceptibility by systematic periods. germplasm evaluation and, eventually, breeding. Diseases and pests Pink disease, caused by Literature |1|Centr o Internacional de Agricul­ Sclerotium (formerly Corticium) salmonicolor, and ture Tropical, 1983-1990. Annual reports, Trop­ root-knot nematode Meloidogyne javanica have ical Pastures Program. CIAT, Cali, Colombia. |2| been reported as major problems in South-East Keiler-Grein, G. (Editor), 1990. Red Internacional Asia. de Evaluation de Pastos Tropicales, RIEPT-Ama- A false rust or wart disease caused by Synchytrium zonia. I Reunión-Lima, Peru, 6-9 Noviembre desmodii was accidentally introduced to South 1990. Documento de Trabajo No 75, 2 Vols. 1119 America with commercial ssp. ovalifolium seed pp. |3|Ohashi , H., 1973.Th e Asiatic species of Des- produced in Sri Lanka. In order to prevent this dis­ modium and its allied genera (Leguminosae). ease from becoming a major constraint in South- Ginkgoana 1: 210-216. |4| Ohashi, H., 1991. Taxo­ East Asia, and as the fungus survives on the seed nomie studies in Desmodium heterocarpon (L.) surface and mainly in the seed debris, all seed lots DC. (Leguminosae). Journal of Japanese Botany should be treated with concentrated sulphuric 66:14-25. |5|Rugayah , 1987. A new status for Des­ acid prior to sowing. modium ovalifolium (Papilionaceae). Reinwartia In tropical America, a stem-gall nematode (Ptero- 10(3): 381-382. |6| Schultze-Kraft, R. & Benavides, tylenchus cedidogenus) can be a serious pest. G., 1988. Germplasm collection and preliminary Harvesting When used as a forage, ssp. ovalifo­ evaluation of Desmodium ovalifolium Wall. lium is usually harvested by grazing animals CSIRO Division of Tropical Crops and Pastures, rather than in cut-and-carry systems. St. Lucia, Australia. Genetic Resources Communi­ Yield The average range of DM yields reported cation No 12.2 0pp . for ssp. ovalifolium grown on acid, poorly fertile R. Schultze-Kraft soils but under adequate soil-moisture conditions is 2-3.5 t/ha per 12 weeks. On more fertile soils, yields as high as 6 t/ha per 12 weeks can be Desmodium heterophyllum (Willd.) DC. obtained. Liveweight gain per animal from a ssp. ovalifolium-bsLsed pasture is moderate to low, but Prodr. 2:33 4(1825) . animal production per hectare can be high, LEGUMINOSAE because of the high carrying capacity of such pas­ 2n = 22 tures. Synonyms Hedysarum heterophyllum Willd. Seed yields range from 150-300 kg/ha given ade­ (1802),Meibomia heterophylla (Willd.)Kuntz e (1891). quate soil moisture. Vernacular names Hetero, Spanish clover Genetic resources The genetic base of commer­ (En). Indonesia: sisik betok (Indonesian), heuheu- cially available ssp. ovalifolium is narrow. Howev­ langan (Sundanese), sukut jareman (Javanese). er, the species is well represented, in the form of Malaysia: rumput sisek naga, rumput sisek betok, a large number ofquit e variable ecotypes collected rumput telinga tikus. Cambodia: trôm' prei, smau in Thailand, Malaysia and Indonesia, in the two 'âm'bôk. Laos: padông môd liinz. Thailand: ya- major collections of tropical forage legume germ- maengmi (northeast). Vietnam: hah the,tràn g qu'a plasm at CIAT (Colombia) and ATFGKC (CSIRO, di diêp (southern). DESMODIUM 111

Origin and geographic distribution Hetero is on the lower surface. Inflorescence a few-flowered native to tropical Asia and adjacent islands, occur­ axillary or terminal raceme to 6 cm long, and/or ring naturally in Mauritius, India, Nepal, Sri leaf-opposed clusters of 1-3 flowers; pedicel 10-25 Lanka, Burma, Thailand, Indo-China, Malesia, mm long; flower about 5m m long, purple or white. China and Taiwan. It is adventive to the South Pa­ Pod narrowly oblong, 10-20 mm x 3-5 mm with cific and naturalized in parts ofnorth-easter n Aus­ 3-6 broadly oblong or quadrate articles 3-4 mm tralia. long, densely covered with minute hooked hairs, Uses Hetero is primarily used as a legume for the isthmus between the articles ca. fthe width of heavily grazed pastures in the wet tropics, and is the pod, dehiscing along the lower margins when recommended as such in Malaysia. It is also used ripe. Seed transversely elliptical, about 1.5 mm x as a cover crop, e.g. in pepper gardens in Sarawak. 2mm . Hetero isself-compatibl e but normally cross- It is not suitable for cut-and-carry systems. pollinated. Cultivar 'Johnstone' was released in Properties Nitrogen concentration in new Australia in 1973. growth of hetero is about 2.8-3.0%. When grown Ecology Hetero is adapted to the humid tropics in mixtures with pangola grass (Digitaria eriantha with annual rainfall exceeding 1500 mm, and to a Steudel) it increased the N concentration of the wide range of soils from sands to clays. It is grass from 0.86% to 1.12 %. reported to be less tolerant of soil acidity than D. Botany A prostrate or ascending perennial heterocarpon (L.) DC. It is intolerant of fire and undershrub, with woody rootstock. Stems up to 1.5 salinity, and susceptible to frost. m long, freely branching and covered with spread­ Agronomy Seed production of hetero is not ing reddish-brown hairs. Leaves trifoliolate, on easy, owing to poor synchrony within the crop lower parts often unifoliolate; petiole 5-15 mm (associated with the indeterminate nature of the long; leaflets obovate or elliptical, usually 10-25 plant and the two inflorescence types) and the fra­ mm x 8-14 mm, laterals smaller than the termi­ gility ofrip e pods which are carried within or close nals, rounded or indented at the tip, hairless on to the upper levels of the herbage. Therefore, it is the upper surface, sparsely covered with long hairs commonly established vegetatively by cuttings. However, because hetero requires a specific strain of Bradyrhizobium, it is preferable to plant out sods which contain vegetative material, seed and Bradyrhizobium. Fresh seed has some hard-see- dedness, which may be broken by scarification. Seed should be sown no deeper than 5 mm and should be inoculated with the appropriate Brady­ rhizobium, if possible. Early growth has been noted as slightly unpalatable, but this is an advan­ tage as it promotes successful establishment. Hetero occurs as a component of native pastures in parts of Indonesia and Fiji, where its poor pro­ ductivity as a naturally occurring species can be improved by applying P fertilizer. Unlike most tropical pasture legumes, hetero tolerates heavy grazing. It has an excellent ability to spread natu­ rally, both bymean s ofit s freely-branching stolons and its free-seeding habit, and competes well against low-growing weeds. Reports on its toler­ ance to shade are confusing, but recent experience suggests that, when adequately fertilized, it is one of the most shade-tolerant of the pasture legumes. Hetero is sensitive to cercospora leaf-spot (Cercos- pora sp.) and to foliar blight (Rhizoctonia solani), which is frequent in humid regions of the tropical American lowlands. Wilt (Sclerotium rolfsii) has Desmodium heterophyllum (Willd.) DC. -1, flower­ caused seedling mortality in greenhouse tests and ing and fruiting branch; 2,fruit; 3, seeds. little leaf (mycoplasma-like organisms) has caused 112 FORAGES damage, but these diseases are considered to be (1775),Aeschynomene spicata Poiret (1797), Desmo­ unimportant under grazing. Root-knot nematodes dium canum Schinz &Thellun g (1913, nom. illeg.). (Meloidogyne arenaria, M. incognita) can also Vernacular names Kaimi, kaimi clover, creep­ cause damage. ing beggar weed (En). Pega-pega (Port). Since hetero can withstand heavy grazing, it has Origin and geographic distribution Original­ the ability to persist in mixtures with vigorous ly from southern United States to Uruguay and creeping grasses where twining and shrubby Argentina, this species is now widely distributed legumes would be grazed out. However, persis­ in the wet tropics and some areas of the wet tence has been poor on soils of poor fertility, par­ subtropics. It is abundant onvolcani c soilsi n Haw­ ticularly those deficient in trace elements, and in aii, Fiji, Vanuatu and other Pacific Islands. ungrazed or lightly grazed stands with tall-grow­ Uses A potentially useful pasture species in ing grasses such as guinea grass (Panicum maxi­ mixed grass/legume pastures, especially under mum Jacq.). Cattle grazing north-eastern Austra­ conditions of heavy grazing or low soil fertility. lian pastures based on hetero and either Digitaria Reported to be a promising pasture species in Fiji eriantha or Brachiaria decumbens Stapf had live- and Vanuatu but not widely recommended else­ weight gains of over 750kg/h a per year. where in the Pacific region. Genetic resources and breeding Collections Properties Kaimi is not toxic to livestock, but of D. heterophyllum are held by ATFGRC (CSIRO, palatability is reduced by tannins. Nitrogen con­ Australia) and CIA T (Colombia). No breeding has centrations of 2.0-2.5% have been reported. been carried out in this species. Botany Ascending to erect perennial herb or Prospects The relatively unusual ability of low shrub to 60 cm tall, rarely to 3 m, but in a hetero to withstand and spread under heavy graz­ grazed pasture situation it is typically prostrate ing and its tolerance of shade suggests that it is and less than 20 cm tall. Stem trailing, fibrous to likely to be more widely used in the humid tropics. woody, sometimes beneath the soil surface, root- Literature |1|Bogdan , A.V., 1977. Tropical pas­ ture and fodder plants. Longman, London, pp. 343-344. |2| Halim, R.A. (Editor), 1989. Grassland and forage production in South-east Asia. Pro­ ceedings of first meeting of the regional working group on grazing and feed resources of South-east Asia, 27 Feb - 3 Mar, 1989. FAO, Rome. 216 pp. |3j Knaap-van Meeuwen, M.S., 1962. Preliminary revisions of some genera of Malaysian Papiliona- ceae 5. - A census of the genus Desmodium. Rein- wardtia 6: 251-252. |4| Lenné, J.M. & Stanton, J.M., 1990. Diseases of Desmodium species. Trop­ ical Grasslands 24: 1-14. |5| Ohashi, H., 1973. The Asiatic species of Desmodium and its allied genera (Leguminosae). Ginkgoana 1: 239-241. |6| Par­ tridge, I.J. , 1986. Effect of stocking rate and super­ phosphate level on oversown fire climax grassland ofmissio n grass (Pennisetum polystachion) in Fiji. Tropical Grasslands 20:166-173. |7iSkerman , P.J. , Cameron, D.G. &Riveros , F., 1988. Tropical forage legumes. FAO, Rome. pp. 271-274. J.B. Hacker &J.K . Teitzel

Desmodium incanum DC.

Prodr. 2:33 2(1825) . LEGUMINOSAE 'In = 22 Desmodium incanum DC. - 1, habit leafy plant; 2, Synonyms Hedysarum racemosum Aublet flowering branch; 3,flower; 4,fruit; 5, seed. DESMODIUM 113 ing readily at the nodes; root system deep, well- considered essential. It grows with a range of pas­ branched. Leaves trifoliolate, stipulate, petiolate; ture grasses,bu t isusuall y associated with stoloni- stipules at least partially connate abaxially, 3-11 ferous or rhizomatous species. mm x 1-3 mm, long persistent; petioles up to 3.5 D. incanum is reported to be sensitive to Peanut cm long; leaflets very variable, mostly elliptical, Mottle Virus and may act as a source for this virus terminal leaflets 2-9 cm x 1.5-4.5cm , lateral ones for cultivated groundnuts in the United States. up to 6c m x 3cm ; leaflets on upper leaves tending Several fungal diseases have been recorded on D. to be narrower than those on lower leaves, dark incanum, as well as little leaf and desmodium mo­ green on the upper surface and often with a paler saic virus, but they rarely cause serious problems. streak along the midrib, paler and densely pilose Seedlings may be damaged by cutworms, and in beneath. Inflorescence a terminal or axillary Hawaii the rose beetle and the cyst nematode raceme, up to 20 cm long; flowers solitary or in fas­ Heterodera trifolii may cause damage. cicles of 3; pedicel 4-10 mm long; flower blue, red The stoloniferous habit of this species enables it or purple, the standard up to 6 mm long. Pod up to withstand heavy grazing. A 30-40 day rest peri­ to 4 cm long, the upper margin straight, the lower od between grazings has been recommended, but strongly indented, covered with hooked hairs, the species can persist under close continuous with up to 8article s of 3.5-5 mm x 2-3.5 mm. Seed grazing. It is amongst the more persistent legume oblongoid to kidney-shaped, 1m m x 0.5 mm, light species under heavy grazing. D. incanum can yield brown. up to 6500 kg/ha of DM but is normally considered D. incanum is a very variable species and has been to be low-yielding. Seed yields may exceed 200 kg/ described many times; hence its nomenclature is ha, but lower yields are more usual. Under lenient quite complicated. The texture of the leaflets and defoliation, and particularly with high P input, it the characters of the fruits are less variable. is reported to be lower-yielding and to fix less N Initial growth is slow, but later growth rates than the more widely grown D. intortum (Miller) improve, provided moisture is adequate. Trailing Urban. stems begin to develop some 6month s after sowing. Genetic resources and breeding No cultivars In regions with a pronounced seasonal climate, are available. Collections are maintained at growth during the dry season is slow. At higher ATFGRC (CSIRO, Australia), EMBRAPA (Bra­ latitudes, flowering occurs when daylengths shor­ zil), CIAT (Colombia), and USDA (Fort Pierce, ten. It is naturally spread by animals and humans, Florida, United States). D. incanum is self-fertile, the hooked hairs on the pod adhering to fur or clo­ but some outcrossing is believed to occur. It has thing. been hybridized with D. uncinatum (Jacq.) DC, Ecology D. incanum grows best in regions with but the hybrid is sterile. annual rainfall of 1500-3000 mm, but will persist Prospects D. incanum is a species which could and spread in areas receiving 1000 mm. Optimum warrant further investigation for South-East temperatures are reported to be 30°/25°C during Asian pastures since it tolerates heavy grazing or the growing season. The species is tolerant of light cutting. Early results from Hawaii were extremely frosts and temporary flooding. In regions where it promising, but interest in the species has not been is naturalized, it is particularly common along maintained. roadsides, wasteland and other disturbed ground. Literature '1| Cameron, D.G., Jones, R.M., Wil­ D. incanum is apparently best adapted to fertile, son, C.P.M., Bishop, H.G., Cook, B.G., Lee, CR. & neutral to slightly alkaline soils,bu t may be grown Lowe, K.F., 1989. Legumes for heavy grazing in on a wide range of soil types from sands to light coastal subtropical Australia. Tropical Grass­ clays, pH 4.0-8.0. lands 23: 153-161. |2! Imrie, B.C., Jones, R.M. & Agronomy D. incanum is propagated by seed. Kerridge, P.C., 1983. Desmodium. In: Burt, R.L., Seed is often hard and may require treatment with Rotar, P.P., Walker, J.L. &Silvey , M.W. (Editors): concentrated sulphuric acid for 10minute s before The role of Centrosema, Desmodium and Stylo- sowing. It is recommended that the seed be inocu­ santhes in improving tropical pastures. Westview lated with the special Desmodium strain oi Brady- Press, Boulder, Colorado, United States, pp. rhizobium. It requires a well-prepared seed-bed for 97-140. 3 Lenné, J.M. &Stanton , J.M., 1990. Dis­ the most reliable establishment but may also be eases of Desmodium spp. - a review. Tropical broadcast into an existing pasture. Recommended Grasslands 24: 1-14. ;4i Nicolsen, D.H., 1978. Ille­ sowing rate is 5 kg/ha at ca. 5 mm depth. It res­ gitimate 'basionyms', impact on priority and au­ ponds to lime and P, but these are not normally thor citation or, the rise of Desmodium incanum 114 FORAGES and fall of D. canum (). Taxon 27(4) 365-370. |5| Schubert, B.G., 1980. Desmodium. In: Dwyer, J.D. (Editor): Flora of Panama. Part 5: Family 83, Leguminosae subfamily Papilionoi- deae. Annals ofth e Missouri Botanical Garden 67: 640-643. |6|Skerman , P.J., Cameron, D.G. & Rive- ros, F., 1988.Tropica l forage legumes. FAO, Rome, pp. 263-267. J.B. Hacker

Desmodium intortum (Miller) Urban

Symb. Antill. 8:29 2(1920) . LEGUMINOSAE 2w = 22 Synonyms Hedysarum intortum Miller (1768), Desmodium aparines (Link) DC. (1825). Vernacular names Greenleaf desmodium (En). Pega-pega (Sp). Philippines: karikut-ritkut (Ba- gobo). Thailand: thua kleen leap. Origin and geographic distribution Greenleaf desmodium is native to the Americas, from south­ ern Mexico to as far south as southern Brazil. Fol­ lowing widespread testing as a forage legume, it is now naturalized in small areas of the higher rainfall subtropics and elevated tropics. In Souht- East Asia it occurs most in Papua New Guinea, the Philippines and Thailand. Uses Greenleaf desmodium is mostly utilized as a pasture legume in mixed sowings with grasses. Desmodium intortum (Miller) Urban -1, flowering It can also be cut for use as fresh fodder or for hay and fruiting branch; 2,flower; 3, fruit. or made into silage. Properties Nitrogen concentrations in top to 30 cm long; flowers pink to purple, 8 mm long, growth range from 2-4.2%. In one sampling the borne in pairs within the axils of caducous bracts. N concentration in leaves (3.7%) was double that Pod 15-50 mm x 3-4 mm, curved, up to 12-articu- of the stems (1.7%). Phosphorus concentrations in late, covered with short hooked hairs and hence leaf material range from 0.10-0.45%. Tannin adhering to animals and clothing, slightly levels in leaves range from 3-9% which may indented along the upper margin, more strongly account for the somewhat low in vitro digestibility indented along the lower margin between the 3-7 range of 55-60%. mm long articles, breaking up at maturity. Seed Botany A robust taprooted perennial herb with reniform, 2m m x 1.3 mm. trailing stems and stolons to several metres long. D. intortum is closely related to D. uncinatum Stems form roots if in contact with moist soil and (Jacq.) DC, together forming a complex which is may scramble, but not twine, over the surrounding taxonomically not well separated and in which vegetation; they are grooved, often reddish in col­ more names are involved (e.g. D. aparines ((Link) our, densely covered with short hooked hairs, DC). glandular, sticky to the touch. Leaves with three Greenleaf desmodium is self fertile but flowers leaflets; petiole up to 5 cm long; leaflets usually may require tripping for pollination to occur. ovate, 3-12 cm x 1.5-7 cm, the lateral ones Seedlings develop slowly and require favourable smaller than the terminal, often with sparse conditions during the first weeks of growth. reddish-brown marks on the upper surface, cover­ Once established, greenleaf desmodium makes vig­ ed with ascending hairs on both surfaces. Inflores­ orous growth. In the higher latitude tropics and cence a quite dense terminal or axillary panicle up in the subtropics it has a long growing season DESMODIUM 115 before flowering late in the wet season. It does not ronment, 'Greenleaf may only persist for 5-10 spread readily from seed but individual plants can years. spread quite a long distance by means of stolons. 'Greenleaf is usually grazed by cattle, although Well-known cultivars are 'Greenleaf (Australia) it can be used in a cut-and-carry system. Annual and 'Tengeru' (Tanzania). DM yields of 12-19 t/ha have been recorded from Ecology Greenleaf desmodium usually requires 'Greenleaf grown in pure stands or in mixtures more than 1100 mm annual rainfall. It can be with grasses. Liveweight gains from 'Greenleaf/ grown at sea-level in the subtropics, but favours grass pastures usually range from 250-600 kg/ha elevated areas (500-2000 m) in the tropics. Hence per year, although higher gains have also been it has shown promise in the wetter Australian recorded. Given low grazing pressure, liveweight subtropics and in elevated tropical areas as in gains of over 200kg/hea d per year can be obtained Papua New Guinea, Thailand and the Philippines. from good 'Greenleaf' pastures. Milk production It is tolerant of shade, but not adapted to the low­ from Jersey cows grazing pure 'Greenleaf' was land tropics where the major plantation crops are only 7.7 kg/day. This may have been associated grown. It is susceptible to heavy frosts and also to with difficulties in achieving a high intake from extended dry spells during the growing season. It the open sward; higher milk production figures will grow satisfactorily on a range of soils, from have been obtained from 'Greenleaf/grass pas­ sandy soils to clay loams, provided the pH(H20) tures. is not less than 5.0 (preferably 5.5) and they are Genetic resources and breeding Germplasm not saline. It is intolerant of fire. collections are available at ATFGRC (CSIRO, Agronomy 'Greenleaf is sown by seed into a Australia) and CIAT (Columbia). There is little well-prepared seed-bed, or more rarely established variation within the species in most agronomic by rooted cuttings. Seed may be drilled to a depth characters, but there is variation in resistance to not exceeding 10 mm, or broadcast; seeding rates nematodes. This suggests that, where this is a are generally 1-2 kg/ha. It does not establish satis­ problem, genetic advances can be made. Breeding factorily when oversown into an existing pasture. in Australia to improve seedling vigour, by hybri­ Mechanically-harvested seed exhibits negligible dization with the related D. sandwicense E. Meyer hard-seededness, but hand-harvested seed may and selection for large seed size, was discontinued. benefit from treatment with sulphuric acid for five Prospects Greenleaf desmodium has a restrict­ minutes, followed by washing. Seed should be ed range of adaptation and will be further restrict­ inoculated with a specific 'desmodium' culture of ed by its inability to withstand sustained heavy Bradyrhizobium. It is generally sown with a com­ cutting or grazing. However, within the limited panion grass such as Setaria sphacelata (Schu­ areas and farm systems to which it is suited, it is macher) Stapf & C.E. Hubbard ex M.B. Moss and a productive legume. sometimes together with other tropical pasture Literature !1: Bogdan, A.V., 1977. Tropical pas­ legumes. Greenleaf desmodium requires moderate ture and fodder plants. Longman, London, pp. to high levels of P, S, K and Mo for growth. It has 344-348. 2 Imrie, B.C., Jones, R.M. & Kerridge, a higher requirement for Mo than most tropical P.C., 1983.Desmodium . In: Burt, R.L., Rotar, P.P., legumes. Walker, J.L. &Silvey , M.W. (Editors): The role of Several fungal diseases have been reported on Centrosema, Desmodium and Stylosanthes in greenleaf desmodium but usually they have had improving tropical pastures. Westview Press, minimal impact on persistence and production. Boulder, Colorado, United States, pp. 97-140. 3 Little leaf, caused by a mycoplasma-like organism Jones, R.M., 1989.Productivit y and population dy­ also occurs but usually only affects isolated namics of silverleaf desmodium (Desmodium unci- plants. In Australia, the most important pests are natum) and greenleaf desmodium (Desmodium weevil larvae of Amnemus and Leptopius spp. intortum) and two D. intortum x D. sandwicense which damage or even sever taproots and larger hybrids at two stocking rates in coastal south-east adventitious roots, making 'Greenleaf more sus­ Queensland. Tropical Grasslands 23: 43-55. i4i ceptible to soil moisture stress. Lenné, J.M. &Stanton , J.M., 1990.Disease s of Des­ It will usually be eliminated from a pasture by fre­ modium species. Tropical Grasslands 24: 1-14. 5 quent close cutting or heavy grazing, as this Skerman, P.J., Cameron, D.G. &Riveros , F., 1988. removes axillary and terminal buds and reduces Tropical forage legumes. FAO, Rome. pp. 274-282. the plant's ability to recover. Even with appro­ J.B. Hacker priate grazing management and a suitable envi­ 116 FORAGES

Desmodium triflorum (L.) DC.

Prodr. 2:33 4(1825) . LEGUMINOSAE 2n = unknown Synonyms Hedysarum triflorum L. (pro parte) (1753),Desmodiumparuifolium Blanco (1845), Mei- bomia triflora (L.) Kuntze (1891). Vernacular names Three-flowered beggarweed (Hawaii) (Am).Indonesia : daun mules (Indonesian), jukut jarem (Sundanese), delilan (Javanese). Malaysia: rumput barek sisek putih, sisek tenggil- ing. Philippines: kaliskis-dalag (Tagalog), himbis- puyo (Visaya), gumadep (Ifugao). Cambodia: smau kaè lolook.Thailand : ya-klethoi,ya-tanho i (central), ya-tansai (norhtern). Vietnam: trang qu'a ba hoa. Origin and geographic distribution D. tri­ florum is pan-tropical. In the Asian region, its dis­ tribution includes India, Sri Lanka, Burma, Thai­ land, Indo-China, China, Taiwan, Malaysia, Indonesia, the Philippines, Papua New Guinea, Australia and the Pacific Islands. Uses D. triflorum is widespread as a naturally occurring forage component of short grasslands. It is also used as a green manure and ground cover. Properties Nitrogen concentrations of 2-3% have been recorded. Dry matter production is low. It isresistan t to heavy grazing. Botany Mat-forming prostrate annual or peren­ Desmodium triflorum (L.) DC. - 1, habit; 2,part of nial herb, with woody rootstocks and long thick­ branch; 3,flower; 4, fruit. ened taproot; stem 8-50 cm long, much branched, sometimes rooting at the nodes, white hairy. collections are present at ATFGRC (CSIRO, Aus­ Leaves trifoliolate, occasionally unifoliolate; leaf­ tralia) and CIA T (Colombia). lets obovate-oblong, obovate or obcordate, 4-14 Prospects Because D. triflorum produces so lit­ mm x 4-12 mm, rounded to emarginate at the tle dry matter, it is unlikely to be considered in apex, rounded to cuneate at the base, more or less plant improvement programmes. hairy beneath. Flowers 1-3 in leaf axils, about 5 Literature |1| Ohashi H., 1973.Th e Asiatic spe­ mm long; pedicel 5-10 mm long; corolla usually cies of Desmodium and its allied genera. (Legumi­ pink to purple. Fruit flat, 6-18 mm x 2-3 mm, 3-5 nosae) Ginkgoana 1: 245-249. •%•• Skerman P.J., articled, indéhiscent, glabrous to uncinulate- Cameron, D.G. &Riveros , F., 1988.Tropica l forage pubescent; the upper suture straight, the lower legumes. FAO, Rome. pp. 285-286. !3| Verdcourt, margin constricted between the articles which are B., 1979.Amanualo f New Guinea legumes. Botany 2-3.5 mm long and wide with a strong reticulation Bulletin No 11. Office of Forests, Division of Bota­ ofraise d nerves. Seed quadrangular to nearly orbi­ ny, Lae, Papua New Guinea, p.409 . cular in outline, ca. 1.2 mm x 1.7 mm. B.C. Pengelly Ecology D. triflorum is widespread throughout the wet tropics and appears to be well adapted to a wide range of soils. It is usually found in heavily Desmodium uncinatum (Jacq.) DC. grazed or closely cut areas. Agronomy D. triflorum is resistant to heavy Prod. 2:33 1(1825) . grazing, but itproduce s little drymatter . However, LEGUMINOSAE it provides forage of reasonable N status to ani­ 2M = 22 mals. It is not cultivated. Synonyms Hedysarum uncinatum Jacq. (1798), Genetic resources and breeding Germplasm Meibomia uncinata (Jacq.) Kuntze (1891). DESMODIUM 117

Vernacular names Silverleaf desmodium, sil- hooked hairs, sticky to the touch. The stems form verleaf Spanish clover (En). Indonesia: semanggi. roots if in contact with moist soil and may scram­ Philippines: pega-pega (Tagalog), koer (Bagobo). ble but not twine through the surrounding vegeta­ Thailand: enieo. tion. Leaves with 3leaflets ; petiole up to 5c m long; Origin and geographic distribution Silverleaf leaflets usually ovate, 2-10 cm x 1-4.5 cm, dark desmodium is indigenous to the Americas, from green with a characteristic narrow elliptical sil­ northern Argentina to Mexico. Since first released very shiny area about the midrib above, covered as a pasture plant in 1962,i t now occurs in isolated on both surfaces with ascending hairs. Inflores­ localities in the more humid regions ofth e subtrop- cence a terminal or axillary raceme up to 50 cm ics and elevated tropics, including South-East long, sometimes branching at lower floral nodes; Asia. flowers pink, 7-12 mm long, borne in pairs within Uses Silverleaf desmodium is used as a pasture the axils of caducous bracts. Pod 10-50 mm x 3-4 legume in combination with a range of tropical mm, curved, upt o 12-articulate, covered with short pasture grasses. It can be made into hay. hooked hairs and hence adhering to passing ani­ Properties Nitrogen concentrations in whole mals and clothing, slightly indented on the upper top growth have ranged from 1.9-3.7% , and N con­ margin and deeply indented along the lower mar­ centrations in the leaves were approximately dou­ gin between the 5 mm long articles, breaking up ble those of the stems. Tannin levels of 3.6% have at maturity. been measured in the leaves. Silverleaf desmodium is not vigorous in the seed­ Botany Robust perennial herb with trailing ling stage, but in warm conditions and in the stems to several metres long, ascending to about absence ofmoistur e stress, grows vigorously there­ 1 m at flowering, densely covered with short after. In the subtropics and elevated tropics, it flowers at the end of the wet season, earlier than Desmo­ dium intortum (Miller) Urban. It is self-fertile, but flowers may require tripping. The only cultivar of this species is 'Silverleaf, released in Australia in 1962. Taxonomically D. uncinatum belongs to a not well separated complex ofspecie s (including the closely related D. intortum). Ecology Silverleaf desmodium requires an an­ nual rainfall exceeding 1000 mm, well distributed through the growing season, and has some toler­ ance offloodin g and shallow water tables.I t grows at sea-level in the subtropics but favours eleva­ tions of500-200 0 mi nth e tropics. It has little toler­ ance ofsalinit y or moisture stress during the grow­ ing season, but is more cold-tolerant than most other tropical pasture legumes. It will grow on a range of soils from sands to clay loams, but not heavy clays. It requires a soil of at least pH(H20) 5.0, preferably 5.5. It will recover from moderate fires, but burning is not recommended. Agronomy Ideally, 'Silverleaf should be sown into a well-prepared seed-bed, but it has also suc­ cessfully been sod-seeded into an existing pasture following treatment with a herbicide. It is nor­ mally sown with pasture grasses such as Setaria sphacelata (Schumach.) Stapf & Hubbard ex M.B. Moss. Seed should be inoculated with a special Bradyrhizobium strain to ensure effective nodula- Desmodium uncinatum (Jacq.) DC. - 1, flowering tion. 'Silverleaf has a requirement for P, K, S and and fruiting branch; 2,flower; 3, fruit. Mo, and these should be applied where they are 118 FORAGES naturally in short supply. Dichanthium annulatum (Forssk.) Several fungal diseases have been reported on 'Sil- Stapf verleaf, but they have usually not restricted its production and persistence. Little leaf, caused by Fl. Trop. Afr. 9:178(1917) . a mycoplasma-like organism also occurs, but GRAMINEAE usually only affects isolated plants. It is also sus­ In = 20 (diploid), 40 (tetraploid), 60 (hexaploid) ceptible to root-knot and other nematodes. In Aus­ Synonyms Andropogon annulatus Forssk. (1775). tralia, the most important pests are weevil larvae Vernacular names grass, sheda grass of Amnemus and Leptopius species, which damage (En). Philippines: alabang-grass. Thailand: ya or even sever taproots and larger adventitious hindi, ya waen. Vietnam: song tha'o to. roots, increasing its susceptibility to moisture Origin and geographic distribution D. annu­ stress. latum is a grass occurring in tropical and North Frequent cutting or grazing to < 10 cm will result Africa, through the Middle East to Malesia, south­ in disappearance of the legume from a pasture. ern China, and the Pacific Islands. It has been Even with appropriate grazing management and introduced to southern Africa, tropical America a suitable environment, 'Silverleaf may only per­ and Australia. In the it is sist for 3-10 years. Herbage is normally grazed. mainly found between latitudes 8° and 28°. The species is not always particularly palatable to Uses An excellent and widely used fodder grass stock, which may take some time to get used to it. which is sought out by stock from a mixture of Annual DM yields of 1.6-15 t/ha have been grasses and is preferred to all others. In India it recorded from 'Silverleaf' and 'Silverleaf/grass is promising for grazing or hay-making after pastures. Studies have shown that nodulated 'Sil­ reseeding into degraded grassland. verleaf in a grass/legume pasture can increase soil Properties Nitrogen concentrations of D. annu­ nitrogen by 90-160 kg/ha per year. latum range from 0.4-1.7%, depending on the Genetic resources and breeding Germplasm stage of growth. It can support 7 sheep per ha in collections are held by ATFGRC (CSIRO, Austra­ India. It gives a good 'standing hay'. lia) and CIAT (Colombia). There have been no Botany A perennial grass, often tufted, spread­ breeding programmes with this species. ing and stoloniferous, with erect, branched or rar­ Prospects Silverleaf desmodium has a restrict­ ely geniculate culms up to 1 m tall, hairy at the ed range of adaptation and will be further restrict­ nodes. Leaf-sheath terete, 3-4 cm long, sparsely ed by its inability to withstand sustained heavy hairy at the margins; ligule truncate, 2-4 mm long, grazing or cutting. However, within the limited membranous; leaf-blade linear, 3-30 cm x 2-8 areas and farm systems to which it is suited, it is mm, often hairy above, at the margins and in the a productive legume. throat. Inflorescence composed ofu p to 15subdigi - Literature 1| Bogdan, A.V., 1977. Tropical pas­ tate, shortly peduncled racemes each 3-7 cm long; ture and fodder plants. Longman, London, pp. spikelets paired, one pedicelled, one sessile, alter­ 350-354. |2| Imrie, B.C., Jones, R.M. & Kerridge, nately arranged along a filiform axis; the lowest P.C., 1983.Desmodium . In: Burt, R.L., Rotar, P.P., 1-2 pairs similar, male or neuter, awnless and per­ Walker, J.L. &Silvey , M.W. (Editors): The Role of sistent; all other pairs dissimilar and deciduous; Centrosema, Desmodium and Stylosanthes in sessile spikelet 2-5 mm long, bisexual, upper improving tropical pastures. Westview Press, lemma with a 1-2.5 cm long, kneed and twisted Boulder, Colorado, United States, pp. 97-140. |3; awn; pedicelled spikelet male or neuter; lower Jones, R.M., 1989.Productivit y and population dy­ glumes pubescent to villous in lower half, upper namics of silverleaf desmodium (Desmodium unci- half with long bulbous-based hairs especially on natum) and greenleaf desmodium (Desmodium the margins. Caryopsis obovoid. intortum) and two D. intortum x D. sandwicense D. annulatum is a very variable species, morpholo­ hybrids at two stocking rates in coastal south-east gically as well as cytologically. Several varieties Queensland. Tropical Grasslands 23: 43-55. -4| have been distinguished, mainly based on size and Lenné, J.M. &Stanton , J.M., 1990.Disease s of Des­ number of spikes, on hairiness ofth e spikelets, and modium species. Tropical Grasslands 24: 1-14. !5 on ploidy level. Skerman, P.J., Cameron, D.G. &Riveros , F., 1988. Flowering commences early in the growing season Tropical forage legumes. FAO, Rome. pp. 286-296. and continues throughout the year. In India, rec­ J.B. Hacker ommended selections include 'Marvel 8', and lines 485, 487 and 490. DlGITARIA 119

Genetic resources andbreedin g It is unlikely that substantial germplasm collections are being maintained. In India some breeding programmes on D. annulatum arebein g carried out. Prospects D. annulatum will have a place in grazing lands in drier areas. It is a promising spe­ cies as it is drought-tolerant, persistent and resis­ tant to heavy grazing. The prospects of selecting leafy strains could be investigated. Literature |1|Gilliland , H.B., Holttum, E.E.& Bor, N.L., 1971. Grasses of Malaya. In: Burkill, H.M. (Editor): Flora ofMalaya . Vol3 . Government Printing Office, Singapore, pp. 282-286. |2; Häf- liger, E. & Scholz, H., 1980.Gras s weeds 1. Ciba- Geigy Ltd., Basel, Switzerland, p. 39. |3|Holm , J., 1971. Feeding tables. Nutrition Laboratory of Thai-German Dairy Project, Livestock Breeding Station, Chiangmai, Thailand, p. 25. |4| Skerman, P.J. & Riveros, F., 1990. Tropical grasses. FAO, Rome. pp.328-332 . C. Manidool

Digitaria ciliaris (Retzius) Koeler

Descr. Gram.: 27(1802) . GRAMINEAE 2re = 54 Synomyms Digitaria marginata Link (1821), D.- sanguinalis auct., non Scop. (e.g.i n Heyne, 1927), Dichanthium annulatum (Forssk.) Stapf-1, flower­ D. adscendens Henrard (1934). ing plant; 2, ligule; 3,spikelet; 4, caryopsis. Vernacular names Tropical finger grass, sum­ mer grass (Australia) (En). Crab grass (Am). Indo­ Other species of the genus, occurring in Malesia, nesia: jelamparam, suket ceker ayam (Javanese), are often also good fodder grasses, e.g. D. carico- jampang jemprak (Sundanese). Malaysia: caker sum (L.)A . Camus andD. mucronulatum Jansen. ayam, bamboo grass. Philippines: baludyangan Ecology D. annulatum is adapted to the drier (Tagalog), saka-saka (Ilokano). Thailand: yah-tin- areas ofth etropica l and subtropical zones.I t does nok, ya-plongkhaonok (central). Vietnam: tue not thrive inwaterlogge d places nori nshad y envi­ hinh leo. ronments, but is tolerant of grazing, burning and Origin andgeographi c distribution D. ciliaris saline soils. is widely distributed throughout the tropics and Agronomy D. annulatum canb e established by subtropics, especially in Asia. It is much less com­ seed or by rooted tillers planted at 60c m spacing. mon in Africa. It is a good seed producer, but slow growing at the Uses Although aba d weed in cropping areas, D. seedling stage. It is highly palatable when young ciliaris provides good forage, assists in protecting and fresh. Frequent grazing is required to main­ soil against erosion and provides material for tain it ina leafy condition, although cattle will eat mulch or compost. the stems. In Thailand, observations of a mixture Properties D. ciliaris is a palatable grass, espe­ of this species and Caribbean stylo (Stylosanthes cially when young. Penfeedin g trials inth e United hamata (L.) Taub.) on road edges indicate that it States showed that it had a similar digestibility may be possible to improve its quality as a forage and higher N concentration than sudan grass (Sor­ by oversowing with a legume. Dry matter yields ghum x drummondii (Steud.) Millsp. & Chase or of 1.4-2.0 t/ha have been obtained. It can be pre­ pearl millet (Pennisetum americanum (L.) K. served as 'standing' or cutha ybu tno t as silage. Schum. exLeeke) . 120 FORAGES

occurrence of glassy bristles, and the spacing of the nerves in the lower lemma. D. ciliaris is an agressive and opportunistic col­ onizer of bare, disturbed or newly cleared habitats due to its prolific seed production, vigorous growth and rapid development of rooted stolons. Seeds germinate in spring or summer in temperate or subtropical areas or with the onset of rains in the tropics. Ecology In the tropics, D. ciliaris grows at alti­ tudes from sea-level up to 1600 m (Indonesia) or 2000m (Papu a New Guinea).I t grows best in moist, sandy or loamy soils and is very responsive to high soil fertility. Agronomy D. ciliaris is propagated vegetati- vely or by seed. If newly ripened seed is being used for sowing, treatment with alternating tempera­ tures and scarification helps to reduce dormancy. Germination is also stimulated by exposure to light. The spikelet scales act as a barrier to oxygen entry for up to 4month s storage time. A relatively high N level in the substrate seems to be needed for optimal germination. Seed dormancy can last up to 7 months. Good seed-bed preparation and sowing rates of 2-3 kg/ha of seed help to ensure good establishment. Planting with rooted cut­ tings, at spacings of about 10 cm x 10 cm, gives more rapid establishment than sowing seed. D. ciliaris is usually eaten by grazing animals, but Digitaria ciliaris (Retzius) Koeler - 1, flowering it can be cut for stall feeding or for silage. It is tol­ plant; 2, ligule; 3, spikelets; 4, caryopsis in two erant of close cutting or grazing. It may produce views. several seed crops per year if periodically grazed or mown. Its presence in pastures is usually a sign Botany Tufted annual, rarely short-lived peren­ of newly established pastures, recent cultivation nial; culms erect, ascendent or prostrate, 50-100 or disturbance, or deterioration of perennial cmlong , often rooting at the lower nodes and form­ grasses. It is usually a minor component or is not ing loose mats.Leaf-sheat h loose, variously pilose; found in vigorous stands ofperennia l grasses. Pure ligule membranous, truncate, up to 3 mm long; stands have produced DM yields of 10-12 t/ha per leaf-blade lanceolate to linear, 2-25 cm x 3-13 year. One disadvantage of D. ciliaris is that it is mm, short bristly on both sides or glabrous. Inflor­ an alternative host of the viruses that produce escence consisting of 2-10 racemes, each up to stripe disease and black-streak dwarf disease in 15(-22) cm long, digitately arranged or in 2-3 rice. whorls; peduncle up to 40 cm long; rachis charac­ Genetic resources and breeding D. ciliaris is teristically winged, serrate, spicules at least 0.05 a very variable species, but it is unlikely that sub­ mm long; spikelets binate, homomorphous, lanceo­ stantial germplasm collections are being main­ late, 2.5-3.5 mm long, smoothly hairy; two florets tained. There are no known breeding programmes. per spikelet, the upper bisexual, the lower neutral. Prospects D. ciliaris will continue to be a useful Caryopsis oblongoid, about 2 mm long, golden natural forage in specific situations, usually asso­ brown to grey. ciated with disturbance or cropping. It is unlikely A very variable species, about which much confu­ that it will be deliberately propagated more widely sion exists in the taxonomie literature where it has than at present. In Malaysia, where D. ciliaris pre­ been given more than 50 different names. Four vails as a weed under rubber and oil palm, sheep variable characters are responsible: the hairs on grazing will reduce its weed impact. the rachis, the ciliate frill around the spikelets, the Literature ilj Holm, L.G., Plucknett, D.L., Pan- DlGITARI A 121

cho, J.V. & Herberger, J.P., 1977. The world's worst weeds: distribution and biology. The East- West Center, University of Hawaii Press, Hono­ lulu, Hawaii, United States, pp. 249-253. |2| Sker- man, P.J. & Riveros, F., 1990. Tropical grasses. FAO, Rome. pp. 348-350. |3|Soerjani , M., Koster- mans, A.J.G.H. &Tjitrosoepomo , G., 1987.Weed s of rice in Indonesia. Balai Pustaka, Jakarta, pp. 402-404. i4j Veldkamp, J.F., 1973. A revision of Digitaria Haller (Gramineae) in Malesia. Blumea 21: 32-35. B.B. Baki, C.P.Che n &LB . Ipor

Digitaria eriantha Steudel

Flora 12:46 8(1829) . GRAMINEAE In = 18, 27, 36, 45, 50, 54 Synonyms Digitaria smutsii Stent (1924), D. decumbens Stent (1930), D.pentzii Stent (1930), D. valida Stent (1930),D. pentzii Stent var. stolonifera (Stapf) Henrard (1950); D. eriantha Steudel with 4 subspecies:ssp .eriantha, ssp.pentzii (Stent) P.D.F. Kok (1981), ssp. stolonifera (Stapf) P.D.F. Kok (1981),ssp . transvaalensis P.D.F. Kok(1981) . Vernacular names Common finger grass, digit grass (also used for other spp.),pangol a or pongola grass (Digitaria decumbens) (En). Digitaria eriantha Steudel 1, flowering plant; 2, Origin and geographic distribution D. erian­ part of raceme; 3, spikelet. tha is a species of subtropical southern Africa. A single clone, commonly referred to as pangola sward. Stolon internodes with or without hairs. grass, isno wwidesprea d in grazing areas through­ Culms erect or ascending, sometimes rooting at out the world's humid tropics and subtropics, the nodes, to 1.5 m tall. Leaf-sheath scabridulous, including South-East Asia, having been extensi­ glabrous to hairy; ligule subtriangular, 2-4 mm vely planted from the 1960s to the 1980s. Many long, shortly ciliate; leaf-blade linear, 5-60 cm x other lines have been distributed for evaluation. 1.5-12mm , glabrous orhairy , butminutel y scaber- Uses Iti sutilize d extensively asa grass for graz­ ulous on both surfaces. Inflorescence a racemose ing, mostly with N fertilization rather than a com­ panicle, composed of 3-14 erect racemes 6-18 cm panion legume. It is also utilized as aha y crop. long, borne in loose whorls and with some racemes Properties D. eriantha is often considered to be single on an axis up to 7 cm long; spikelets 2-4 one of the higher quality tropical grasses. Pangola mm long, conspicuously hairy; lower glume to 0.5 grass has been extensively studied but there is mm long, the upper § as long as the spikelet and relatively little information on other genotypes. appressed hairy; lower lemma minutely hairy, Pangola grass has an N concentration of characteristically with 7 smooth rather than sca­ (0.5-)1.5-2.0(-4)%. Phosphorus concentration brous nerves. Many clones are sterile or almost can be too low for livestock on low-P soils in the sterile. absence of P fertilizer. Dry matter digestibility Growth and development In subtropical varies between 45-70%. Pangola grass has relati­ areas, pangola grass flowers in mid-summer; flow­ vely high concentrations ofN a in its tissues, com­ ering time differs with other genotypes, with culti- pared with many other tropical grasses. var 'Premier' (ssp. eriantha in the sense of Kok) Description More or less robust perennial, flowering in late spring and again in autumn. growing either as a dense tussock, with or without Other botanical information D. eriantha is extended stolons or as a continuous stoloniferous extremely diverse but it is probable that 80% of 122 FORAGES agronomic publications are concerned with a sin­ and 'Advance' require a reasonable seed-bed for gle genotype. Performance data should therefore establishment and an absence of serious competi­ not be generalized for the species as a whole. D. tion in the early stages. Establishment is most sat­ eriantha is closely related to the tropical D. milan- isfactory on sandy to sandy-loamy soils. Once es­ jiana (Rendle) Stapf, from which it may only be dis­ tablished, however, they have the capacity to tinguished by the absence of scabrosity on nerves thicken up and spread from the sown area. D. of the lower lemma, a characteristic which may be eriantha is an aggressive species and does not gen­ difficult to determine, even with a lens. Frequently erally combine well with pasture legumes. in the agronomic literature, accessions are de­ Husbandry Once established, D. eriantha toler­ scribed simply as Digitaria sp., which is of little ates heavy stocking rates. On infertile soilsi t bene­ help when documenting species characteristics. fits from heavy fertilizer application, especially N. In the taxonomie literature, the tremendous varia­ Stoloniferous cultivars can become sodbound and bility of this species has resulted in numerous may benefit from periodic renovation by disking. names and subclassifications (see Synonyms). Diseases and pests Susceptibility to the rust Much more research is needed to unravel the D. Puccinia oahuensis a widespread disease occur­ eriantha complex. In 1981 Kok distinguished 4 ring inth e Americas and Australia, varies between subspecies for the South African taxa, but in 1984 genotypes, with 'Transvala' showing some resis­ he reduced them to synonyms of the whole species tance. The most serious disease of D. eriantha is complex. Two seed-producing cultivars, 'Premier' pangola stunt virus, a dwarfing disease which has and 'Advance', belonging to D. eriantha ssp. erian­ seriously reduced the usefulness of pangola since tha (in the sense of Kok), have recently been it was first reported in Surinam in 1960. The dis­ released in Australia. Pangola grass belongs to D. ease istransmitte d by an , Sogatella furcifera, eriantha ssp.pentzii (in the sense of Kok), and cul­ or in Australia by the related S. kolophon. 'Trans­ tivars 'Taiwan', 'Transvala' and 'Slenderstem' vala' has some resistance to this disease but trials have been developed in the United States from in Australia on a number of lines of digit grasses southern African introductions of this subspecies. failed to show any resistance. Ecology Accessions of ssp.pentzii (in the sense Insects which cause damage include spittlebugs of Kok) (pangola, 'Slenderstem') are recommended ( flavopicta, T. humeralis, Prosapia in Malaysia and the Philippines for poorly drained bicincta) in Brazil and Taiwan, Rhodes grass mealy soils. They are tolerant of flooding, and any areas bug () (also in Brazil and where the grass dies out are rapidly re-invaded by Taiwan), chinch bug ( leucopterus) in stolons from the surrounding sward. Pangola Taiwan, sugar-cane aphid ( flava) in the Ca­ grass is intolerant of shade and is therefore not ribbean region and Taiwan, army worms suitable for integrating with plantation agricul­ (Laphigma spp., Spodoptera spp. and Mods spp.), ture, but can grow on a wide range of soil types and the root-knot nematodes Belonolaimus longi- from sands to heavy clays. Cultivars 'Premier' and caudatus and Platylenchus brachyurus in the 'Advance' are adapted to sub-humid subtropical Americas. conditions and they have markedly better cool- Harvesting D. eriantha is usually grazed, but is season growth than other cultivars. In general, the also suited to cut-and-carry feeding systems or hay seed-producing lines of the species tend to be bet­ making. ter adapted to sandy loam soils as seedling estab­ Yield Herbage DM yields as high as 36 t/ha per lishment can be difficult on heavier soils. Howev­ year have been reported for pangola grass, but pro­ er, once satisfactorily established, they also grow duction is more normally in the range 11-22 t/ha vigorously on clays. per year. Beef production from N fertilized pan­ Propagation and planting Stoloniferous geno­ gola can exceed 1000 kg/ha per year. Grown with types are planted vegetatively. Actively growing tropical legumes, 'Transvala' digit grass produced swards are allowed to become stemmy and are then an average liveweight gain of 350kg/h a under con­ cut and the material spread on a cultivated surface tinuous grazing at a stocking rate of 2.6 steers/ha at 0.5-2 t of green matter per hectare. This is then in Malaysia. From ssp. eriantha (in the sense of disked in, or trampled in by cattle if the ground Kok) in South Africa, beef production ranged from is too wet for implements. In warm moist environ­ 84-110 kg/head per season at a stocking rate of 7.5 ments, pastures establish rapidly by this method animals per hectare. Milk yields of 6000 kg/ha per and in general weeds are suppressed by the planted year have been obtained from well-fertilized pan­ grass. Seed-producing cultivars such as 'Premier' gola grass pastures. DlGITARIA 123

Genetic resources Germplasm collections are ences in leaf Na concentration of 0.01-2.30% of available at ATFGRC (CSIRO, Australia). the dry matter were shown to be associated with Breeding There is a continuing interest in provenance, the high Na ecotypes being of coastal selecting and breeding D. eriantha and the closely African origin. Differences in leaf digestibility related D. milanjiana, with which it is often con­ were also demonstrated. fused. Botany Perennial with erect or geniculately as­ Prospects Future work is likely to result in the cending culms to 2.5 m tall from a loosely tufted commercialization of seed-producing cultivars more or less rhizomatous base, sometimes stoloni- which should extend the usefulness of this impor­ ferous. Culm nodes glabrous, rarely hairy. Leaf- tant species. blade 15-30 cm x 3-13 mm, the blade and sheath Literature |1| Blair, G.J., Ivory, D.A. & Evans, more or less hairy to densely villous. Inflorescence T.R. (Editors), 1986. Forages in Southeast Asian a digitate or subdigitate panicle with 2-18 and South Pacific agriculture. ACIAR Proceed­ racemes on an axis up to 6 cm long, the racemes ings Series No 12. ACIAR, Canberra. 202 pp. |2| 5-25 cm long; spikelets paired on a triquetrous Bogdan, A.V., 1977. Tropical pasture and fodder winged rachis, lanceolate, usually 2.5-3 mm long; plants. Longman, London, pp. 111-122, 124-126. lower glume 0.2-0.5 mm long,th e upper ^-f as long ;3|Halim , R.A. (Editor), 1989.Grasslan d and forage as the spikelet; lower lemma as long as the spike- production in South-east Asia. Proceedings of first let, 7-nerved, more or less hairy and with shortly meeting of the regional working group on grazing ciliate margins, the nerves characteristically sca- and feed resources in South-East Asia, 27 Feb - 3 brid and often with stiff spreading brown glassy Mar, 1989.FAO ,Rome .216pp . |4|Kok , P.D.F.,1984 . bristles. Caryopsis ellipsoid, greyish-brown. Studies on Digitaria () 1. Enumeration of In subtropical latitudes D. milanjiana flowers species and synonymy. South African Journal of Botany 3: 184-185. j5| Oram, R.N., 1990. Register of Australian herbage plant cultivars. CSIRO, Australia, pp. 105-106. J.B. Hacker

Digitaria milanjiana (Rendle) Stapf

Fl. Trop. Afr. 9:43 0 (1919). GRAMINEAE 2re = 18, 36,5 4 Synonyms Digitaria swynnertonii Rendle (1911), D. endlichii Mez (1921), D. setivalva Stent (1924),D. mombasana CE. Hubbard (1926), D.pole- vansii Stent ssp.peterana Henrard (1950). Vernacular names Digit grass, (woolly) finger grass, Milanje finger grass (En, not restricted to this species).Thailand : ya mardi digit (for 'Mardi'). Origin and geographic distribution D. milan­ jiana naturally occurs in tropical East and south­ ern Africa, from Ethiopia and Somalia southwards to South Africa. It has been introduced as a fodder grass to other tropical areas, including several countries in South-East Asia. Uses D. milanjiana is used as a forage for graz­ ing by cattle and goats. It has also proved to be useful in banana plantations in tropical Australia for the control of burrowing nematodes. Properties Nitrogen concentrations of the cul- Digitaria milanjiana (Rendle) Stapf- 1, habit leafy tivar 'Mardi' ranging from 1.3-3.0% have been culm; 2, inflorescence; 3,dorsal view of two spikelet recorded in Malaysia. In Australia, genetic differ­ types; 4,ventral view of two spikelet types. 124 FORAGES freely during the growing season. In the tropics, should be grazed relatively frequently to prevent flowering time appears to be more seasonal and them from becoming too stemmy. A grazing trial seed set is often poor. It is extremely variable in in Malaysia on 'Mardi' indicated that higher stoc­ growth habit and with regard to presence or king rates could be maintained better on rotatio- absence of setae (spiny bristles) on the lemmas. nally grazed pastures than on set-stocked pas­ Genotypes which have these setae are frequently tures. The optimal stocking rate in Malaysia, referred to as D. setivalva. The essentially tropical when fertilized with 150 kg/ha per year of N, is D. milanjiana may be distinguished from related 20-40 goats/ha, giving a liveweight gain of45 0 kg/ subtropical taxa (e.g. D. eriantha Steudel) by the ha per year. Also in Malaysia, liveweight gains of presence of scabrosity along the outer nerves of cattle of from 700-1200 kg/ha per year have been the lower lemma. Due to the extreme variability, recorded from the same cultivar given N fertilizer, the taxonomy of D. eriantha is complicated. In the although with a high level of year-to-year varia­ literature, both narrow and wide species concepts tion. Milk yields of 16.4kg/da y have been obtained have resulted in numerous names and subdivi­ from Friesian cows grazing a genotype selected for sions. More research is needed to solve all prob­ high leaf digestibility in Australia, in the absence lems of this complex. of any supplementary feeding. Cultivar 'Mardi' has been released in Malaysia Genetic resources and breeding Germplasm and 'Jarra ' in tropical Australia. collections are held by ATFGRC (CSIRO, Austra­ Ecology D. milanjiana occurs naturally in a lia). Selection trials have identified promising wide range of habitats which are mostly subject accessions in Australia and Malaysia. Experimen­ to some degree of disturbance, at altitudes up to tal breeding studies in Australia have contributed 2000 m, with an annual rainfall of 400-800 mm. It to an understanding of variation in a number of is well-adapted to the humid lowlands of Malaysia morphological, developmental and chemical attri­ and cultivar 'Mardi' (generally referred to as D. butes. setivalva) issuitabl e for peat and alluvial soils. The Prospects D. milanjiana is a species with con­ cultivar is recommended for smallholdings and siderable potential. Its extreme variability leads to commercial enterprises on well-drained, fertile the possibility of developing improved cultivars soils and can tolerate brief periods of moisture for both the wet and seasonally dry tropics. Culti­ stress. It is neither shade-tolerant nor tolerant of vars with higher seed yields than 'Mardi' are waterlogging. It is reported to combine well with needed and there are reasonable prospects for Leucaena leucocephala (Lamk) de Wit on alluvial achieving this as seed production is very variable soils in Malaysia. between accessions. Ecotypes have been shown to differ in seed-dor­ Literature 1: Clayton, W.D. & Renvoize, S.A., mancy depending on provenance. Accessions from 1982.Graminea e (Part 3).In :Polhill , R.M. (Editor): regions with a protracted dry season have an Flora of tropical East Africa. A.A. Balkema, Rot­ extended period ofdormanc y which may be broken terdam, pp. 647-649. 2! Hacker, J.B., 1988. Poly­ by high temperature. ploid distribution and seed dormancy in relation Agronomy 'Mardi' is vegetatively propagated. to provenance rainfall in the Digitaria milanjiana Stolons are planted into a cultivated seed-bed and, complex. Australian Journal of Botany 36: provided the seed-bed is kept moist, intervening 693-700. ;3| Halim, R.A. (Editor), 1989. Grassland spaces are rapidly filled in. Vegetative propaga­ and forage production in South-East Asia. Pro­ tion should be carried out in the first half of the ceedings of first meeting of the regional working growing season, so as to allow sufficient time for group on grazing and feed resources in South-East the cuttings to establish. Asia, 27 Feb - 3 Mar 1989. FAO, Rome. 216 pp. ;4! Some accessions of D. milanjiana produce ade­ Lowe, K.F., Moss, R.J., Cowan, R.T., Minson, D.J. quate quantities of seed for commercial seed pro­ & Hacker, J.B., 1991.Selectin g for nutritive value duction. The conditions necessary for good estab­ in Digitaria milanjiana. 4. Milk production from lishment from seed are likely to be similar to those an elite genotype compared with Digitaria erian­ required by other small-seeded tropical grasses. tha ssp. pentzii (pangola grass). Australian Jour­ D. milanjiana issusceptibl e to pangola stunt virus, nal of Experimental Agriculture 31: 603-608. but this is not a major problem in Malaysia. J.B. Hacker & C.C.Wong Annual DM yields of 25 t/ha have been recorded in Malaysia for 'Mardi' in trials which were har­ vested every 6 weeks. Swards of D. milanjiana ECHINOCHLOA 125

Echinochloa colona (L.) Link

Hort. Berol. 2:20 9(1833) . GRAMINEAE In = 36,48 , 54,7 2 Synonyms Panicum colonum L. (1759). Echino­ chloa crus-galli P. Beauv. ssp. colona (L.) Honda (1923).Th e specific epithet is also spelled 'colonum'. Vernacular names Jungle rice, awnless barn­ yard grass (En). Indonesia: rumput jajagoan kecil (Sundanese, Javanese), rumput bebek (Indone­ sian), tuton (Javanese). Malaysia: padi burung, rumput kusa-kusa. Philippines: pulang-puwit (Tagalog), guinga (Visaya), dakayang (Ilokano). Burma: myet-thi, pazun-sa-myet. Laos: khauz nôk. Thailand: ya noksichomphu (central), ya-nokkhao (northeast). Vietnam: co' long vu'c. Origin and geographic distribution E. colona is widely distributed throughout the tropics and subtropics, including South-East Asia. Uses E. colona is used as forage and is grazed by all kinds ofstock . During famines, the seeds are eaten by humans. In rice fields, jungle rice can be a serious problem because the young plants closely resemble young rice plants. It can also be a weed in other crops. Properties Nitrogen concentrations of E. co­ lona range from 1.0-2.5%, depending on the growth stage. Echinochloa colona (L.) Link - 1,flowering plant; Botany A tufted annual grass with erect or as­ 2, ligule; 3,two views of'a spikelet. cending culms up to 1 m tall, often rooting at the lower nodes. Leaf-sheath 3-4 cm long, glabrous, year, commencing 3-4 weeks after emergence, and often reddish; ligule absent; leaf-blade linear-acu­ seeds abundantly. minate, 5-30 cm x 2-8 mm, sometimes marked Ecology E. colona grows in swampy places in en­ with purple bars. Inflorescence up to 15 cm long, vironments receiving from 400t o 1200m m rainfall composed of 3-10 short racemes up to 3 cm long; per year and at altitudes ranging from sea-level to racemes neatly 4-rowed, simple, commonly half 2000 m. It is adapted to full sunlight or partial their length apart and appressed to the axis but shade and grows on loam, silt and clay soils, but sometimes subverticillate and spreading; spikelets it does not tolerate dry periods. numerous, ovate-elliptical to subglobose, 1.5-3 Agronomy E. colona is established by seeds or mm long, pubescent; lower floret male or barren, rooted tillers. It requires high soil moisture levels acute to cuspidate; upper lemma 2-3 mm long, to get good establishment. It isver y palatable, even sharply pointed. Caryopsis ellipsoid to subglobose. when flowering. Light stocking rates and long The absence of a ligule, the purplish-tinged leaves intervals between grazing are recommended. It can and the neatly 4-rowed racemes are characteristic be cut 3 to 4 times during the wet season when of this species. Less constant are the awnless soft flower heads are in full bloom. The common prac­ indumented spikelets - some forms tend to inter- tice in many countries of South-East Asia is to grade with E. crus-galli (L.) P. Beauv. which has remove whole plants out of rice fields where it is awned spikelets. regarded as a weed. The collected material is then E. colona is a fast growing grass. Seeds germinate fed to the ruminants. It can be conserved as hay but quickly with the onset of the first rains and it can will take several days to dry as it is very succulent. cover other cultivated forages in the first few Genetic resources and breeding It is unlikely weeks. Axillary shoots develop during the second that substantial germplasm collections are being week after emergence. It flowers throughout the maintained. 126 FORAGES

Prospects E. coloria is a valuable grass. Selec­ tion could be made to obtain accessions that pro­ duce higher yields when grown on upland soils as these soils are easier to work than the swampy or lowland areas. Literature [1|Departmen t ofLivestoc k Develop­ ment, 1986.Repor t on feed analysis [in Thai]. Tech­ nical Bulletin No 13-0116-29. Division of Animal Nutrition, Department ofLivestoc k Development, Bangkok. |2|Gilliland , H.B., Holttum, R.E. & Bor, N.L., 1971. Grasses of Malaya. In: Burkill, H.M. (Editor): Flora of Malaya. Vol. 3. Government Printing Office, Singapore, pp. 167-168. |3| Häf- liger, E. & Scholz, H., 1980. Grass weeds 1. Ciba- Geigy Ltd., Basel, Switzerland, p. 54. |4| Mehra, K.L. & Fachrurozi, Z., 1985. Indonesian economic plant resources: forage crops. Lembaga Biologi Nasional - LIPI, Bogor, No 31.p . 14 |5| Skerman, P.J. & Riveros, F., 1990. Tropical grasses. FAO, Rome. pp. 370-372. C. Manidool

Echinochloa crus-galli (L.) P. Beauv.

Ess.Agrost. : 53,161(1812) . GRAMINEAB 2ra = 42, 48, 54 Synonyms Panicum crus-galli L. (1753). Echinochloa crus-galli (L.) P. Beauv. - 1, flowering Vernacular names Barnyard millet, chicken- plant; 2, ligule; 3, short-awned spikelet; 4, long- panic grass, cock's foot (En). Crête-de-coq, pattes awned spikelet. de poule (Fr). Indonesia: jajagoan (Sundanese), jawan (Javanese). Malaysia: padi burung, rumput linear-acuminate, 5-50 cm x 0.5-2 cm, glabrous. kekusa basar. Philippines: bayokibok (Tagalog), Inflorescence erect or nodding, 6-22 cm long, lagtom (Bikol), marapagay (Ilokano). Burma: green or purple-tinged, composed of 9-12 racemes, myet-ihi. Cambodia: smao bek kbol. Thailand: ya- 2-4(-10) cm long and sometimes branched; plonglaman, ya-khaonok (central). Vietnam: cô' racemes untidily 2-several-rowed, hispidulous on long vue, song chong. the back, scaberulous on the margins; spikelets Origin and geographic distribution Barnyard crowded, hispid, ovoid-ellipsoid, 3-4 mm long millet is widespread throughout the tropics, e.g. excluding the awns; lower floret barren with Africa, India, Burma, Thailand, Indo-China, Male- lemma acuminate or awned, awn up to 0.5-l(-5) sia and Australia, and extends to the subtropics cm long; upper lemma pointed, shiny. Caryopsis and some warm temperate regions. ovoid, 2-3 mm long, brownish, dorsally flattened, Uses Barnyard millet is used as fodder but it is with longitudinal ridges on the convex surface. also regarded as the world's worst weed in paddy An extremely variable species which frequently rice. However, in times of scarcity the grain is has been split into various varieties and forms, eaten by people. sometimes even into different species. This com­ Properties Barnyard millet is a palatable and plex still needs thorough investigation. succulent feed which produces copious quantities The species flowers over a wide range of photoper­ ofseed. iodic conditions. Flowering is quicker and more Botany Arobus t annual, tufted grass with erect abundant under short-day circumstances, but or geniculate culms, up to 1.5 tall, often rooting under long-day conditions the vegetative and and branching near the base. Leaf-sheath 9-13 cm generative parts are much more robust. long, glabrous or hairy; ligule absent; leaf-blade Ecology Barnyard millet is adapted to wet ERAGROSTIS 127

places and waterlogged land, and grows very Eragrostis tenella (L.) P. Beauv. ex vigorously in hot wet conditions from sea-level to Roemer & Schultes 2500 m altitude. It grows best in rich moist soils with high N content, but it can also thrive on Syst. Veg. 2:57 6(1817) . sandy and loamy soils. GRAMINEAE Agronomy Barnyard millet is propagated by 2n = unknown seed or rooted tillers. Sometimes the seeds have a Synonyms tenella L. (1753), P. amabilis L. dormancy period (in Japan 4-8 months; in the Uni­ (1753), Eragrostis amabilis (L.) Hook. & Arnott ted States 4-48months) . Optimum temperature for (1840). germination ranges from 32-37°C. Seeds germi­ Vernacular names Bug's egg grass, Japanese nate under water provided it is not more than 15 love-grass (En). Indonesia: suket emprit-empritan cm deep. It needs very moist conditions for estab­ (Javanese), jukut karukuan (Sundanese), luh- lishment by rooted tillers. Permanent flooding is buluhan (Madura). Laos:hnhaa z nhung. Thailand: recommended if vigorous stands are to be main­ ya harng krarork. Vietnam: xuân tha'o min. tained. As a weed in rice, fertilizer applications Origin and geographic distribution E. tenella favour the growth of E. crus-galli more than they occurs throughout the tropics of the , do the rice crop. Rice yields can be severely also in South-East Asia. It has been introduced to reduced, especially when more than 20 plants/m2 tropical America. are present. Uses E. tenella is grazed by cattle and water buf­ Hand harvesting is a common practice where the faloes in traditional feeding systems. It is a weed grass grows in swampy places. The upper parts of of minor agricultural importance. the stem above water are cut and carried to feed Properties It isregarde d as a low quality forage to animals. It can be made into hay but is difficult because of its low leaf/stem ratio. The grain is said to dry as it is very succulent. Several cuts are pos­ sible if plants are kept submerged in water. In paddy fields where barnyard millet isth e main spe­ cies, yields of 4-11 t/ha of green material are obtained. It is relished by cattle and water buffa­ loes. Genetic resources and breeding No data are available, but observations suggest that there are different strains which may occur in different hab­ itats. It is unlikely that any substantial resource collections are being maintained. Prospects As it grows in wet areas, barnyard millet can supply feed at the times when other dry­ land grasses are not available. Agronomic studies should be conducted on mixtures of rice and barn­ yard millet to find out how to optimize the yield of barnyard millet without unduly reducing the yield of rice. Literature |1| Bor, N.L., 1960. The grasses of Burma, Ceylon, India and Pakistan. Pergamon Press, London, p. 310. |2| Gilliland, H.B., Holttum R.E. &Bor , N.L., 1971. Grasses of Malaya. In: Bur- kill, H.M. (Editor): Flora of Malaya. Vol. 3. Gov­ ernment Printing Office, Singapore, pp. 168-169. !3|Holm , L.G., Plucknett, D.L., Pancho, J.V .& Her - berger, J.P., 1977.Th e world's worst weeds: distri­ bution and biology. The East-West Center, Univer­ sity of Hawaii Press, Honolulu, Hawaii, United States, pp. 32-40. |4| Skerman, P.J. & Riveros, F., Eragrostis tenella (L.) P. Beauv. ex Roemer & 1990.Tropica l grasses. FAO, Rome.pp . 373-375. Schultes - 1,flowering plant; 2, ligule; 3, spikelet; C. Manidool 4, floret. 128 FORAGES to be very nutritious. Eragrostis unioloides (Retzius) Nees ex Botany A small tufted annual grass, with erect Steudel or ascendent culms, up to 60 cm tall. Leaf-sheath 2-4 cm long, light purple, hairy at the mouth; Syn. PI. Glum. 1:26 4(1854) . ligule a ring of fine soft hairs; leaf-blade narrowly GRAMINEAE linear with broad base and acute tip, up to 13 cm 2n = unknown x 7 mm, glabrous, often tinged purple. Inflores­ Synonyms Poa unioloides Retzius (1789). cence a terminal panicle, up to 35 cm long, open, Vernacular names Indonesia: padang togu sometimes hairy at the axils, the branches spread­ (Batak), rumput udang (Indonesian). Malaysia: ing and bearing oblong not sticky yellowish ramput kolam padang. Thailand: ya-khaipu glands; spikelets 4-8-flowered, up to 3.5 mm long, (northern). Vietnam: xuân tha'o do'. strongly compressed, breaking up from the apex, Origin and geographic distribution E. unio­ allflower s bisexual or the upper ones rudimentary; loides is native to South-East Asia. It is also found palea-keels ciliate. Caryopsis ellipsoidal, 0.5-0.7 in the Indian subcontinent, Sri Lanka and the Pa­ mm long, golden-brown. cific Islands. Based on the habit of the inflorescence (spreading Uses E. unioloides is used as a forage. Its abun­ or more contracted), two botanical varieties have dance in irrigated rice fields makes it also useful been distinguished. for grazing after the rice harvest and as a green Seeds of E. tenella germinate very quickly at the manure. beginning of the rainy season. It flowers profusely Properties No information on chemical compo­ throughout the year, producing a very small sition is available, but its quality is likely to be amount of leaf. low because of its low leaf/stem ratio. Ecology E. tenella is common on open waste Botany Annual or short-lived perennial grass, ground and is adapted to sandy soils from sea-level up to 1400m altitude. It can be found on old walls, roadsides and dykes, usually in areas with a pro­ nounced dry season. Agronomy It is not deliberately sown but spreads naturally by seed. It grows quickly with the onset ofth e first rains and thus produces green feed early in the season. It can be cut by hand, but as it has a very weak root system farmers normally dig up whole plants and feed them to livestock. Yields are low. It is fed green but could be dried and conserved. Genetic resources and breeding It is unlikely that substantial germplasm collections are being maintained. Prospects It is unlikely that E. tenella will ever be deliberately established as a forage crop. Literature jlj Gilliland, H.B., Holttum, R.E. & Bor, N.L., 1971. Grasses of Malaya. In: Burkill, H.M. (Editor): Flora of Malaya. Vol. 3. Govern­ ment Printing Office, Singapore, pp.71-73 . {21Häf - liger, E. & Scholz, H., 1981. Grass weeds 2. Ciba- Geigy Ltd., Basel, Switzerland, p. 75. |3j Holm, J., 1971. Feeding tables. Nutrition Laboratory of Thai-German Dairy Project, Livestock Breeding Station, Chiangmai, Thailand. 14! Mehra K.L. & Fachrurozi, Z., 1985. Indonesian economic plant resources: forage crops. Lembaga Biologi Nasio- nal - LIPI, Bogor. p.17 . C. Manidool Eragrostis unioloides (Retzius) Nees ex Steudel - 1, flowering plant; 2,ligule; 3,inflorescence; 4, spikelet. Ficus 129 tufted, with culms 10-80 cm tall, erect or genicula- Ficus subcordata Blume tely ascending, sometimes rooting from the lower nodes. Leaf-sheath about 1.5 cm long, striate, pur­ Bijdr.: 440(1825) . plish, hairy at the mouth; ligule a fringe of hairs; MORACEAE leaf-blade linear with broad base and acute top, 2n = 26 2-20 cm x 2-8 mm, flat or inrolled. Inflorescence Synonyms Ficus garciniifolia Miquel (1867), F. a loose or contracted, terminal, usually stiffly calophylloides Elmer (1911), F. fairchildii Backer erect panicle, up to 20c mlong ; spikelets 8-60-flow- (1947). ered, ovoid to oblongoid, 4-16 mm x 2.5-4 mm, Vernacular names Indonesia: bunut lengis strongly compressed, on up to 15 mm long pedun­ (Bali), wunut (Javanese), sipadi (Sumatra). Philip­ cles, usually yellowish but reddish-purple tinged; pines: marabotum (Bagobo), balete (Tagalog), tibi lower florets all fertile, upper ones caducous, but (Bikol). Thailand: sai (Nakhon Si Thammarat). florets falling in succession from the base up; Origin and geographic distribution F. subcor­ glumes very densely packed, keel scabrid. Caryop- data is widespread in South-East Asia, from north­ sis obovoid to ellipsoid, laterally compressed, ca. ern Vietnam to the New Hebrides. It is also occa­ 0.7m m long, orange-brown. sionally cultivated in this area. Flower heads emerge 6-8 weeks after seedling Uses The foliage of F. subcordata is used as a emergence and plants flower throughout the year. feed supplement during the wet season and as the It is a very variable species, the main variation sole diet during the dry season for ruminants in being in the annual to perennial type and in the some dryland farming areas. The young fruit can degree of stoloniferous habit. be fed to ruminants. The wood is used as fuel for Ecology E. unioloides can grow from sea-level brick and limestone kilns, and the smaller up to 1250m altitude , in open or moderately shaded branches are used for household firewood. The areas, in swampy or paddy fields, roadsides and mature stem is used for farmyard posts. The bark cultivated land. isuse d for making string for farm tools.Th e timber Agronomy E. unioloides is grazed but does not is not hard enough for building houses, making withstand heavy grazing. In northern Thailand, farm implements or woodcarving. The tree is used villagers may pull up whole plants and feed them as shade for livestock, for storing crop residues, fresh to cattle. for reclamation of denuded land, for protecting Genetic resources and breeding It is unlikely soil on sloping land and as a windbreak. that substantial germplasm collections are being Properties The leaves contain 1.2-1.8% N, maintained. crude fibre 26-30%, N-free extract 42-47%, ash Prospects Although this species contributes to 8-11%, total digestible nutrient 33-35%, and the feeding of livestock, it is not of great impor­ massic energy of DM is 10000-19000 kJ/kg. Cut tance. stems exude white latex but this does not affect the Literature |1| Backer, CA. & Bakhuizen van palatability ofth e leaves. den Brink, R.C., 1968. Flora of Java. Vol. 3. Description Strangling deciduous tree, without Wolters-Noordhoff, Groningen, the Netherlands, aerial roots, up to 30m tall and 70 cm in diameter; p. 531. |2I Gilliland, H.B., Holttum, R.E. & Bor, branching starts 2 m above the ground and twigs N.L., 1971. Grasses of Malaya. In: Burkill, H.M. are brownish-grey; in shallow soil the lateral roots (Editor): Flora of Malaya. Vol. 3.pp .66-68 . |3IHäf - near the soil surface can spread 4-7 m away from liger, E. & Scholz, H., 1981. Grass weeds 2. Ciba- the base of the trunk. Bark whitish-grey, slightly Geigy Ltd., Basel, Switzerland, p. 78. |4| Mehra, smooth and fissured, flexible and durable, 10-17 K.L. & Fachrurozi, Z., 1985. Indonesian economic mm thick; inner bark whitish, exuding white sap. plant resources: forage crops. Lembaga Biologi The blunted spearhead-like bud extends from the Nasional - LIPI, Bogor, No 31. p.18 . node while the leaf is still intact. Leaves alternate, C. Manidool oblong, ovate-oblong, or elliptical, 9-20 cm x 4-10 cm, with a prominent light green midrib and a light green petiole of 2-5 cm length; leaf margin entire; leaf-blade broadly cuneate or rounded at base, pointed at apex, smooth to hairy, purple when young, light green beneath and dark green above when mature. Fruit a short-ellipsoid fig, 3-5 cm x 2-2.5 cm, solitary, occasionally in pairs, 130 FORAGES

Besides F. subcordata, in dryland farming areas in Bali (Indonesia), farmers plant also 3 other Ficus species vegetatively: - F. stricta Miquel ('bunut bingin' in Bali): ever­ green tree, up to 18m tal l with deeply fluted bole and large spreading crown; leaves elliptical, 7-12 cm x 3-6 cm, with numerous almost paral­ lel secondary nerves; figs globose, 1.5 cm in di­ ameter, orange; Indo-China, Malaysia, Indone­ sia, the Philippines. - F. elastica Roxb. ex Hörnern, ('bunut lulub' in Bali): large strangling fig developing many aer­ ial roots from the trunk and main branches; young parts reddish; leaves elliptical, 7-15 cm x 4-7 cm,large r in saplings;fig s oblongoid, 1.25 cm x 0.8 cm, yellow. Formerly used for rubber production (India rubber tree); originally distri­ buted from India through Indo-China and Malaysia to Sumatra; now cultivated all over the tropics, mainly as an ornamental. - F. drupacea Thunb. ('bunut bulu' in Bali): big tree with brown woolly hairs on buds, twigs and underside of leaves; leaves oblong-elliptical, 11-23 cm x 4-11 cm, tri-nerved at base; figs oblongoid, 2-3 cm x 1-2 cm, yellow; widely dis­ tributed from India, throughout South-East Ficus subcordata Blume - 1, habit tree; 2, fruiting Asia to Australia. branch. Ecology In Indonesia, F. subcordata grows well in dryland and hilly areas with annual rainfall green when young, gradually turning from yellow ranging from 900-2500 mm, mean daily maximum to reddish-brown or black when ripe. Seeds small, temperatures from 26-39°C and altitudes ranging hard and numerous. Weight of a fresh fruit ranges up to 800 m. It tolerates a wide range of soil types, from 10-20 g, and there are 1000-2000 ripe seeds growing well on limestone-based soil and on slop­ per g. ing land of 25c m soil depth. Growth and development Under natural con­ Propagation and planting Even though F. sub­ ditions, reproduction starts when the tree is 5-6 cordata can be propagated by seed or layering, years old. Flowering and fruiting take place dur­ propagation by cuttings is most commonly prac­ ing the dry season when the tree sheds its old tised by farmers. For direct planting, healthy and leaves.Th e mature fruit falls off when the new sea­ straight 2-year-old stems of 5-10 cm in diameter son's leaves are fully expanded. Under favourable and 1.5-2.5 m length are cut from the parent tree. conditions the small seeds will germinate in 3-4 The end to be planted should be even and free from weeks. Under natural conditions, the seeds may splitting, and any leaves and twigs should be germinate in a crack in other trees and develop removed. Each cutting should be planted in a pre­ into a stunted tree (known in Bali as 'bunut pang- pared hole of 25 cm depth and 15 cm width, then gang'). Once the aerial rootsreac h the ground they covered with soil in such a way that the planted can develop into a normal tree that can kill the stem cannot move. Direct planting should be car­ host. ried out at the onset of the rainy season, since Other botanical information F. subcordata is planting during the wet season causes the buried subdivided into two varieties by Corner: var. sub­ cambium to rot. Twelve weeks after planting 75% cordata, the typical variety; and var. malayana of the cuttings can develop buds and about 70% Corner, with large subcylindrical figs 3.5-5 cm x survive after 52 weeks. Twenty-six weeks after 2-2.5 cm, and elliptical to narrowly obovate, thick planting a cutting, there can be 8-13 main leaves, 11-16 cm x 4-7 cm, occurring in Peninsu­ branches of 45-55 cm length and 10-12 leaves per lar Malaysia and in northern Borneo. branch. For planting in nurseries, twigs with FLEMINGIA 131

50-100 cm length are inserted in 10-15 cm of soft species rather than of regional importance. How­ and moist soil. Such cuttings are not ready for ever, once established, it produces a year round transplanting until the roots are well developed. supply of fodder for ruminants in dryland farming Since F. subcordata seed isver y small, it is prefera­ areas. F. subcordata is one of the fodder trees inte­ bly sown under nursery conditions. After develop­ grated in the 'Three Strata Forage System' cur­ ment ofth e cotyledons and a few secondary leaves, rently under test to increase the productivity of seedlings should be transplanted into pots. Plant­ dryland farming in Indonesia. In this system the ing into permanent sites is carried out during the 1st, 2nd and 3rd strata consist of grass and herba­ rainy season when the plants are 6-12 months old. ceous legumes, shrub legumes and fodder trees, re­ It can be planted at 5-10 m spacing when used as spectively. Establishment of F. subcordata from fence border and at 10 m x 10 m within and be­ seeds, so as to increase its distribution and role as tween rows spacings when used as windbreak or a multipurpose tree, should be explored. fodder bank. Literature ill Backer, CA. & Bakhuizen van Husbandry When planted as a cutting, it can den Brink, R.C., 1965.Flor a of Java. Vol. 2. Noord- be lopped 3year s after planting. Complete defolia­ hoff, Groningen, the Netherlands, pp. 20-35. |2.' tion can be carried out before the end of the dry Corner, E.J.H., 1965. Check-list of Ficus in Asia season every year, or partial defoliation twice dur­ and Australasia with keys to identification. The ing the wet season and twice during the dry season. Gardens' Bulletin Singapore 21: 1-186. 13'd e Guz­ For accumulation of in situ fodder, the tree should man, E.D., Umali, R.M. &Sotalbo , E.D., 1986. Non- be lopped 3-4 months before the dry season so that Dipterocarps. Ficus. In: Umali, R.M., Zamora, the foliage iswel l developed during the dry season, P.M., Gotera, R.R. &Jara , R.S. (Editors): Guide to otherwise the leaves will shed during the period Philippine flora and fauna. Vol. 3. Natural offlowe r and seed formation. Resources Management Center and University of Many species can be grown with F. subcordata the Philippines, Manila, pp. 123-130. |4| Kochum- including grasses such as Cenchrus ciliaris L., men, K.M., 1978. Moraceae, Ficus. Inf Ng, F.S.P. Panicum maximum Jacq. and Urochloa mosambi- (Editor): Tree flora of Malaya. A manual for For­ censis (Hack.) Dandy; herbaceous legumes such as esters. Vol. 3.Longma n Malaysia, Kuala Lumpur, Stylosanthes hamata (L.) Taub., Stylosanthes sea- pp. 135-162. |5| Nitis, I.M., Lana, K., Suarna, M., bra Vogel, and Centrosema pubescens Benth.; Sukanten, W., Putra, S. & Arga, W., 1989. Three shrub legumes such as Leucaena leucocephala strata system for cattle feeds and feeding in dry­ (Lamk)d e Wit and Gliricidia sepium (Jacq.) Kunth land farming area in Bali. Final report to IRDC, exWalp . and fodder trees such as Lannea coroman- Canada. 252pp . !6iPanday , Kk., 1982.Fodde r trees delica (Houtt.) Merrill and Hibiscus tiliaceus L. and tree fodder in Nepal. Swiss Development Co­ Diseases and pests F. subcordata is generally operation, Berne and Swiss Federal Institute of resistant to diseases and pests. Eed ants (Phagiole- Forestry Research, Birmensdorf, Switzerland. 107 pis longipis) and black ants (Formica fusa) living pp. |7| Sukarji, N.W., Puger, A.W. & Nitis, I.M., in the foliage cause no ill effect to the host, but 1990. Draft performance of Bali steers fed grass, may inconvenience the farmer lopping the tree. shrub legume and fodder tree [Indonesian]. Natio­ Harvesting and yield Fruits of F. subcordata nal Seminar on Bali cattle, September 20-22,1990, are not regularly harvested since they lack com­ Denpasar, Bali. 5pp . mercial or socio-cultural value. The annual fodder I.M. Nitis and firewood DM yields of a 3-year-old F. subcor­ data range from 12-20 kg and 30-65 kg per tree respectively. As the tree grows older, the annual Flemingia macrophylla (Willd.) Merr. fodder and firewood DM yields can increase up to 140-225 kg and 240-350 kg per tree respectively. Philip. Journ. Sei. 5:130 (1910). The foliage of a 25-year-old tree can feed one ani­ LEGUMINOSAE mal of25 0k g live weight for 20-30 days. 2n = 22 Genetic resources and breeding Germplasm Synonyms Flemingia congesta Roxb. ex Ait. f. has been collected but there are no breeding pro­ (1812), F. latifolia Benth. (1852), Moghania macro­ grammes of Ficus species in South-East Asia. phylla (Willd.) Kuntze (1891). Germplasm collection and breeding of Ficus spe­ Vernacular names Indonesia: apa apa (Java­ cies has been initiated in Nepal. nese), hahapaan (Sundanese), pok kepokan Prospects F. subcordata is a location-specific (Madura). Malaysia: serengan jantan, beringan. 132 FORAGES

Philippines: laclay-guinan (Tagalog), gewawini (Ifugao), malabalatong (Pampanga). Laos: thwàx h'èè h'üad, hom sam muang (Xieng Khouang), thoua huat (Vientiane). Thailand: mahae-nok (northern), khamin naang, khamin ling (central). Vietnam: tóp mo' lato, cây dau ma (Vinh phu), cai duoi chon (Thuan Hai). Origin and geographic distribution F. macro- phylla originates from and is widely distributed in South-East Asia and in Taiwan, southern China, India, Sri Lanka and Papua New Guinea. It has been introduced and naturalized in East, Central and West Africa. Uses F. macrophylla is grown in hedges and pro­ vides forage and also mulch for associated food crops grown in alley cropping systems. It is also grown in terraces to control soil erosion and is used to provide dry season forage in the savanna zone of Nigeria. In Malaysia, Indonesia, Sri Lanka, West Africa and Madagascar the plant is used as a cover crop and as a shade crop in young plantations of cacao, coffee, banana and rubber. It is one of the sources of the Arab dye which is called 'warrus' or 'waras'. 'Waras' is a purple or orange-brown coarse powder, consisting of the glandular hairs rubbed from dry Flemingia fruits, capable of dying silk but not wool or cotton, and which is called flemingin. It is a minor host of the Flemingia macrophylla (Willd.) Merr. - 1, flower­ Indian lac insect. The stems are a source of flavo- ing branch; 2,fruiting branch. noid compounds. In Indonesia and Malaysia the leaves are used as a source of medicine. In the la with greenish elliptical standard and distinct Ivory Coast it is used to reduce nematodes in pine­ parallel red veins, wings narrow and much shorter apple plantations and as green manure and mulch. than the keel, light purple at the apex. Pod oblong, Properties Leaf nutrient contents (based on inflated, 8-15 mm x 5mm , covered with fine glan­ DM) are: N 2.3-3.8%, P 0.19-0.25%, K dular hairs, dehiscent, dark brown, 2-seeded. Seed 1.00-1.40%, Ca 0.55-0.75%, and Mg 0.18-0.28%. globular, 2-3 mm in diameter, shiny black. In vitro digestibility of the leaves is much lower Nodulation is often difficult to locate, partly than that oîLeucaena leucocephala (Lamk) de Wit. because the nodules are very small. The leaves decompose relatively slowly and are Ecology F. macrophylla can be found from sea- useful as mulch material. Under humid tropical level up to 2000 m altitude, within a wide range conditions, losing half of an initial quantity of 4 of rainfall patterns, from sub-humid to humid t/ha of dry matter took 53days . (1100-2850 mm/year). It cannot tolerate long dry There are 55 seeds/g. spells and is capable of surviving on very poorly Botany Woody, deep-rooting, tussock-forming drained soils with waterlogging. Its natural habi­ shrub, 1-4 m tall. Young branches greenish, tat is along watercourses, both on clay and lateri- ribbed, triangular in section, silky. Old stems tic soils, as well as under drier conditions such as brown, almost round in section. Leaves digitately in fields infested with Imperata cylindrica (L.) trifoliolate; petioles narrowly channelled, slightly Raeuschel. The plant tolerates shade and poor winged, up to 10 cm long; leaflets lanceolate, 6-16 soils. cm x 4-7 cm, papery, veins covered with silky Agronomy F. macrophylla is propagated by hairs, dark green. Inflorescence a dense axillary seed. Since untreated seed ishardseeded , scarifica­ raceme, subspiciform, sessile, 2.5-10 cm long, tion is usually required to increase the germina­ silky; bracts ovate, 3-6 mm long; calyx green, 6-13 tion percentage of seed prior to sowing. A good mm,pal e velutinous with 5lanceolat e lobes; corol- weed-free seed-bed should be prepared, and the GLIRICIDIA 133

necessary fertilizers for a particular soil worked nese Society of Grassland Science, Nishihasuno, in prior to sowing or banded beneath the row of Japan, pp. 1-3. \%Budelman , A., 1988. Leaf dry seed. Seed should be inoculated with a suitable matter productivity of three selected perennial strain of Bradyrhizobium such as CIAT 4203 or leguminous species in humid tropical Ivory Coast. 4215. Planting density varies according to the pro­ Agroforestry Systems 7: 47-62. 3 Budelman, A., jected use of the stand. In Indonesia seed is often 1991. Woody species in auxiliary roles: live stakes sown in rows 90 cm apart with 3-4 seeds planted in cultivation. Royal Tropical Institute, every 60 cm along the row. Good weed control is Amsterdam. 151 pp. !4i Rubber Research Institute required during the first six months after sowing, of Malaysia, 1975. Species and varieties of Fle­ since the plants are relatively slow to establish. mingia in Malaya. Planters Bulletin 61: 78-85. 5 Once established, they require little attention. Schultze-Kraft, R., 1985. Development of an inter­ Under humid, lowland tropical conditions in the national collection of tropical germplasm for acid Ivory Coast, with 10000 plants/ha and 9 regrowth soils. In: Proceedings of the XV International cycles of3 month s each, an average annual produc­ Grasslands Congress, August 24-31, Kyoto, tion of 12 t/ha of leaf DM has been achieved, al­ Japan. Science Council of Japan and Japanese So­ though typical yields in South-East Asia may be ciety of Grassland Science, Nishihasuno, Japan, closer to 8t/ha . Plants can be cut more frequently pp. 1-3. 61 Skerman, P.J., Cameron, D.O. & Rive- than every 3 months, but preferably not at inter­ ros, F., 1988.Tropica l forage legumes. FAO, Rome, vals of less than 40 days. They will survive under pp. 561-562. this cutting regime for many years. A. Budelman &M.E . Siregar Insect pests such as the fly Agromyza sp. reduce seed production by laying eggs in green pods. In Malaysia, spraying with Endrex (1:800) once every Gliricidia sepium (Jacq.) Kunth ex two weeks after flowering has begun gives effec­ Walp. tive control. Genetic resources and breeding Germplasm Repertitorium bot. syst. 1:67 9(1842) . collections are located at CIAT (Colombia), the LEGUMINOSAE Research Institute for Animal Production (Ciawi, 2n = 20 Indonesia), LBN/LIPI (Bogor, Indonesia), Synonyms Gliricidia maculata (Kunth) Kunth ATFGRC (CSIRO, Australia), IBPGR (Bangkok, ex Walp. (1842). Thailand) and ICRISAT (Hyderabad, India). Vernacular names Gliricidia, mother of cocoa, Prospects F. macrophylla has excellent coppic­ quick stick (En). Indonesia: gamal (Javanese), li- ing capacity and is showing promise when used in riksidia (Indonesian). Philippines: balok-balok hedges to provide forage or mulch to associated (Tagalog), apatot (Bikol), kukuwatit (Pangasi- food crops in alley cropping. As a feed F. macro­ nan). Laos: kh'è: nooyz, khê fàlangx. Thailand: phylla is inferior to species such as Leucaena leuco- khae-. Vietnam: anh dào gi'a, sât thu, hông cephala (Lamk) de Wit and Gliricidia sepium mai. (Jacq.) Kunth ex Walp. since it is less easily Origin and geographic distribution Gliricidia digested. Owing to its slow decomposition, the is a native of the seasonally dry Pacific Coast of mulch has long-term effects in weed control, mois­ Central America from sea-level to 1200 m, but has ture conservation and soil temperature reduction. been long cultivated and naturalized in tropical Furthermore, the species is useful as a cover crop Mexico, Central America and northern South in perennial plantations, as a shelter belt, in ero­ America, up to 1500 m altitude. The species was sion control, and in planted fallows for soil im­ also transported to the Caribbean and later to provement. Improvement of the plant's early de­ West Africa. It was introduced to the Philippines velopment and its integration into alley cropping by the Spaniards in the early 1600s, and to Sri systems and planted fallows deserve priority in Lanka in the 18th century; from there it reached research. other Asian countries including Indonesia (about Literature |1| Asare, E.O., 1985. Effect of fre­ 1900),Malaysia , Thailand and India. quency and height of defoliation on forage yield Uses In the past, gliricidia forage was often col­ and crude protein content of Flemingia macro­ lected from trees which had been planted for other phylla (flemingia). In: Proceedings ofth e XV Inter­ purposes, but increasing attention is now given to national Grasslands Congress, August 24-31, options for cultivating it specifically for fodder Kyoto, Japan. Science Council of Japan and Japa­ production either in cut-and-carry or grazing sys- 134 FORAGES tems. It provides useful forage in the form of leaves, green stem and bark, and is commonly used to supplement poor quality, low protein roughage, especially in dry seasons when it may become a major source offee d for goats and cattle in dryland cropping areas. Its forage has been reported to be toxic to horses, but clear confirmation is lacking. Leaf meal can also be fed to poultry and rabbits. The tree is widely used for many other purposes. The wood is often utilized as firewood, charcoal or as posts and farm implements, and locally for furniture, construction purposes and railway sleepers. The species is extensively used in differ­ ent cropping systems, e.g. as a shade tree in tea, cocoa or coffee plantations, as live stakes to sup­ port vanilla, black pepper and yams (in West Africa), as a hedge, a green manure crop in inter­ cropping systems with arable crops;i t is also being tested in alley-cropping systems. The tree has also been planted to reclaim denuded or lands invested with Imperata cylindrica (L.) Raeuschel. Seeds, bark, leaves and roots may be used as a rodenticide and pesticide after fermentation; flowers serve for honey production. The tree is often planted as an ornamental. In the Philippines juice of the leaves, bark and roots is used to alleviate itches and wounds. Properties The leaves contain from 3-5% N, Gliricidia sepium (Jacq.) Kunth ex Walp. - 1, leaf; 13-30% crude fibre and 6-10% ash. Digestibility 2,flowering branch; 3,fruiting branch. ranges from 48-77%. Forage quality varies with age, plant parts, season and genotype. It is highest of thin foliage. Leaves alternate, pinnate, 15-40 in the youngest leaves;wit h maturity N concentra­ cm long, with slender, yellow-green, finely hairy tions decrease slightly and crude fibre increases. rachis; leaflets 7-17 per leaf, opposite except in In 3-month-old growth, gliricidia bark had lower upper part of rachis, elliptical or lanceolate, 3-6 N concentrations than the leaves, but higher cm x 1.5-3 cm,rounde d or cuneate at base, acumi­ levels than the stem. Palatability can be a problem nate at top, thin, dull green and glabrous above, as the forage contains some anti-nutritional fac­ grey-green and often pubescent beneath; petiole 5 tors, with 1-3.5% flavonol and 3-5% total phe­ mm long. Flower in a 5-12 cm long, axillary nols on a DM basis. Ruminants unaccustomed to raceme, about 2 cm long, on an 8-12 mm long, it may not eat the foliage when introduced to it slender pedicel; calyx campanulate, 5-toothed, for the first time. However, once initial aversion light green tinged with red; corolla whitish-pink has ended, animals will eat a high proportion in or purple, with a broad standard, turned back and their diet for extended periods of time, especially yellowish near the base, 2 oblong, curved wings, when mixed with other species. In some cases it and a narrow keel; stamens 10, white, 9 united in has been observed that day-old wilted leaves are a tube and 1 separate; pistil with stalked, narrow preferred to fresh leaves; also silage is more palat­ red ovary and whitish curved style. Pod narrow, able than fresh foliage. Leaves also contain caro­ flat 10-15 cm x 1.2-1.5 cm, yellow-green when tene. immature, turning yellowish-brown, shortly There are about 80 seeds/g. stalked and with a short mucro, splitting open at Description A small deciduous tree up to 12 m maturity. Seeds 4-10 per pod, ellipsoidal, ca. 10 tall with a short trunk up to 50 cm in diameter, mm long, shiny, dark reddish-brown. with smooth or slightly fissured, whitish-grey to Wood characteristics Light-brown sapwood light brown bark, often branching from the base; and dark-brown heartwood turning reddish on the mature tree has an irregular spreading crown exposure to air, hard, coarse-textured with irregu- GLIRICIDIA 135 lar grain. It is very durable and -resistant. the onset oflowe r temperatures during winter, and The dense wood, with a volumetric mass up to 750 plants are less resistant to frosts. However, it is kg/m3, is difficult to work. Wood does not reach more tolerant of waterlogged conditions than leu­ large dimensions, the bole seldom has a diameter caena. The tree resprouts after fire. of more than 40 cm and a length of 8 m; normally Propagation and planting Gliricidia is easily dimensions are smaller, especially if the tree is propagated from seed or cuttings. Direct seeding regularly coppiced. In young coppices the wood is is not often used and potted plants or bare rooted less dense, with a volumetric mass of about 500 kg/ stock are raised in nurseries. Freshly harvested m3. seed or seeds that have been preserved in cold stor­ Growth and development Seeds germinate in age have a germination percentage of 80-90%. 7-10 days. Early seedling growth is slow, but once Seed pretreatment is not necessary and seeds may established, growth isfas t (upt o 3m/year) . Flower­ be sown directly. Nursery stock can be trans­ ing and fruiting take place during the dry season, planted after 10-12 weeks. It can be propagated when the tree has shed its leaves if it has not been very easily by large cuttings, 3-6 cm thick and cut. Flowers are insect-pollinated and the species 0.5-2 mlong ;th e bark may be incised to assist root­ is outcrossing. Pods ripen 40-55 days after flower­ ing. Cuttings should be taken from mature ing, seeds are mature when pods turn yellow- branches with brownish-green bark and planted brown; fruiting isrelativel y uniform with about 20 fresh. Rooting starts 6-7 weeks after planting. days from first to last seed dispersal. In its native Plants grown from cuttings may have 50-150 nod­ area seed production is usually abundant and can ules after 3months , compared with 20-70 nodules be predictably timed. In more humid zones, shoot after 6month s on plants from seed. Trees obtained growth tends to be continuous and the evergreen from cuttings are often more shallow-rooted than tree flowers only sporadically on the basal parts trees grown from seed. of twigs from which the leaves have fallen. Gliricidia may be planted in hedges spaced 4-10 Other botanical information G. maculata has m apart with 10-50 cm between trees in the rows, been used extensively as a synonym for G. sepium. or as live fences with 20-100(-200) cm spacing. In Recently G. maculata has been proposed as a dis­ fodder plots, spacings of 0.25 m x 1.0 m or larger tinct species with a different natural geographic may be used; yields of leaves are little affected by distribution, i.e. Yucatan , northern planting densities ranging from 5000-40000 trees/ Guatemala and Belize. It differs from G.sepium by ha. Sometimes trees are used at wide spacings (e.g. having white flowers rather than whitish-pink or 10 m x 10 m) over pasture lands. Where animals purple, and it has smaller pods and seeds. Most are grazed in young plantations, young trees must gliricidia planted as an exotic can be attributed to be protected. Growth on degraded lands may be G. sepium, but former introductions of G. maculata stimulated by inoculation with an appropriate cannot be ruled out. strain of Bradyrhizobium and fertilization. When Ecology In its native range the climate is relati­ the tree is used as a live post for black pepper or vely uniform sub-humid with an annual rainfall of vanilla, the crops can be planted at the same time 900-1500 mm and a five-month dry period. The spe­ asth e tree.Suc h livepost s provide some protection cies has been introduced successfully in more against climatic stress. In woodlots spacings of 1.5 humid zones with up to 3500 mm annual rainfall m x 2m to 2m x 2.5m are common. and no marked dry season. In its native range, Husbandry Fodder from gliricidia can be pro­ mean annual temperature varies from 20-29°C, duced in various systems. Protein banks consist of maximum temperature below 42°C ; light night dense plots which are periodically harvested for frost is tolerated, but not prolonged frost. Glirici­ fodder or grazed; hedges may be planted in existing dia occurs naturally in early and middle successio- pasture lands and interplanted with grasses; living nal vegetation types on disturbed sites such as fences may be established around pasture lands or coastal sand dunes, river banks, floodplains and agricultural fields and homesteads; or spaced trees swidden lands. It tolerates a wide range of soil may be used in pastures to enhance livestock pro­ types, both alkaline and acidic, but prefers free duction by providing fodder and shade. In Sri drainage. It is more tolerant of acid and poorly fer­ Lanka the tree has also been integrated in pastures tile soils than leucaena (Leucaena leucocephala under coconut to produce dry season fodder. In (Lamk) de Wit), but will respond to fertilizer appli­ Bali (Indonesia), gliricidia has been incorporated cation on such soils. It is not as well adapted to in a 'Three Strata Forage System' consisting of a the subtropics as leucaena; leaves are shed with strip of 5 m wide, in which fodder trees (e.g. Ficus 136 FORAGES subcordata Blume, Hibiscus tiliaceus L., Lannea ing is possible. During the dry season, harvesting coromandelica (Houtt.) Merrill), shrub legumes should be less intensive (every 3-4 months) than (e.g. gliricidia) and grasses (e.g. Cenchrus ciliaris during the rainy season (every 2-3 months). L.) and herbaceous legumes (e.g.Stylosanthes spp.) Regrowth should be 1-2 m in height before each are combined; cattle feed mainly on the grass- harvest. To obtain fodder during the dry season legume stratum in the wet season, the shrub trees should be cut about 3month s before the onset legumes in the mid-dry season and leaves from the of this season. If trees are uncut for more than 4-6 fodder trees in the late dry season. Variable months before the dry season commences, they will amounts offodde r may also be collected from plan­ shed their leaves during the dry season. Cutting tations established for other primary purposes heights commonly range from 0.5-2 m. Trees such as erosion control or green manure contour grown in contour lines and fodder banks are plantings in cropped fields. usually cut lower than those cultivated in living The influence of gliricidia on pasture grass pro­ fences or as shade trees in pastures, where brows­ duction depends on the grass species, site condi­ ing cattle may interfere with regrowth. tions and management. Even if grass production Forage from gliricidia is usually cut by hand and is decreased, gliricidia production may compen­ left on the ground for grazing or carried to pad­ sate for this loss with the total production being docks or stalls. Acceptable silage can be prepared more evenly distributed over the year. A wide using standard techniques; the chopped forage range of grasses and legumes can be grown may be mixed with grasses or maize and additives together with gliricidia, including Panicum maxi­ such as molasses and sugar cane or formic acid mum Jacq. var. trichoglume Robijns, Cenchrus (0.85%) should be added to provide fermentable ciliaris, Urochloa mosambicensis (Hack.) Dandy, carbohydrate. Stylosanthes scabra Vogel, and Stylosanthes ha- In woodlots, first harvesting can be after 3-4 mata (L.) Taub. When fed as a supplement to poor years, giving wood yields of 8-15 m3/ha. Subse­ quality grass, it improved the survival of ewes and quent coppicing isdon e every 2-3 years and yields lambs, lambing percentage, and birth weight and up to 40% more than the first harvest. growth of lambs. It is normally recommended that Yield Under average conditions, yields of3- 4 kg when feeding cattle, goats and sheep, gliricidia be DM per tree per harvest may be achieved. In fodder used at levels of 10-30% fresh weight mixed with plots, annual yields of 9-16 t/ha of leaf DM or up either grass, straw or other roughages. Levels of to 43 t/ha fresh leaves have been obtained. In 2-4% should be used when it is fed to poultry as Nigeria gliricidia hedgerows interplanted with 4 a replacement for lucerne (Medicago sativa L.). rows of Panicum grasses yielded 20 t/ha per year Diseases and pests Few diseases and pests have of mixed DM, which was sufficient to feed 3 head been recorded on gliricidia, and only sporadically of cattle. do these cause noticeable damage. Very little In woodlots, when coppiced every 2-3 years, wood information is available on the effect of pests and yield varies from 10-20 m3/ha. Wood production diseases on fodder quality and palatability, but in in living fences has been reported at 9 m3/km per Indonesia it was observed that buffelgrass (Cen­ year. All harvested produce isusuall y used locally. chrus ciliaris) was adversely affected by an attack Genetic resources Major germplasm collec­ of Aphis craccivora on gliricidia. When inter­ tions have been made by the Oxford Forestry Insti­ cropped the tree may either positively or negati­ tute (United Kingdom), CATIE (Turrialba, Costa vely affect crop pests. In several cases the tree has Rica), and ILCA Humid Zone Programme (Ibadan, been reported to control pests, e.g. in Sri Lanka Nigeria). The Oxford Forestry Institute adminis­ termite damage to tea and in the Philippines stem- ters an international network of provenance eval­ borer damage to rice were minimized. In India, on uation involving 29provenance s from 8Sout h and the other hand, the tree was found to enhance the Central American countries. The ILCA Humid transmission of (Aphis craccivora) causing Zone Programme has developed a high-yielding rosette disease in groundnuts. bulk composite from four Costa Rican accessions. Harvesting The first harvest can be as early as Breeding There are no known breeding pro­ 6-8 months on plants grown from cuttings and grammes. Early introductions in many countries 12-16 months on plants grown from seedlings. usually had a very narrow genetic base and dis­ There should be only one or two harvests per year tinct indigenous ecotypes have evolved in several during the first 2 years. Trees must be well-estab­ areas. Recent provenance evaluations indicate lished, 18-24 months old, before regular harvest­ significant differences in growth rates. Further HETEROPOGON 137 rapid genetic gains can be expected, as seed pro­ speargrass (En). Tanglehead (Am). Herbe polisson duction starts early, superior types can be cloned (Fr). Indonesia: bejeng-benjeng, merakan (Java­ and production cycles are short. nese). Philippines: sibat-sibatan (Tagalog). Thai­ Distinct selection programmes for high-yielding, land: ya-nuatrusi (central), ya-lem (northern). palatable fodder cultivars and for arboreal culti- Origin and geographic distribution The natu­ vars combining wood and foliage production are ral distribution of speargrass is pantropical/sub- desirable. tropical. It is most likely of Gondwanan origin Prospects Gliricidia is showing considerable with a predominant distribution in Australia, the promise as a fodder species throughout the tropics. Indian subcontinent and Africa. However, it may It is a multipurpose tree well adapted to humid be found throughout South-East Asia and the Pa­ areas and acid and infertile soils. Although most cific and in South and Central America in sporadi­ fodder is produced in the wet season, the tree can cally occurring suitable habitats. be managed to provide fresh leaf during the dry Uses Speargrass is used as part of the naturally season. Favourable properties include its versatil­ occurring savanna pasture resources for domestic ity in production systems and the ease with which livestock production and wildlife. However, pas­ it can be grown together with grasses or herba­ tures containing speargrass are not suitable for ceous legumes. Its prospects may be further wool-producing sheep. The most severe limitation enhanced by further selection programmes and de­ to animal production from speargrass pastures is velopment of innovative production systems such the wide fluctuation of growth and herbage quali­ as the 'Three Strata Forage System'. It has excel­ ty between wet and dry season. lent properties for various forms of agroforestry, Properties In unimproved pasture its N concen­ as well as for site reclamation, including suppres­ tration ranges from 2.5% in very young green ma­ sion of such obnoxious weeds as Imperata cylin- terial from the early flush of the growing season drica (L.) Raeuschel. to as low as 0.3% in dry forage by the end of the Literature jl| Chadhokar, P.A., 1982. Gliricidia dry season. Digestibility varies similarly from maculata, a promising legume fodder plant. World 60% down to 40% . The sharply pointed and barbed Animal Eeview 44:36-43 . I2|Devendra , C. (Editor), seeds can be a problem to both humans and ani­ 1990.Shrub s and tree for farm animals. Pro­ mals, particularly woolly sheep. ceedings of a workshop in Denpasar, Indonesia, Botany Atufte d perennial, 0.5-1.5 mtall , rather 24-29 July 1989.IDRC , Ottowa, Canada. 349 pp.|3 | variable in habit with stems erect to geniculate at Glover, N., 1989. Gliricidia production and use. the base, often branched above, particularly at NFTA, Waimanalo, Hawaii. 44pp . |4|Lindsay , F.J., flowering, flattened towards the base. There is a 1982. Gliricidia maculata, a review. International tendency to being weakly rooted late in the dry Tree Crops Journal 2: 1-14. |5| Palmer, B., Bray, season. Stem nodes smooth and hairless. Leaves R.A., Ibrahim, T. & Fulloon, M.G., 1989. Shrub green or bluish, basal and on the culm; leaf-sheath legumes for acid soils. In: Crasswell, E.T. & Push- smooth, compressed, keeled, striate, sometimes parajah, E. (Editors): Management of acid soils in with a few hairs near the ligule; ligule a short mem­ the humid tropics of Asia. ACIAR Monograph No branous rim; leaf-blade linear, 3-30 cm x 2-8 mm, 13. ACIAR, Canberra, pp. 36-43. |6|Withington , D., folded in the lower part, becoming flat, slightly Glover, N. &Brewbaker , J., 1987.Gliricidi a sepium rough to the touch with a few long hairs, particu­ (Jacq.) Walp.: management and improvement. larly towards the base, apex blunt, almost canoe- Nitrogen Fixing Tree Association Special Publica­ shaped. Inflorescence a simple raceme of spikelet tion 87-01. NFTA, Waimanolo, Hawaii. 255pp . pairs arranged intw o rows, 3-8 cmlon g (excluding K.F. Wiersum &I.M . Nitis the awns), the outermost of these are pedicellate and overlapping and enclosing the innermost ses­ sile spikelets; at the base of the raceme the lower­ Heteropogon contortus (L.) P. Beauv. most spikelets are similar and unawned, male or ex Roemer & Schultes neuter, while the uppermost pairs are of dissimilar bisexual awned sessile and male or neuter Syst. Veg. 2:83 6(1817) . unawned pedicellate spikelets;th e base ofth e spik­ GRAMINEAE elet has a pronounced callus which is strongly 2n = 20,40 , 44,60 , 80 bearded upwards, which at maturity disarticulates Synonyms Andropogon contortus L. (1753). obliquely to form a very sharp point, the whole Vernacular names Speargrass, black or bunch structure forming a most effective barb; the awns, 138 FORAGES

this could be a short-lived perennial form growing in a difficult environment. The late-flowering forms have higher yields of leaves and are more responsive to increased soil fertility. Ecology As a typical member of the grass Andropogoneae Dumort., speargrass is a'savanna grass in seasonally warm wet and cooler dry cli­ mates ofmoderat e rainfall. It isrelativel y drought- tolerant and intolerant of prolonged waterlogging or high salinity. Its main range of distribution is in areas of 600-1100 mm of annual rainfall with some extension below this (on deep sands of drai­ nage lines) and above this (on well-drained sites). It will grow on a wide range of well to moderately free-draining soil types. Speargrass appears not to have been a climax dominant but it has become so, particularly in Australia, India and Africa, through habitat disturbance such as heavy graz­ ing and burning. Agronomy In its naturally occurring state in savanna, speargrass has a relatively short period of growth and adequacy for good livestock nutri­ tion.Thi s causes alarg epar t ofth e annual biomass production to become unusable and left as dry standing material. Commonly this material is burnt off during or at the end of the dry season so that the new season's growth may come away Heteropogon contortus (L.) P. Beauv. ex Roemer & cleanly. This procedure leads to increasing domi­ Schultes - 1, flowering plant; 2, ligule; 3, two pairs nance of speargrass in the pasture, especially in of heterogamous spikelets. conjunction with grazing. However, speargrass can be reduced to a minor component of pastures which tangle conspicuously at maturity, are an ex­ when subjected to sustained overgrazing, particu­ tension of the inner lemma of the fertile spikelet larly on poorly fertile soils or where high stocking and 5-12 cm long;the y are geniculate, twisted and rates are sustained through supplementary feed­ hygroscopically active, enabling them to bore into ing. Up to an 8-fold improvement of livestock pro­ the ground. Caryopsis cylindrical, 3.5-4,5 mm duction from pastures based- on naturally occur­ long, grooved, whitish. ring speargrass can be achieved by introducing Speargrass exhibits a high degree of embryo and tropical legumes such as Stylosanthes spp. or Ma- seed-coat dormancy in freshly ripened seed. This croptilium atropurpureum (DC.) Urban in the sub­ declines over the dry season to a high level of ger­ tropical zones. This improvement comes from both mination at the opening rains of the next wet sea­ increased carrying capacity and rate of growth of son, following which the germination falls off rap­ the livestock. However, care must be taken, espe­ idly with little or no survival beyond one year. cially in the more strongly seasonal tropics, not Seeds which are burrowed into the soil are in a to overgraze speargrass as a result ofth e increased favourable environment for germination. Flower stocking rates. initiation is obligately to facultatively short-day, The following smut diseases of the inflorescences the former being characteristic ofth e intertropical have been reported: Sorosporium antheseriae, S. forms, which flower late in the wet season, and the caledonicum and Sphacelothica monilifera. latter of the subtropical forms which flower early Annual DM yields can range from less than 500 to mid-season. Speargrass is an apomict and this kg/ha in very dry years to 8000 kg/ha in very good has ledt o a considerable amount oflocalize d varia­ years when associated with an effective legume or tion, resulting in the proliferation of local species on fertile soils. and varieties in the early botanical literature. Genetic resources and breeding It is unlikely An annual form has been reported from India, but that substantial germplasm collections of spear- HYMENACHNE 139

grass are being maintained. There are no breeding Uses The foliage is used as fodder. The pith of programmes. While there is considerable genetic the culms is used for making lamp wicks. Some­ variability for selection, there are two serious times the plant becomes a troublesome weed in irri­ obstacles to breeding: it is an obligate apomict and gated rice. it has a most unpractical seed morphology for easy Properties H. acutigluma is very palatable. seed production. Nitrogen concentrations in 6 samples from Thai­ Prospects If speargrass is not overgrazed dur­ land ranged from 0.8-1.7%, but concentrations of ing the growing season, it can perform very well 2.5% N in whole plants and 3.6% in leaves have as the basic component of native or improved pas­ been recorded in Surinam. The average intake of tures in the tropical and subtropical savanna zone. dry matter by sheep in Thailand was 506 g/head It ismor e likely to beuse d in extensive commercial per day. beef cattle production than by smallholders. Botany Aperennial , aquatic grass, erect in shal­ Literature |1| Bogdan, A.V., 1977. Tropical pas­ low water, in deep water with very long floating ture and fodder plants. Longman, London, pp. stolons which may reach 4-6 m; culms and stolons 150-152. |2|Bor , N.L., 1960. The grasses of Burma, hollow or filled with a white spongy pith, branched Ceylon, India and Pakistan. Pergamon Press, and rooting at nodes to form a bunch of feathery Oxford, pp. 163-165. |3|Langdon , K.A.& Fullerton, rootlets; culms erect, up to 1 m tall. Leaf-sheath R., 1978. Micotaxon 6: 421-456. |4| Lazarides, M., up to 9 cm long, hairy at the margins; ligule mem­ 1980. The tropical grasses of southeast Asia. Pha- braneous, 1-2.5 mm long; leaf-blade linear to lan­ nerogamarum Monographiae Tomus XII, J. ceolate, 1.5-40 cm x 1-3.5 cm, shiny dark green, Cramer, Vaduz, pp. 46-47. |5| Skerman, P.J. & amplexicaul at the base, tapering towards the Riveros, F., 1990.Tropica l grasses. FAO,Rome . pp. apex, glabrous except at the margins at the base. 446-449. |6|Tothill , J.C. &Hacker , J.B., 1983. The grasses of southern Queensland. University of Queensland Press, Brisbane, pp. 258-260. |7i Toth- ill, J.C., Nix, H.A., Stanton, J.P. & Russell, M.J., 1985. In: Tothill, J.C. & Mott, J.J. (Editors): The ecology and management of the world's savannas. Australian Academy of Science, Canberra, pp. 125-144. J.C. Tothill

Hymenachne acutigluma (Steudel) Gilliland

Gard. Bull. Sing. 20:31 4(1964) . GRAMINEAE In = unknown Synonyms Panicum acutigluma Steudel (1854), Hymenachne pseudointerrupta C. Mueller (1861), H. myurus sensu Burkill, Diet.: 1234 (1966), H. amplexicaulis sensu Backer, F.Jav a 3: 561(1968) . Vernacular names Wick grass, dal grass (En). Indonesia: rumput kumpai (Indonesian), jujuket (Sundanese), blem bem (Javanese). Malaysia: rum- put kumpai. Philippines: lagtom (Bikol). Thailand: ya plong. Vietnam: bâc nhon. Origin and geographic distribution H. acuti­ gluma is widespread in wet habitats throughout South-East Asia. It is abundant in peat swamps in southern Thailand and is also found in India, Hymenachne acutigluma (Steudel) Gilliland - 1, Bangladesh, Burma, Indo-China, China, Japan, part ofculm with leaves; 2,ligule; 3,flowering culm; Australia and Polynesia. 4,two views ofa spikelet. 140 FORAGES

Inflorescence a terminal, spike-like panicle reach­ and nutritive value, and on selection of better cul- ing 55 cm in length, branched at the base but tivars, is warranted. remaining very narrow, cylindrical, 1-3.5 cm Literature 1 Bor, N.L., 1960. The grasses of wide; number of branches varying, 1-10 cm long; Burma, Ceylon, India and Pakistan. Pergamon spikelets narrow, long pointed but awnless, 3-5.5 Press, London, pp. 313-314. 12, Manidool, C, 1989. mm long, with one bisexual floret (the upper one) Natural grassland and native grasses of Thailand and one neuter (the lower one), green. Caryopsis [inThai]. Technical Bulletin No 1301-26-32. Divi­ ellipsoid, up to 1.5 mm long. sion of Animal Nutrition, Department of Live­ Aver y variable species in its leaves, inflorescences stock Development, Bangkok, p. 31. j3:Oram , R.N., and spikelets. The species has long been thought 1990. Register of Australian herbage plant culti- to be identical to the American H. amplexicaulis vars. CSIRO, Australia, pp. 107-108. I41 Soerjani, (Rudge) Nees, causing confusion in botanical liter­ M., Kostermans, A.J.G.H. & Tjitrosoepomo, G., ature. 1987. Weeds of rice in Indonesia. Balai Pustaka, New shoots grow from the old plants at the begin­ Jakarta, No 3521. pp.432-433 . ning of the rainy season. Very few seeds are pro­ C. Manidool duced in Thailand, although it sets seed during autumn in Australia. It flowers from the middle of the rainy season onwards. Cultivar 'Olive' has Imperata cylindrica (L.) Raeuschel been released in Australia. Ecology H. acutigluma thrives in open swamps Nomencl. bot., ed. 3:10 (1797). and ditches from sea-level up to 1200 m altitude. GRAMINEAE It is adapted to acid soils in peat swamps having 2n = 20 pH 4-4.8. It is intolerant of shade. It grows best Synonyms Lagurus cylindricus L. (1759), Impe­ in high rainfall regions. It can tolerate water up rata arundinacea Cyr. (1792), I. cylindrica (L.) to 1 m deep, but is better adapted to seasonal im­ Raeuschel var. major (Nees) C.E. Hubbard (1940). mersion than to permanent water. Vernacular names Blady grass (Australia), Agronomy Propagation by rooted stolons and cogongrass, satintail (En). Paillotte (Fr). Indone­ division of rootstocks is best. Good soil moisture sia: alang-alang, ilalang (Indonesian), kambengan is needed for its establishment; cuttings should be (Javanese). Malaysia: lalang, alang-alang. Papua planted deeply in saturated soils. It is not drought- New Guinea: kunai. Philippines: kogon (Tagalog), tolerant, but has been observed to recover from gogon (Bikol), bulum (Ifugao). Burma: kyet-mei. seedlings when flooding recommenced after Cambodia: sbö':w. Laos: hnha:z kh'a:. Thailand: extended drought periods in Australia. Topgrowth ya-kha. Vietnam: co'tranh. of H. acutigluma above water level can be cut sev­ Origin and geographic distribution A native eral times throughout the rainy season. The tops of the Old World tropics that is widely distributed are then carried to animals. In Thailand it is also throughout the tropics and subtropics of South- grazed by cattle or water buffaloes when flood East Asia, Africa, the Indian subcontinent and water recedes.I n Australia it is established behind Australia. It occurs to a lesser extent in North, ponding banks where it tolerates deeper water Central and South America. It also occurs in warm than para grass (Brachiaria mutica (Forssk.) temperate areas and has been recorded at latitudes Stapf) and is only used for grazing. In Thailand, of 45°i n and Japan. it is estimated that a minimum yield of 24 t/ha of Uses /. cylindrica, in an early stage of growth, fresh grass can be obtained from peat swamps. It iswidel y used as feed for ruminants, including cat­ does not make a good hay as the succulent stems tle, buffaloes, goats and wild herbivores. It is tradi­ take a long time to dry. tionally used in the countryside as roofing materi­ Genetic resources and breeding There is a al as it is readily available and durable. It has also noticeable variation within the species. It is un­ been used to make paper. It is useful for mulching likely that any substantial germplasm collections or controlling soil erosion, and the rhizomes are are being maintained, although small collections used medicinally in a decoction to purify the blood are held by ATFGRC (CSIRO, Australia). and as a diuretic. The rhizomes contain a fair Prospects It is a good source of feed for live­ amount of starch and a kind of beer can be made stock in lowland areas that are subject to lengthy from them. It is regarded as a serious weed in crop­ periods of flooding, especially when no other quali­ ping systems. ty forage is available. Research on its agronomy Properties /. cylindrica is regarded as a low IMPERATA 141 quality grass. Nitrogen concentrations may only branches ascending close to the main axis espe­ remain above 1 % for some 6week s where it grows cially at anthesis; spikelets paired, bisexual, 3-6 rapidly, but can remain above this level for up to mmlong , 1-flowered, at thebas e with a dense whorl 20 weeks with slower growth at higher latitudes of silky white hairs up to 2.5 cm long; pedicels un­ and altitudes. Very young growth may have diges­ equal, up to 1m m long; glumes equal, 3-9-nerved; tibilities of 70%, falling to below 40% after 150 lower floret reduced to a hyaline lemma; stamens days.Wher e there isopportunit y for selective graz­ 2, anthers orange to purple; stigmas 2,purple . Car­ ing, other species are usually selected ahead of it. yopsis ellipsoid, ca. 1m m long, brown. Description Aggressively rhizomatous, robust Growth and development I. cylindrica is a pro­ perennial with white, deeply buried (up to more lific seed producer and the light seeds are readily than 1 m),branched , fleshy, scaly rhizome, forming dispersed by wind. There isn o dormancy and seeds loose to compact tufts of leaves. Culms erect, may give 95% germination within one week after 0.1-1.2(-3 m) tall, l-4(-8)-noded, unbranched, harvest. The optimum temperature for growth is solid, usually hairy at the nodes. Leaf-sheath with about 30° C with negligible growth at 20° C and ciliate margins, lower ones broad and leathery, 40°C . Some individual plants flower frequently, overlapping, the upper ones finally splitting into some never flower, and other are intermediate. thin fibres; ligule membraneous, truncate, up to 1 The rhizomes are highly competitive and pene­ mm tall; leaf-blade linear-lanceolate, 10-180 cm x trate the roots of other plants, causing rot or 5-25 mm,flat , erect, spreading or drooping, pilose death. at the base, when old with hard, serrate, cutting Other botanical information Based on e.g. ap­ edges and a stout whitish midrib. Inflorescence a pearance ofleaf-blade s (rolled or flat), size of spike- spiciform panicle, cylindrical, 6-30 cm x 2 cm, its lets and hairiness of the node, several botanical varieties have been distinguished in the taxono­ mie literature. All varieties, however, intergrade so much that individual specimens are often unidentifiable. It seems best, therefore, to ignore the varietal classification, with the understanding that there are a number of imperfectly separable geographical variants. Cultivars have never been developed as research activities are much more di­ rected towards the eradication of/, cylindrica than towards its cultivation! Ecology /. cylindrica is often found in areas receiving more than 1000m m rainfall, but has been recorded in sites receiving from 500-5000 mm. It can withstand waterlogging but not continuous flooding. It grows at altitudes from sea-level up to 2000 m in several countries and has been recorded at 2700 m in Indonesia. It has been estimated that it covers some 500millio n ha worldwide including 200millio n ha in South-East Asia. Whereas I. cylindrica may have originally been re­ stricted to very low fertility and acid soils in the tropics, it has become widely spread through man's intervention, particularly following slashing and burning of forest lands. Its resistance to burning is associated with its vigorous underground rhi­ zomes, but seedlings also establish after burning. It is found in a wide range of habitats including dry sand dunes of seashores and deserts, as well as swamps and river valleys. It grows in grassland, Imperata cylindrica (L.) Raeuschel - 1, habit; 2, cultivated areas, and plantations. It quickly ligule, 3, inflorescence; 4, spikelet; 5, two views of invades abandoned farmland and occurs on rail­ a caryopsis. way and highway embankments and in deforested 142 FORAGES areas. It isregarde d as a light-loving plant and will legumes, this will only take place very slowly. not persist under heavy shade in plantations. Al­ Care should be exercised that replacement of this though it grows in a wide range of soil types with species does not lead to invasion of inedible weeds widely differing fertility levels, it grows most and/or increased soil erosion. vigorously in wet soil ofreasonabl e fertility. It has Literature |1| Bor, N.L., 1960. The grasses of been reported to grow on soils with pH ranging Burma, Ceylon, India and Pakistan. Pergamon form 4.0-7.5. It can even tolerate very hot, steamy Press, Oxford, pp. 169-171. ;2| Falvey, «LL., 1981. and sulphurous conditions near an active volcanic Imperata cylindrica and animal production in fumarole or vent. South-east Asia: a review. Tropical Grasslands 15: Propagation and planting I. cylindrica is sel­ 52-56. |3|Holm , L.G., Plucknett, D.L., Pancho, J.V. dom propagated deliberately, but spreads by rhi­ & Herberger, J.P., 1977. The world's worst weeds: zomes and seed. If rhizomes are cut by cultivation, distribution and biology. The East-West Center, they can establish from pieces with as few as 2 University of Hawaii Press, Honolulu, Hawaii, nodes. United Sates, pp. 62-71. j4i Santiago, A., 1965. Husbandry I. cylindrica is favoured by burning, Studies on the autecology ofI .cylindrica . Proceed­ which can lead to virtually monospecific swards. ings of the Ninth International Grassland Con­ If it is to be used for thatching, swards are left gress, Säo Paulo, Brazil, pp. 499-502. |5| Skerman, ungrazed until after the mature growth is P.J. & Riveros, F., 1990. Tropical grasses. FAO, removed. It can be eliminated by heavy continuous Rome. pp. 368-373. |6| Soerjani, M., 1970. Alang- grazing, where it may be replaced by inedible alang, Imperata cylindrica (L.) Beauv., pattern of weeds; vigour will be reduced by close, frequent growth as related to its problem of control. BIO- cutting and rhizomes can be destroyed by frequent TROP Bulletin 1.Regiona l Centre for Tropical Bi­ intensive cultivation. If 7.cylindrica is being regu­ ology, Bogor. 88pp . larly used for grazing, it has been suggested that N.O. Aguilar it be rotationally grazed when 15-25 cm high. Because of its aggressiveness and low quality, it is widely regarded as a weed and research has been Ischaemum ciliare Retzius carried out on improving its quality through intro­ ducing legumes or even replacing it with other Observ. bot. 6:3 6(1791) . grasses. However, it is not easy to maintain herba­ GRAMINEAE ceous legumes with /. cylindrica, though some pro­ In = 36 mising results have been achieved with stylo (Sty- Synonyms Ischaemum indicum auct. (non losanthes guianensis (Aublet) Swartz) and with (Houtt.) Merrill, 1938), /. aristatum auct., (non L., leucaena (Leucaena leucocephala (Lamk) de Wit). 1753). Diseases and pests There are no records of Vernacular names Batiki blue grass, smut- major diseases or pests on I. cylindrica. grass (En). Indonesia: blembem (Javanese), rum- Harvesting I. cylindrica is either grazed or cut put padang (Billiton). Malaysia: rumput gerek tel- for thatch when mature. It islittl e used as a species inga. Thailand: ya wai (eastern), ya-yonhu for cut-and-carry grazing systems. (southern). Yield Reported DM yields of I. cylindrica range Origin and geographic distribution Batiki from 2-11 t/ha per year. Weight gains of 0.25 kg/ blue grass is native to South and South-East Asia. head per day have been recorded from I. cylindrica It was introduced to West Africa, southern Eu­ pastures in the Philippines and Papua New Gui­ rope, Australia and the Pacific Islands. nea. In each case, gains from alternative, improved Uses It isuse d as forage and also as a cover grass pastures were two or three times higher. Very low to bind soil and reduce erosion. A dwarf form is liveweight gains of 0.04 kg/head per day were used for lawns. recorded in northern Thailand. Properties Nitrogen concentrations ranged in Genetic resources and breeding There are no experiments from 2.2% at 3 weeks to 1.7% at 6 known breeding programmes on this species. weeks, 1.3% at 8 weeks and to below 1.0% in Prospects Although I. cylindrica is a low-quali­ mature forage. There have been reports that it ty forage, it is an important forage resource in taints milk. South-East Asia and is often inadequately appre­ Botany Aperennial , spreading or tufted stoloni- ciated. Although there is potential for improving ferous grass rooting at the nodes, with erect or its productivity through growing associated with geniculate culms up to 60 cm tall, often branching ISCHAEMUM 143

3 varieties with one variety divided into 3 subvar- ieties, but the whole complex needs a thorough investigation with modern experimental taxono- micalmethods .Th e species has for a long time been considered to be /. aristatum L., but this has gla­ brous nodes, an unlobed lower glume and only its sessile spikelets are awned. Since 1938thi s species has been known as I. indicum (Houtt.) Merrill. In 1991, however, Veldkamp discovered that the basionym for /. indicum, Phleum indicum Houtt., was another species {Polytrias indica (Houtt.) Veldk.) and consequently the correct name for this species became I. ciliare. Ecology Batiki blue grass is an opportunistic invader of open or disturbed habitats. It is adapted to a wide range of rainfall regimes, including waterlogged areas in the wet tropics receiving over 2000m m annual rainfall and to seasonally dry monsoonal areas of India. It has been reported as having some shade tolerance. It tolerates acid soils and soils with poor fertility, but responds to appli­ cations of fertilizer on such soils. Agronomy Batiki blue grass can readily be es­ tablished from rooted cuttings and a spacing of 30 cm x 30c mha s been recommended. Establishment from seed is much slower and requires good seed­ bed preparation. It is usually harvested by grazing Ischaemum ciliare Retzius - 1, habit flowering animals, but can be cut and fed fresh or used to plant; 2, habit stoloniferous plant part; 3, ligule; 4, make hay and silage. It should be cut at booting pair of spikelets. as quality ispoo r if left for longer. Four cuts a year are possible. and hairy at the nodes. Leaf-sheath 3-6 cm long, In Malaysia flowers are often infected with a smut often tight, glabrous or sparsely hairy; ligule mem­ fungus resulting in diseased inflorescences that do branous, truncate, 1-2 mm long, entire or lacerate; not emerge from the uppermost leaf-sheaths. leaf-blade narrowly lanceolate, up to 20c m x 1 cm, Dry matter yields of Batiki blue grass have ranged acuminate at apex, usually sparsely to densely from 3-20 t/ha per year depending primarily on hairy. Inflorescence well exserted, composed of soil fertility and fertilizer application. When two terminal, closely opposed or somewhat diver­ grown with Stylosanthes guianensis (Aublet) gent racemes 2-10 cm long; spikelets paired, bisex­ Swartz in Sarawak, the mixture yielded 14.5 t/ha ual, one sessile, one pedicelled, alternately on one ofDM. side of the triangular rachis; rachis and pedicel Genetic resources and breeding Although it hairy along angles; lower glume winged at apex, is a very variable species, it is unlikely that sub­ forming 2 large obtuse lobes; upper lemma deeply stantial germplasm collections are being main­ notched, excurving into a long kneed and twisted tained and there are no breeding programmes. awn 8-15 mm long. Prospects Batiki blue grass is a useful forage Seeds have a dormancy period and germination plant in several areas of South-East Asia and the improves after 9-10 months storage. Seedling Pacific Islands. It is a competive grass and resists growth is vigorous. weed invasion. It is palatable, well grazed and can It is a short-day plant, flowering throughout the persist under heavy grazing. There are good pros­ year in Malaysia, in June-July in Thailand, and pects for its use, especiallly in plantation crops in July-August and January-February in India. and in moist areas. Germplasm collections of this Seed set is often poor and the grass spreads prima­ species are needed so that its variability can be rily by stolons. assessed and utilized in agronomic studies for the /. ciliare is a very variable species, subdivided into release of new cultivars to farmers. 144 FORAGES

Literature lij Bor, N.L., 1960. The grasses of Burma, Ceylon, India and Pakistan (excluding Bambuseae). Pergamon Press, Oxford, pp. 180-182. |2|Dunsmore , J.R. &Ong , C.B., 1969. Pre­ liminary work on pasture species and beef produc­ tion in Sarawak, Malaysia. Tropical Grasslands 3: 117-121. |3|Gilliland , H.B., Holttum, E.E. & Bor, N. L., 1971. Grasses of Malaya. In: Burkill, H.M. (Editor): Flora of Malaya. Vol. 3. Government Printing Office, Singapore, p. 263. |4| Häfliger, E. &Scholz , H., 1980.Gras s weeds 1. Ciba-Geigy Ltd., Basel, Switzerland, p. 64. |5>Ng , T.T., 1976. Perfor­ mance of some tropical grass-legume mixtures in Sarawak. Malaysian Agricultural Journal 50: 400-410. |6|Ranacou , E., 1986.Eevie w of research and recorded observations on pastures in Fiji (1920-1985). III. Batiki blue grass (Ischaemum indicum). Fiji Agricultural Journal 48: 24-29. |7| Skerman, P. J. & Riveros, F., 1990. Tropical grasses. FAO, Rome. 474-477. |8| Veldkamp, J.F., 1991. Miscellaneous notes on Southeast Asian Gramineae. Blumea 36:180-181. B.B. Baki, LB.Ipo r &C . Manidool

Ischaemum magnum Rendle

J. Bot. 32:102(1894) . GRAMINEAE Ischaemum magnum Rendle - 1,flowering culm; 2, 2n = unknown leaf; 3, ligule; 4,pair of spikelets. Synonyms Ischaemum laeve Ridley (1905). Vernacular names Malaysia: rumput melayu, without, seldom with small awns; each raceme rumput tembaga kasar. with spikelets arranged in pairs, one sessile, one Origin and geographic distribution I. mag­ pedicelled, on one side ofa triangular hairy rachis; num occurs in restricted areas of Peninsular spikelets similar but the pedicelled spikelet has a Malaysia, Singapore, Borneo, Thailand and Burma lower glume with a narrow wing all down one side. between latitudes 20°N and 5°S. It flowers and sets seed throughout most of the Uses I. magnum is not well recognized as a for­ year, but the peak flowering time in Malaysia is age grass for grazing but its young growth is readi­ in October-November. The maturation of the ly eaten by free-ranging cattle and goats. spikelets is not synchronized and seeds easily shat­ Properties Herbage obtained from cutting ter. This makes seed harvesting difficult and also experiments contained 0.6-1.3% N, 0.10-0.20% P means that it has the potential tob e a weed in culti­ and 1.2-2.6% K depending on the age of regrowth vated areas. and soil fertility. There are about 400 seeds/g. Ecology I. magnum is found abundantly in both Botany A perennial tussock grass with stout full sunlight and under shade and is a common culms,u p to 2m tal l and a strong root system. Leaf- component of natural pastures. The optimum tem­ sheath terete, up to 16 cm long, fringed with fine perature for growth in its natural habitat is hairs; ligule a prominent membrane fused with the 30-35°C (day) and 22-24°C (night) with 12 hours auricles, 4 mm long; leaf-blade large, lanceolate- daylength varying by only 30 minutes throughout acuminate, up to 30 cm x 1.8 cm, abruptly nar­ the year. It grows in areas receiving 2000-4000 mm rowed to a petiole at the base, often hairy near the annual rainfall. It is highly tolerant of flooding base, the margins scabrid. Inflorescence composed and waterlogged conditions and hence is found in of two terminal, well-exserted, one-sided racemes ditches and canals. It is adapted to a wide range closely appressed together, 9-18 cm long, usually of soils ranging from sands, marine peats to heavy ISCHAEMUM 145

clays. It grows aggressively and can compete with tilizer nitrogen and pasture legumes. Tropical common weeds including Imperata cylindrica (L.) Grasslands 10: 179-185. ,!3| Skerman, P.J. & Rive- Raeuschel. It can also survive short, dry periods. ros, F., 1990. Tropical grasses. FAO, Rome. pp. Agronomy I. magnum can be established by 478-479. seed or by vegetative propagation. Seed is sown T.T. Ng into a prepared seed-bed and usually about 30% germinates. Rooted tillers are planted about 1 m apart in wet soil. Seedling growth is slow, but once Ischaemum muticum L. established, the plants are able to regrow despite severe slashing or annual burning. Sp. PL: 1049(1753) . No serious pests and diseases have been recorded GRAMINEAE but prédation by birds reduces seed set. 2n = unknown Herbage is well grazed by cattle when it is young Synonyms Ischaemum repens Roxb. (1820), 7. but palatability declines towards maturity. Grass glabratum Presl (1830),Andropogon repens (Roxb.) swards containing I. magnum are usually burnt Steud. (1854). yearly to obtain new growth. It is compatible with Vernacular names Seashore centipede grass, Centrosema pubescens Benth. and Stylosanthes drought grass (En). Indonesia: suket resap (Java­ guianensis (Aublet) Swartz. nese), rumput kerupet (Bangka). Malaysia: rum- Legumes such as Centrosema pubescens, Desmo- put kemarau, rumput tembaga jantan. Thailand: dium heterocarpon (L.)DC . ssp.ovalifolium (Prain) ya-waitham (eastern). Vietnam: môm trui. Ohashi and Stylosanthes guianensis tend to domi­ Origin and geographic distribution 7. muti­ nate the grass under continuous grazing. Herbage cum is indigenous to South and South-East Asia. can be cut for stall feeding but it is usually grazed It is widely distributed in Malaysia, particularly by free-ranging animals. Under heavy grazing or near the sea and in sandy places, and in Thailand, regular cutting it develops a more prostrate Indonesia, the Philippines, Sri Lanka, India, growth habit. Burma, and some islands in the western Pacific In a 3-year cutting trial in Malaysia, the mean Ocean. It has been introduced to West Africa and annual DM yield of pure swards of I. magnum, fer­ Australia. tilized with 200, 25 and 190 kg/ha of N, P and K, Uses I. muticum provides fodder, but is also used respectively, and cut every 8-10 weeks, was 13.7 to protect soil from erosion (especially coastal t/ha. When grown with the legumes Centrosema sand dunes) and to make compost and mulch. It is pubescens, Pueraria phaseoloides (Roxb.) Benth. a weed in many annual and perennial crops. and Macroptilium atropurpureum (DC.) Urban, Properties 7. muticum is a palatable and nutri­ total DM yields were about 10 t/ha. These DM tious grass and Nconcentration s of1. 5-2. 4 % have yields were similar to those from comparable been measured. swards of Brachiaria decumbens Stapf, although Botany A leafy, much branched, stoloniferous the total yield of N from I. magnum swards was spreading perennial, rooting at the nodes where slightly less. Yields declined markedly when the they touch the soil, sometimes scrambling several cutting interval was reduced to 4weeks . m high among and over shrubs; stolons long, up Genetic resources and breeding It is unlikely to 5 m or more, with numerous joints and up to 6 that any germplasm collections are being main­ mm in diameter, covered by slightly overlapping tained and there are no selection or breeding pro­ pale or purple leaf-sheaths with extremely short grammes with ƒ. magnum. leaf-blades, resembling culm-sheaths in bamboos; Prospects I. magnum has many advantages flowering culms up to 60 cm tall, glabrous, often including hardiness, persistence and productivity red. Leaf-sheath ciliate along outer margin; ligule in harsh environments, as well as acceptability by truncate, small, shortly ciliate; leaf-blade ovate- livestock. Research into all aspects of the agro­ lanceolate to linear, 2-18 cm x 0.5-2 cm, apex nomy of 7.magnum would be warranted. acute, base cordate, narrowing into a short Literature |1| Gilliland, H.B., Holttum, R.E. & pseudo-petiole. Inflorescence terminal, composed Bor, N.L., 1971. Grasses of Malaya. In: Burkill, of 2raceme s closely pressed together, each 1.5-5.5 H.M. (Editor): Flora of Malaya. Vol. 3. Govern­ cm long, rarely fully exserted; spikelets arranged ment Printing Office, Singapore, pp. 261-262. |2| in pairs, one sessile and one pedicelled, on one side Ng, T.T. &Wong , T.H., 1976. Comparative produc­ of a triangular rachis, straw-coloured; the sessile tivity of two tropical grasses as influenced by fer­ spikelet glabrous, lower glume distinctly reticula- 146 FORAGES

cum although it isoccasionall y infested with smut. High DM yields of up to 25t/h a per year have been measured. It is usually harvested by grazing ani­ mals and only occasionally by cutting. Both fresh and dry foliage can be used for composting or mulch. Genetic resources and breeding It is unlikely that substantial germplasm collections are being maintained and there are no breeding pro­ grammes. Prospects /. muticum isa useful forage in specif­ ic situations, but it is uncertain whether it will be specifically cultivated in the future. Literature |1| Backer, CA. & Bakhuizen van den Brink, R.C., 1968. Flora of Java. Vol. 3. Wolters-Noordhoff, Groningen, the Netherlands, p. 592. |2jBarnes , D.E. &Chan , L.G., 1990. Common weeds of Malaysia and their control. Ancom Ber- had, Shah Alam, Malaysia, p.22 . |3|Gilliland , H.B., Holttum, R.E. & Bor, N.L., 1971. Grasses of Malaya. In: Burkill, H.M. (Editor): Flora of Malaya. Vol. 3. Government Printing Office, Singapore, pp. 255-257. LB. Ipor &B.B . Baki

Ischaemum muticum L. - 1, habit leafy plant part; 2, ligule; 3,flowering culm; 4,pair of spikelets. Ischaemum rugosum Salisb.

.tely nerved near the apex, enfolding the spikelet Icon. Stirp. Rar.: 1,fig . 1(1791) . base, upper glume with a strongly keeled acute GRAMINEAE apex; lower floret male or hermaphrodite, upper 2n = 18, 20 floret hermaphrodite; the pedicelled spikelet Synonyms Ischaemum colladoa Sprengel (1825), hairy, lower glume chartaceous, with a winged /. segetum Trinius (1832). keel, upper glume subchartaceous; the upper Vernacular names Wrinkle duck-beak, sara- lemma of both spikelets short-awned, awn mattagrass, wrinkled grass (En). Indonesia: suket included in spikelet. blembem (Javanese), jukut randan (Sundanese). Ecology I. muticum is an opportunistic and Malaysia: rumput mele (Sarawak), rumput ekor agressive colonizer of open or disturbed habitats chawi, rumput colok cinia (). and can develop into huge thickets in drainage ca­ Philippines: tinitrigo (Tagalog), daua (Subanum), nals and ditches. On the edges of secondary forest, gulong lapas (Pangasinan). Burma: ka-gyi-the- plants grow very large, scrambling among bushes. myet. Cambodia: smao srauv. Thailand: ya daeng It tolerates wet conditions, especially flooding, (central), ka-du-ai-nu, ya-noksichomphu. Viet­ and short dry periods. It is usually found in areas nam: môm u. receiving more than 1500m m rainfall annually. Its Origin and geographic distribution I. rugo­ presence is sometimes regarded as indicative of sum is indigenous to tropical Asia and is widely poor soil fertility. distributed throughout the tropics. Agronomy /. muticum can be propagated by Uses /. rugosum is used as forage. It is a serious stem cuttings and by seed. Newly harvested seed weed in many crops, particularly in rice fields. It is dormant, so scarification is desirable to improve also provides suitable material for compost and germination. Planting with stem cuttings gives mulch. In times of scarcity the grain is eaten by faster establishment than sowing seed. people. Although adapted to infertile conditions, it res­ Properties The forage quality of /. rugosum de­ ponds to fertilizer application on such soils. clines quickly with age of the material. Nitrogen There are no serious diseases and pests of I. muti­ concentration ranges from 0.5-1.2%. ISCHAEMUM 147

been distinguished: var. rugosum, with developed pedicelled spikelets, and var. segetum Hackel, with much reduced pedicelled spikelets; they are not separated geographically. Seeds germinate early in the wet season and grow vigorously. Swards may form a dense mass of sturdy culms 25-30 weeks after germination. Ecology /. rugosum is an opportunistic and effective coloniser of open, disturbed or newly cleared areas! Although a sun-loving plant it can persist in sites receiving only 30-35% of full sun­ light. It occurs at altitudes of up to 2400 m in the Philippines. I. rugosum is particularly well adapted to wet sites, and is often found in rice fields or low lying areas that are periodically flooded. It tolerates acid soils of pH(H20) 4.0. Agronomy I. rugosum can be established by seeds or rooted culms. Newly harvested seeds appear to have innate or induced dormancy, and scarification is desirable to reduce hard-seeded- ness. When sowing seed, good seed-bed prepara­ tion assists in obtaining good establishment and seedling growth. Planting rooted culms at a 25 cm x 25 cm spacing will result in faster sward devel­ opment than using seed. Although it "will persist on soils ofpoo r fertility, it will respond to fertilizer application. Ischaemum rugosum Salisb. - 1, habit flowering I. rugosum has no major diseases or pests. Infesta­ plant; 2, ligule; 3,pair ofspikelets. tions by Puccinia spp. are commonly observed; they may reduce the quantity and quality of feed Botany A vigourous perennial or annual (in available, but control by chemicals is not war­ strongly desiccating soil) tufted grass, sometimes ranted. It is an alternative host of the viruses caus­ with stilt roots, rooting at the nodes, with erect, ing rice and maize leaf blight galls and of Piricu- slanting or ascending, often much branched culms, laria sp. up to 1.5 m tall. Leaf-sheath long-auricled, ciliate It can be grazed, but will not persist when conti­ along outer margin, densely soft hairy on node; nuously grazed by large ruminants at high stoc­ ligule a brownish truncate membrane, 1-7 mm king rates. When it is growing in water, the tops long; leaf-blade linear, 10-40 cm x 1-4 cm, apex above water level can be cut and fed to animals. acute, rarely hairy with long slender hairs. Inflor­ It is usually fed when fresh, but can be ensiled or escence terminal, well exserted, composed of 2 dried and conserved for dry season feeding. racemes that are firmly appressed together and Genetic resources and breeding It is a very interlocked when young, separating when mature, variable species, but it is unlikely that substantial each 3-12 cm long; spikelets binate, dissimilar, germplasm collections are being maintained. one sessile at the abaxial side of the rachis, one Prospects I. rugosum will continue to provide stalked at the adaxial side, provided with a short useful forage in existing feeding systems. As it is blunt hairy callus; sessile spikelet 5-6 mm long, a very variable species, there is potential for 2-flowered, lower floret male or neuter, upper one selecting for better forage types and for studies on bisexual; lower glume strongly transversely its agronomic requirements and grazing manage­ 5-7-ribbed, and winged above;uppe r lemma deeply ment. 2-cleft and with 1-2 cm long awn which is twisted Literature |1| Bor, N.L., 1960. The grasses of basally; pedicelled spikelet more or less reduced, Burma, Ceylon, India and Pakistan. Pergamon pedicel up to 2m m long, hairy, confluent with the Press, London, pp. 184-185. !2! Gilliland, H.B., callus. Caryopsis ovoid, 2m m long, brown. Holttum, R.E. & Bor, N.L., 1971. Grasses of It is a very variable species. Two varieties have Malaya. In: Burkill, H.M. (Editor): Flora of 148 FORAGES

Malaya. Vol. 3. Government Printing Office, Singapore, p. 259. |3|Holm , L.G., Plucknett, D.L., Pancho, J.V. &Herberger , J.P., 1977. The world's worst weeds: distribution and biology. The East- West Center, University of Hawaii Press, Hono­ lulu, Hawaii, United States, pp. 295-298. |4| Mani- dool, C, 1989. Natural grassland and native grasses of Thailand [in Thai]. Technical Bulletin No 1301-26-32. Division of Animal Nutrition, Department of Livestock Development, Bangkok, p. 19. |S!Mehra , K.L. & Fachrurozi, Z., 1985. Indo­ nesian economic plant resources: forage crops. Lembaga Biologi Nasional - LIPI, Bogor, No 31. p. 21.|6 | Soerjani, M., Kostermans, A.J.G.H. & Tji- trosoepomo, G., 1987. Weeds of rice in Indonesia. Balai Pustaka, Jakarta. No 3521. pp. 440-441. B.B. Baki &C . Manidool

Ischaemum timorense Kunth

Kévis. gramin. 1:369 ,t.9 8 (1830). GRAMINBAE 2re = 20,3 6 Synonyms Ischaemum macrurum Stapf ex Bid- ley (1925). Vernacular names Centipede grass, lucuntu grass (En). Indonesia: jukut jampang manggung Ischaemum timorense Kunth - 1, flowering plant (Sundanese), rumput apet (Sumbawa). Malaysia: part; 2, ligule; 3, pair of spikelets; 4, caryopsis in rumput sarang buaya. Vietnam: môm timor. two views. Origin and geographic distribution I. timor­ ense is indigenous to tropical Asia and is widely Inflorescence terminal, composed of 2(-3) closely distributed from India to Malesia and Polynesia. opposed racemes, each 2-15 cm long; spikelets in Uses I. timorense is a fodder species of minor pairs, one sessile, one pedicelled; alternately on importance in South-East Asia. It is useful for pro­ one side of the triangular rachis; spikelets similar, tecting soil against erosion and in providing mate­ 2-flowered, green or tinged with purple, lower rial for mulch, but can be a weed in annual and floret male, upper floret bisexual; sessile spiklet perennial crops. Particularly in Indonesia it is a 5-7 mm long, at the base swollen and stipe-like and common weed in rainfed rice. white-hairy, lower glume with two acute lobes at Properties When well fertilized with N, P and apex, upper glume with a short 2-3 mm long awn, K, three-week-old herbage had a N concentration upper lemma 2-lobed with a 10-16 mm long awn of 3.2%, falling to 2.6% at 6 weeks. The N concen­ in the middle. Caryopsis ellipsoid, 1-2 mm long. tration of leaves was approximately double that of The flowering season in Indonesia is April- the stems. November. Botany A spreading stoloniferous perennial or Ecology As /. timorense establishes readily from annual, creeping and rooting in lower part, erect, seed and spreads byroote d stolons,i t is an opportu­ slanting or scrambling in upper part with culms nistic colonizer of bare or disturbed areas. Hence up to 1 m high. Leaf-sheath 3-6 cm long, white- it is common along roadsides, terraces, ditches and hairy on the nodes and often with long hairs at the forest margins, and it is a weed in agricultural mouth, the outer margin and the back; ligule a crops. It is found most frequently in areas with short fringed membrane, sometimes long ciliate; 800-2000 mm annual rainfall, but is not adapted to leaf-blade lanceolate to linear-lanceolate, 2-16 cm either dry or waterlogged sites or to heavy soils. It x 3-15 mm, base obtuse or petiole-like, apex grows under full or partial (30-50%) sunlight. It is acute, adpressed long-hairy, or glabrous above. tolerant ofhig h soil acidity and poor soil fertility. LEPTOCHLOA 149

Agronomy /. timorense can be established by India, China, Japan, Australia and from East to sowing 3-6 kg/ha into a well prepared seed-bed. South Africa. If recently harvested seed is used, scarification Uses L. chinensis is used as fodder. Normally it may be required to break dormancy. It can also be is regarded as a weed in paddy fields but livestock established by spreading cut grass on to the sur­ owners collect and feed it to their animals. In East face ofmois t soil and disking it in. Africa the grain is used as famine food. There are no serious diseases or pests of I. timor­ Properties It is highly palatable and very leafy, ense, although spikelets are often infected by smut. but there is no information on its nutritive value. I. timorense is usually harvested by grazing ani­ Botany An aquatic or semi-aquatic tufted annu­ mals and only rarely by cutting. It is well grazed al or perennial, with stout to slender, erect or geni­ by cattle, horses and sheep. Some form of resting culate culms up to 1.5 m tall, often rooting at the or rotational grazing may be required so that suffi­ lower nodes. Leaf-sheath loose, subglaucous, cient seeding can take place to ensure long-term smooth, up to 10 cm long; ligule a fringed, hairy persistence. It is not a particularly vigorous grass, membrane, 1-2 mm long; leaf-blade linear, up to although it is more productive in very fertile situa­ 50 cm x 1 cm, long-attenuate, flat or folded, sca- tions. Yields of3 0t/h a per year offres h forage have brid above.Inflorescenc e 10-60 cmlong , composed been obtained. Both fresh and dry foliage can be of numerous slender racemes scattered along an used for composting and mulching, and it can be elongate central axis; racemes flexuous, 2-13 cm conserved,a s hay. long, erect or laxly ascending; spikelets 3-7 flow­ Genetic resources and breeding It is unlikely ered, narrowly elliptical-oblong, 2-3 mm, subses- that there are substantial germplasm collections sile, often purplish, disarticulating above the and there are no known breeding programmes. glumes and between the florets; glumes unequal, Prospects Although /. timorense is a useful for­ age in specific situations, its dry matter yield is too low to warrant cultivation. Literature |1|Bogdan , A.V., 1977. Tropical pas­ ture and fodder plants. Longman, London, p. 161. •2| Gilliland, H.B., Holttum, R.E. &Bor , N.L.,1971 . Grasses of Malaya. In: Burkill, H.M. (Editor): Flora of Malaya. Vol. 3. Government Printing Office, Singapore, pp. 264-265. |3I Mehra, K.L. & Pachrurozi, Z., 1985. Indonesian economic plant resources: forage crops. Lembaga Biologi Nasio- nal-LIPI, Bogor, No 31.p .22 . |4|Soerjani , M., Kos- termans, A.J.G.H. & Tjitrosoepomo, G., 1987. Weeds of rice in Indonesia. Balai Pustaka, Ja­ karta, pp.442-443 . LB. Ipor, B.B. Baki &C.P . Chen

Leptochloa chinensis (L.) Nees

Syll.Eatisb. 1:4 (1824). GRAMINEAE 2« = 40 Synonyms Poa chinensis L. (1753). Vernacular names Indonesia: timunan (Java­ nese), bebontengan (Sundanese), jangkiri (Flores). Philippines: palay-maya (Tagalog), karukauâyan (Bikol).Thailand : ya-yonhu, ya-dokkhao (central), ya-metnga (northeast). Vietnam: côduó i phung. Origin and geographic distribution L. chinen­ sis originated in tropical Asia and is distributed Leptochloa chinensis (L.) Nees - 1, habit flowering throughout South-East Asia, Burma, Sri Lanka, plants; 2, ligule; 3, spikelet. 150 FORAGES scabrid on the back of the nerves; lemmas hairy Leucaena leucocephala (Lamk) de Wit on the nerves, awnless. Caryopsis ellipsoid-oblon- goid, 6-9 mm long, brown, smooth or wrinkled. Taxon 10:5 3(1961) . L. chinensis is an abundant seed producer. The LEGUMINOSAE seedlings develop quickly so that they can keep In = 104 pace with the rising level of flood water and thus Synonyms Leucaena glauca (Willd.) Benth. survive. It flowers over most of the year. (1842), Leucaena latisiliqua (L.)Gilli s (1974). The related, more widely spread L. panicea (Ret- Vernacular names Leucaena (En). Indonesia: zius) Ohwi (L. polystachya Benth. sensu Burkill) lamtoro, klandingan (Javanese), petai cina (Indo­ is eaten by cattle when young. It differs by the lon­ nesian). Malaysia: petai belalang, ipil-ipil. Papua ger leaf-sheaths (upt o 13.5cm) ,th e papillate-pilose New Guinea: kunai. Philippines: ipil ipil, elena hairs on the leaves and the 2-5-flowered spikelets. (Tagalog), palo-maria (Bikol), kariskis (Ilokano). L. panicea much resembles the American L. filifor- Cambodia: ka-thum' théét. Laos: kathin, kh'oonz mis (Lamk) P. Beauv. koong khaaw. Thailand: krathin (general), to-bao Ecology L. chinensis grows from near sea-level (southern). Vietnam: keo dâu, bo chét. up to 1400 m altitude. It is adapted to moist, Origin and geographic distribution Leucaena swampy places in open habitats, especially if dis­ evolved in the Guatemalan centre of origin, as a turbed, on heavy or light soils. probable tetraploid hybrid of diploid species in Agronomy L. chinensis ispropagate d by seed or that region. Two major forms are found. The 'com­ by rooted tillers. Adequate soil moisture is the mon' shrubby form grows up to 8 m tall and is evi­ main factor affecting the growth of this aquatic dently indigenous to the Yucatan Peninsula. The plant, although it has been observed that improved arboreal 'Salvador' type grows to 16m an d appears soil fertility in rice fields also results in better to have originated in the regions of Salvador, Gua­ growth. temala and Honduras. Both forms were distributed It is harvested by cutting forage above the water widely throughout Mexico and Central America level in paddy fields or by grazing. to northern South America prior to 1500 AD. The No yield data are available, but in open waste common form was probably brought by Spanish places it should givereasonabl e yields.I t can make galleons to the Philippines in the early 1600s, from good hay but when cut it is normally collected and whence it was pantropically distributed in the 19th fed green to livestock. Century. The Salvador forms are more recent in Genetic resources and breeding It is unlikely distribution and are known by names such as 'lam­ that any germplasm collections are being main­ toro gung' in Indonesia, 'giant ipil-ipil' in the Phi­ tained. lippines and 'subabul' in India. Leucaenas are Prospects L. chinensis will continue to be a use­ found throughout South-East Asia; on many ful supplementary feed source in lowland areas. islands common leucaenas dominate the vegeta­ Literature |1| Gilliland, H.B, Holttum, R.E. & tion on coralline soils. Bor, N.L., 1971. Grasses of Malaya. In: Burkill, Uses Leucaenas are versatile multipurpose H.M. (Editor): Flora of Malaya. Vol. 3. Govern­ trees. In South-East Asia they usually provide a ment Printing Office, Singapore, pp.73-75 . |2|Häf - combination of fodder, fuelwood, posts, shade, liger, E. & Scholz, H., 1981. Grass weeds 2. Ciba- food and green manure. Foliage is fed to ruminant Geigy Ltd., Basel, Switzerland, p. 94. |3| Mehra, animals as browse or by cut-and-carry methods, or K.L. & Fachrurozi, Z., 1985. Indonesian economic is milled for poultry and pelleted for export. Wood plant resources: forage crops. Lembaga Biologi is cut for home fuelwood and used in industries Nasional - LIPI, Bogor, No 31.p . 24. |4| Soerjani, such as ceramics or electrical power generation; M., Kostermans, A.J.G.H., Tjitrosoepomo, G., it is also converted into charcoal. Increasing use 1987.Weed s ofric e in Indonesia. Balai Pustaka No is made of the wood for posts and props, in chip­ 3521, Jakarta, pp. 446-447. board and plywood, for paper pulp, and for furni­ C. Manidool ture and parquet flooring. In Asia people eat the young green shoots prior to leaflet unfolding, but in the Americas the green seeds are eaten. Alley cropping involves planting leucaena hedges on contours at intervals of 3-6 m with crops in be­ tween. Hedges provide a high-nitrogen green ma­ nure, protect against soil erosion and can be har- LEUCAENA 151

vested for fodder or wood. ty may also be reduced. In most countries in South Other uses include living fences, support systems and Central America, the United States, Mexico, for vines like pepper and passionfruit, shade trees Indonesia, the Philippines and India, rumen mic­ for coffee and cocoa and ornamental plantings. robes can completely detoxify mimosine and DHP. Throughout the tropics leucaenas provide a major Isolation ofspecifi c bacteria has enabled the DHP- nitrogen-fixing component of lowland forests, degrading bacteria to be transferred successfully notably on wasteland where they are often a pri­ to Australia, Africa and China with beneficial mary source of fixed nitrogen in the ecosystem. effects for ruminants ondiet s containing much leu­ The dried seeds are widely used for ornamentation caena. and household items. The trees occasionally exude a gum very similar Production and international trade Leu­ to gum , with similar uses and properties; caena leaf meal is milled, pelleted and shipped sterile hybrids exude copiously. internationally in a very variable annual volume, Description Shrub or tree up to 18m tall, forked largely to Japan and Europe for animal feed. when shrubby or after coppicing, with greyish Demand is estimated to be up to 1 million t/year, bark and prominent lenticels. Leaves bipinnate far exceeding production, with world prices simi­ with 4-9 pairs of pinnae, variable in length up to lar to those for pellets or hay. Prices in local 35 cm, with a large gland (up to 5 mm) at base of markets in Asia vary widely for both fodder and petiole; leaflets 11-22 pairs per pinna, 8-16 mm x wood. However, leucaena is the primary legumi­ 1-2 mm, acute. Flowers numerous, in globose nous feed in large regions ofIndonesi a and the Phi­ heads with a diameter of 2-5 cm, white, calyx ca. lippines, and the trees are a major source of fuel- 2.5 mm, corolla ca. 5 mm, stamens (10 per flower) wood in these and other countries. Most and pistil 10m m long, anthers pilose, dehiscing at production is on communal lands or small farms. dawn. Pod 14-26 cm x 1.5-2 cm, pendant, brown Attempts to commercialize production on large plantations (1000 ha or more) for dendrothermal energy in the Philippines have not been a great success. Properties Leucaena foliage is noted for its good digestibility and high protein values. Typical values for 'browse fraction' of foliage include 55-70% digestibility, 3-4% N, 6% ether extract, 6-10% ash, 30-50% N-free extract (neutral deter­ gent fibre 20%), 1.5-2.5% tannins, 0.8-1.87% Ca and 0.23-0.27%) P. However, the Na levels are invariably low: 0.01-0.05%). The seeds and leaves contain galactomannan gums that block protein extraction and possibly its utilization by animals; they may potentially have useful biomedical pro­ perties. Leucaena also contains the toxic amino-acid mimosine which has antimitotic and depilatory effects on animals. It occurs in high concentra­ tions in the growing tips (8-12%), young leaves (4-6 %) and young pods and seeds (4-5 %). For this reason leucaena leaf cannot safely be included in rations for non-ruminants at a level above 5% on a DM basis. The mimosine ingested by ruminants is converted to 3-hydroxy-4(IH)-pyridone(DHP) by plant enzymes and rumen bacteria. The DHP is goi­ trogenic and, if not degraded, can result in low serum thyroxine levels, ulceration of the oesopha­ gus and reticulo-rumen, excessive salivation, poor appetite and low liveweight gains especially when Leucaena leucocephala (Lamk) de Wit - flowering the diet contains more than 30% leucaena. Fertili­ and fruiting branch. 152 FORAGES

at maturity. Seeds 18-22 per pod, 6-10 mm long, thrives under irrigation regimes similar to those brown. used for crops like maize (i.e. > 1200 mm/year). Wood characteristics Leucaena produces a Leucaena favours soils with pH > 5,an d has a low medium hardwood with specific gravity between tolerance of free Al. It performs best on coralline 0.5 and 0.6 and a moisture content which varies and other calcareous soils, but can be found on between 30-50% depending on maturity. Heating saline soils and on alkaline soils up to pH 8. Leu­ values (bone-dry) average 19250 kJ/kg. Bark is caena is not suited to acid soils with pH(H20) < thin. The wood turns well, matures to a golden- 4.5 or to waterlogged conditions. brown colour and is hard enough for flooring. It Propagation and planting Leucaena estab­ accepts preservatives well but does not resist ter­ lishes fairly slowly, particularly in competition mites. Pulp yields are high (50-52 %), lignin levels with weed species and when grown on soils which low, fibres short (1.1-1.3mm) ;pape r quality gener­ are acid or low in nutrient status. For best results, ally is considered excellent. leucaena should be planted on well-drained soils Growth and development Leucaena sets pods with a pH above 5.5 and with a deep soil profile. cyclically every 6-8 months if moisture is suffi­ This is particularly important if productivity dur­ cient, and this is associated with suppression of ing the dry season is required. vegetative growth during fruiting. Arboreal culti- Seeds must be scarified to improve germination, vars have been selected for greatly reduced flower­ usually by placing the seeds for 3minute s in water ing. Fruits ripen in 10-15 weeks. The flowers are at 80° C and then allowing them to cool. Inocula­ self-fertile and most seed results from self-pollina­ tion using peat cultures of improved strains such tion (this is not true for related species with In = as CB81 or CB3060 (TAL 1145) is important for 52 or 2n = 56). Seeds have a hard seed-coat and early nodulation and growth. In the absence of survive for a long time in the soil. Seedlings pro­ peat inoculants, the soil under well-established duce a single strong taproot in the first month, fol­ stands of leucaena could be used to promote early lowed later by feeder roots. Nodulation is often establishment. This may also promote early infec­ slow, but occurs within 2months . Rates of growth tion by mycorrhiza. It is important not to sow the usually increase after 3 months, continuing seed too deeply (below 2 cm) and, where possible, linearly for 3-4 years. Coppiced stems sprout 5-15 competing species should be controlled either by branches, depending on diameter of the cut sur­ slashing or by appropriate chemicals. For forage, face, and 1-4 stems dominate after a year of seeds are usually sown in rows 1-5 m apart with regrowth. Individual leaves persist from 4-6 a seeding rate of 5-7 kg/ha using fertilizer where months and fold at night or under stress. necessary to correct known soil deficiencies. In Other botanical information The common and cut-and-carry systems, closer plant spacing gives giant forms of L. leucocephala are distinguished higher yields of leucaena but in grazed situations taxonomically as L. leucocephala var. leucocephala the wider row spacings of 2-5 m are more appro­ (common form, shrubby, small plant parts, pubes­ priate to enable the grass to grow between the rows cent shoot tips) and L. leucocephala var. glabrata to prevent soil 'plugging' during wet weather. Leu­ Rose (giant form, arboreal, with large plant parts, caena can also be established by raising seedlings glabrous shoots). Intermediate types are referred in the nursery in long narrow containers (3 cm x to as the 'Peru' form. The giant or 'glabrata' form 15cm) ,accommodatin g the strong taproot without gives the highest yields of fodder with infrequent coiling. Transplanting is done when seedlings are cutting. The best known cultivars in South-East 3-5 months old, preferably after a month in the full Asia are 'K8', 'K29', 'K67', 'K156' (a cultivar of L. sun. Bare-rooted seedlings can be transplanted diversifolia (Schlecht.) Benth.) and 'K636' which effectively if shoot and roots are topped. resulted from research work in Hawaii, and the Establishment is often more successful on areas cultivar 'Cunningham' in Australia. Psyllid-resis- which were previously under grassland than on tant cultivars 'KX1', 'KX2' and 'KX3' are interspe­ areas which were previously cropped and there­ cific hybrids and are becoming popular in Asia. fore contain large numbers of weed seeds. Despite weed competition, leucaena isofte n able to survive Ecology Leucaena is found from lowlands up to because of its ability to tolerate shade, thereby 1000 m elevation, but new hybrids such as 'KX3' eventually growing above the weed canopy, pro­ greatly extend this range to cooler climates. Leu­ vided the area is not closely grazed or mown. caena generally requires annual rainfall of 650-1500 mm, but can be found in drier and wetter Husbandry For grazing situations, leucaena sites depending on competitive vegetation. It can be grown with many grasses.I n the subtropics, LEUCAENA 153 rhodes grass (Chloris gayana Kunth) and setaria Harvesting In cut-and-carry systems, the plants (Setaria sphacelata (Schumach.) Stapf & Hubbard are cut back to 0.5-1.0 m height every 6-8 weeks ex M.B. Moss) have been suitable companion during the growing season and fed fresh to rumi­ grasses. Pangola grass (Digitaria eriantha Steu- nants. In many situations, leucaena is fed as a pro­ del), guinea grass (Panicum maximum Jacq.), sig­ tein supplement to grass or cereal straws. Some­ nal grass (Brachiaria decumbens Stapf) and Sabi times the feeding of leucaena improves the intake grass (Urochloa mosambicensis (Hack.) Dandy) are ofth e basal diet, but generally it improves the total suitable in the tropics. intake of DM, protein and digestible nutrients. Leucaena is very palatable and stands can be easi­ The extent of the improvement depends on the ly weakened by heavy continuous grazing. Several amount of leucaena that is fed. rotational grazing strategies have been successful, In some of the Indonesian islands (particularly including a simple 2 paddock system of 4 weeks Timor),fres h leucaena may form alarg e part of the grazing / 4 weeks rest and a 4 paddock system of diet of tethered animals intended for slaughter. 2week s grazing / 6week s rest. The main principle Banana pseudostems are also fed to provide water. is to move the animals before they graze new leu­ Such a diet is grossly deficient in Na, and salt sup­ caena regrowth. Specific paddocks ofleucaen a can plementation is required for good production. also be used in conjunction with areas of native Wood harvest periods range very widely, from 1-8 pasture, using the rotational principle to prevent years, depending on size of desired product and overgrazing of the palatable leucaena. When ade­ harvesting equipment. Machetes are commonly quate leucaena is available, cattle should be cap­ used in Asia, but handsaws and chainsaws can also able ofweigh t gains ofabou t 1 kg/head per day pro­ be used. vided the Na level in the diet is adequate. Yield Yields of forage vary with soil fertility, Diseases and pests Diseases of leucaena are rainfall, altitude and cutting management from few and include seedling rots such as Phytoph- 1-15 t/ha of DM per year. Total yields are reduced thora drechsleri and Fusarium semitectum that by frequent cutting, though leaf yield per day may attack primarily under waterlogged conditions. vary little between cutting at 6, 8 or 12 weeks. Until the mid-1980s leucaena was relatively free Highest yields are obtained under wet tropical of serious diseases and pests. However, devastat­ lowland conditions on deep well-drained, neutral ing effects of the leucaena psyllid {Heteropsylla to alkaline soils. Although leucaena is drought- cubana), a small aphid-like sucking insect, have tolerant, yields in the dry season are low unless been experienced in many areas where leucaena the plants have access to sub-surface water or are is grown for fodder. Psyllid damage is rarely seen irrigated. in leucaena's centre of origin in South and Central Wood yields compare favourably with the best America, and damage caused by the psyllid has tropical trees, with height increments of 3-5 decreased with time in other areas. Populations of m/year and wood increments of 20-60 m3/year for these insects fluctuate through the season and can arboreal varieties. cause losses of over 50% in yield. Attempts to use Handling after harvest Fodder is commonly predatory and parasitic insects for control have fed fresh or provided as a browse. Sun-drying is been variably successful. This pest is now limiting practised for leaf pelleting and marketing, often the further development of leucaena for forage in by placing branches over treliiswork or on asphalt some areas. Other leucaena species show resis­ to allow the leaflets to drop.Woo d handling issimi ­ tance to psyllids, and have been used in the breed­ lar to that of other fuelwood or pulpwood species. ing ofresistan t hybrids 'KX1', 'KX2' and 'KX3'. Genetic resources Two major collections are On plants allowed to grow tall, attacks ofsof t scale held at NFTA (Hawaii) and ATFGRC (CSIRO, (Cocus longulus) and an associated sooty mould Australia). They comprise all 14Leucaena species can be serious. Seed crops can also suffer yield re­ and total 1800 accessions derived largely from 7 duction through attacks on the inflorescences and expeditions to Mexico and Central or South Amer­ young pods from the larvae of the moth Ithome las- ica. They are identified by K numbers (Hawaii) or sula. In some areas, notably Central America, bru- CPI numbers (Australia). Naturalized populations chid beetles can seriously reduce or destroy seed of leucaena in Asia show limited genetic variation crops. and are not recommended for production as they Seedlings can suffer attack from cutworms and ter­ are out-yielded by improved cultivars. mites, but, provided there is an adequate stand den­ Breeding Breeding of leucaena is in progress at sity, subsequent production is usually not reduced. NFTA (Hawaii) and CSIRO (Australia). The key 154 FORAGES objective is to incorporate psyllid resistance from 1-114. |9|Pound , B. &Martine z Cairo, L., 1983.Leu ­ other Leucaena species (primarily L. diversifolia caena, its cultivation and use. Overseas Develop­ and L. pallida Britton &Eose ) into agronomically ment Administration, London. 287 pp. |10| Taka- desirable forms of L. leucocephala. hashi, M. & Ripperton, J.C., 1949. Koa Prospects Over the last 2decades , leucaena has (Leucaena glauca). Its establishment, culture and been one of the most promising forage legumes in utilisation asa forag e crop.Universit y ofHawai iAg ­ South-East Asia. The arrival of the psyllids has ricultural Experiment Station Bulletin 100.pp . 1-56. curbed the previous enthusiasm. However, partial R.J. Jones, J.L. Brewbaker &C.T . Sorensson control of the problem with natural or introduced predators, and the prospect of new lines or hybrids more tolerant or resistant to the psyllid, have Lotononis bainesii Baker renewed interest in this and related species. Newly-bred cultivars widen the climatic range of FI. Trop. Afr. 2:6 (1871) . leucaena to the highlands and subtropical regions, LEGUMINOSAE with great cold tolerance characterizing some new 2n = 36 hybrids ('KX3').Improve d bole shape ('K636'), psyl­ Vernacular names Lotononis (En). Thailand: lidresistanc e ('KX1', 'KX2'),lo w mimosine content thua-lotononit. ('KX3') and increased vegetative vigour are among Origin and geographic distribution The natu­ other advances in breeding. Improved alley crop­ ral distribution of lotononis is confined to south­ ping methods of managing leucaena have been de­ ern Africa, mainly between latitudes 20° S and veloped in Africa and Indonesia. These are 30° S in , South Africa and . expected to improve crop yields in association There are now isolated occurrences in many coun­ with leucaena and aid in the stabilization of shift­ tries as a result of experimentation. It is natura­ ing systems of agriculture and of fragile tropical lized in areas of subtropical Australia, particular­ soils. ly along roadsides. Literature |1|Brewbaker , J.L., 1987. Species in Uses Lotononis is used as a pasture legume, the genus Leucaena. Leucaena Research Reports mainly in the subtropics. 7: 6-20. |2j Brewbaker, J.L., 1987. Leucaena: A Properties Lotononis forage is of better quality genus of multipurpose trees for tropical agrofor- than most tropical legumes, particularly in terms estry. In: Steppler, H.A., Nair, P.K.R. (Editors): of N concentration which ranges from 1.5-4.0%. Agroforestry; a decade of development. ICRAF, Digestibility is good for a tropical legume, and Na Nairobi, Kenya, pp. 289-323. |3| Hill, G.D., 1971. concentrations also tend to be higher. Seeds are Leucaena leucocephala for pastures in the tropics. very small with 3000-4000 seeds/g. Herbage Abstracts 41: 111-119. |4| International Botany A stoloniferous, prostrate, creeping, Development and Research Centre, 1983. Leu­ short-lived perennial, with a primary taproot and caena research in the Asia-Pacific region. Pro­ smaller adventitious roots arising from stolons. ceedings of Singapore Workshop, Nitrogen Fixing The plant canopy is seldom more than 30 cm high, Tree Association (NFTA) & IDRC, Ottawa, but stems may reach 1.8 m length. Leaves usually Canada. 192 pp. |5|Jones , R.J., 1979. The value of trifoliolate; petiole up to 7.5 cm long; stipules Leucaena leucocephala as a feed for ruminants in leafy, up to 1 cm long; leaflets polymorphic, oblong the tropics. World Animal Review 31: 13-22. |6| to lanceolate, up to 6.5 cm x 1.5 cm, glabrous or Jones, R.J., 1985.Leucaen a toxicity and the rumi­ slightly pubescent, margins entire. Flowers about nai degradation of mimosine. In: Seawright, A.A., 1 cm long in head-like inflorescences 1-3 cm long Hegarty, M.P., James, L.F. & Keeler, R.F. (Edi­ with few to more than 20 flowers; peduncle up to tors): Plant toxicology. Proceedings of the Austra- 27 cm long; calyx hairy; corolla yellow to reddish. lia-U.S.A. Poisonous Plants Symposium, Bris­ Pod linear-oblong, 8-12mm x 2mm ,densel y white bane, Australia. May 14-18, 1984. pp. 111-119. |7| hairy, with long persistent style. Jones, R.J. & Megarrity, R.G., 1986. Successful Lotononis flowers early in the wet season. transfer of DHP-degrading bacteria from Haw­ Cultivar 'Miles' has been released in Australia. aiian goats to Australian ruminants to overcome Ecology As a sown pasture species, lotononis is the toxicity of Leucaena. Australian Veterinary best suited to areas receiving 700-1200 mm annual Journal 63:69-78 . |8| Oakes, A.J., 1968. Leucaena rainfall in the subtropics or tropical highlands, al­ leucocephala; description, culture, utilization. Ad­ though it occurs naturally in areas with rainfall vancing Frontiers of Plant Science (India) 10: as low as 250mm/year . It ison e ofth e best subtropi- MACROPTILIUM 155

rainfall late in the wet season. Lotononis responds to applied P, but will persist on soils of low P sta­ tus. Lotononis is most susceptible to diseases during very wet periods if growth is lush. It is susceptible to Cercospora leaf-spot, Botrytis flower blight, Sclerotium rolfsii, Fusarium and Pythium root and stolon rots and legume little leaf. Genetic resources and breeding Cultivar 'Miles' is available commercially. A germplasm collection is held by ATFGRC (CSIRO, Australia), but preliminary testing suggests that there is no substantial variation within this collection. Prospects Lotononis will not be suited to most of South-East Asia, but could be a useful pasture component in the humid subtropics and tropical highlands. Literature |1| Bryan, W.W., 1961.Lotononi s bai­ nesii Baker. A legume for the subtropics. Austra­ lian Journal of Experimental Agriculture and Ani­ mal Husbandry 1: 4-10. :2| Bryan, W.W., Sharpe, J.P. &Haydock , K.P., 1971.Som e factors affecting the growth of lotononis (Lotononis bainesii). Aus­ tralian Journal of Experimental Agriculture and Animal Husbandry 11: 29-34. |3| Jones, R.M. & Evans, T.R., 1976.Soi l seed levels ofLotononi s bai­ nesii, Desmodium intortum and Trifolium repens Lotononis bainesii Baker -1, f'lowering and fruiting in subtropical pastures. Journal of the Australian branchlet; 2,flower; 3, fruit. Institute of Agricultural Science 43: 164-166. j4| Pott, A. &Humphreys , L.R., 1983.Persistenc e and cal legumes in terms of winter growth and is resis­ growth on Lotononis bainesii-Digitaria decum- tant to light frosts. It grows best on well-drained bens pasture. 1. Sheep stocking rate. Journal of sandy soils under moderate to heavy grazing, par­ Agricultural Science, Cambridge 101:1-7. |5| Sker- ticularly with more open grass swards. It has good man, P.J., Cameron, D.F. &Riveros , F., 1988.Trop ­ tolerance of low levels of available soil P and Mo. ical forage legumes. FAO, Rome. pp. 319-327. Agronomy Seed is often hard-seeded and scari­ R.M. Jones fication may be required before sowing. Lotononis can be established by sowing into a seed-bed or broadcasting into undisturbed swards using a Macroptilium atropurpureum (DC.) seeding rate of 0.5-1.0 kg/ha. As seed is very small Urban it is best sown on top of cultivated soil and then rolled in so that it is not buried too deeply. Sowing Symb. Antill. 9:45 7(1928) . in the hottest months should be avoided. Seedling LEGUMINOSAE emergence is often slow and erratic. 2re = 22 Inoculation with a specific Bradyrhizobium strain Synonyms Phaseolus atropurpureus DC. (1825). such as CB 376 is essential. Lotononis is usually Vernacular names Siratro, atro, purple bean sown as a component of a mixture of legumes. It (En). Thailand: thua-sirato. persists poorly with lightly grazed, vigorous grass. Origin and geographic distribution This spe­ The productivity of lotononis tends to be erratic, cies occurs naturally from to Col­ but once established in a suitable site, it is rarely ombia and northern Brazil. The greatest diversity lost completely. Recovery is aided by a large seed is in Mexico. Being one of the first commercially 2 bank in the soil of up to 100000 seeds per m . The available tropical legumes, siratro cultivar 'Sira­ best lotononis years are those with good rainfall tro' is now widespread throughout the tropics and early in the wet season; there is less response to subtropics. 156 FORAGES

Uses Siratro is primarily used as a pasture surface remains continuously moist for some legume with introduced or natural grasses but weeks; trailing stems rarely greater than 5m m di­ could be useful as a cover crop or as a fallow crop ameter but may exceed 5m in length. Leaf trifolio- in shifting cultivation. Siratro is used for revege- late, often 2-3-lobed, dark green and slightly hairy tation and stabilization of earthworks, and in road on the upper surface, silvery and very hairy on the and railway cuttings and embankments. It has lower surface; petiole up to 5 cm long; leaflets shown promise as a leguminous forage crop sown ovate to rhomboid, 2.5-8 cm x 2.5-5 cm, often in conjunction with upland rice or after harvest­ asymmetrically lobed. Inflorescence a raceme; ing of lowland rice. peduncle 10-30 cm long with 6-10 deep purple Properties Nitrogen concentrations range from flowers, about 2 cm long, crowded at the apex; 2-4%, higher than some other tropical legumes calyx campanulate, 5-lobed; standard orbicular such as Stylosanthes spp. Digestibility of young with 2 small basal auricles; wings long stipitate, material ranges from 60-70% , but in older materi­ longer than standard and keel; keel petals apically al with a lower leaf percentage this may decrease spiralled, basally adnate to the staminal tube. Pod to 45%. It tends to have higher P concentrations linear, straight, 4-8 x 4-6 mm, adpressed pubes­ than Stylosanthes spp. with levels usually above cent, usually with 10 seeds, dehiscing violently 0.2%, reflecting its higher P requirement. Sodium when ripe, throwing seed for up to 5 m. Seed concentrations are usually less than 0.02%. It is oblong-ovoid, 3-4 mm long, light brown, black or a palatable legume though cattle prefer to eat mottled, the latter usually with greater hard-see- grass early in the growing season and siratro later dedness. in the growing season. There are 70-90 seeds/g. Growth and development Siratro sown into a Description A perennial herb with twining seed-bed grows vigorously and will flower and pro­ stems arising from a taproot ofu p to 2c m diameter. duce seed in the first year. When broadcast into Stems close to the ground may develop secondary undisturbed pasture or emerging from seed in roots, usually on light textured soils when the soil undisturbed grassland, growth is slower and seed production is unusual in the first year. The half- life of established plants is 1-4 years. Siratro is a short-day plant. The main flowering period is in the late wet season, but flowering also occurs in the early and mid wet season. The onset of flower­ ing during the wet season partially relates to onset of moisture stress. Under light grazing pressure in higher rainfall areas and on sandy soils,siratr o de­ velops stolons.Thes e can live for several years and attain 5 mm diameter. Siratro plants survive dry periods by shedding leaves and then through death of stems to reduce the area oftranspirin g leaves. Other botanical information Only one culti- var of siratro, 'Siratro', is presently available. It was developed in Queensland about 1960 from a cross between two Mexican introductions. Ecology Siratro grows best under rainfall regimes ranging from 700-1500 mm, but in Mexico has been collected from sites receiving annual rainfall as low as 250 mm. It is not suited to the wet tropics or to the very hot, dry tropics, nor to elevations above 1600 m in the tropics or to lati­ tudes higher than 30°. Top growth is killed by frost, but the taproots survive frosts in subtropical regions. Growth is optimal at day/night tempera­ tures of 30/25°C . It grows best on light to medium textured soils and does not persist on poorly Macroptilium atropurpureum (DC.) Urban drained sites. It requires a soil available P level flowering and fruiting vine; 2,flower; 3, seed. of at least 10 mg/kg (bicarbonate extraction) or MACROPTILIUM 157 preferably 15 mg/kg. It tolerates a pH(H20) range than 30% of dry matter in grazed legume/grass of5-8 . There is considerable variation within sira- pastures. Siratro can yield up to 1t/h a of seed, al­ tro and some characters, such as an erect growth though seed yields are usually 100-300 kg/ha. habit and fewer days to flowering, are related to High seed yields can be obtained from hand har­ drier areas of origin. vesting seed from plants grown on wire trellises. Propagation and planting Siratro is estab­ Good siratro pastures can give the same animal lished by seed. It is not as hard-seeded as many production as grass-only pastures fertilized with tropical legumes, but scarification may still be 100kg/h a of N in 700-1000 mm rainfall areas. required. Establishment isfaste r and more reliable Genetic resources The main germplasm collec­ when sown into a seed-bed, but it can be established tions are held at ATFGRC (CSIRO, Australia) and into undisturbed pasture or with minimal cultiva­ CIAT (Colombia). tion if conditions are favourable. Seeding rates of Breeding A plant breeding programme to incor­ 2-6 kg/ha are recommended when sown alone. Sira­ porate rust resistance into siratro, based at CSIRO, tro nodulates freely with native cowpea rhizobia Brisbane, is nearing completion. There is scope for and there are no records ofnodulatio n failure. further development to select and breed cultivars Husbandry In permanent grasslands siratro ofsiratr o for specific regional adaptation. must be allowed to seed since recruitment of new Prospects Although siratro is not as persistent plants to replace older plants is essential for long- as initially thought, it is still a very useful legume. term persistence. Soil seed banks of over 200seeds / Its ability to provide high yields of quality materi­ m2 are desirable for persistence; levels below 50 al,fi x N and improve grass growth and quality, and seeds/m2 may lead to extinction. Siratro responds its fairly wide area of adaptation, mean that fur­ well to P fertilizer, if required. Liming may be ther work on the development of new cultivars is required if soil pH(H20) falls below 5.0. justified. Diseases and pests The main diseases of siratro Literature 1! Carangal, V.R., Tengco, P.L., are leaf blight (Rhizoctonia solani), which is more Miah, N.I.& Topar k Ngarm, A., 1988.Roo d and for­ serious in the wetter end of its zone of adaptation age crop intercropping under rainfed lowland con­ and limits the use of siratro in the wet tropics, and ditions. Proceedings of the Crop-Animal Systems rust (Uromyces appendiculatus). Rust is not a Research Workshop, Serdang, Malaysia. MARDI/ lethal disease but has been shown to reduce the IRDC/ARFSN, Serdang, Malaysia, pp. 453-471. 12. yield and seed set of 'Siratro' by 30%. There are Gutteridge, R.C., 1985. Survival and regeneration different races of the rust pathogen but some lines of four legumes oversown into native grasslands of siratro have resisted a range of races under ex­ in northeast Thailand. Journal ofApplie d Ecology perimental conditions. Violet root rot (Rhizocto­ 22:885-894 . |3| Hutton, E.M., 1962.Siratr o - a trop­ nia crocorum) has led to death oftaproots . It is sus­ ical legume bred from Phaseolus atropurpureus. ceptible to halo blight (Pseudomonasphaseolicola) Australian Journal of Experimental Agriculture which, although of minimal consequence to sira­ and Animal Husbandry 2: 117-125. |4| Jones, R.J. tro, can act as a source ofinfectio n to nearby crops & Jones, R.M., 1976. The ecology of siratro based of Phaseolus spp. Seedlings of siratro can be killed pastures. In: Wilson, J.R. (Editor): Plant relations by larvae of bean fly (Melanagromyza phaseoli). in pastures. CSIRO, Melbourne, Australia, pp. Siratro is resistant to root-knot nematodes (Meloi- 353-367. !5: Jones, R.M. & Bunch, CA., 1988. The dogyne javanica). effect of stocking rate on the population dynamics Harvesting Siratro isusuall y harvested by graz­ of siratro in siratro (Macroptilium atropurpu- ing animals, but can be cut for stall feeding, or for reum)-setaria (Setaria sphacelata) in south-east hay or silage.Siratr o pastures can be continuously Queensland. Parts I and II. Australian Journal of or rotationally grazed but cannot withstand sus­ Agricultural Research 39: 209-219, 221-234. |6< tained heavy grazing. Ifcut , ares t period of at least Oram, R.N., 1990. Register of Australian herbage 6 weeks and a stubble height of at least 10 cm is plant cultivars. CSIRO, Australia, p. 239. |7I Sker- suggested. man, P.J. , Cameron, D.G. &Riveros , F., 1988.Trop ­ Yield Under cutting, legume DM yields of 7 t/ha ical forage legumes. FAO, Rome. pp. 328-337. |8i have been measured in pure swards and 5 t/ha in 't Mannetje, L. & Jones, R.M., 1990. Pasture and siratro/grass swards, although siratro yield de­ animal productivity of buffel grass pastures with clines markedly with more frequent cutting. DM siratro, lucerne or nitrogen fertilizer. Tropical yields of over 5t/h a have been reported when used Grasslands 24: 269-281. as an intercrop legume.I t seldom contributes more R.M. Jones &L . 't Mannetje 158 FORAGES

Macroptilium lathyroides (L.) Urban

Symb. Antill. 9:45 7(1928) . LEGUMINOSAE 2re = 22 Synonyms Phaseolus lathyroides L. (1763). Vernacular names Phasey bean (En). Wild pea bean (Am). Indonesia: kacang batang. Thailand: thua-phi. Origin and geographic distribution Phasey bean originated in tropical America and is now widely distributed, although usually of only minor importance, throughout Central and South Amer­ ica, the Caribbean, Australia, South-East Asia and parts of Africa. Uses Phasey bean is primarily used as a forage plant, but can be used as a cover crop in rotations and as green manure. Properties Nitrogen concentrations range from 4% in young vegetative growth to 1 % when most leaves have fallen off and the sample is primarily old green stem. Similarly, digestibility can range from 70-40%. There are 110-120 seeds/g. Botany Herbaceous annual or less commonly short-lived perennial, erect with variable branch­ ing, 0.5-1.5 m tall, with base of stem becoming somewhat woody, sometimes becoming trailing or twining, especially under shade. Stems terete, Macroptilium lathyroides (L.) Urban -1, flowering with long deciduous reflexed hairs.Leave s trifolio- and fruiting branch; 2, flower. late; leaflets ovate-lanceolate to elliptical, 3-8 cm x 1-3.5 cm, not lobed. Inflorescence a semi-erect field. It is strictly self pollinated. The only cultivar or erect raceme up to 15 cm long, borne on pedun­ is 'Murray', developed in Australia, which is taller cle up to 40 cm long; pedicels very short; flowers and more robust than most ofth e lines evaluated. red-purple, about 1.5 cm long. Pod subcylindrical, Ecology Phasey bean is adapted to a wide rain­ 5-10 cm x 3mm , containing 18-30 seeds, abruptly fall range of500-300 0 mm. The optimum day/night dehiscent. Seeds oblongoid to rhomboidal, 3 mm temperature for growth is 35/20°C. Plants are long, mottled light and dark grey-brown. killed by frost but usually seed before frosting. It Sometimes a subdivision of the species into two is adapted to acid and alkaline soils, and a wide varieties is made: range of soil textures from sand (given reliable - var. lathyroides: leaflets narrowly lanceolate, rainfall) to heavy clay. It is tolerant of waterlog­ often more or less lobed; only in tropical Amer­ ging and poor drainage and frequently grows in ica; drains along the edges of roads. Drought survival - var. semierectum (L.)Urba n (synonym Phaseolus is achieved by soil seed reserves. semierectus L.): leaflets ovate to elliptical, not Agronomy Phasey bean is easy to establish lobed; this variety is the widely distributed for­ from seed and seeding rates of 3-10 kg/ha are rec­ age plant. ommended. Establishment in undisturbed soil, Phasey bean is day-neutral for flowering and can particularly in existing pasture, is rarely success­ flower throughout the wet season but under ful, except under favourable moisture conditions shaded and humid conditions flowering and seed with scarified seed. In subtropical regions, it can production are depressed and the plant assumes a be sown during spring and summer when moisture twining growth habit. There is evidence that twin­ conditions are favourable. Seedlings nodulate ing is initiated by shading whereas flowering is freely with native cowpea rhizobia, and seedling depressed by high humidity, even though these en­ growth is vigorous when sown into a prepared vironmental factors often occur together in the seed-bed. MACROPTILIUM 159

Seedlings are susceptible to attack from bean fly Macroptilium longepedunculatum (Melanagromyza phaseoli) and plants are suscepti­ (Benth.) Urban ble to Phaseolus virus 2, root-knot nematode (Meloidogyne javanica), especially on sandy soils, Symb. Antill. 9:45 7(1928) . and to mildew. LEGUMINOSAE It can be cut or grazed, but does not tolerate sus­ 2n = 22 tained heavy cutting or grazing which can greatly Synonyms Phaseolus longepedunculatus Ben­ reduce or even eliminate seeding. If grazing ani­ tham (1837),P. campestris Bentham (1837,no n Ben­ mals have scope for selection, they may avoid graz­ tham 1859). ing phasey bean till after seeding. Vernacular names Llanos macro (En). DM yields of 13 t/ha at 90 days after sowing have Origin and geographic distribution The natu­ been reported. Although phasey bean seeds readi­ ral distribution of llanos macro is from southern ly,seedlin g recruitment is usually poor and so per­ Brazil to northern Mexico. It was taken into culti­ sistence is poor in permanent pastures after the vation outside tropical America only recently, e.g. second year. Recruitment can be aided by rough in Australia. cultivation. Hence its main use is as a component Uses M. longepedunculatum is a pasture legume of a mixture to give grazing in the first year but for monsoonal lowland regions. It can be grazed it also shows some promise as a one year legume during the growing season, or used as afodde r crop leyo r as aforag e intercrop. Plants arekille d by fire, to make hay, as saved standover fodder or as green but stands can recover through seedling germina­ manure. tion after fire. Good hay can bemad e iflos s of leaves Properties Nitrogen concentrations range from is avoided. Harvested seed yields are typically 1.4-3.0% during the growing season but decline 200-250 kg/ha but much seed is lost through dehis­ to 1-1.4% during the dry season. DM digestibility cence and the true seed yields would be higher. of plant material during the growing season Genetic resources and breeding There are no ranges from 57-67 %,fallin g to 47-53 % during the breeding programmes on phasey bean. Germplasm dry season. Phosphorus concentrations range up collections are held byATFGR C (CSIRO, Australia) to 0.23% during the growing season and from and CIAT (Colombia). There is known variation for 0.06-0.13% during the dry season. Sulphur con­ mildew resistance and in the intensity of branching. centrations range up to 0.21% during the growing Prospects It is likely that the potential for pha­ season and from 0.06-0.10% during the dry sea­ sey bean lies in its use in short leys and as an inter­ son. There are approximately 300 seeds/g. crop, or to provide first-year feed when sown as Botany Short-lived perennial twining herb to 60 part of a mixture for permanent pastures. Further cm high with trailing stems; vegetative parts exploitation of the variability within this species shortly pilose, hairs of the stem retrorse to spread­ may make it more suited to these roles. ing. Leaf pinnately 3-foliolate; petiole, petiolule Literature |1| Cameron, D.G., 1985.Tropica l and and rachis grooved above and closely ribbed; subtropical pasture legumes 8: Phasey bean (Ma­ petiole 24-45 mm long; lateral leaflets ovate with croptilium lathyroides). The predecessor of Sira- asymmetrical bases, 2-4 cm x 1-3 cm; terminal tro. Queensland Agricultural Journal 111: leaflet broadly lanceolate to deltoid, 2.5-5 cm x 211-214. |2j Elich, T.W., 1988. Studies on seed pro­ 1-3 cm. Inflorescence an axillary raceme, 20-40 duction in phasey bean (Macroptilium lathyr­ cm long; flowers shortly pedicelled, solitary or in oides). CSIRO, Australia, Division of Tropical pairs and threes;standar d greyed orange, concave, Crops and Pastures, Technical Memorandum No obovate, emarginate, ca. 14m m long;wing s purple, 58.9 pp . |3| Oram, R.N., 1990.Registe r of Australian partly adhering to the keel petals, limb orbicular, herbage plant cultivars. CSIRO, Australia, p. 240. ca. 15 mm long, apex curved and slightly twisted. j4| Paltridge, T.B., 1955. Sown pastures for south­ Pod linear, 3-7 cm x 2-3 mm, containing 10-18 east Queensland. CSIRO, Australia, Bulletin No seeds; valves twisted on drying. Seed small, flat­ 274. pp. 106-107. |5| Skerman, P.J., Cameron, D.G. tened ovoid, mottled light and dark grey. &Riveros , F., 1988.Tropica l forage legumes. FAO, Llanos macro has a juvenile phase during which Rome. pp. 337-345. |6[ Thro, A.M. & Shock, C.C., floral initiation will not take place. It flowers and 1987. Performance of subtropical forage legumes seeds prolifically, producing up to 240 kg/ha of in Louisiana, south-central United States. Trop­ seed. Freshly harvested seed has a very variable ical Agriculture 64: 297-304. (20-80) percentage of hard seed. R.M. Jones &L . 't Mannetje M. longepedunculatum is a new species to cultiva- 160 FORAGES

using seeding rates of 2-4 kg/ha. A good seed-bed preparation is desirable for establishment. It can be sown as amixe d pasture or a pure stand. It nodu- lates freely and effectively with natural cowpea type rhizobia and does not need inoculation. Llanos macro grows best with good soil fertility, but will persist at low soil fertility levels after being established with minimal non-N fertilizer as required. After periods of wet weather small patches of dead leaves (leaf blight) are caused by Rhizoctonia spp. The areas involved are only small and no control measures are required. DM yields of llanos macro in well-fertilized pure swards reach 5-7 t/ha. In mixed swards, they can be 3-4 t/ha. It is usually consumed fresh or as a saved standover fodder. Seed formation is essen­ tial for long-term persistence, particularly in the year of establishment. It should not be severely defoliated early in the wet season. Genetic resources andbreedin g Llanos macro 'Maldonado' is new to cultivation, and seed pro­ duction is increasing. Small quantities of seed are available for sale. A germplasm collection is held by ATFGRC (CSIRO, Australia). There isscop e for further development of cultivars as some lines have different flowering times than 'Maldonado' and some lines are amphicarpic. Macroptilium longepedunculatum (Benth.) Urban Prospects 'Maldonado' is recommended for - 1, flowering branch; 2, two types of leaves; 3, tropical monsoonal areas with annual rainfall be­ flower; 4, infructescence; 5, seed. tween 1100 and 1600 mm. It may grow in wetter monsoonal environments. Other cultivars may be tion and has not been closely studied. It is closely developed for cooler, subtropical regions. related to M. atropurpureum (DC.) Urban, M. gra­ Literature |lj Cameron, A.G., 1990. Maldonado cile (Poepp.) Urban and M. domingense Urban, but - a new legume for the Top End. Department of further biotaxonomic studies are needed. There is Primary Industry and Fisheries, Darwin, Austra­ only one cultivar, 'Maldonado', released in the lia. Agnote No 429.2 pp . j2|Lackey , J.A. & D'Arcy, Northern Territory of Australia. 'Maldonado' is a W.G., 1980. Macroptilium. In: Dwyer, J.D. et al. late flowering cultivar, flowering from mid-April (Editors): Flora of Panama 5(5). Leguminosae. in northern Australia. It appears to be a short-day Annals of the Missouri Botanical Garden 67: plant, but the critical photoperiod has not been 724-728. |3| Maréchal, R., Mascherpa, J.M., Stai- determined. nier, F., 1978.Etud e taxonomique d'un group com­ There are a number of lines of M. longepeduncula­ plex d'espèces des genres Phaseolus et Vigna tum, some of which flower earlier than 'Maldo­ (Papilionaceae) sur la base de données morpholo­ nado', and some of which are amphicarpic. giques et polliniques, traitées par l'analyse infor­ Ecology In its region of origin llanos macro matique [Taxonomie study of a group-complex of occurs generally at low altitudes (up to 250 m) in species of the genera Phaseolus and Vigna (Papi­ areas with rainfall between 950-1300 mm per year. lionaceae) based on morphological data and on It is adapted to tropical monsoonal regions with pollen morphology, through computer analysis]. strongly seasonal rainfall between 1100-1600 mm. Boissiera 28: 151-157. ,4| Ross, B.J. & Cameron, It is not adapted to the humid tropics. It is tolerant A.G., 1991. Pasture legume evaluation on seaso­ ofdrought , burning, waterlogging and flooding for nally flooded soils in the Northern Territory. Trop­ up to three months. ical Grasslands 25: 32-36. Agronomy Llanos macro is propagated by seed A.G. Cameron r

MACROTYLOMA 161

Macrotyloma axillare (E. Meyer) Verde.

Kew Bulletin 24:40 2(1970) . LBGUMINOSAE 2n = 20 Synonyms Dolichos axillaris E. Meyer (1836). Vernacular names Axillaris, perennial horse gram (En). Origin and geographic distribution Axillaris originates in sub-Saharan Africa, from Ethiopia to Senegal and south to the Transvaal and Natal in South Africa (16° N to 31° S latitude). Var. axillare extends to Madagascar and Arabia, and var. gla- brum (E.Meyer ) Verde, to Madagascar, Mauritius and Sri Lanka, but var. macranthum (Brenan) Verde, is more restricted and is only found from Tanzania to Zimbabwe. Cultivar 'Archer' has become naturalized in small areas of eastern Aus­ tralia and Papua New Guinea. Uses Axillaris is used as a perennial sown pas­ ture legume in Australia where it is grazed during the growing season or used asstand-ove r feed early in the dry season. Properties In Australia N concentrations of whole tops of 'Archer' fell from 2% in the early wet season to 1 % in the dry season. Phosphorus concentrations, on a high-P soil, fell from 0.25-0.30% (early wet season) down to 0.12-0.17% (dry season). Seed contains about 4% N and 0.45% P. Macrotyloma axillare (E. Meyer) Verde. -1, flower­ The plant is not toxic, but has a bitter taste which ing and fruiting branch; 2,flower; 3, fruits. may cause stock to reject it until they get used to it. It sometimes taints the meat of lambs when fed of short dense hairs. Pod shortly stipitate, linear- as a large part of their diet for a long time, and oblong, 3-8 cm x (5-)6-8 mm, glabrous or pubes­ so could taint milk in similar circumstances. cent, containing (5-)7-8(-9) seeds. Seed rounded Across a range of accessions, there were 50-200 or ellipsoid, compressed, 3-4.2 mm x 2.5-3 mm x seeds/g (80-90 in 'Archer'). 0.6-1.5 mm, smooth and hard, buff to dark red with Description Perennial climbing or trailing sparse to very dense black mottling. twining herb with a strong woody taproot and Growth and development Flowering is rootstock. Stems do not root at the nodes; stems delayed in the year of sowing, but not subse­ cylindrical, glabrescent or hairy, > 5 m long if quently, which is indicative of a juvenile phase climbing, otherwise usually < 3.5m ;th e main axis during which flowering is inhibited. This also is strongly ascending on young plants. Leaves tri- seems to apply to the regrowth ofindividua l shoots foliolate; leaflets entire, elliptical, ovate-lanceo­ on mature plants. Flowers are cleistogamous but late, ovate or subrhombic, 1.1-7.5 cm x 0.7-4.2 honey bees can transfer pollen between them once cm, rounded to acuminate and mucronulate at the they are open. Pods usually mature 4-8 weeks apex, rounded to subacute at the base, glabrous to after flowering and shatter readily once dry. pubescent, somewhat glossy above and paler be­ Other botanical information Three varieties neath; stipules ovate-lanceolate, 2-5 mm long, arerecognized ; var. axillare and var.glabrum have veined. Flowers axillary in 2-4(-10)-flowered flowers 1.2-1.5(-2) cm long; those of var. macran­ clusters; peduncles very short or absent; standard thum are larger, (1.5-)2-2.4 cm long. The first two oblong-elliptical, 1-2.4 cm x 0.6-1.5 cm, whitish varieties may be distinguished by stem indûment: or greenish-yellow with a crimson or purplish spot var. axillare has dense, spreading hairs and var. inside near the centre; stigma surrounded by a ring glabrum has sparse, adpressed hairs although in- 162 FORAGES termediate types are fairly common. 'Archer', ing. 'Archer' will not grow well on soils poor in P which isth eonl y cultivar, released in Queensland, or Swithou t fertilizer. Compared with centro(Cen- Australia, isclos e tovar . glabrum. trosema pubescens Benth.), it has a higher require­ Ecology Axillaris is essentially a tropical or ment for K,bu tsimila r requirements forCu . subtropical short-day plant from grassland, wood­ Diseases andpest s There aren oseriou s disease land, forest margins and coastal dunes. It is com­ or pest problems with axillaris. Individual plants mon in disturbed places such as fallow land and may be stunted and killed by the mycoplasma roadsides in itsnatura l range. It prefers relatively 'legume little leaf', but the productivity of the mild conditions (optimum day/night temperature sward is scarcely affected. It issusceptibl e to root- for growth is about 26/21°C ) so is better adapted knot nematodes (Meloidogyne spp.) but control to low altitudes in the subtropics and higher alti­ measures are unpractical. tudes, up to 2250 m, in the tropics. Although top Harvesting Axillaris isharveste d bygrazin gan ­ growth is killed by frost, it regrows from the root- imals rather than bycutting , although hay making stock in spring in the subtropics. It is drought- is possible provided care istake n to conserve leaf. hardy but not very productive if rainfall is less Its major strength lies in providing better stand- than 800 mm/year. It grows best on well-drained over feed in the early part of the dry season and soils such asalluvia l sandy loams andbasalti c clay faster growth in the early part of the wet season loams, andi ti smoderatel y shade-tolerant. Iti sno t than most other tropical legumes which shed their well adapted to heavy clays, or infertile, acid leaves more readily and only grow slowly in the (pH(H20) < 5.5)solodi c soils, andi t will not toler­ early wet season. Thus it is often sown on ridges ate waterlogging or saline soils. aboveth efros t line.Rotationa l grazing isbest , and Natural ecotypes have been collected under a swards should not be grazed lower than 15 cm range of conditions: rainfall 700-1400 mm/year; because axillaris will notwithstan d frequent close growing season 140-365 days; mean maximum grazing. temperature of the hottest month of the growing Yield In Australia, axillaris has yielded 16t/h a season 26-31°C ; mean minimum of the coldest of DM under irrigation and 3-4 t/ha without irri­ month of the growing season 11-24°C; and mean gation. In India, green yield of an axillaris/green minimum of the coldest month of the dry season panic (Panicum maximum Jacq. var. trichoglume 3-18°C. Robijnsmixtur e (39t/ha ) exceeded that ofth e grass Propagation and planting Hand harvested and alone (24 t/ha). DM yields of fertilized swards threshed seedma y bever y hard andrequir e scarifi­ under regular cutting at three sites in the high­ cation. Mechanically harvested or threshed seed lands (1200-1500 m) of northern Thailand ranged is usually satisfactory for sowing without further between 1.5-7.6t/h ai nth esecon d year. treatment to break hard-seededness. Seed yields of 100-150 kg/ha are common. Yields In prepared seed-beds, seed may be drilled 1-2.5 of more than 550 kg/ha are possible from small cm deep andlightl y covered, or else broadcast and areas, especially if the sward is harvested just harrowed. Axillaris canals o beestablishe d in nat­ before thepod s start to shatter andi sthe n allowed woodland by broadcasting seed mixed with to dry.Large-scal e seed crops are difficult to han­ fertilizer into the ashes after burning atth e begin­ dle with normal heading equipment because the ning ofth e wetseason . Recommended sowing rates pods are borne closely intermixed with the long are 2-4 kg/ha in pure stands and 0.5-1 kg/ha in tough stems andshatte r assoo n asthe y are dry. mixtures. Inoculation isno tusuall y necessary, but Genetic resources The present world collec­ the rhizobium strain CB 1024, or its equivalent, tion of axillaris is held at ATFGRC (CSIRO, Aus­ should be used if inoculation is required. Early tralia) and CIAT (Colombia). It contains a fair growth is rather slow. Axillaris combines well representation of material from Kenya to South with a wide range of grasses and is often sown in Africa, but the remainder of the natural range is mixtures with other twining or scrambling poorly represented. legumes. Breeding There are no known breeding pro­ Husbandry Axillaris competes very well with grammes. Its main limitations of pod shattering, weeds once established and has been deliberately which increases the cost of seed production, lack used to control shrubby weeds byallowin g the axil­ of long-term persistence under continuous heavy laris to grow without grazing from winter into grazing and limited adaptation to relatively mild spring in Australia. Axillaris will usually overtop climates could possibly beovercom e byfurthe r col­ weedy grasses by early autumn in the year of sow­ lection from other parts ofth e natural range or by MEDICAGO 163 genetransfe r from related species (e.g. indéhiscent Uses Lucerne is the world's most important for­ pods from M. uniflorum (Lamk) Verde). age crop and is cultivated mainly to provide feed Prospects The main strengths ofaxillari s are its for ruminants, but is also used as an ingredient of drought and disease resistance, its vigorous rations for poultry and pigs. Protein is extracted growth in summer, and the valuable grazing it pro­ from the forage for consumption by humans and vides in early winter and early spring in the a wide range of animals. Seeds are sprouted for subtropics. However, unless its climatic adapta­ human consumption. tion can be broadened and its resistance to heavy Properties Lucerne provides highly palatable, grazing improved, then it is only likely to play a highly nutritious forage, hay, silage and pellets. minor, though locally very useful, role. It is show­ At the early flowering stage, when it is commonly ing promise in Bolivia, Brazil, China (Hainan harvested, the dried forage contains 2.5-4.0% N Island), Ghana, Guatamala, India, Panama, the and is 60-70% digestible. There are 400-1000 Philippines, and Thailand, and could play a useful seeds/g. role in upland areas in most tropical countries. Botany A herbaceous perennial with deeply Literature |1| Blumenthal, M.J., O'Rourke, penetrating taproot. Stems procumbent, ascen­ P.K., Hilder, T.B. &Williams , R.J., 1989. Classifi­ ding to erect, arising from woody base, branching cation of the Australian collection of the legume at base and rising to 30-80(-120) cm. Leaf trifolio- Macrotyloma. Australian Journal of Agricultural late; stipules triangular, 5-15 mm long, pubescent Research 40: 591-604. |2| Cameron, D.G., 1986. on lower surface, glabrous onuppe r surface, joine d Tropical and subtropical pasture legumes. 10. at base, coarsely-toothed; petiole pubescent, 5-30 Axillaris (Macrotyloma axillare): a legume with mm long; leaflets narrow, oblong to ovate or obo- limited roles. Queensland Agricultural Journal vate, 8-28 mm x 3-15 mm, dentate near apex, gla­ 112: 59-63. |3| Gillett, J.B., Polhill, R.M. & Verd- brous on upper surface, slightly pubescent on court, B., 1971.Macrotyloma . In: Milne-Redhead, lower surface. Inflorescences in dense racemes E. &Polhill , R.M. (Editors): Flora of tropical East Africa. Leguminosae 4 - Papilionoideae 2. Crown Agents for Oversea Governments and Administra­ tions, London, pp. 581-594. |4| Luck, P.E., 1965. Dolichos axillaris - prospects in Queensland. CSIRO Australia, Division of Plant Industry, Plant Introduction Review 1(3):50a-51a . |5|Oram , R.N., 1990. Register of Australian herbage plant cultivars. CSIRO, Australia, pp. 177-178. I6| Sker- man, P.J. , Cameron, D.G. &Riveros, F., 1988.Trop ­ ical forage legumes. FAO, Rome. pp. 346-349. LB. Staples

Medicago sativa L.

Sp. PL: 778(1753) . LEGUMINOSAE 2n = 32 (tetraploid in cultivated forms), In = 16(diploi d in wild forms). Vernacular names Lucerne (En), alfalfa (Am). Luzerne (Fr). Philippines: galeria (Visaya). Thai­ land: thua-anfanfa. Origin and geographic distribution Lucerne is believed to have originated in the highlands of , Asia Minor and north-western , and/or . It has been cultivated for thousands of years and introduced to most countries of the world. It is only a minor species Medicago sativa L. -1, flowering branch; 2, flower; in South-East Asia. 3, infructescence; 4, fruit. 164 FORAGES containing 10-35 flowers, on peduncles 1-5 cm in the tropics. The most significant insect pests long;pedice l 1.5-2 mm long; calyx 5-lobed, 3-6 mm worldwide are the alfalfa weevil (Hypera postica) long, tube and pointed teeth about equal in length; and several aphid species. corolla purple or blue, rarely white, with some yel­ Lucerne is usually cut at intervals of 4-8 weeks low-flowered plants in cultivars containing M. fal- while it is growing, but can be grazed provided ade­ cata genes. Pod curled through 2-5 coils of 3-10 quate rest periods are allowed. Defoliation at the mm in diameter, indéhiscent, not spined, contain­ early flowering stage is preferable as this coin­ ing 2-6 seeds. Seed yellow to brown, kidney- cides with the development of new stems from shaped to ovoid, 1-2.5 mm x 1-1.5 mm. basal buds.Longevit y isreduce d under continuous The taxonomy of M. sativa is confused and the cul­ grazing. Frequent cutting is common in the tropics tivated forms may well be a tetraploid of hybrid but reduces quantity and stand life. origin. Because M. sativa is part of a species com­ In parts ofth e world where lucerne is a major crop, plex including M. falcata L., M. glutinosa M. v. annual DM yields of 10-20 t/ha are obtained. In Bieb. and M. glomerata Balbis, many hybrid forms South-East Asia, lucerne isusuall y fed green to ru­ have erroneously been given specific names. minants. It is a high quality feed but cattle grazing Most cultivars flower in long days (> 10-12 hours) green or freshly cut lucerne may suffer bloat (tym­ but vary in their quantitative response, which panitis). depends also on temperature. In suitable areas, in­ Genetic resources and breeding Significant dividual plants are long-lived (up to 20 years or germplasm collections are held in many countries. more). In tropical regions, plants persist for only The internationally recognized centre for Medi- 1-5 years. Regeneration from seed is usually poor cago germplasm isICARD A (Aleppo, ). Other in uncultivated ground. Soil seed reserves are low major collections are held by the USDA (Ames, because plants are rarely allowed to set seed. Iowa, United States) and the South Australian Lucerne ecotypes vary considerably, particularly Department of Agriculture (Adelaide, Australia). in winter dormancy. Diversity has been increased Commercial and government-funded breeding pro­ by natural and artificial hybridization. There are grammes are in progress in many countries. Dis­ hundreds of cultivars. ease resistance is a key objective. Ecology Lucerne is adapted to temperate and Prospects Lucerne will continue to have a mi­ Mediterranean climates and to heavy, neutral or nor role in South-East Asia. It will be restricted alkaline soils. Suitable cultivars can survive to suitable soils receiving adequate rainfall in ele­ extremely cold winters and hot, dry summers. It vated areas. It will have greater use in suitable is drought-resistant because of its deep root sys­ sites in the subtropics. tem, but its water use efficiency is poor. It cannot Literature ill Hanson, A.A., Barnes, D.K. & tolerate waterlogging. In the tropics, lucerne is Hill, R.R., 1988. Alfalfa and alfalfa improvement. adversely affected by hot, humid conditions which Monograph No 29. Crop Science Society of Amer­ promote several major diseases, by the absence of ica, Madison, Wisconsin, United States. 1084 pp. winter cold and by acid and waterlogged soils. 12, Leach, G.J. &Clements , R.J., 1984. Ecology and Agronomy Lucerne is propagated from seed, at grazing management of alfalfa pastures in the seeding rates of 2-12 kg/ha, sown at a depth of subtropics. Advances in Agronomy 37:127-154. 31 5-15 mm into cultivated seed-beds, usually in pure Lesins, K., 1976. Alfalfa, lucerne. In: Simmonds, stands. Control of existing vegetation is essential. N.W. (Editor): Evolution of crop plants. Longman, The time ofplantin g varies to suit local conditions. London, pp. 165-168. |4| Lesins, K.A. & Lesins, I., Inoculation of seed with suitable strains oîRhizo- 1979.Genu s Medicago (Leguminosae): a taxogene- bium is recommended where lucerne has not pre­ tic study. W. Junk, The Hague, Netherlands, pp. viously been grown. It is commonly irrigated and 95-108. heavily fertilized with P and K fertilizers to R.J. Clements enhance forage yield. Many diseases and insect pests attack lucerne. The most important diseases in the tropics are root and crown rots caused by fungal pathogens such as Phytophthora megasperma and Colletotrichum trifolii. Some cultivars, notably 'Trifecta', have improved resistance to these diseases. Plant para­ sitic nematodes are probably important pathogens MlCROSTEGIUM 165

Microstegium ciliatum (Trinius) A. upper part erect or scrambling, up to 2m tall . Leaf- Camus sheath 4-5 cm long, glabrous to long-hairy; ligule up to 2 mm long, glabrous to densely appressed- Ann. Soc.Linn . Lyon 68: 201(1921) . hairy at the back, not ciliate; leaf-blade narrowly GRAMINEAE lanceolate, up to 25 cm x 2.5 cm, narrowed at base In = 40 and acuminate at apex, glabrous, but hairy near Synonyms Pollinia ciliata Trin. (1832). the base. Inflorescence composed of 2-22 subdigi- Vernacular names Indonesia: banyonan tem- tate racemes, each 3-16 cm long; spikelets in alter­ baga (Javanese), jukut canklor (Sundanese). Phi­ nate pairs, one sessile, one pedicelled, along a flat­ lippines: bariri magwakat (Han.). Laos: hnhaaz tened axis; spikelets with small callus which has nhung. Thailand: ya kom ka. a few stiff setose hairs; lower floret male or neuter; Origin and geographic distribution M. cilia­ upper lemma with a long kneed and twisted smooth tum occurs widely throughout South-East Asia; it awn of up to 1.5 cm length. Caryopsis oblongoid, is also found in India, Sri Lanka, southern China, 1.5 mm long, brown. Taiwan and Japan. A very variable species. In the Flora of Java this Uses M. ciliatum isuse d for cattle and water buf­ taxon is considered as a complex of 3specie s which faloes through grazing or cut-and-carry in traditio­ are separated by the number of racemes (M. rufi- nal farm systems. spicum (Steud.) A. Camus: normally 2-5; M. cilia­ Properties It is regarded as a good quality for­ tum s.S.: 3-22; M. montanum (Nees) Henrard: age. 5-12). Botany A perennial grass growing in dense M. ciliatum grows slowly out in the open. In shady masses with branched culms, in lower part pro­ environments growth continues until September. strate or creeping and rooting at the nodes, in It flowers from September through the wet season in Thailand. Ecology It is adapted to light shade at altitudes up to 2400m in high rainfall areas, preferably with 2000 mm per year or more. It is common in rubber plantations, in disturbed and cultivated sites, and in open places and along the margins of rain for­ ests. It grows in patches and its trailing culms scramble over other low herbs. It is said to be able to suppress Imperata cylindrica (L.) Raeuschel, hence the name 'kom ka' meaning, in Thai, that it weakens Imperata cylindrica. Agronomy M. ciliatum is established by rooted stems spaced at 40 cm x 60 cm. Wider spacing allows excessive growth of weeds that interfere with M. ciliatum which is slow to establish. Newly planted stands should be kept weed free. A cutting height of 10-15 cm is recommended as it does not tolerate close defoliation. DM yields of 2t/h a have been obtained from 4cut s per year. It is a palatable species and is offered fresh to cattle or buffaloes, but it does not tolerate frequent grazing. It is not conserved. Genetic resources and breeding Although it is a variable species, it is unlikely that substantial germplasm collections are being maintained. Prospects M. ciliatum will continue to be an important source of feed in plantations. Research objectives would be to increase its yield and to de­ velop mixtures with legumes. Microstegium ciliatum (Trinius) A. Camus - 1, Literature il1 Backer, CA. & Bakhuizen van leafy culmpart; 2, inflorescence; 3,pair of spikelets. den Brink, R.C., 1968. Flora of Java. Vol. 3. 166 FORAGES

Wolters-Noordhoff, Groningen, the Netherlands, pp. 589-590. 121Gilliland , H.B., Holttum, R.E. & Bor, N.L., 1971. Grasses of Malaya. In: Burkill, H.M. (Editor): Flora of Malaya. Vol. 3. Govern­ ment Printing Office, Singapore, pp. 248-249. !3| Manidool, C., 1989. Natural grassland and native grasses of Thailand [in Thai]. Technical Bulletin No 1301-26-32. Division of Animal Nutrition, Department of Livestock Development, Bangkok, pp. 24-25. C. Manidool

Mikania cordata (Burm.f.) B.L. Robinson

Contrib. Gray Herb. 104:6 5(1934) . COMPOSITAE 2n = unknown Synonyms Eupatorium cordatum Burm.f. (1768), Mikania volubilis (Vahl) Willd. (1803), M. scandens Hook.f. (1881),no n (L.)Willd . (1803). Vernacular names Mile-a-minute (En). Indone­ sia: brodjo lego (Javanese), hila hitu lama (Ambon), blukar (Sumatra). Malaysia: ceroma, akar lupang, selaput tunggul. Philippines: bikas (Bagobo), detidid (Igorot), uoko (Bontok). Thai­ land: khikaiyan. Origin and geographic distribution M. cor­ Mikania cordata (Burm.f.) B.L. Robinson -1, flow­ data is native to tropical Asia and Africa and is ering plant part; 2, flower-head; 3, achene with widespread throughout South-East Asia and the flower. Pacific Islands. Uses Leaves of M. cordata constitute a highly with short hairs. Leaves cordate or triangular- palatable forage, especially to sheep.The y are also ovate, 1-12 cm x 1-6 cm, base cordate or shortly used as a poultice for swellings (Taiwan), itches contracted, margin crenate-dentate, sinuate or (Malaysia) and wounds (Indonesia). In South entire, apex acutely acuminate, subglabrous, glan- Africa it is used as a cure for snake and scorpion dularly spotted beneath petiole 1-8 cm long. bites. Inflorescence composed of peduncled heads, com­ Properties The DM digestibility is about 50% bined into small dense corymbs, at the top of short, and N concentrations range from 2.6-3.4%, Ca side branches and in the axils of leaves; peduncle concentrations from 1.5-1.9% and P concentra­ of corymbs very variable in length; peduncle of tions from 0.6-0.9 %. It has high concentrations of heads up to 6m m long; heads 6-9 mm x 1.5-2 mm, Cu, about 18 mg/kg. It has been reported from 4-flowered; involucre of 4 elliptical-oblong bracts; Malaysia that M. cordata contains phenolic or fla- corolla campanulate, 5 mm long, yellowish-white, venoid substances that inhibit the growth of rub­ 5-6-lobed; style with 2 long exserted branches, ber, tomato and tropical kudzu (Pueraria phaseo- each 2.5 mm long, white. Fruit a linear-oblongoid, loides (Roxb.) Benth.), and depress nitrification in 4-ribbed achene, 2-3 mm long, glandular, black- soils. brown; pappus hairs 3-4 mm long, white to It can be an important weed in cropped land and reddish. areas under young plantation crops. M. cordata spreads by seed or by rooting at nodes Botany Scandent perennial herb, often forming which are in contact with soil. Flowering occurs a dense tangled mass. Stem subterete or irregu­ throughout the year. Seeds are produced in large larly angular, ribbed, up to 6 m x 2-3 mm, inter- numbers and the pappus enables effective wind dis­ nodes 6-14 cm long, nodes thickened, sometimes persal over long distances. With its rampant MIMOSA 167 growth it can rapidly smother young tree crops 50% of the diet. There is evidence that these prob­ and other plants, hence the common name 'mile-a- lems relate to the high Cu concentrations in M. minute'. It can rapidly form a tangled mass to a cordata. There is very little information about its depth of 0.6-1 m. If undisturbed, it often spreads productivity. Yields of4 t/h a ofD M have been mea­ in massive circular patterns. sured in Mauritius. There are no records of animal This species was long considered to be conspecific production where it wasth e solefeed , but good ani­ with its allies in (M. scandens (L.) mal production levels have consistently occurred Willd.) and in South America (M. micrantha in plantations where it has been an appreciable Kunth). Distinctive characters are: component of the understorey. - M. scandens: heads 5-6 mm long; involucral Genetic resources and breeding It is unlikely bracts lanceolate-oblong, long-acuminate and that any germplasm collections of M. cordata are very acute at apex; corolla purple; achene longer being maintained. than 2mm . Prospects M. cordata is an aggressive forage of - M. micrantha: heads 4-5 mm long; involucral high acceptability to both large and small rumi­ bracts oblong-elliptical, shortly acute at apex; nants. Further study should be given to its agro­ corolla white; achene less than 2m m long. nomy and to feeding systems using this species so In Malesia 4 forms of M. cordata have been distin­ that its potential as forage can be better exploited, guished, mainly based on leaf form, leaf indumen­ while at the same time aiding in the biological con­ tum and leaf margin characteristics. In West trol of a serious weed. Africa 2varietie s have been distinguished, mainly Literature |1! Barnes, D.E. & Chandapallai, based on peduncle length and length of involucral M.M., 1972. Common weeds of Malaysia and their bracts. control. Ansul (M) Sdn. Bhd. Kuala Lumpur, p.93 . Ecology M. cordata is adapted to hot, humid !2 Devendra, C, 1979. Malaysian feedingstuffs. tropical environments with 1500m m or more annu­ MARDI, Kuala Lumpur, p.29 . ;3j Henderson, M.R., al rainfall and plenty of sunlight, at altitudes rang­ 1974. Malayan wild flowers 1. Dicotyledons. The ing from sea-level to 2000 m or more. Hence it is Malayan Nature Society. Kuala Lumpur, p. 233. commonly found in young secondary jungle, forest |4|Holm , L.G., Plucknett, D.L., Pancho, J.V. &Her - clearings, abandoned ground, secondary regrowth berger, J.P., 1977.Th e world's worst weeds: distri­ areas, ravines, mountain slopes, roadsides, water­ bution and biology. The East-West Center, Univer­ courses, fallow lands, low-lying areas along sity of Hawaii Press, Honolulu, Hawaii, United streams and rivers and open plantations. However, States, pp.320-327 . 5| Koster, J.T., 1935.Th e Com- it can also persist with reduced vigour in planta­ positae of the Malay . 11. Mikania. tions. Although it may even be found under closed Blumea 1:503-510 . |6iPerry , L.M., 1980. Medicinal canopies of 4-5 year-old rubber and oil palm, it is plants of east and southeast Asia: attributed pro­ markedly etiolated with little vigour. It is rarely perties and uses. MIT Press, Cambridge, Massa­ found in 5-15 year-old plantations. chusetts, United States, p. 103. \T Wong, R., 1964. Agronomy M. cordata can be a devastating Evidence for the presence of growth inhibitory weed in crops of tea, coconut, cocoa, rubber, oil substances in Mikania cordata (Burm.f.) B.L. palm, coffee, and sugar cane, and can Robinson. Journal ofRubbe r Research Institute of smother leguminous cover crops. Spraying with Malaya 18:231-242 . herbicides can reduce its vigour and spread. C.P. Chen&Y.K. Chee It has been reported asbein g susceptible to parasit­ ic growth ofdodder , Cuscuta chinensis Lamk in Sri Lanka and C.australis R. Br. in Fiji and Malaysia. Mimosa pudica L. It is very palatable to livestock, particularly to sheep. Where present, it is the first species to be Sp. PL: 518(1753) . eliminated when sheep graze pastures. It is equally LEGUMINOSAE acceptable when either grazed or cut and fed as 2n = 52 a supplement to other forages. There are no Synonyms records of it being used for hay or silage. Sheep var. tetrandra (Humb. & Bonpl. ex Willd.) DC: should not be fed forages where it is the main com­ Mimosa tetrandra Humb. & Bonpl. ex Willd. ponent. Instances of abortion, death of newborn (1806). lambs and of older sheep have been recorded in var. unijuga (Duchass. & Walp.) Griseb.: rubber plantations where it comprised more than Mimosa unijuga Duchass. & Walp. (1850). 168 FORAGES

Vernacular names Sensitive plant (En). Sensi­ tive (Fr). Indonesia: putri malu (Indonesian), bujang kaget (Sundanese), kucingan (Javanese). Malaysia: memalu, malu-malu. Philippines: maka- hiya (Tagalog), torog-torog (Bikol), Babain (Ilo- kano). Cambodia: smau bânla, bânkrap. Laos:f a:z langab, th'üb nhub. Thailand: maiyarep (central), ya-ngap (south). Vietnam: xâu hô', trinh nû'. Origin and geographic distribution Probably M. pudica originated in South America but is now spread pantropically. It occurs throughout South- East Asia, usually along roadsides and in waste­ lands. Uses Young stems and leaves are useful as for­ age. When the prickles on the stem and the fruits become too hard, they can cause intestinal inflam­ mation in animals. It is often considered as a nox­ ious weed. In areas where other leguminous plants establish with difficulty, M. pudica can be of use as a cover crop or green manure. On Java the plants are believed to induce sleep. In Malaysia pounded leaves are applied as a dressing to relieve swellings. Properties Very littlei sknow n about the forage quality ofsensitiv e plant. It is likely that the quali­ ty of any sample would vary considerably with the leaf/stem ratio. There are appoximately 110 seeds/ g- Mimosa pudica L. - 1, flowering and fruiting Botany Annual or perennial herb, sometimes branch; 2, leaflet; 3, flower; 4, fruit; 5, one-seeded woody at base, up to 1.5 m tall, often prostrate or segment offruit; 6,seed. straggling. Stem usually sparsely armed with prickles up to 5 mm long, glabrous to densely his­ Willd.) DC. (corolla in bud densely grey puberu­ pid. Leaves bipinnate, unarmed, sensitive; petiole lous; heads in bud not or sparsely bristly; stipules 1.5-5.5 cm long; rachis very short, giving the 2 4-8 mmlong) ;var . uni)uga (Duchass. &Walp.)Gri- pairs of pinnae a subdigitate position; leaflets seb. (corolla in bud glabrous; heads in bud not 10-26 pairs, linear-oblong, 0.6-1.5 cm x 1-3 mm, bristly; stipules 4-8 mm long). margins setulose. Inflorescence a globose head, Ecology Sensitive plant is common in waste­ about 1 cm in diameter, 1-5 together per axil; land, disturbed areas and overgrazed sites with peduncle up to 4 cm long; flowers lilac, pink or moderately to poorly fertile soils. It occurs in the blue-purple; calyx about 0.2m m long, corolla 2m m wet tropics and tolerates waterlogging but is not long; stamens 4.Po d flattened-oblongoid, 1-1.8 cm well adapted to the seasonally dry tropics. It is x 3-5 mm, several together in a cluster, densely regarded as a weed in dryland field crops, in setose, prickly on margins only, consisting of 3-5 rainfed wetland rice and in plantation crops where one-seeded joints which break away from the per­ it is reasonably tolerant of shade. It occurs over sistent sutures. Seed suborbicular to ellipsoidal, a wide range of soil types, but iron chlorosis has flattened, 2.5-3 mm long, light-brown, surface been noted on coralline soils with a high pH. finely granular. Agronomy Sensitive plant is not deliberately The species flowers throughout the year and it can sown, but its persistence and spread is aided by its complete its life cycle in 3 months. Four varieties high seed set. It tends to invade pastures of declin­ are distinguished: var. pudica (only known from ing soil fertility and is less common where soil fer­ the sterile type specimen, no distinction possible); tility is good and pastures are not overgrazed. Sen­ var. hispida Brenan (corolla in bud densely grey sitive plant has been eliminated from pastures puberulous; heads in bud densely bristly; stipules where soil fertility is very poor and grazing pres­ 8-14 mm long); var. tetrandra (Humb. &Bonpl . ex sure is very high. It grows with a wide range of NEONOTONIA 169

grasses, including signal grass (Brachiaria decum- and mainland Asia and Indonesia. It can currently bens Stapf) provided it is not too vigorous. It can be found in many humid tropical and subtropical hamper the establishment of improved species. regions of the world since its widespread introduc­ Sensitive plant is more readily accepted if it is tion for use as forage. grazed continuously rather than rotationally. Uses Glycine is used as a grazed pasture legume Observations suggest that it can at times stimulate in Australia in the subtropical and tropical high- better growth of associated grasses. There are no altitude regions, and for grazing and hay in Brazil. records of pests and diseases and yield. It is used in small areas of Papua New Guinea Genetic resources and breeding There are no where it volunteers as a fallow crop in abandoned known germplasm collections of sensitive plant. gardens and is used as a cover crop and for woody Prospects Although sensitive plant can provide weed control in overgrazed pastures. useful forage, it will continue to be primarily Properties Nitrogen concentrations ranging regarded as a weed. While more appropriate man­ from 2-4.2% have been measured in the leaves, agement may promote the use ofit , it would be pref­ with DM digestibilities of between 55-61%. Al­ erable to grow species that are more readily though oestrogenic substances are present, no tox­ accepted by grazing animals or more suited to cut- icity problems have been recorded. and-carry feeding systems. Botany Perennial twining or climbing herb with Literature jlj Brenan, J.P.M., 1955. Notes on a woody base, 0.5-4.5 m long. Taproot long, nodu- Mimosoideae: 1. Mimosa. Kew Bulletin 1955(2): lating, rootstock thick and woody. Stems slender 184-189. |2| Moody, K., Munroe, CE., Lubigan, and well branched; a dense crown may develop E.T. & Palier, Jr., E.C., 1984. Major weeds of the under grazing; stolons in contact with moist soil Philippines. University of the Philippines at Los often root at the nodes. Leaves pinnately trifolio- Banos, College, Laguna, the Philippines. |3| Niel­ sen, J., 1981. Légumineuses-Mimosoïdées. Mimosa pudica L. In: Aubréville, A. & Leroy, J.-F. (Edi­ tors): Flore du Cambodge, du Laos et du Vietnam. Vol. 19. Muséum National D'Histoire Naturelle, Laboratoire de Phanérogamie, Paris, pp. 35-38.14| Skerman, P.J., Cameron, D.G. &Riveros , F., 1988. Tropical forage legumes. FAO, Rome. pp. 352-353. |5jSoerjani , M., Kostermans, A.J.G.H. &Tjitrosoe- pomo, G., 1987. Weeds of rice in Indonesia. Balai Pustaka, Jakarta, pp. 330-331. |6j Verdcourt, B., 1979. A manual of New Guinea legumes. Botany Bulletin No 11. Office of Forests, Division of Bota­ ny, Lae, Papua New Guinea, pp. 148,151. R.M. Jones &N.O . Aguilar

Neonotonia wightii (Wight & Arnott) Lackey

Phytologia 37:21 0(1977) . LEGUMINOSEAB In = 22 (diploid), 44. Synonyms Notonia wightii Wight & Arnott (1834), Glycine javanica auct. mult., non L. (1753), G.wightii (Wight &Arnott ) Verde. (1966). Vernacular names Glycine (En). Brazil: soja perene. Thailand: thua peelenian soibean. Origin and geographic distribution The natu­ ral distribution of glycine ranges from Africa to Neonotonia wightii (Wight & Arnott) Lackey - 1, Asia. It occurs as far south as the wetter parts of flowering and fruiting vines; 2, flower; 3, fruit; 4, southern Africa, in East Africa, Ethiopia, India seeds. 170 FORAGES

late; petiole 2-12 cm long; leaflets ovate or ellipti­ able strain of Bradyrhizobium. Seed should be cal, 1-15 cm x 1-12.5 cm, glabrous to densely vel­ sown at a depth of less than 2 cm. Where weeds vety onbot h sides.Raceme s axillary, 2-60 cm long, become a problem, high slashing is the best con­ 20-150-flowered; peduncle 2-10 cm long; flower trol. white or violet, 4.5-11 mm long. Pod cylindrical, To allow build-up of a soil seed reserve and to en­ 1.5-5 cm x 2.5-5 mm, straight or slightly falcate, able re-establishment, the pasture should be thinly septate, glabrous to densely hairy, allowed to seed down in its first year. As with other 3-8-seeded. Seed cylindrical, compressed, 2-4 mm twining legumes, glycine is susceptible to heavy x 1.5-3 mm x 1-1.5 mm, reddish-brown. grazing so it should be leniently grazed and rest Three subspecies are recognized: ssp. wightii (pod periods should be allowed at flowering time if densely hairy, flower 4.5-7.5 mm long), ssp. pseu- under continuous grazing. Because of its ability dojavanica (Taubert) Lackey (pod glabrous, flower to climb, glycine has the potential to become a 4.5-7.5 mm long), and ssp. petitiana (A. Rich.) weed among tree crops, although this is unlikely Lackey (pod densely hairy, flower 7.5-11 mm where it would be regularly cut or grazed. long), and within these a number of varieties; the Glycine combines well with a number of grasses species as a whole is extremely variable, but due and has been particularly successful with a range to the occurrence of intermediates, the practical of Panicum cultivars and with setaria (Setaria value of the botanical subclassifications can be sphacelata (Schumacher) Stapf &Hubbar d ex M.B. questioned. Moss). Several cultivars have been developed throughout Under wet conditions, the leaves can be damaged the world, their selection being based on such fac­ by leaf blight (Rhizoctonia), leaf-spot (Cercospora) tors as cold tolerance, maturity, drought tolerance and sclerotinea (Sclerotinea sclerotiorum). On the and tolerance to high Mn levels. The cultivars Atherton tableland in Queensland, the webworm released in Australia are 'Tinaroo', a diploid late- (Oncopera sp.) can severely damage tops. Control flowering variety with high cold tolerance, 'Coo­ is seldom attempted. Glycine may yield up to 4-5 per', a diploid early-flowering type which tolerates t/ha of DM per year when sown with a grass and excess moisture and drought stress, 'Clarence', the up to 10 t/ha per year if grown alone. Forage is earliest flowering cultivar and the first variety to usually grazed directly by animals but where start spring growth, and 'Malawi', a tetraploid mown for hay, care should be taken not to cut too which is less branched than the other cultivars but low as this could damage the crown or low stolons. adapted to more acid and less fertile soils. Genetic resources and breeding Germplasm is Ecology Glycine is a short-day plant. Optimum available from ATFGRC (CSIRO, Australia). day/night temperatures for growth and seed pro­ Because glycine plants have a short-day photope- duction of glycine are in the range 22-27/16°C riod response, seed production in tropical regions with a base temperature for growth of 13°C , lower will be limited for some accessions and cultivars. than for most tropical legumes. It has limited frost However, material with origins in the high alti­ tolerance, but leaf damage occurs in all cultivars. tudes of tropical Africa should be reproductive in It isadapte d to dry to humid (800-1800 mm) subtro­ similar environments inAsi a and it isthi s material pical or high altitude (up to 3000 m) tropical which would need to be developed for the region. regions with relatively deep, well drained, usually Prospects In South-East Asia, glycine will be basalt-derived soils, and prefers a pH above 6.0. It limited to deep, fertile and well-drained soils in is not adapted to very wet or waterlogged condi­ tropical high-altitude areas. Such areas are tions. It is not as drought tolerant as siratro (Ma- limited and this species is unlikely to be widely croptilium atropurpureum (DC.) Urban), but more used. Currently no selection, breeding or develop­ so than centro (Centrosema pubescens Benth.). ment studies are being carried out. Agronomy Propagation is by seed which, espe­ Literature il| Bogdan, A.V., 1977. Tropical pas­ cially if hand harvested, often has a high propor­ ture and fodder plants. Longman, London, pp. tion ofhar d seed. Mechanical or acid scarification 357-364.12! Cameron, D.G., 1984.Tropica l and sub­ (concentrated sulphuric acid for 25 minutes and tropical pasture legumes 4. Glycine (Neonotonia then washed) is usually required for successful wightii): an outstanding but soil specific legume. establishment. Because seedling growth and nodu- Queensland Agricultural Journal 110:311-316 . '3| lation are slow, a weed-free seed-bed is favoured Duke, J.A., 1981. Handbook of legumes of world for establishment. Sowing rates of 2-6 kg/ha are economic importance. Plenum Press, New York, recommended with the seed inoculated with a suit­ pp. 88-90. |4|Oram , R.N., 1990.Registe r of Austra- OTTOCHLOA 171

lianherbag e plant cultivars. CSIRO,Australia , pp. 179-182. j5| Verdcourt, B., 1971. . In: Milne-Redhead, E. &Polhill , R.M. (Editors): Flora of tropical East Africa. Leguminosae 4 - Papilio- noideae 2. Crown Agents for Oversea Govern­ ments and Administrations, London, pp. 528-533. B.C. Pengelly &A.K . Benjamin

Ottochloa nodosa (Kunth) Dandy

J. Bot. 69:5 5(1931) . GRAMINEAE 2n = unknown Synonyms Panicum nodosum Kunth (1833), Hemigymnia multinodis Stapf (1920), H.fusca Rid­ ley (1925). Vernacular names Slender panic grass (En). Indonesia: suket pring-pringan (Javanese). Malay­ sia: rumput pait, rumput rawa, rumput sarang buaya. Philippines: banig-usa, kawakawayanan (Tagalog), bariri-magwakat (Han.). Laos: chaax kh'aa. Thailand: ya-laman (central), ya-khui- phaikhon (western). Origin and geographic distribution 0. nodosa occurs throughout South-East Asia and also in India, Burma and Sri Lanka. It has been intro­ duced to Mexico and parts ofAfric a and Australia. Ottochloa nodosa (Kunth) Dandy - 1,habit flower­ Uses 0. nodosa is used as a forage for cattle and ing plant part; 2, ligule; 3, spikelets. sheep, particularly in rubber and oil-palm planta­ tions in South-East Asia, where it also can become Ecology O. nodosa occurs from sea-level up to a troublesome weed if ungrazed. 600 m, mainly in forest, rubber, or oil-palm planta­ Properties Nitrogen concentrations range typi­ tions. It also occurs in rice fields and disturbed cally between 1.1 % and 1.3% , with DM digestibili­ habitats. ties between 38% and 50%. DM intakes by sheep Agronomy O. nodosa spreads naturally, but it of 350-400 g/head per day have been measured in can be planted by seeds or rooted culms. It is Thailand. usually grazed by cattle or sheep but can be cut Botany Perennial grass with slender, decum­ byhan d or mowers and fed to animals. It is modera­ bent or scandent culms, more than 1 mtall , rooting tely palatable but not tolerant of heavy grazing or at basal nodes. Leaf-sheath shorter than the inter- frequent cutting, so stocking rates should not node, ciliate at its margins; ligule a shallow fim­ exceed 1.5 beast/ha and cutting intervals of 8-9 briate membrane, about 0.5 mm tall; leaf-blade weeks are preferred. narrowly lanceolate, up to 15 cm x 12 mm, gla­ Genetic resources and breeding It is unlikely brous or with scattered bulbous-based hairs. that substantial germplasm collections are being Inflorescence of paniculately arranged racemes, maintained. up to 30 cm long; lower branches often in a whorl; Prospects O. nodosa is regarded as a trouble­ largest branches with ca. 10 spikelets, grading to some weed and as a fodder of moderate value, espe­ solitary spikelets at apex of a branch; spikelets cially in plantations, but more information is elliptical, 2m m long, with glumes shorter than the needed about its management and utilization. spikelets; lower floret neuter, upper floret her­ Literature jlj Gilliland, H.B., Holttum, R.E. & maphrodite. Bor, N.L., 1971. Grasses of Malaya. In: Burkill, The inflorescence is extremely variable in number H.M. (Editor): Flora of Malaya. Vol. 3. Govern­ ofbranche s and spacing ofspikelets . In Peninsular ment Printing Office, Singapore, pp. 143-144. 21 Thailand it flowers from September to May. Manidool, C, 1989. Natural grassland and native 172 FORAGES grasses of Thailand [in Thai]. Technical Bulletin No 1301-26-32. Division of Animal Nutrition, Department of Livestock Development, Bangkok. 45pp . |3|Mehra , K.L. &Fachrurozi , Z., 1985. Indo­ nesian economic plant resources: forage crops. Lembaga Biologi Nasional - LIPI, Bogor, No 31. p. 28. |4| Santos, J.V., 1986. Grasses. In: Umali, R.M., Zamora, P.M., Gotera, R.R. &Jara , R.S.(Edi ­ tors): Guide to Philippine flora and fauna. Vol. 4. Natural Resources Management Center and Uni­ versity of the Philippines, Manila, p. 100. C. Manidool

Panicum maximum Jacq.

Ic.pl. rar. 1:2,1.13(1781). GRAMINEAE In = 18, 32, 36, 48 Synonyms Panicum polygamum Swartz (1788), non Forsskal (1775). Vernacular names Guinea grass, buffalo grass, green panic grass (En). Herbe de Guinée, panic élevé (Fr). Indonesia: rumput banggala (Indone­ sian), suket londo (Javanese), rebha luh-buluhan (Madura). Malaysia: rumput kuda, rumput beng- gala. Thailand: ya-kinni (central). Vietnam: co' kê to. Origin and geographic distribution Guinea Panicum maximum Jacq. 1,habit flowering plant; grass is East African in origin, but is now widely 2, ligule; 3, spikelet. cultivated throughout the tropics and subtropics as a forage. It was introduced from Africa into the tussock grass, rhizomatous at base or rooting at West Indies before 1756,bu t for production of bird the lower nodes, glabrous to hairy. Low/medium seed rather than forage. It reached Singapore in height forms 1-1.5 m tall, tall forms 2.5-3.0 m or 1876 and the Philippines in 1907 and is now widely more. Leaf-sheath glabrous; ligule membranous, distributed throughout South-East Asia. 4-6 mm long,fringed ; leaf-blade linear to narrowly Uses Guinea grass is a palatable and good quali­ lanceolate, up to 60 cm x 2 cm, usually glabrous ty tropical grass used as forage for ruminants in except in hairy forms (e.g. cultivar 'Makueni'); grazed pastures or in cut-and-carry systems. Gui­ midrib pale, vanishing near the tip, other nerves nea grass forage is also dried and ground for use distinct. Inflorescence usually a large, well in mixtures with legumes as leaf meal, mainly for extended pyramidal panicle up to 40 cm long by non-ruminants such as chickens and pigs. It can 25 cm across with lower primary branches about be conserved as hay or ensiled. It is also used as 20 cm long and arranged in a whorl; spikelets soli­ medicine for heartburn by the under the tary, numerous, narrowly ellipsoidal, 3-4.5 mm name 'berita'. long, green or tinged with purple, acute; lower Properties In vitro digestibilities of 64%, 58%, glume 1-2 mm long, thin, clasping, obtuse, 54% and 50% have been measured in regrowth of 3-nerved; upper glume 3-4.5 mm long, thin, ages 2, 4, 6 and 8 weeks. Comparing young leafy narrowly boat-shaped, 5-nerved, acute; lower flor­ with older stemmy herbage, similar trends occur ets male, lemma similar to upper glume; upper flor­ in N concentrations which can fall from over 2% ets hermaphrodite, lemma tranversely rugose, to below 1%, and in metabolizable energy which boat-shaped, pointed, palea fitting inside lemma, declines from 11-7 MJ/kg. both ca. 2 mm long, anthers and stigma purple. There are 1000-2000 seeds/g. Seed production apomictic. Description An erect or ascending perennial Growth and development Poor seed produc- PANICUM 173

tion, as low as 50 kg/ha in the wet tropics, has dis­ available. Fertilization immediately before or couraged the commercial production of guinea after planting or sowing can be important because grass seed. Environments with longer daylengths of the slow growth of seedlings in the first month, and dry seasons are more conducive to successful when they can be easily overwhelmed by agressive seed set. Stems of guinea grass root freely from weeds, and it is essential in soils which are poor. nodes when in contact with moist soil but, as a Husbandry After establishment, a guinea grass result of its erect growth habit, such rooted creep­ pasture should be allowed to flower and set seed ing stolons are rarely seen in the field. and then be given a light grazing before the seed Other botanical information There are many drops. Guinea grass combines well with herba­ guinea grass cultivars commercially available. ceous legumes, but it is not suited to sustained fre­ Australia is the main producer of guinea grass quent and close grazing. Guinea grass is very res­ seed; the forms available include common guinea ponsive to chemical fertilizers, especially N. Dry and coloniâo and cultivars 'Hamil', 'Riversdale', matter yield responses under cutting are obtained 'Gatton' and 'Makueni'. Although the selection of with N applications up to 800kg/h a per year. How­ these cultivars was primarily done within Austra­ ever, a more efficient use of N is obtained with lia, they are adapted to the different environments 100-300 kg/ha N per year. in South-East Asia. The coloniâo form was Diseases and pests Ergot can be a problem in selected in Brazil. Variety trichoglume Robijns is seed production during wet periods. No major dis­ treated separately in this volume. eases or pests of guinea grass have been docu­ Ecology Guinea grass is indigenous to the more mented. fertile soils in Africa where it is often found grow­ Harvesting Fresh foliage can be grazed directly ing close to the trunks of thorny Acacia spp. The by animals or can be cut for stall feeding. The N grass benefits from the trees because of the N they concentration of grazed guinea grass can be 30% fix and the shade they provide. Guinea grass higher than in cut material. Although usually fed thrives best under high rainfall conditions and a fresh, guinea grass can be conserved as hay or minimum annual rainfall of 1100-1300 mm has silage. However, chopping, wilting and the addi­ been suggested. Cultivar 'Hamil' is more tolerant tion of 4% molasses are recommended before of waterlogged conditions than other cultivars. silage making. Guinea grass is adapted to fertile soils or where Yield In Malaysia, annual DM yields range from appropriate fertilizer is applied. Generally it does 16-30 t/ha per year on sedimentary soils, 19-30 not thrive on oxisols or ultisols unless they have t/ha per year on peat soils and 1.5-9 t/ha per year been heavily limed. Extensive natural areas of vig­ on bris (coastal marine sand) soils when cut at 6-8 orous guinea grass in the tropics are usually only weekly intervals. Well managed guinea grass- found on fertile dark clay soils with a high Ca con­ legume pastures can support cattle at 1000 kg/ha tent, which are rare in South-East Asia. liveweight, gaining 450kg/h a per year. When ferti­ Guinea grass responds to moderate shading. At lized with 300kg/h a Npe r year, the animal biomass about 30% shade its yield exceeds that obtained can be increased to 1400 kg/ha liveweight with an in full sunlight. Even in 50% shade, it can still annual gain of 800kg/ha . Similar liveweight gains maintain about 50% of its normal production. to N fertilized guinea grass pastures can be Because of this shade tolerance, it is well suited achieved with guinea grass-leucaena pastures. to agroforestry situations as inrubbe r and coconut Milk production of 8500 kg/ha milk per year has plantations, or to growing with leguminous forage been recorded on guinea grass with N fertilizer trees such as leucaena (Leucaena leucocephala using 2.5 Holstein-Friesian cows per ha yielding (Lamk) de Wit) which provide partial shade and 11.3kg/co w per day. Guinea grass-glycine (Neono- the essential N. tonia wightii (Wight & Arnott) Lackey) pastures Propagation and planting Sowing, usually at produced 8220 kg/ha per year using Friesian cows 2-3 kg/ha of seed into a well-prepared seed-bed, is producing 12.7kg/day , at a slightly lower stocking the best way to establish large areas of guinea rate. grass. A fine soil surface and rolling encourage Genetic resources Large collections of guinea germination and establishment. Vegetative plant­ grass have been assembled by ATFGRC (CSIRO, ing of 2-3 rooted tillers per clump in a triangular Australia) and CIAT (Colombia). These have pro­ planting pattern at 40 cm spacing can expedite vided a wide range of morphological forms and establishment, and is suitable for small-scale pas­ edaphic adaptations. ture development, especially when seed is not Breeding Selection and breeding of guinea 174 FORAGES grass for specific ecological conditions has been Panicum maximum Jacq. var, carried out by a number ofresearc h organizations, trichoglume Robijns including CIAT (Colombia) and EMBEAPA (Bra­ zil). Japanese scientists have shown increasing in­ Mem. Inst. Roy. Col. Beige 1(6):3 1(1932) . terest in breeding guinea grass for the more diffi­ GRAMINEAE cult climatic conditions prevailing in that In = 32 country. As a result, cultivar 'Natsukaze' has been Synonyms Urochloa maxima (Jacq.) R. Webster registered. var. trichoglume (Robijns) R. Webster (1987). The discovery of a few sexual types of guinea grass Vernacular names Green panic, slender gui­ in very large populations of this usually apomictic nea grass, fine-leaved guinea grass (En). grass has made it possible for plant breeders to de­ Origin and geographic distribution Green velop cultivars suited to different environments. panic is native to tropical and subtropical Africa Apomicts like guinea grass have a considerable and has been widely distributed, amongst others reservoir of 'fixed' variability which can be to India, Japan, Australia, Sumatra, Fiji, and the released by crossing with sexual types now avail­ United States. able. Using crosses between these sexual types and Uses Green panic is used in pastures for grazing other vigorous lines, a breeding programme to de­ and for hay or silage making. It can also be grown velop guinea grass cultivars tolerant of acid soils as ley pasture between two crops, e.g after a paddy was carried out in Colombia and Brazil. This pro­ rice crop;i t isa n important grass in sub-humid sub­ gramme has led to the release of cultivar 'Vence- tropical regions with summer rainfall. dor' by EMBRAPA in 1990. This cultivar thrives Properties Green panic is apalatabl e grass with in soils with pH < 5, but has also been shown to an N concentration of green material between have better than usual shade and cold tolerance. 1.4-3.6% ofDM , depending on stage of growth and At the Rubber Research Institute in Malaysia, soil fertility or fertilization rate. Phosphorus con­ 'Vencedo r' wa s found to be one ofth e highest yield­ centrations of0.2-0.4 % ofD M have been recorded ing tropical grasses at 55-80 % light transmission. in green material. Dry matter digestibility ranges Prospects Guinea grass is unquestionably one between 50-65%, depending on age of material ofth e best tropical grasses and will be increasingly and time ofyear . There are about 2400 caryopses/g. used in South-East Asian pastures. With the trend Botany A bunched, tussock-forming perennial towards agroforestry, guinea grass will become grass with ascending habit, up to 2 m tall; the more important in associations with trees as crown expands by short horizontal stems; root sys­ diverse as rubber and leucaena. More use could be tem consists of fine, richly branched, shallow made ofit , particularly when mixed with leucaena, roots. Stem slender, 6-8 noded. Leaves fine and for feeding as leaf meal to non-ruminants. soft; leaf-sheath sparsely hirsute; ligule a ring of Literature |1|Bogdan , K.V., 1977. Tropical pas­ downy hairs; leaf-blade velutinous on lower sur­ tures and fodder plants. Longman, London & New face. Inflorescence a loose panicle, reaching well York. pp. 181-191. |2| Eng, P.K., Chen, C.P. & 't above the leaves; spikelets solitary or paired, Mannetje, L., 1978.Effect s of phosphorus and stoc­ oblongoid, 2.7-3.5 mm long; glumes puberulent, or king rate on pasture and animal production from strigose to velutinous; lower lemma 5-nerved, a guinea grass legume pasture in Jahore, Malay­ hairy; palea of lower floret fully developed, ellipti­ sia. 2:Anima l live weight change. Tropical Grass­ cal; upper lemma mucronate, coarsely transver­ lands 12:198-207. |3|Gilliland , H.B.,Holttum , R.E. sely rugose. & Bor, N.L., 1971.Grasse s of Malaya. In: Burkill, Green panic seed reaches its maximum germina­ H.M. (Editor): Flora of Malaya. Vol. 3. Govern­ tion rate after storage for up to 18 months. It has ment Printing Office, Singapore, pp. 132-134. |4j vigorous seedlings, is fast growing and responds Sato, H., Shimizu, N., Nakagawa, H. & Nakajima, quickly after rain. Its flowering period extends K., 1990. A new registered cultivar 'Natsukaze' of throughout the wet season. Guinea grass. Japanese Agricultural Research In the literature a distinction is not always made Quarterly 23:196-201. between green panic and guinea grass (Panicum C.P. Chen &E.M . Hutton maximum Jacq.) The only registered cultivar is 'Pétrie' from Australia. Ecology Green panic is adapted to humid (10-12 humid months) and sub-humid (6-9 humid months) subtropical climates with hot wet summers and dry PANICUM 175

pared seed-bed. Green panic seed can also be sown into the ashes after burning existing vegetation, as long as there is adequate moisture in the sub­ soil.I t can either besow n on the surface and rolled, or sown 1c m deep and covered. Seeding rates (2-7 kg) depend on quality of the seed. Sowing beneath a mulch of crop residues improves emergence and decreases the germination of weeds. Green panic grows well in association with: Stylo- santhesspp., CentrosemapubescensBenih., Chloris gayana Kunth, Medicago sativa L., annual Medi- cago spp., Desmodium intortum (Miller) Urban, D. uncinatum (Jacq.) DC. and Macroptilium atropur- pureum (DC.) Urban. Husbandry Green panic is sown as a sole crop on fertile soil and it responds well to fertilizer application of N, P and/or S on poor soils. When grown in association with a legume P and S fertiliz­ er will be beneficial on poor soils. Periods of heavy grazing are tolerated. Seed formation once every two years is necessary for long-term persistence. Green panic tolerates an occasional fire. It is suit­ able for continuous as well as rotational grazing. Diseases and pests Green panic has no serious diseases or pests. / Harvesting Green panic can be harvested by grazing or cutting to 5 cm above ground level 6 Panicum maximum Jacq. var. trichoglume Robijns weeks after emergence. For hay, green panic is - 1,habit leafy plant; 2, inflorescence. usually cut at the beginning ofth e flowering stage. Yield Dry matter yields ranging from 5 t/ha per cool winters, in contrast to guinea grass, which is year (without fertilizer) to 35 t/ha per year with adapted to humid tropical climates. However, complete fertilizer have been recorded for green green panic has also shown good performance in panic. It was one of the highest-yielding grasses northern Sumatra and the Philippines. Optimal tested in northern Sumatra. annual rainfall isbetwee n 560an d 1800mm . Green Seed yield varies from 30-200 kg/ha. Liveweight panic is moderately drought-tolerant. It does not gains of 140-180 kg/ha were recorded in south­ tolerate flooding for more than two weeks. Opti­ eastern Queensland. In northern Queensland, Jer­ mal temperature for growth is 30°C . It is less frost- sey cows and Friesian cows produced respectively tolerant than Rhodes grass (Chloris gayana 2500 kg and 4200 kg milk per lactation on green Kunth). Frost kills top growth, but regrowth panic in association with Neonotonia wightii occurs in spring. Green panic is a quantitative (Wight & Arnott) Lackey 'Tinaroo'. In Hawaii short-day species. It tolerates shade and even green panic in combination with Leucaena leucoce- grows right up to the trunks of trees and shrubs. phala (Lamk) de Wit produced nearly 101mil k and Green panic prefers a slightly acid to neutral soil 400k g of beef per ha/year under irrigation. (pH 6.5-7.0), but tolerates acid or alkaline soils Genetic resources and breeding Green panic (pH 5.0-8.0). It does best on deep, very fertile is a pseudogamous apomictic grass with limited loams, but also performs well on sandy loams of sexuality. Germplasm of green panic is held at reasonable fertility. Deep sands are unsuitable. ATFGRC (CSIRO, Australia). There are no known Green panic has an intermediate tolerance of high breeding programmes with green panic. levels of Mn in the soil. Prospects Green panic is a useful pasture grass Propagation and planting Dehulling of the for sub-humid subtropical regions and probably of seed increases the percentage and rate of germina­ little prospect in the humid tropics because of the tion because it removes the germination inhibitors availability of other species. in the bracts. Seeds are small and need a well pre­ Literature |1 Cowan, R.T., Byford, I.J.R. & 176 FORAGES

Stobbs, T.H., 1975.Effect s of stocking rates and en­ ergy supplementation on milk production from tropical grass-legume pastures. Australian Jour­ nal of Experimental Agriculture and Animal Hus­ bandry 15:740-746 . \2\ Delaney, N.E., 1975. Green panic iswidel y accepted. Queensland Agricultural Journal 101: 729-735. !3IPlucknett , D.L. 1970. Pro­ ductivity of tropical pastures in Hawaii. Proceed­ ings 11th International Grassland Congress, Surfers Paradise, Australia, pp. A38-A49. I4| Sil- vey, M.W. & Jones, R.M., 1990. Permanent pas­ tures on a brigalow soil: changes in pasture yield and composition during the first five years. Trop­ ical Grasslands 24: 282-290. |5| Skerman, P.J. & Riveros, F., 1990.Tropica l grasses. FAO,Rome . pp. 266-274. L. 't Mannetje &S.M.M . Kersten

Panicum repens L.

Sp. PL ed. 2:8 7(1762) . GRAMINEAE 2n = 40 Synonyms Panicum convolutum P. Beauv. ex Sprengel (1825). Vernacular names Torpedo grass (Am). Panic rampant (Fr). Indonesia: benda laut (North Suma­ Panicum repens L. - 1, habit flowering plant; 2, tra), suket balungen (Javanese), ramput kuma- ligule; 3,spikelet in two views. ranting (Minang). Malaysia: kerunong padi, metu- bong, telur ikan. Philippines: luya-luyahan at the margins near the throat; leaf-blade linear- (Tagalog), luy-a-luy-a (Visaya), maralaya (Ilo- acuminate, 7-25 cm x 2-8 mm, flat or rolled when kano). Burma: myet-kha. Cambodia: smau phluk, dry, often stiff and pungent, ascending close to the chhlong. Thailand: ya-chanakat (central), ya- stem; ligule a shallow membrane, 0.5 mm high, khaemman (northeast), ya-onoi (northern). Viet­ fringed with whitish hairs.Inflorescenc e a narrow­ nam: co' cu'a gà, co' ông. ly oblong panicle, 5-20 cm long, sparsely to moder­ Origin and geographic distribution Torpedo ately branched, branches usually ascending; spik- grass is widespread in Indonesia, Malaysia, Thai­ elets narrowly elliptical, ca. 3 mm long, acute, land and other South-East Asian countries and it often tinged with purple; lower glume clasping the also occurs in wetter areas throughout the tropical base of the spikelet, ^ the length of the spikelet; and subtropical regions of the world. upper glume as long as the spikelet; lower floret Uses Torpedo grass is used as a forage for graz­ male, upper floret bisexual. Caryopsis glossy ing or cutting. It isregarde d as an excellent native white. forage in South-East Asia, being both nutritious Flowering starts 3-4 weeks after seedling emer­ and palatable. It is also used to control erosion on gence and continues throughout the year, but seed sandy soils, but it can be a serious weed in crops. production is poor. Rhizomes develop so fast that Rhizomes from sandy soils are sometimes dug up they give a dense sward within 5-6 months. and fed to cattle as they are very palatable. Ecology Torpedo grass can grow up to 2000 m Properties Nitrogen concentrations in torpedo altitude in the tropics on humid or marshy places, grass range from 0.7% to 3.8 %. on moist open or partially shaded meadows, in Botany A perennial grass with long, sharp paddy fields, along lagoons, canals and roadsides, pointed rhizomes and often also surface stolons; and on sandy soils on the coast. It is extrememly culms erect or decumbent, up to 120 cm tall, often tolerant of acid soils. It cannot stand permanently from a knotty base. Leaf-sheath 4-7 cm long, hairy flooded conditions. PASPALUM 177

Agronomy Torpedo grass is established by seed (Madura). Malaysia: rumput kerbau, rumput ala or rhizomes. A seeding rate of about 10 kg/ha is negri. Philippines: kulape (Tagalog), kauat-kauat used in some regions, sowing seed on the soil sur­ (Visaya), kalo-kawayan (Ilokano). Thailand: ya- face with only a light covering. Ploughing nomnon, ya-hep (southern). Vietnam: co san cap. enhances subsequent spread. It is very palatable Origin and geographic distribution Original­ and extremely tolerant of heavy grazing. Frequent ly from the American tropics, P. conjugatum is and close cutting or grazing is recommended to naturalized throughout South-East Asia and in maintain plants in a leafy condition. It shows little many tropical countries of the world. It is abun­ response to liming in acid peat soils. Irrigated pas­ dant in Indonesia, the Philippines and the Pacific tures, fertilized with 100 kg/ha of N at each of 5 Islands. cuts, have yielded 100 t/ha per year of green mat­ Uses P. conjugatum is used as a forage for graz­ ter. On abandoned rice fields it has yielded 60 t/ha ing or in cut-and-carry systems, and is rated as per year of green matter. On peat soils, it can yield very important as a natural pasture grass in coco­ up to 11t/h a of DM per year. nut plantations. It is occasionally used as a lawn As a weed it is difficult to control because of the grass and is also regarded as an important weed long-living and deep penetrating rhizomes; it may in rice and plantation crops. spread into improved pastures competing with and The Iban ofBorne o use leaf decoctions in the treat­ choking out desirable species. ment of wounds and sores, and in the Sepik area Genetic resources and breeding Observations of Papua New Guinea crushed spikelets are used suggest that torpedo grass isa variabl e species, but for the same purpose. it is unlikely that any substantial germplasm col­ Properties At the pre-flowering stage, the N lections are being maintained. concentration in P. conjugatum ranges from Prospects Torpedo grass is a useful local forage 1-2.2%. It is stated that only the young stage of grass. Selection should aim at obtaining acces­ the grass is suitable for grazing since- the fruits sions with more leaf, less stem, and less agressive tend to stick in the throats of livestock and choke rhizomes. them. The presence of a haemostatic glucoside, Literature 1 Bor, N.L., 1960. The grasses of which reduced the time for blood clotting by 50% , Burma, Ceylon, India and Pakistan. Pergamon has been reported for this species. Wet fruits may Press, London, p. 330. |2|Manidool , C, 1989. Natu­ become very irritating as they easily stick to one's ral grassland and native grasses of Thailand [in legs and clothing. Thai]. Technical Bulletin No 1301-26-32. Divi­ Botany A vigorous, creeping perennial with sion of Animal Nutrition, Department of Live­ long stolons, rooting at nodes, with culms ascen­ stock Development, Bangkok, p. 22. |3| Manidool, ding to erect, 40-80(-100) cm tall, branching, C, Vagama, V.& Poositikul , A., 1985. Digestibility solid, slightly compressed. Leaf-sheath strongly ofPanicu m repens from organic soils area in Pikul- compressed, usually 30-50 mm long, ciliate on the thong village [in Thai]. Annual Eeport on Animal margins; ligule collar-shaped, about 1 mm long; Production Research. Department of Livestock leaf-blade linear or lanceolate-acuminate, 8-20 cm Development, Bangkok. \4\Mehra , K.L. & Fach- x 5-12 mm, glabrous to sparsely pubescent. Inflor­ rurozi, Z., 1985. Indonesian economic plant escence well exserted with two or occasionally resources: forage crops. Lembaga Biologi Nasio- three diverging racemes, 7-16 cm long; spikelets nal - LIPI, Bogor, No 31. p. 31. [5. Skerman, P.J. solitary, imbricate, flattened ovate, up to 2 mm & Riveros, F., 1990. Tropical grasses. FAO, Rome, long, with long hairs on the margins; lower glume pp. 543-545. absent, upper glume with a fringe of long hairs (1 C. Manidool mm) along its margin. Caryopsis broadly ovoid, plano-convex, about 1m m long, dark brown. The germination percentage of P. conjugatum seed Paspalum conjugatum Bergius is usually low. Flowering commences 4-5 weeks after seedling emergence and it continues to Act. Helv. Phys. Math. 7:129, t.8 (1762). flower year round. New shoots develop at every GRAMINBAE rooted node. In = 20,40 ,8 0 Ecology P. conjugatum grows from near sea- Vernacular names Buffalo grass, carabao level up to 1700 m altitude in open to moderately grass, sour paspalum (En). Indonesia: jampang shaded places. It is adapted to humid climates. It pahit (Sundanese), paitan (Javanese), klamaran is found under plantation crops and also along 178 FORAGES

such as improving DM yield and seed production, is desirable. Literature ;1| de Koning, R. & Sosef, M.S.M., 1985. The Malesian species of Paspalum (Grami- neae). Blumea 30:279-318. |2|Gilliland , H.B., Holt- turn, R.E. &Bor , N.L., 1971. Grasses of Malaya. In: Burkill, H.M. (Editor): Flora of Malaya. Vol. 3. Government Printing Office, Singapore, pp. 180-182. |3i Holm, J., 1971. Feeding tables. Nutri­ tion Laboratory of Thai-German Dairy Project, Livestock Breeding Station, Chiangmai, Thai­ land. |4|Manidool , C, 1989.Natura l grassland and native grasses of Thailand [in Thai]. Technical Bulletin No 1301-26-32. Division of Animal Nu­ trition, Department of Livestock Development, Bangkok. 45 pp. !5

Paspalum dilatatum Poiret

Lamk, Tabl. Encycl. 5:3 5(1804) . Paspalum conjugatum Bergius - 1,habit flowering GRAMINEAE plant; 2, ligule; 3, spikelet. 2n = 40 (sexual, yellow-anthered biotype), 45 (interspecific hybrid), 50 (apomict, common culti- stream banks, roadsides and in disturbed areas on var), 70 (hybrid) a variety of soils, often growing gregariously. Synonyms Paspalum platense Sprengel (1825), Agronomy P. conjugatum is propagated from P. ovatum Trinius (1826), Digitaria dilatata prostate culms, using 2-3 nodes per cutting. Close (Poiret) Coste (1906). grazing is required to keep it in palatable stage as Vernacular names Dallis grass, Australian palatability declines rapidly after flowering. It is grass, paspalum (Australia)(En). Paspalum dilaté eaten more readily by water buffaloes than by cat­ (Fr). Indonesia: rumput australi. Philippines: tle. In a mixture with Imperata cylindrica (L.) lawa-lawa (Tagalog), halanaw (Bikol), sakata Eaeuschel, it will dominate the latter ifth e pasture (Visaya). Thailand: ya-daenlit. Vietnam: co' san is grazed heavily. Close cutting and heavy grazing dep. are recommended since it istoleran t of defoliation, Origin and geographic distribution Dallis and because this prevents seed head maturity, grass is native to the humid subtropics of Brazil, resulting in higher quality regrowth. Cut feed can Argentina and Uruguay. Now it is widely distri­ be conserved as hay. Under a coconut plantation buted in coastal subtropical Australia, the south­ without any fertilizer a yield of 19 t/ha of green eastern part of the United States, India, some Afri­ material has been obtained. Yield increased to 30 can countries, Madagascar, Hawaii, Fiji and t/ha of green material following application of Malesia. 15-15-15 N:P:K fertilizer at 310 kg/ha. Uses Dallis grass is used in pastures for grazing, Genetic resources and breeding It is unlikely hay and silage-making. It is better suited for graz­ that substantial germplasm collections are being ing than for cutting, because of its tendency to maintained. lodge. It can also be sown into the stubble after Prospects Because of the wide distribution of a rice crop. Dallis grass gives excellent protection P. conjugatum, research on agronomic aspects, against erosion. PASPALUM 179

Properties Dallis grass is very palatable at framing the spikelet; lower floret with sterile early stages of growth, but this decreases rapidly lemma; upper floret bisexual. Caryopsis suborbi- with increasing age. The N concentration of green cular, plano-convex, up to ca. 2m m long. material ranges from 0.8-3.8% of the DM and the Growth and development Early growth is DM digestibility between 50-63%, both depend­ slow, but once established growth is vigorous and ing on plant maturity, soil fertility and environ­ many leaves are produced in a short time. Dallis mental conditions. There are 570-750 seeds/g. grass has its fastest growth rate early in the grow­ Description A leafy, tufted perennial with clus­ ing season, but growth slows down during mid­ tered stems arising from very short, creeping rhi­ summer and autumn. During the cool period in zomes, forming a sod under grazing, rooting up to subtropical regions and at higher altitudes the 1.2 mdeep .Ste m ascending to erect, 50-150 cm tall, plants are dormant. Flowering occurs throughout rooting at the nodes when in contact with the soil. the growing period and seeds scatter as soon as Leaf-sheath slightly keeled, basal ones often they are ripe. With declining soil fertility, the pilose; ligule bluntly triangular, up to 6 mm long; vigour of Dallis grass declines and it is then often leaf-blade linear, flat, (5-)12-16(-39) cm x 3-13 invaded by Axonopus affinis Chase. mm, usually with some up to 6 mm long hairs at Other botanical information P. dilatatum can the base of the ligule, otherwise glabrous. The be divided into two varieties, var. dilatatum and inflorescence consists of 3-5(-9) alternate var. pauciciliatum Parodi. The latter has In = 40 racemes, the lowest 5-13 cm long, the others grad­ chromosomes and is more prostrate. It has slightly ually shorter upwards; spikelets paired, usually smaller and less hairy spikelets, more spikelets per imbricate, in rows along one side of a flattened raceme and its lemma and palea have three veins axis, ovate, 3-4 mm long; lower glume absent; compared with nine in var. dilatatum. This variety upper glume as long as the spikelet, with a fringe is better adapted to light and poorly drained soils of white, ca. 2 mm long hairs along the margin than common Dallis grass. Important cultivars of var. dilatatum are 'Raki' from New Zealand, 'Charu' from Uruguay and 'Natsugumo' from Japan. Ecology Dallis grass is adapted to permanently humid subtropical climates (10-12 humid months) with over 1000 mm annual rainfall mainly during the warm season. Optimum temperatures for leaf growth, tillering and flowering are 30°C, 27°C and 22.5° C respectively. Seed production is inhibited at temperatures below 13°C. Dallis grass grows on lowlands as well as at altitudes up to 2000 m. It is one of the most frost tolerant subtropical grasses. It is a long-day plant, with an optimal day- length of 14hours . Dallis grass has little shade tol­ erance. It requires fertile soils and performs best on alluvial soils or basaltic clays. Optimum pH(H20) is between 5.5 and 7.0. Dallis grass does not tolerate salinity. Propagation and planting Dallis grass seed has some post-harvesting dormancy which can be broken by removal ofth e glumes. It is usually sown at a seeding rate of 6-10 kg/ha, either broadcast on the surface with a light covering, or drilled to a depth of 1-1.5 cm,preferabl y just before the start ofth e rainy season. After a rice crop it can be sown into the stubble without seed-bed preparation. Because Dallis grass is a slow starter, it is often established in mixtures with rapidly growing Paspalum dilatatum Poiret - 1, habit leafy plant; grasses such as Chloris gayana Kunth, which can 2, ligule; 3,inflorescence; 4, spikelet. compete better with weeds. Other species growing 180 FORAGES in association with Dallis grass are Trifolium its ability to combine with Trifolium repens and repens L., T. semipilosum Fresen. and Vigna par­ to withstand heavy grazing, Dallis grass is a desir­ ken Baker. able species, particularly in areas where it is Husbandry Dallis grass tolerates high stocking naturalized. However, it requires careful manage­ rates and frequent defoliation, which also pre­ ment because of its susceptibility to ergot. vents it from flowering and consequently ergot in­ Literature ill Bogdan, A.V., 1977.Tropica l pas­ fection, which can cause poisoning. It can be ture and fodder plants. Longman, London and New grazed continuously or rotationally. Lenient graz­ York. pp. 197-202. |2, de Koning, R. & Sosef, ing, not shorter than 5-8 cm, results in up to three M.S.M., 1985. Malesian species of Paspalum. Blu- times more forage than heavy grazing. Dallis grass mea 30:293-294 . 3;Shaw , N.H., Elich, T.W., Hay- responds well to fertilization. dock, K.P. & Waite, R.B., 1965. A comparison of Diseases and pests Dallis grass isver y suscepti­ seventeen introductions of Paspalum species and ble to ergot (Clavicepspaspali). This fungus infects naturalised P.dilatatu m under cutting at Samford, the inflorescence and excretes a substance which south-eastern Queensland. Australian Journal of is toxic to cattle and can occasionally cause abor­ Experimental Agriculture and Animal Husbandry tion. Ergot is also the most important factor affect­ 5: 423-432. |4 Skerman, P.J. & Riveros, F., 1990. ing seed quality. Other diseases are anthracnose Tropical grasses. FAO, Rome. pp. 579-584. 5;Wat ­ (Colletotrichum graminicola) and leaf blight (Hel- son, V.H. &Burson , B.L., 1985.Bahi a grass, Carpet minthosporium micropus), but these are generally grass and Dallis grass. In: Heath, M.E., Barnes, unimportant. Dallis grass pastures are sometimes R.F. &Metcalf , D.S.(Editors) : Forages, the science invaded by white-grubs (Lepidiota caudata and of grassland agriculture. Mississippi, United Rhopaea paspali), which destroy the roots and States, pp. 255-262. reduce productivity. L. 't Mannetje &S.M.M . Kersten Harvesting For hay or silage making Dallis grass can be cut every 4-6 weeks, preferably before the grass is 30 cm tall and before flowering. Paspalum distichum L. Seed is best harvested about 21 days after flower­ ing when inflorescences are brown, but before seed Syst. Nat. ed. 10,2 :85 5(1759) . shattering becomes severe. GRAMINBAE Yield Total DM yield ranges from 3-15 t/ha, 2n = 40, 48, 60 depending on soilfertilit y or fertilization. High an­ Synonyms Paspalum paspalodes Scribner imal production can be obtained from Dallis grass (1894). under good management. In southern Australia Vernacular names Ginger grass, knot grass, stocking rates of 25 sheep/ha can be maintained. (salt-)water couch grass (En). Indonesia: rumput With irrigation, optimal fertilization and by italia, asinan (Javanese), lamhani (Sundanese). including winter-growing species, milk yields of Philippines: malit-kalabaw (Tagalog), bakbaba 10000 kg/ha per year have been recorded. (Ilokano), gagayut (Wfumo). Thailand: ya-sakat- Seed yields are variable: 90-500 kg/ha. namkhem (northern). Vietnam: co' san dôi. Handling after harvest Seed should be dried Origin and geographic distribution A com­ immediately after harvest at 60° C or spread in a mon weed of the tropics, subtropics and warm tem­ thin layer and dried to 7-10% moisture content. perate regions of unknown origin. In Malesia P. Genetic resources and breeding A germplasm distichum probably has been introduced. collection of Dallis grass is held at ATFGRC Uses Considered as a valuable pasture grass on (CSIRO, Australia). Seed is produced in Australia, alluvial flats, P. distichum is used as a fodder and the United States and Uruguay. Dallis grass is is relished by water buffaloes. On stream banks it mainly apomictic by apospory and pseudogamy. is useful as a soil binder. In direct-seeded rice it The sexual type is less common. Crossing an apo- is a serious weed that grows vigorously under mict with a female sexual species gives an interspe­ favourable conditions. cific hybrid with 2n = 50. The first generation Properties Nitrogen concentrations of up to reproduces sexually and seed set increases pro­ 1.5% have been measured, with Ca and P concen­ gressively. trations of 0.22% and 0.14% respectively. It is Breeding work is being done to increase ergot and moderately palatable. anthracnose resistance. Botany A stoloniferous perennial, sometimes Prospects Due to its palatability, productivity, tufted, with leafy stolons rooting at the nodes, and PASPALUM 181

swamps, in polluted shallow water, along irriga­ tion ditches, etc.,u p to 1700m altitude . It can stand high salinity and tolerates waterlogged conditions and periodic flooding in salt swamps and by tidal waters. Agronomy P. distichum can be established by seed or stolons. It requires minimum land prepara­ tion and is easily established by placing stolons in holes made in moist soil. The plant remains green throughout the year if growing in water. Light grazing with long rest periods of 80-90 days is rec­ ommended since the rooted stolons are sometimes floating. Harrowing of dry surface soils will later help the spreading stolons to root and so form a denser sward. It can be cut and fed to animals, but normally villagers let their animals graze it. Observations suggest that it does not give high DM yields. It can be made into hay but is not suit­ able for silage. Genetic resources and breeding It is unlikely that substantial germplasm collections are being maintained. Prospects P. distichum should be included when selecting for species that are tolerant to saline soils. / Literature l! de Koning, R. & Sosef, M.S.M., 1985. The Malesian species of Paspalum (Grami- neae). Blumea 30: 295-297. 2! Manidool, C, 1989. Paspalum distichum L. - 1, habit flowering plant; Natural grassland and native grasses of Thailand 2, ligule; S,spikelet in two views. [inThai] . Technical Bulletin No 1301-26-32. Divi­ sion of Animal Nutrition, Department of Live­ ascending or erect culms 10-60(-100) cm long. stock Development, Bangkok, pp. 26-32. 3' Sker- Leaf-sheath usually hairy along the margins; man, P.J. & Riveros, F., 1990. Tropical grasses. ligule truncate or slightly lobed, 0.5-3 mm long; FAO, Rome. pp. 565-568. 4i Soerjani, M., Koster- leaf-blade linear, 6-10(-20) cm x 3-5(-9) mm, at­ mans, A.J.G.H. & Tjitrosoepomo, G. (Editors), tenuate-acute, usually with some white hairs at 1987. Weeds of rice in Indonesia. Balai Pustaka, the immediate base of the ligule, otherwise gla­ Jakarta, pp. 460-461. brous. Inflorescence composed of 2 subopposite C. Manidool racemes, each 2-8 cm long; spikelets solitary, the upper ones usually imbricate, oblongoid, 3-4 mm long, with acute apex; lower glume often reduced Paspalum notatum Flueggé or absent; upper glume pubescent, sometimes leathery. Caryopsis ovoid, plano-convex, ca. 1.5 Gram, monogr., Paspalum: 106(1810) . mm long. GRAMINEAE Flowering is throughout the year. Seeds appear to In = 20 (sexual), 30,4 0 (apomicts) have a period of dormancy; they germinate best at Vernacular names Bahia grass, Pensacola 20-30°C. From India obligate apomixis has been Bahia grass (En, Am). Indonesia: rumput pencasi- reported for P. distichum. lan (Timor). Thailand: ya-bahia. Vietnam: co san The species closely resembles P. vaginatum Swartz dâû. with which it is often confused; P. vaginatum does Origin and geographic distribution Bahia not extend into temperate regions, has at least two grass is indigenous to the Americas, particularly strictly opposite racemes in the inflorescence, and in southern Brazil. It is now widely distributed in its upper glumes are glabrous. the southern United States, Central and South Ecology P. distichum occurs in wet marshlands, America, and occurs in restricted areas in Austra- 182 FORAGES

lia, Asia, and Africa, extending into and even linear, 5-20(-50) cm x 2-10 mm, sometimes hairy beyond the subtropics. on margins; ligule membraneous, 0.4 mm tall, Uses Bahia grass is primarily used as forage for hairy from behind. Inflorescence composed of 2 grazing, but also to protect slopes and terraces (rarely up to 5) terminal racemes, each 4-12 cm from soil erosion. It is also used as a lawn grass long; spikelets solitary in 2 rows on a narrow and provides suitable material for compost or rachis and broadly ellipsoid, 2-4 mm long; florets mulch. It is sometimes regarded as a 'weed' grass 2,th e lower reduced to an empty lemma. Caryopsis when it invades pastures of preferred species. ovoid, 3m m long, flattened on one side, yellowish- Properties In vitro digestibilities of 65-70% green, glossy. have been measured in young growth, falling to Bahia grass is a polymorphic species in which 40-50% in older material. Similar declines have three varieties have been distinguished: var. nota­ been measured in N concentrations, with old mate­ tum, 2n = 40, tetraploid apomict, with short nar­ rial containing only 0.5% N. Palatability declines row leaves, mainly occurring in southern United markedly once seeding occurs or leaves mature. States; var. latiflorum Doell., 2n = 40, tetraploid, The digestibility of 6-week-old material in Queens­ usually apomict, vigorous, widespread throughout land (Australia), was slightly higher (60%) under Central and South America; and var. saureae Pa- 50% shade than it was in full sunlight (55 %). rodi, 2n = 20, sexual diploid, long narrow leaves, There are 300-550 seeds/g. endemic to northern Argentina. There are several Botany A low-growing creeping perennial with cultivars, the best known being 'Pensacola', stolons and stout rhizomes. The tough stolons selected from var. saureae in the United States. close to the ground have short internodes and root Other selections have been released as cultivars freely from the nodes, forming a dense sod. Culms in the United States, Brazil and Australia, and cul- erect or geniculate, up to 75 cm tall. Leaf-blade tivar 'Tifthi-1' was bred in the United States. Several studies have been made into the fixation ofatmospheri c Nb ymicro-organisms , such as Azo- tobacter paspali, in association with Bahia grass. Most reports indicate fixation of only small amounts of N, up to 20 kg/ha per year, but one reported fixation of 90 kg/ha per year. Ecology Bahia grass is a vigorous aggressive grass that spreads vegetatively and by seed. Viable seed is spread through dissemination in faeces. It prefers sub-humid to humid subtropical climates with annual rainfall ranging from 800 to 2000 mm. The optimum temperature ranges for growth (25-30°C maximum and 20°C mean) are slightly lower than those of grasses best suited to the low­ land humid tropics. Top growth is killed by frost but established plants can tolerate temperatures down to -10°C. It is very tolerant of shade and 35% higher yield has been measured under 50% shade than in full sunlight. It is best suited to sandy or light-textured soils but can grow on a wide range of soil types. It has good drought toler­ ance and has also been known to survive 36 days of flooding, and has some salt tolerance. Agronomy Bahia grass can be established from seed or vegetatively, from pieces ofrhizome s or sto­ lons planted closely at spacings of 15-25 cm. Hand- harvested seed has a high proportion of hard-see- dedness so the germination percentage is initially low, but improves progressively with up to 3 years Paspalum notatum Flueggé 1, habit flowering of storage. Hammer milling of seed or treatment plant; 2, ligule; 3, spikelet. with sulphuric acid also improves germination. PASPALUM 183

Sowing rates of 2-5 kg/ha are recommended. For Literature |1| Agata, W., 1985. Studies on dry optimum establishment, seed is sown into a fully matter production of Bahia grass (Paspalum nota- prepared seed-bed to a depth of less than 1cm , fol­ tum) sward. I. Characteristics of dry matter pro­ lowed by rolling. duction during the regrowth period. In: Proceed­ Bahia grass isver y persistent and competitive and, ings of the XV International Grassland Congress, given very fertile soil, can be reasonably produc­ August 24-31, 1985, Kyoto, Japan. Science Coun­ tive. Frequent defoliation close to ground level is cil of Japan and Japanese Society of Grassland not only tolerated but is desirable to keep the Science, Nishinasuno, Japan, pp. 1235-1236. 21 sward leafy and acceptable to animals. It is more Blue, W.G., 1988. Response of Pensacola Bahia difficult to maintain legumes with Bahia grass grass on a Florida Spodosol to nitrogen sources than with most tropical grasses, partly because it and times of application. Proceedings Soil & Crop is so competitive and partly because Bahia grass Sciences Society ofFlorid a 47:139-142. ;3. Bogdan, pastures have to be closely and regularly grazed. A.V., 1977. Tropical pasture and fodder plants. Good results have been achieved with prostrate Longman, London, pp. 205-212. \4\ Marousky, F.J. legumes, especially with white clover {Trifolium & West, S.H., 1988.Germinatio n of Bahia grass in repens L.) in the subtropics, but also with Vigna response to temperature and scarification. Jour­ parkeri Baker and perennial Arachis spp. nal of the American Society for Horticultural Cultivars differ in their susceptibility to a stinging Science 113: 845-849. ;5; Shaw, N.H., Elich, T.W., nematode (Belonolaimus longicaudatus), but Haydock, K.P. &Waite , R.B., 1965. A comparison Bahia grass is resistent to root-knot nematodes of seventeen introductions of Paspalum species and has been used in rotation with crops suscepti­ and naturalized P.dilatatu m under cutting at Sam- blet o these nematodes to reduce populations. Seed ford, south-eastern Queensland. Australian Jour­ yields can be reduced by Paspalum ergot (Claui- nal of Experimental Agriculture and Animal Hus­ cepspaspali), although 'Pensacola' is resistant. bandry 5:423-432 . * Skerman, P.J. &Riveros , F., Herbage yields of Bahia grass are not particularly 1990.Tropica l grasses. FAO, Rome. ppf571-575. high. Yields of 3-8 t/ha of DM can be expected B.B. Baki, LB. Ipor &C.P . Chen given moderate fertility and a suitable climate, with extreme yields ranging from 1t o 20 t/ha, the latter from heavily fertilized and irrigated swards. Paspalum plicatulum Michaux In the United States, Bahia grass pastures ferti­ lized with N, usually 100-200 kg/ha, produce Fl. Bor. Amer. 1:45(1803). 400-600 kg/ha of liveweight gain per year and can GRAMINEAE carry about 5 head per hectare. In comparative 2« = 20 (diploid), 30,40 ,6 0 studies, gains per head are usually slightly lower Synonyms Paspalum undulatum Poiret (1804), than recorded from pastures of Cynodon spp. Bahia Panicumplicatulum (Michaux) Kuntze (1898). grass is usually harvested by grazing animals. It Vernacular names Plicatulum, brown seed is not suited for making hay or silage as yields are paspalum, brown top paspalum (En, Am). Herbe à low when it is in a young leafy stage and as quality cheval (Fr).Thailand : ya-phlikhathiulum. is poor if it is left for a longer growth period to Origin and geographic distribution Plicatu­ achieve higher yields. It is a heavy seeder and seed lum occurs naturally in South and Central Amer­ yields of 100-350 kg/ha have been reported. ica. It is distributed throughout the world from Genetic resources and breeding A major col­ Africa (Kenya, Ivory Coast) to Australia, the Uni­ lection ismaintaine d by the USDA at Tifton, Geor­ ted States, (Fiji) and South-East Asia gia. Hybrid lines of Bahia grass have been pro­ (including the Philippines, Thailand, Indonesia, duced in the United States, but there are no Malaysia and China). current breeding programmes. Uses Plicatulum is used as a pasture grass for Prospects The potential for Bahia grass is grazing and silage or hay making, as an under- South-East Asia has yet to be assessed. Although storey in coconut plantations and in ley pastures not ideally adapted to the lowland tropics, it has after harvesting a paddy rice crop. potential for use in plantation crops because of its Properties Nitrogen concentration ranges from tolerance to moderate shade. Where it is well 0.8-2.0% of the DM and DM digestibility from adapted, the benefits of its persistence and compe­ 39-50% of whole plant material and 50-70% of titiveness must be balanced against its tendency green leaves. Plicatulum is a palatable grass with to have lower yield, quality and acceptability. a high nutritive value at the young leafy stage of 184 FORAGES

growth. 'Hartley' maintains a high nutritive value od at the end of the wet season. Seed is ready for after frosting, in contrast to 'Rodds Bay'.Ther e are harvesting 21day s after panicle emergence. Seed­ 750-1200 seeds/g. lings have a juvenile period of 50-60 days in which Description A tufted, decumbent perennial, they will not flower. growing to a height of 1.2 m, sometimes with a Other botanical information Three cultivars short rhizome. Leaf-sheath keeled, usually gla­ have been released in Queensland, Australia, viz. brous; ligule membranous, 1-3 mm tall; leaf-blade 'Rodds Bay', with narrower leaves than the other folded at the base, linear 10-50(-85) cm x two cultivars; 'Hartley', more prostrate growing 3-7(-10) mm, often slightly hairy on the upper sur­ and leaves with wavy margins towards the base; face towards the base, in the tropics often hairy 'Bryan' is intermediate to these two cultivars and on both surfaces. Inflorescence a panicle com­ is the hairiest one. posed of (5-)10-13(-19) racemes 2-10 cm long; Ecology Plicatulum is adapted to tropical and spikelets in pairs (one of the pair sometimes unde­ subtropical climates with over 750 mm of annual veloped), obovoid-ellipsoidal, 2-3 mm x 1.5-2 rainfall. Plicatulum survives dry seasons of 5-6 mm; lower glumes absent; lower lemma with short months. 'Rodd's Bay' has the best drought toler­ transverse wrinkles just inside the slightly raised ance. Temporary waterlogging is also tolerated. margin. Caryopsis as large as the spikelet, dark- Plicatulum does not grow at temperatures below brown, shiny. 10°C and top growth is killed by frost, although Growth and development Plicatulum has a regrowth occurs in spring. Optimal temperature slow seedling growth, but rapid growth rate once for germination and growth is between 20-35°C. established. It is a short-day plant, with a critical The grass can grow on a wide range of soils; it is photoperiod of 13hour s and a short flowering peri- very tolerant of low pH and of high Al concentra­ tions. Plicatulum is tolerant of fire, but it is not adapted to shade. Propagation and planting Plicatulum is pro­ pagated normally by seed, but can also be planted vegetatively. The seed can remain dormant for sev­ eral months after maturity. This is caused by per­ meability restrictions imposed by the lemma and the palea. Dormancy can be broken by chilling at 7°C for 30 days. Full land preparation to a fine seed-bed is necessary and the recommended seed­ ing rate is 2-3 kg/ha. The seed can be broadcast on the surface and covered by rolling, or drilled to a depth of 1-1.5 cm.. Plicatulum can be grown in association with Macroptilium atropurpureum (DC.) Urban, Stylosanthes spp., Desmodium intor- tum (Miller) Urban and Trifolium repens L. Husbandry Plicatulum responds well to N fer­ tilizer, but it can also grow and persist on poor soils. Plicatulum can be grazed continuously or rotationally. Diseases and pests Plicatulum isver y resistant to ergot and no other diseases or pests are known to affect it. Yield Dry matter yield varies from 8.5-24 t/ha per year. With irrigation and complete fertilizer (NPK) and cutting intervals of five weeks, DM yield per cut of2. 5t/h a can be obtained. In Fiji DM yield under coconut was 9.5 t/ha per year, while in Malaysia the yield under oil palm Paspalum plicatulum Michaux- 1,habit f lowering was negligible. Seed harvest yields 200-300 kg/ha. plant; 2, ligule; 3, inflorescence; 4, two views of a Annual liveweight gains of beef cattle of up to spikelet. 250-300 kg/ha have been measured in the sub- PASPALUM 185

humid subtropics in Queensland. Origin and geographic distribution P. scrobi­ Genetic resources and breeding Plicatulum is culatum occurs throughout the Old World tropics an aposporous apomict (2n = 40), or sexual (2n = and is occasionally cultivated elsewhere. In India 20). A germplasm collection of plicatulum is main­ it is often also cultivated. In South-East Asia it is tained at ATFGRC (CSIRO, Australia). Seed is a common grass. also produced in Queensland. There are no known Uses P. scrobiculatum provides useful forage but breeding programmes with plicatulum. isals o regarded as a weed of annual and plantation Prospects Plicatulum has good prospects for crops. Occasionally it is used as compost or mulch. use in open pastures in subtropical and tropical In India it is cultivated for the grain. regions of South-East Asia because of its persis­ Properties Nitrogen concentrations of 1-5% tence even on infertile soils and its ability to com­ during the growing season and below 0.5% in stan- bine with legumes. dover seed have been measured. It is a very palat­ Literature 111 Bisset, W.J., 1975. Plicatulum able grass except when it is frosted. The cultivars finds a place in coastal pastures. Queensland Agri­ grown for grain in India are said to contain com­ cultural Journal 101: 603-608. 2, Bogdan, A.V., pounds in the grain which are toxic to man and 1977. Tropical pasture and fodder plants. Long­ to animals. The fungi Sorosporium paspali and man, London and New York. pp. 213-214. |3: Uredo paspali-scrobiculari are almost invariably Bryan, W.W. &Shaw , N.H., 1964. Paspalum plica­ present in the outer husks of the grain and are tulum Michx. Two useful varieties for pastures in thought to be responsible for its toxicity. In Aus­ regions of summer rainfall. Australian Journal of tralia it has not been toxic to grazing animals. Experimental Agriculture and Animal Husbandry There are 300-600 seeds/g. 4:17-21. ;4I Skerman, P.J. &Riveros, F., 1990.Trop ­ Botany A tufted perennial, not stoloniferous, ical grasses. FAO, Rome. pp. 579-584. |5| 't Man­ with ascending to erect, sometimes branching netje, L., 1961. Key on vegetative characters of culms up to 135 cm tall. Leaf-sheath glabrous or Paspalum species. CSIRO, Australia, Division of hairy at the margins; ligule collar-shaped, up to 1.8 Tropical Pastures, Technical Paper No 1.pp . 7-8. mm long; leaf-blade linear, up to 53 cm x 2.5 cm, L. 't Mannetje &S.M.M . Kersten flat or folded length-wise, narrowed towards the base, hirsute at base, along the margins and above the ligule. Inflorescence composed of 1-14 alter­ Paspalum scrobiculatum L. nate or subopposite racemes each up to 15c m long; peduncle 0.5-2 mm in diameter; rachis flat, Mant. PI. 1:29 (1767). straight to slightly zigzag, winged,; spikelets soli­ GRAMINEAE tary, imbricate or not, usually suborbicular, 2-3 2re = 20, 40, 60 mm long; lower glume usually absent; upper glume Synonyms 5-13-nerved, sterile lemma 5-9-nerved. Caryopsis var. auriculatum: P. auriculatum Presl (1830), P. obovoid to ovoid, ca. 2m m long. zollingeri Steudel (1853); P. scrobiculatum is a polymorphic species, possibly var. bispicatum: P. commersonii Lamk (1791), P. an aggregate swarm of apomicts. Although the cartilagineum Presl (1830); variability is quite continuous, at present 5 varie­ var. horneri: P. horneri Henrard (1935); ties are distinguished with the following charac­ var. lanceolatum: P. lamprocaryon K. Schum. teristics: (1895). - var. scrobiculatum: culms 8-20(-55) cm long; Vernacular names cauline leaf-sheaths hairy at margins, rarely var. scrobiculatum: kodo or kodra millet (En). glabrous; leaf-blades (4-)7-20(-30) cm x Indonesia: rebu bawang, rumput kinangan (3-)5-7(-9) mm; peduncle 0.5-1 mm in diame­ (Java), suket krisik (Madura), ter; racemes 1-2, alternate or subopposite, 1.5-7 var. bispicatum: koda grass, scrobic (En). Indo­ cm long; spikelet 2.5-3.3 mm x 2.1-2.6 mm; nesia: jaringan, rumput ketih belalang (Indone­ upper glume with 7, 9 11 or 13 nearves; sterile sian), kumpai batu (Kalimantan). Malaysia: lemma with 7o r 9nerves . Distributed from India pala belang, rumput tulo santadok. Philippines: throughout Malesia to the Pacific. Its occur­ sabung-sabungan (Tagalog), bias-biasin rence in Africa and elsewhere is uncertain. (Visaya), tat-awwa (Bontok). Cambodia: smau - var. auriculatum (Presl) Merrill: culms up to 135 'ânnchien. Thailand: ya-sakhorbik (central). cm long; cauline leaf-sheaths glabrous; leaf- Vietnam: co' san tru'ng, co' tru'hg. blades 20-40(-53) cm x 10-16(-19)mm;pedun- 186 FORAGES

cm x 12-27 mm; racemes 2-4; spikelets 2-2.7 mm long. Distributed only in tropical Africa. P. scrobiculatum flowers freely, and in Malaysia it produces seed during 3-4 months per year. The seeds fall as they mature. P. orbiculare Forster f. much resembles P'. scrobicu­ latum, especially var. bispicatum. Its spikelets are usually broadly obovate with 3 or 5 discolorous nerves in the upper glumes and sterile lemmas. In var. bispicatum the spikelets are usually suborbi- cular and the nerves are 5 or 7, concolorous. In Australia cultivar 'Partridge' of P. scrobiculatum was released in 1966, but it is now rarely culti­ vated. Ecology In Indonesia and Malaysia, P. scrobicu­ latum is found mostly in open, wet cultivated areas, up to 1200 m altitude, and in upland and tidal rice fields. It is commonly found on disturbed sites and is a weed in cropping land. It is well adapted to waterlogged soils and can tolerate flooding, but has only limited drought tolerance. The optimum temperature for growth is 25-27°C. It is a sun-loving plant although it can tolerate and flourish with only 30-50% sunlight, as found in young rubber and oil palm plantations or black pepper farms. It isno t as salt tolerant asth e closely related P. vaginatum Swartz. It can tolerate poor soil fertility if there is little competition, but Paspalum scrobiculatum L. - 1, habit flowering prefers very fertile soils and responds well to fertil­ plant; 2, ligule; S,part of raceme. izer application. Agronomy P. scrobiculatum can be propagated cle 1-1.9 mm in diameter; racemes 3-8, alter­ by seed or by rooted tillers. Newly ripened seeds nate, 8-15.5 cm long; upper glume 7-nerved; ster­ are dormant and mechanical or acid scarification ile lemma with 7 or 9 nerves. Distributed from is desirable to reduce dormancy. It needs a very India throughout Malesia. Its occurrance in the fine seed-bed which is usually prepared by plough­ rest of the world is uncertain. ing, disking and harrowing. It is usually sown - var. bispicatum Hackel: culms 8-70(-130) cm through a cereal drill in rows 1.3 m apart on a well- long; cauline leaf-sheaths glabrous or hirsute; prepared seed-bed. leaf-blades 10-25(-42) cm x 4-9(-15) cm; P. scrobiculatum is susceptible to a root-knot peduncle 0.5-1.4 mm in diameter; racemes nematode (Meloidogyne incognita) and can be (l-)(2-6(-14), alternate, 1.5-9(-ll) cm long; attacked by the Paspalum ergot (Claviceps pas- spikelet 1.8-2.6 mm x 1.5-2.2 mm; upper glume pali). and sterile lemma 5 or 7 nerved. Distributed in It is a very palatable and highly digestible grass tropical Africa, Asia (including Malesia), the during the wet season and retains these character­ Pacific and Australia. istics later into maturity than most other grasses. - var. horneri (Henrard) Koning & Sosef: culms Consequently it could be used intensively during 15-80 cm long; cauline leaf-sheaths hirsute; leaf- the dry season when the nutritive value of other blades 3-6 mm wide; racemes 1-4, alternate or species is generally lower. Strategic intermittent subopposite;spikelets 1.7-2.5mm x 1.2-1.9mm; or rotational grazinghelp s to allow sufficient seed­ upper glume 5-nerved; sterile lemma 5 or ing for regeneration. Experience in Australia sug­ 7-nerved. Distribution is disjunct: in Assam and gests that it will not persist in permanent pastures, in Malesia only. where it is replaced by other perennial grasses. - var. lanceolatum Koning & Sosef: cauline leaf- It is usually harvested by grazing animals but is sheaths enveloping the nodes; leaf-blades 7-21 occasionally cut and fed as green fodder. Although PBNNISETUM 187

usually not as persistent as other perennial trop­ 0°-35°. In South-East Asia, kikuyu grass is re­ ical grasses, results from experiments with both stricted to elevated areas (> 1900 m) mainly in sheep and cattle suggest that it can result in good Papua New Guinea and the Philippines. animal production. Uses Kikuyu grass is used for many purposes Genetic resources and breeding There is con­ including lawns and recreational areas, pasture siderable variation within P. scrobiculatum and for ruminants under intensive grazing and to pre­ closely related Paspalum species which might vent soil erosion. It ispotentiall y a weed on arable even belong to the same species complex. It is un­ land and in irrigation channels. likely that any of the existing germplasm collec­ Properties Nitrogen concentrations range from tions adequately document this variability. There 1.8-4.3% depending on age and fertilizer applica­ are no breeding programmes on P. scrobiculatum. tion. Digestibility of DM varies from 73% (young Prospects The present opportunistic use of P. leaf) to 50% (mature or frosted material). With ad­ scrobiculatum will continue. Studies on the varia­ vancing maturity the rate of decline in nutritive bility within this species and on its agronomic value of kikuyu herbage is slower than that of requirements are necessary to determine its poten­ other tropical grasses. As kikuyu grass is adapted tial as a forage species. Australian experience sug­ to very fertile soils, the concentrations of P, K and gests it may be more suited for use as a short-term S are normally adequate for animal growth. On the forage rather than as a long-lived grass in perma­ other hand Na levels are often low (0.02-0.05%), nent pastures. indicating potential deficiencies for very produc­ Literature |1| de Koning, R. & Sosef, M.S.M., tive lactating ruminants. There are approximately 1985. The Malesian species of Paspalum (Grami- 400 seeds/g. neae). Blumea 30: 279-318. |2| Devendra, C, 1979. Description A perennial, stoloniferous and rhi- Malaysian feedingstuffs. Malaysian Agricultural zomatous grass with short culms, 8-15 cm tall, Research and Development Institute (MARDI), arising from long, prostrate runners, multi- Selangor, Malaysia. 145 pp. |3| Kitamura, M. & Nada, Y., 1986. Preliminary evaluation of 24 grasses introduced into sub-tropical Japan. Jour­ nal of Japanese Society of Grassland Science 32: 278-80. |4|Oram , R.N., 1990.Registe r of Australian herbage plant cultivars. CSIRO, Australia, p. 44. j5| Paltridge, T.B., 1955. Studies on sown pastures for southeastern Queensland. Bulletin No 274. CSIRO, Australia, pp. 15-51. |6| Skerman, P.J. & Riveros, F.,1990 .Tropica l grasses. FAO,Rome . pp. 585-589. \T\ 't Mannetje, L., 1961. Key on vegeta­ tive characters of Paspalum species. CSIRO, Aus­ tralia, Division of Tropical Crops and Pastures, Technical Paper No 1.pp . 6-7. B.B. Baki &LB . Ipor

Pennisetum clandestinum Höchst, ex Chiov.

Ann. Ist. Bot. Roma 8:41 , fig. 5(2 )(1903) . GRAMINEAE 2n = 36 Vernacular names Kikuyu grass (En). Thai­ land: ya-khikhuyu. Origin and geographic distribution Kikuyu grass occurs naturally across the elevated pla­ teaux of East and Central Africa, and has been dis­ Pennisetum clandestinum Höchst, ex Chiov. - 1, tributed to other and islands with (sub­ habit of basal plant part; 2, ligule; 3, inflorescence; tropical, humid climates, between latitudes 4, spikelet. 188 FORAGES branched and rooting at the nodes. Roots prolifer­ izes at sealevel. Kikuyu grass is sensitive to water ate densely in upper 0-15 cm of soil, from older supply; with increasing evaporative demand from rhizomes/stolons, becoming less dense towards the 2-5 mm/day growth is reduced and ceases at a growing point; in friable soils, roots can penetrate water potential of -100 kPa. Kikuyu grass is to 3 m. Leaves arise alternately from multi- adapted to free-draining lateritic red loam soils of branched stolons; leaf-sheath 1-2 cm long, pale moderate or good fertility, often with low;pH(H 20) green, eventually turning brown on older stolons, (< 5),an d is tolerant of high Al and Mn levels. densely hairy; ligule a rim ofshor t hairs; leaf-blade Propagation and planting Kikuyu grass estab­ linear, 1-15 cm x 1-5 mm, tightly folded when lishes vegetatively and from seed. In vegetative young, flattened when older with sparingly hairy propagation, rooted stolons can be dug or midrib. Inflorescence reduced to a cluster of 2-4 ploughed out mechanically, spread out and short spikelets each with fine bristles carried close pressed in to the ground. Kikuyu grass seed (1-2 to lateral runners, concealed within the upper­ kg/ha) is either broadcast or drilled into a pre­ most sheath; spikelets consist of two florets, pale pared seed-bed, usually with companion legumes. or translucent, one of which is usually fertile; sta­ Once introduced to a locality, kikuyu grass mens exserted on slender filaments for a short usually spreads naturally because it offers strong time, often asynchronously with the stigma. Car- competition with other species or because ingested yopsis develops near stem at ground level, dark seed is dispersed in dung by cattle. brown, ovoid, 2.5m m long, pointed. Husbandry After sowing, kikuyu grass forms a Growth and development In established dominant sward within 3-9 months, aided by fer­ swards,kikuy u grass growth iscyclical , dependent tilizer application (N and P) and mowing to reduce on temperature, moisture and fertility. Flowering weeds. It responds vigorously to N fertilizer, yield­ is indeterminate, being stimulated by defoliation ing 15-30 kg of DM per kg N applied during active or heavy grazing. After germination and emer­ growth under a cutting regime, although the res­ gence ofprimar y tillers, kikuyu grass seedlings de­ ponse will vary with soil fertility. It responds to velop a prostrate growth habit as stolons grow P and K if these are in short supply. To improve radially. feeding value and to maintain legumes, the grass Other botanical information Early collec­ needs and responds to intensive grazing which can tions of ecotypes in East Africa were distinguished be either continuous or rotational. In arable land by differences in leaf type and flowering behav­ or irrigation channels it is a weed, and intensive iour. Some strains are female fertile (apomictic control by hand or with herbicides is necessary. through aposporic embryos) others are open-polli­ Diseases and pests In Australia, the major dis­ nated. Several cultivars have been released in Aus­ ease is 'kikuyu yellows' caused by a soilborne path­ tralia. 'Whittet', a robust, upright form, was ogen Verrucalvus flavofaciens; symptoms are selected in New South Wales from seed introduced patches of yellow, chlorotic leaves appearing in from Kenya. 'Breakwell' a densely tillered, pro­ summer. Larvae ofvariou s or scarab beetles strate strain, was derived from a naturalized stand (Rhopea magnicornies) cause temporary damage to at Grafton. 'Crofts' is fine-leafed with a degree of kikuyu grass pastures. cold tolerance. 'Noonan' seeds prolifically and is Harvesting Being prostrate, kikuyu grass is tolerant of the disease 'kikuyu yellows'. suited to grazing, rather than for use as cut-and- Ecology Kikuyu grass occurs naturally on the carry forage. Repeated mowing to promote flower­ margins of forests on the highland plateaux of ing is needed for seed production. Seed is har­ Ethiopia, Kenya and Central Africa (1950-3000 m) vested by removing all material to ground level, receiving 1000-1600 mm rainfall per annum. In threshing and sieving. Kenya at elevations of 2250m ,mea n minimum and Yield Dry matter yields of kikuyu grass vary maximum temperatures range from 2-8°C and from 9-30 t/ha, indicating a potential for high car­ 16-22°C, respectively. Optimum growth for ki­ rying capacities and animal production by dairy kuyu grass is attained at 25/20° C (day/night tem­ or beef cattle. Stocking rates of 1.5-3.0dair y cows/ peratures). The grass withstands short duration ha have been achieved in Australia with high rates frosts, but tissue is killed below -2°C. The present of fertilizer, although productivity per head was geographical distribution of the grass coincides moderate (9.0-16.6 kg milk/day). with mesothermal humid climates, frequently Genetic resources Although kikuyu grass where rain forest was the original vegetation. At shows variation in leaf morphology and flowering higher latitudes (25°-35°) kikuyu grass natural­ behaviour in the wild, original clonal introduc- PENNISETUM 189 tions have resulted in uniform pastures in many use as a fresh fodder or as hay. It is useful for con­ countries. Germplasm collections are available in trolling soil erosion, especially on sloping land. It Australia (New South Wales Agriculture & Fisher­ is considered a weed in Thailand, Fiji and parts ies, Grafton) and ILCA (Ethiopia). of Malaysia. The fully mature culms make good Breeding Improvement in kikuyu disease resis­ paper pulp. tance and cold tolerance has been mainly achieved Properties Nitrogen concentrations in P. poly­ by selection within natural variation of existing stachion, before or at early flowering, are 1.8-2.8% , collections. Suitable breeding methods are not declining to 0.8% in older material. Similarly, P available for this species. concentrations fall from 0.5-0.2%. Experiments Prospects Kikuyu pastures in South-East Asia in Thailand showed a DM digestibility of 60-70% are restricted to certain highland areas in New and a DM intake by sheep of 363-536 g/head per Guinea and the Philippines at present. Widespread day. After flowering, nutritive value and accept­ soil erosion on steep, cultivated land and degraded ability to animals decrease rapidly. catchments in South-East Asia, could be reduced Botany An annual or perennial tufted or spread­ if permanent pastures were established. Kikuyu ing grass, 0.5-3 m tall with culms erect or genicu­ grass could have an important role for land stabili­ late, simple or branched. Leaf-blade linear, 10-50 zation and improved animal nutrition in some of cm x 5-18 mm, drooping, glabrous to soft hairy these situations. above. Inflorescence a cylindrical panicle, 5-25 Literature ilj Bogdan, A.V., 1977. Tropical pas­ cm x 13-26 mm, purple or brownish; spikelets ses­ ture and fodder plants. Longman, London, pp. sile, 3-5 mm long, each enclosed by up to 30 cm 222-229. ;2!Edwards , D.C., 1937.Thre e ecotypes of long involucral bristles, two-flowered, only the Pennisetum clandestinum (Höchst.) kikuyu grass. Empire Journal of Experimental Agriculture 5: 371. 3, Mears, P.T., 1970. Kikuyu (Pennisetum clandestinum) as a pasture grass - a review. Trop­ ical Grasslands 4: 139-152. ;4i Oram, R.N., 1990. Register of Australian herbage plant cultivars. CSIRO, Australia, pp. 73-76. i5i Skerman, P.J. & Riveros, F., 1990. Tropical forage grasses. FAO, Rome. pp. 604-611. P.T. Mears

Pennisetum polystachion (L.) Schultes

Syst. Veg. Mant. 2:146 (1824). GEAMINEAE 2n = 32,36 ,45 , 53, 54,56 ,7 8 Synonyms Panicum polystachion L. (1759), Pen­ nisetum setosum (Swartz) L. Rich. (1805), P. atri- chum Stapf & Hubbard (1933), P. subangustum (Schum.) Stapf &Hubbar d (1933). Vernacular names Feather pennisetum (En). Malaysia: rumput ekor kucing, rumput berus ku- cing. Philippines: ikug-kuting (Tagalog). Laos: hnhaaz khachoon. Thailand: ya-khachonchop. Vietnam: co' duôi voi nho'. Origin and geographic distribution P. poly­ stachion is a native oftropica l Africa and India but is widely introduced and distributed in Indonesia, Malaysia, Thailand, Fiji, Australia and generally throughout the tropics, but rarely beyond 23° N Pennisetum polystachion (L.) Schultes - 1, flower­ and23°S. ing plant; 2,ligule; 3,spikelet with bristles; 4, spike- Uses P. polystachion is grazed and also cut for let without bristles. 190 FORAGES

upper floret fertile; bristles scabrous to densely grazed or cut before flowering. When used as cut- ciliate, 20-50 per spikelet. and-carry forage, it can be cut 2-3 times a year. In northern latitudes (Thailand) it flowers in Cut forage can be fed fresh or conserved as hay, October and in southern latitudes (Fiji) in April. although the succulent stems are slow to dry. If Seedlings grow vigorously early in the rainy sea­ it is prevented from flowering, P.polystachion will son. There are conflicting reports about seed dor­ stay green well into the dry season. When sub­ mancy. Some say that seeds may have a dormancy jected to sustained heavy grazing it will not persist period of 3-6 months and others indicate there is and will be replaced by prostrate species. Legumes no dormancy. One report mentions that the germi­ can be grown with P.polystachion, although fertil­ nation percentage of P. polystachion seed was ization with P may be required. When grown with 6-8% after one year of storage, improving to a suitable legume in Fiji, P.polystachion was eaten 14-22 % after two years. It is a prolific seeder. more readily by cattle. Shade-tolerant legumes P.polystachion is a polymorphic species, very vari­ such as Desmodium heterophyllum (Willd.) DC, able in colour, stiffness and length of the bristles. Desmodium heterocarpon (L.) DC. ssp. ovalifolium In fact it is a complex species, formerly subdivided (Prain) Ohashi, and Centrosema pubescens Benth. into the species mentioned under synonyms, which can be grown with P.polystachion. Also Macropti- are now joined by a continuous pattern of varia­ lium atropurpureum (DC.) Urban and Stylosanthes tion. Sometimes the complex is subdivided into 3 guianensis (Aublet) Swartz have been successfully subspecies (without clearly separating character­ introduced into P. polystachion pastures after istics): burning and with the use of P fertilizer. For pulp - ssp.polystachion: annual plant; involucral bris­ making, it should be cut after the seeds are fully tles 6-45, ciliate; In = 32,36 ,45 ,54 . ripe since pulp quality is best at this stage. - ssp. setosum (Swartz) Brunken: ; In Fiji, P.polystachion yielded 11-13 t/ha per year involucral bristles up to 50, densely to sparsely ofD M when fertilized with N.Steer s grazing unfer­ ciliate; 2n = 53,54 ,56 , 78. tilized P. polystachion in Fiji produced liveweight - ssp. atrichum (Stapf & Hubbard) Brunken: pe­ gains of only 110-80 kg per beast per year within rennial plant; involucral bristles up to 30, sca­ a stocking rate range of 1-3 beasts/ha. Seed yields brous; 2n = 36. of up to 420 kg/ha have been obtained in India. P. polystachion resembles P. pedicellatum Trin. Seeds can be viviparous in very wet weather. and P.purpureum Schumach., but it can be distin­ Genetic resources and breeding It is unlikely guished by the lower lemmas which are often that substantial germplasm collections are being three-lobed, the upper florets which disjoint readi­ maintained and there are no breeding programmes ly and rachises with decurrent wings on the ridge with P. polystachion. below the pedicels. Prospects P.polystachion's ability to adapt, per­ Ecology P.polystachion is a short-day plant and sist, give high yields, and combine with a range mainly adapted to the lowland tropics, although of legumes make this a very useful grass. Research it can be found at altitudes up to 2400 m. It grows is needed into its general agronomy, the best man­ well in high rainfall areas, but can also tolerate agement regime to obtain the highest animal pro­ short dry periods. Its optimal temperature for duction, and how to conserve it as hay or silage. growth is 32-35°C with a minimum of 12°C. It is Literature |1| Baki, B.B., Latiff, A. & Ahmad adapted to a wide range of soil types, from light Puat, N., 1989.Th e genus Pennisetum in Peninsu­ sandy soils to waterlogged clay soils, but it will lar Malaysia with particular reference to P. seto­ not stand prolonged flooding. It often occurs on sum, a new record for the weed flora. Proceedings old farmland and other disturbed places. P. poly­ Symposium on weed management, 7-9 June 1989, stachion tolerates acid (pH(H20) 4.5) as well as al­ Bogor, Indonesia. j2j Bogdan, A.V., 1977. Tropical kaline soils. It is tolerant of shade and poor soil grasses and fodder plants. Longman, London, pp. fertility, and is favoured by regular burning. 231-232. |3j Brunken, J.N., 1979. Morphometric Agronomy P.polystachion isbes t established by variation and the classification of Pennisetum sec­ broadcasting seed on cultivated land, sowing at tion Brevivalvula (Gramineae) in tropical Africa. 3.5-4.5 kg/ha. It is usually not fertilized, but either Botanical Journal of the Linnean Society 79: chemical fertilizer or farmyard manure is some­ 51-64. |4| Häfliger, E. & Scholz, H., 1980. Grass times applied in Fiji and India. There are no weeds 1. Ciba-Geigy Ltd., Basel, Switzerland, p. reports of any major diseases or serious pests. 142. |5| Holm, L.G., Plucknett, D.L., Pancho, J.V. When used as forage, P. polystachion should be & Herberger, J.P., 1977. The world's worst weeds: PBNNISETUM 191 distribution and biology. The East-West Center, University of Hawaii Press, Honolulu, Hawaii, United States, pp. 367-371. |6I Manidool, C, 1977. Nutritive values of ya kachonchop (Pennisetum polystachion) [in Thai]. Poultry Magazine, Poultry Promotion Association of Thailand 25: 1. |7| Partridge, I.J. , 1986. Effect of stocking rate and superphosphate level on an oversown fire climax grassland of mission grass (Pennisetum polysta­ chion) in Fiji. Tropical Grasslands 20: 166-180. |8| Skerman, P.J. & Eiveros, F., 1990. Tropical grasses. FAO, Rome. pp. 616-620. B.B. Baki, C. Manidool &C.P . Chen

Pennisetum purpureum Schumach.

Beskr.Guin.Pl.:44(1827). GRAMINEAE In = 28,56 ,2 7 Vernacular names Elephant grass, napier grass (En). Herbe d'éléphant (Fr). Indonesia: rum- put gajah. Malaysia: rumput gajah. Philippines: buntot-pusa (Tagalog), handalawi (Bokil), lagoli (Bagobo). Thailand: ya-nepia. Vietnam: co' duôi voi. Origin and geographic distribution Of trop­ ical African origin, this grass has been introduced to all tropical regions of the world and is natura­ lized throughout South-East Asia where annual Pennisetum purpureum Schumach. - 1, habit of rainfall exceeds 1000 mm and there is no long dry flowering plant; 2,spikelet surrounded by bristles. season. Uses The main use of elephant grass is as a for­ the lower surface. Inflorescence a dense spike-like age for ruminants. As a naturalized species in panicle, up to 30 cm tall and 30 mm wide, not humid regions of South-East Asia, it is collected including the 16-40 mm long bristles on the spike- by farmers by cutting the whole plant, which is lets; spikelets 5-7 mm long, solitary or in clusters offered to ruminants, mainly buffaloes and cattle, of up to five, of which usually only one is fertile; which are either tethered or confined in stalls. It the lower floret is male or void, the upper bisexual can also be used to provide mulch. and fertile, sometimes male. There is little or no Properties The feeding value is influenced seed formation. mainly by the ratio of leaf to stem and by age. Growth and development Elephant grass is an Nitrogen concentrations of regularly harvested obligate quantitative short-day plant, with a criti­ forage are usually in the range of 2-4%. Young cal photoperiod between 13 and 12 hours. Howev­ leaves may have a digestibility of7 0% ,bu t this val­ er, viability of pollen is poor and this may be the ue declines rapidly with age to less than 55%. main cause of the typically poor seed set. In addi­ Stems are of low digestibility, except when very tion, seedlings are weak and grow slowly, so that young. the grass is usually propagated vegetatively. Description A tall, robust, deep-rooting, erect Under favourable conditions, vegetative material perennial, with short rhizomes. Stem up to 7m tall is fast-growing and the plant can reach a height and 3 cm in diameter, up to 20-noded. The plant of several metres within two months. forms clumps to 1 m across. Leaf-sheath glabrous Other botanical information There are to short bristly; leaf-blade linear with broad base numerous cultivar names in various countries and and acute tip, up to 120 cm x 5 cm, glabrous to three subspecies have been proposed in northern hairy at the base, with a prominent midrib along Africa, but there has been no similar subdivision 192 FORAGES for South-East Asia. In Florida (United States), a Cattle liveweight gains of 1 kg/animal per day dur­ dwarf cultivar 'Mott' of high feed value has been ing the growing season and of 120 kg/animal and developed. 480 kg/ha per year over 3 years were recorded on Ecology Elephant grass is adapted to a humid N fertilized dwarf elephant grass ('Mott') in Flor­ warm environment. However, it can exhibit ida, United States. remarkable drought-tolerance and can survive Genetic resources Because of clonal propaga­ light frost. For acceptable agronomic perfor­ tion, planted stands of P.purpureum are often uni­ mance, the species requires a deep soil of at least form. However, the species contains much varia­ moderate fertility, although it will survive at tion in the extent of hairiness of stem nodes and much reduced productivity on all kinds of soils. It leaf-sheaths, and the size, colour and density of the does not tolerate sustained flooding. In its natura­ panicle. There are also differences in stem thick­ lized state, the grass is found mainly along forest ness and height and leaf size between forms of the edges. grass, but these are greatly influenced by the fertil­ Propagation and planting Vegetative propa­ ity of the soil and by rates of fertilizers. gation is either by dividing clumps of roots and Breeding P. purpureum x P. americanum (L.) stubble, or by stem cuttings consisting of at least Leeke hybrids have been developed in many coun­ three nodes, two of which are buried. This can be tries. They produce larger plants with more tiller­ done by hand or with a sugar-cane planter. Row ing and larger total production. However, this also width ranges from 50-200 cm, the greater dis­ means that more mineral nutrients are required. tances being preferred in drier regions. Distance The hybrid is sterile and therefore must be vegeta- within rows varies from 50-100 cm. Intercropping tively propagated. with cassava and banana is often practised in Prospects Elephant grass is widely grown in home gardens. tropical regions and, with adequate use of fertiliz­ Husbandry For high yields and persistence, ele­ ers, large increases in yield can be expected. By phant grass planted as a crop requires a regular harvesting at a young stage of growth or by using water supply and a rich supply of nutrients. The dwarf cultivars, forage of high feeding value can latter applies particularly when the crop is cut fre­ be obtained. An advantage ofthi s species is its ver­ quently. Nutrient removal per t DM is: N 10-30 satility. It can be grown on a large or small scale; kg, P 2-3 kg, K 30-50 kg, Ca 3-6 kg, Mg and S it lends itself to mechanization but is also suitable 2-3 kg. With annual DM yields between 20 and 40 for smallholder agriculture. t/ha, very large quantities are thus extracted from Literature !l; Bodgan, A.V., 1977. Tropical pas­ the soil. If they are not replenished, yield soon ture and fodder plants. Longman, London, pp. drops and weeds will invade. Although elephant 233-243. ;2\Crowder , L.V. & Chheda, H.R., 1982. grass is not often grown with legumes, it combines Tropical grassland husbandry. Longman, London. well with, for instance, Centrosema pubescens 562 pp. j3| Skerman, P.J. &Riveros , F., 1990. Trop­ Benth. and it can be interspaced with the shrub ical grasses. FAO, Rome. pp. 621-627. |4| Sollen- legume Leucaena leucocephala (Lamk) de Wit. berger, L.E. & Jones, CS., 1989. Beef production Diseases and pests The most common disease from nitrogen-fertilized Mott dwarf elephant grass is a blight caused by Helminthosporium sacchari. and Pensacola bahiagrass pastures. Tropical The best control measure is to use a resistant culti­ Grasslands 23:129-134. var. No major pests have been recorded. L. 't Mannetje Harvesting Elephant grass can be harvested year-round. It is usually offered fresh to animals, but it can also be conserved as silage. However, Pueraria phaseoloides (Roxb.) Benth. preservation is often poor and losses of DM and crude protein can be very large. Best results are Journ. Linn. Soc. 9:125(1865) . obtained by chopping the material, mixing it with LEGUMINOSAE molasses, and compressing and covering it to 2n = 22, 24 exclude air. Synonyms Dolichos phaseoloides Roxb. (1832), Yield Annual DM yields that can be expected in Pueraria javanica (Benth.) Benth. (1865), P. pha­ farm practice may range from 2-10 t/ha for unferti­ seoloides Roxb.wax. javanica (Benth.) Baker (1876). lized or slightly fertilized stands and from 6-40 Vernacular names Tropical kudzu, puero (Aus­ t/ha from grass well fertilized with N and given a tralia) (En). Indonesia: krandang (Javanese), foea basic dressing of P. banga (Ternate). Malaysia: kudzu tropika. Philip- PUERARIA 193

pines: singkamasaso (Tagalog), bahay (Bikol), vaay (Ivatan). Laos: pièd, s'üak pièd. Thailand: thua sian paa (Ang Thong). Vietnam: dâu ma, dâu dai, dâu rung. Origin and geographic distribution Tropical kudzu is indigenous to the lowlands of East and South-East Asia where it is widely cultivated as a cover crop. It has been introduced to other trop­ ical regions and is now naturalized in many areas including the wet tropics of Australia and the Americas. Uses Tropical kudzu is especially important as a component of grazed and ungrazed cover crop mixtures in rubber and oil-palm plantations in South-East Asia, Africa and tropical America. It is used as a pasture legume in South-East Asia, tropical America and Australia. Its ability to smother weeds makes it a useful pioneer legume in combination with other more permanent spe­ cies. It has been planted on sloping sites to reduce soil erosion. In mixtures with other cover crops, it has been used as green manure by periodic incor­ poration into the soil. The tuberous roots are edi­ ble. Strong fibres from the stem are used for rope making. In Malesia the plant is used in traditional medicine to cure boils and ulcers. Properties Tropical kudzu is a very palatable legume, although its wet season palatability is Pueraria phaseoloides (Roxb.) Benth. -1, flowering reported to be low in tropical America. Nutrient branch; 2,flower, frontal and side view; 3,fruit; 4, concentrations typically range from 2-4% N, seeds. 30-40% crude fibre, 0.15-0.45% P and 0.4-1.6% Ca. Seed weight is 80-90 seeds per g. violet central blotch. Pod straight or slightly Description Deep-rooting perennial herb with curved, terete or compressed cylindrical, 4-11 cm climbing or twining, hairy stems. Roots subtuber- x 3-5 mm, thinly clothed with stiff appressed ous. Main stems ca. 6 mm diameter, extending hairs, black when mature and containing 10-20 4.5-10 m, rooting at nodes if in contact with moist seeds. Seed oblong to squarish with rounded soil, lateral stems branching from nodes; young corners, about 3m m x 2 mm, brown to brownish- shoots densely covered with brown hairs. Leaves black. large, trifoliolate, borne on hairy petioles 2-13 cm Growth and development Seedlings of tropical long; leaflets triangular or ovate, 2-20 cm x 2-15 kudzu are only moderately vigorous and may only cm, thin, base broadly cuneate or subrhomboidal grow slowly for the first 3 or 4 months. Neverthe­ and very shallowly lobed, apex acuminate, laterals less, seedling vigour issuperio r to other cover crop oblique usually 6-7 cm long and wide, thinly hairy species such as Centrosema pubescens Benth. (cen­ on upper surface, greyish-green and densely pubes­ tre) and Calopogonium mucunoides Desv. (calopo). cent on lower surface; stipules small, lanceolate, Once established, it is very vigorous and quickly ciliate with long brown hairs. Flowers small, smothers weeds. It can form a tangled mat of vege­ mauve to deep purple, borne in scattered pairs on tation 60-75 cm deep, and climb trees and fences. axillary racemes 15-30 cm long; peduncle about Other botanical information Sometimes 2 13 cm long; bracteoles lanceolate, 1-3 mm long; botanical varieties are distinguished within P. calyx campanulate, 6 mm long, hairy, upper teeth phaseoloides: var. phaseoloides and var. javanica. broad, lateral ones triangular, the lower lanceo­ In this view, var. javanica is a more robust plant late and all terminating in a bristle; standard orbi­ with larger flowers and pods. There is no regis­ cular, 1-2 cm diameter, spurred, greenish on out­ tered Australian cultivar; seed may be traded, but side and white on the inner side with a mauve without cultivar name. 194 FORAGES

Ecology Tropical kudzu is only suited to the and poor persistence. Grazing experiments in humid tropics with an annual rainfall in excess of Malaysia have shown that under continuous graz­ 1500mm . It has a temperature optimum of 32/24°C ing at stocking rates of 2, 4 and 6 local cattle per (day/night) and dry matter yields are reduced by ha, the proportion of tropical kudzu was signifi­ 35% with a change in temperature regime to cantly reduced with increased stocking even after 26/15°C . Few reports are available on photoperiod one year ofgrazin g while the effect ofstockin g rate responses. In Puerto Rico (lat. 18°N), flowering on centro was only evident after three years graz­ and seed set occur in the short daylength period ing. Farmers using grass-legume mixtures have from January to March, suggesting that it may be also reported excellent growth of tropical kudzu a short-day plant. Tropical kudzu has been ranked in the first two years, which thereafter rapidly dec­ highly as being shade-tolerant in comparison with lined under grazing. The lack of persistence of other legume species. Under a 50% shade regime tropical kudzu is probably also influenced by soil in coconut plantations in the Solomon Islands, it physical characteristics and related to the poor de­ was the most productive legume and it even sup­ velopment ofroote d stolons on some soils. pressed the accompanying grasses.Thi s character­ When planted under palm and tree crops on former istic makes it suitable in integrated livestock/ high forest land, some initial control of natural plantation production systems. Under a more than regrowth of forest plants is necessary to establish 50% shade regime, tropical kudzu gives way to tropical kudzu. During the subsequent two years, centro or desmodium (Desmodium heterocarpon tropical kudzu has to be cutlassed or beaten down (L.) DC. ssp. ovalifolium (Prain) Ohashi). to a height of 30 cm. Rings around trees are clean- Tropical kudzu is tolerant of waterlogged sites. It weeded to prevent tropical kudzu from climbing prefers heavy soils and iswel l adapted to acid soils. the trees. It is particularly susceptible to Mg and S deficien­ Diseases and pests Tropical kudzu is remarka­ cies and has moderate to low Ca and P require­ bly free from diseases, but leaf-eating caterpillars ments, but it responds to fertilizer application. can damage ungrazed swards and pod-borers Propagation and planting Tropical kudzu is reduce seed production. usually planted by seed. It has a high proportion Harvesting Usually tropical kudzu is directly of hard seed and germination can be increased by grazed in mixed pastures but can be cut for hay, using either hot water, acid or mechanical scarifi­ silage or for stall feeding as fresh forage. cation. Commercial suppliers often sell scarified Yield Annual DM yields of up to 10 t/ha from seed which is obtained by abrading the seed-coats tropical kudzu swards have been recorded in cut­ in a hexagonal drum, lined with sandpaper, rotat­ ting experiments, with some §-§ of the yield from ing at 7.5rp m for 24 hours. the wet season and ^-\ from the dry season. DM Tropical kudzu will usually nodulate with native yields of up to 23t/h a have been measured in trop­ cowpea rhizobia but inoculation with the appro­ ical kudzu-grass swards, 40% ofthi s being tropical priate strain ofBradyrhizobium, such asRRI M 768 kudzu. In tropical America, tropical kudzu-grass in Malaysia, is recommended when planting in pastures have produced live weight gains of 313kg / new areas. Seed is usually broadcast or drilled in ha per year (with Andropogon gayanus Kunth) and rows 1m apart. It can also be established by over­ 542kg/h a per year (with Panicum maximum Jacq.) sowing into an existing pasture if the pasture is Genetic resources and breeding The largest disked or burnt beforehand. It can be propagated germplasm collection is maintained by CIAT (Col­ vegetatively, one recommendation being to plant ombia) and a smaller collection ishel d at ATFGRC two rooted cuttings, 0.7-1 m long, at each point (CSIRO, Australia). There are no known breeding on a 1-2 m grid. programmes with this species. Standard cover crop mixtures include a 5:4:1 ratio Prospects Tropical kudzu is widely grown of calopo, centro and tropical kudzu or a 4:1 mix­ throughout the humid tropics, especially in plan­ ture of centro and tropical kudzu. Seeding rates tation crops. Its main features as a forage legume for these mixtures are 2-5 kg/ha in the inter-row are its vigorous initial growth on fertile soil and areas between rubber or oil palm trees. its high palatability. Improvements in this species Husbandry Tropical kudzu responds well to should primarily be aimed at improving its persis­ added P; linear responses to at least 50 kg/ha of tence under grazing. P have been obtained on infertile soil. Literature |1| Chin, S.L., 1977. Leguminous Tropical kudzu is very palatable, much more so cover crops for rubber smallholdings. Planters than calopo, and this can lead to selective grazing Bulletin 150: 83-97. |2| Costa, N. de L., 1990. Pue- SACCHARUM 195

raria; Leguminosa forrageira para a produçào de proteina. EMBKAPA-UEPAE Porto Velho, com- unicado Técnico CT/92. 4 pp. |3| Crowder, L.V. & Chheda, H.R., 1982.Tropica l grassland husbandry. Longman, London. 562 pp. !4i Duke, J.A., 1981. Handbook of legumes of world economic impor­ tance. Plenum Press, New York. pp. 214-216. |5| Eng, P.K., Chen, C.P. & 't Mannetje, L., 1978. Effects ofphosphoru s and stocking rate on pasture and animal production from a guinea grass legume pature in Johore, Malaysia. 2. Animal live weight change. Tropical Grasslands 12: 198-207. |6| Pur- seglove, J.W., 1968. Tropical Crops-Dicotyledons. Vol. 1.Longman , London, pp.218,220 . |7| Skerman, P.J., Cameron, D.G. & Eiveros, F., 1988. Tropical forage legumes. FAO,Rome .pp .376-388 . |8|White - man, P.C., 1980. Tropical pasture science. Oxford University Press, New York. 392pp . R.A. Halim

Saccharum spontaneum L.

Mant.pl. alt.: 183(1771) . GRAMINEAE 2« varies from 48t o 128 Vernacular names Indonesia: glagah (Java­ nese), tatebu (Timor). Papua New Guinea: pit-pit. Saccharum spontaneum L.-l, base ofplant; 2, culm Philippines: talahib (Tagalog), tigbau (Bikol), node; 3, ligule; 4, flowering culm; 5,panicle; 6,part sidda (Ilokano). Cambodia: 'âm'pëu préi. Laos: of raceme with (partly separated) spikelets. ph'ông, lau. Thailand: phong, lao. Vietnam: co' lach. cle, 20-60 cm long, axis hirsute; racemes 3-15 cm Origin and geographic distribution S. sponta­ long, usually much longer than the supporting neum originates from and is widely distributed in branches; spikelets paired, one sessile, one pedi- the warmer regions of the Old World, including celled, similar, lanceolate, 3-7 mm long, the callus South-East Asia. bearded with silky white hairs 2-3 times as long Uses It is used by villagers as a fodder for cattle, as the spikelet; glumes ciliate on the margins buffaloes and, in some places, elephants. It is also above. Caryopsis about 1.5 mm long. used to prevent erosion of sandy soils. The foliage S. spontaneum is a polymorphic species of which is used for thatching and the plant is used as an its many forms are commonly grouped into two ornamental and to produce paper pulp. subspecies: Properties Nitrogen concentrations ranging - ssp.spontaneum: leaf-blade narrowed to the mid­ from 0.5-1.4% have been measured in India and rib towards its base;ligul e ca.triangular ; mainly Thailand. It is relished by water buffaloes and ele­ in tropical and warm temperate Asia; phants but is less attractive to cattle. In the Philip­ - ssp. aegyptiacum (Willd.) Hack.: leaf-blade not pines it is claimed that N-fixing bacteria live in narrowed at base; ligule crescent-shaped; symbiosis with S. spontaneum. mainly in Africa and the Middle East. Botany Rhizomatous perennial, with culms 1-4 Seeds germinate poorly, but once germinated the m tall or more, waxy below the nodes. Leaf-sheath seedlings resist harsh conditions and soon develop tight, 20 cm or more long, overlapping, striate, a strong root system. Large tussocks will form often purplish, glabrous; ligule obtuse or triangu­ when grown on river banks or ditches, thereby col­ lar, about 2 mm long, shortly ciliate; leaf-blade onizing large areas. It flowers towards the end of linear-acuminate, 50-90 cm x 5-15(-40) mm, gla­ the rainy season. brous, margins scaberulous. Inflorescence a pani- Ecology S. spontaneum grows from near sea- 196 FORAGES level up to 1700m altitude. It prefers a high rainfall diflora (L.) Desv. (1813). environment, usually in excess of 1500m m per year, Vernacular names Agati sesbania, West-In­ and is adapted to a widerang e ofsoils , from alluvial dian pea (En). Fagotier (Fr). Indonesia: turi (Sun- soils on river banks to sandy soils of old mines. danese), toroy (Maduranese), tuwi (Balinese). Agronomy S. spontaneum is easily propagated Malaysia: sesban, kacang turi (Peninsular), petai by division of rhizomes. The spikelets bear long belalang (Sabah). Philippines: pan (Tagalog), silky hairs and are easily and effectively dispersed gauai-gauai (Visaya), katuday (Ilokano). Cambo­ by wind. It is tolerant of heavy grazing. Regular dia: 'ângkiëdèi. Laos:kh'ê : kha:w. Thailand: khae, grazing or cutting is required to keep plants in a khae-ban. Vietnam: so düa. leafy state sotha t they maintain their palatability. Origin and geographic distribution The exact Burning also helps the plants to produce new country of origin of S. grandiflora is not known shoots for grazing. It is normally grazed by village (India or Indonesia have been suggested) but it is cattle and water buffaloes, but young leaves can considered native to many South-East Asian coun­ also be cut and fed to the livestock. It is expected tries. It is widely distributed through the tropics to give fairly high yields. Young leaves can also from southern Mexico to South America and has be made into hay for the dry season as is done with been planted in southern Florida and Hawaii. It sugar cane tops. In Thailand it is reported to be has been cultivated for at least 140 years in West a host for a downy mildew fungus that badly affects Africa and more recently in East Africa. maize plants. Uses The leaves and fruits are used asforag e and Genetic resources and breeding It has been green manure. The tree can be used as an ornamen­ thought that S. spontaneum can cross naturally tal, a shade tree, a windbreak, a living fence, a live with sugar cane to produce a hybrid known as S. support for crops like pepper and vanilla, and for sinense Roxb. which is almost completely sterile. the reforestation of eroded areas. In South-East S. spontaneum is a very variable species but it is Asia it has long been used for fuelwood. The young unlikely that any germplasm collections are being leaves, flowers and tender pods are used as a vege­ maintained. table for salads, curries and soups. The tree is used Prospects For forage purposes, cultivars with as a pulp source for the paper industry, especially better palatibility and nutritive status are required. in East Java. The light wood is used in floating S. spontaneum has goodprospect sfo r usei n control­ fishing nets. The extracts from leaves, flowers, ling soil erosion and reclaiming mine waste. bark and roots are known for their traditional me­ Literature |1| Backer, CA. & Bakhuizen van dicinal uses (e.g.th e root is a well known medicine den Brink, R.C., 1968. Flora of Java. Vol. 3. for ). The bark, when cut or damaged, Wolters-Noordhoff, Groningen, the Netherlands, exudes a clear gum which has some potential in p. 585. |2| Bor, N.L., 1960. The grasses of Burma, the food and non-food industries. Ceylon, India and Pakistan. Pergamon Press, Lon­ Properties Many reports indicate that S. gran­ don, p.214 . |3|Gilliland , H.B.,Holttum , R.E. & Bor, diflora is a very palatable fodder with a high feed­ N.L., 1971. Grasses of Malaya. In: Burkill, H.M. ing value for ruminants. The N concentration of (Editor): Flora of Malaya. Vol. 3. Government the seeds isu p to 6.5% and it ranges from 3.0-5.5 % Printing Office, Singapore, pp. 224-226. |4| Holm, in the foliage. This makes it a very suitable supple­ J., 1971. Feeding tables. Nutrition Laboratory of ment for poor quality roughages. The DM digesti­ the Thai-German Dairy Project, Livestock Breed­ bility of foliage ranges between 65% and 73%; it ing Station, Chiangmai, Thailand, p. 26. |5j Sker- generally has a low crude fibre content (5-18%) man, P.J. & Riveros, F., 1990. Tropical grasses. and a relatively high P concentration (0.30- FAO, Rome. pp. 649-651. 0.45%>). The information on anti-nutritional fac­ C. Manidool tors is limited and although the foliage contains saponins and tannins it has no known toxic reac­ tion to ruminants. However, caution should be Sesbania grandiflora (L.) Poiret used in feeding it to monogastric animals as it has caused mortality in chickens. S. grandiflora has Lamk, Encycl. Méth. Bot. 7:127(1806 ) ('Sesban'). 14-20 seeds/g. The wood is white and soft and not 3 LEGUMINOSAE durable. Its low specific gravity of 0.42 kg/dm 2n = 24 makes it also a poor fuelwood. Synonyms Robinia grandiflora L. (1753), Ae- Description A loosely branching tree, up to 15 schynomene grandiflora (L.) L. (1763), Agati gran­ m tall and about 30 cm in diameter. Roots are nor- SESBANIA 197

tube 10-12 cm long, curved for most of its length; ovary and style glabrous. Pod linear to slightly fal­ cate, 20-60 cm x 6-9 mm with broad sutures, 15-50 seeded, septa 7.5-10 mm apart, glabrous, hanging vertically, indéhiscent. Seed subreni- form, 6.5m m x 5m m x 2.5-3 mm, dark brown. Growth and development One of the charac­ teristics of S. grandiflora is its rapid early growth, reaching heights of up to 2 m in 12 weeks, 4-5 m in one year and about 8 m in 3 years. It is able to produce ripe pods nine months after planting. S. grandiflora seeds lose their viability after about one year when stored at ambient conditions. It improves soil fertility, although the ability to fix N may be suppressed by nematodes or high acidity of the soil. It has responded to the inoculation of soil with Vesicular-Arbuscular Mycorrhizal (VAM) fungi (Glomus fasciculatum and Glomus mossaeae). This treatment could facilitate the introduction of this species to P-deficient soils. The tree has a lifespan of about 20 years. It is not wind resistant. Other botanical information A closely related species, Sesbania formosa (F. Mueller) N. Bur- bridge, is native to northern Australia^ Ecology S. grandiflora is only suitable for the lowland tropics, up to 800 m above sea-level, as it Sesbania grandiflora (L.) Poiret - 1, flowering isfrost-sensitiv e and cannot tolerate cool tempera­ branch; 2,flower bud; 3,flower; 4, fruits. tures over an extended period. Although it has been successfully grown in arid areas with only mally heavily nodulated with large nodules. The 800 mm annual rainfall it is best adapted to places tree can develop floating roots and aerenchyma with an annual rainfall between 2000-4000 mm. It tissue. Stems tomentose, unarmed. Leaf pinnately can be grown in a wide range of soils including compound, up to 30 cm long, including a petiole those that are poor and waterlogged. It tolerates of 7-15 mm long; the rachis slightly pubescent or saline and alkaline soils and has also some toler­ glabrous; leaflets 20-50, in pairs opposite to alter­ ance to acidic soils down to pH(H20) 4.5. S. grandi­ nate on the same leaf, oblong to elliptical, 12-44 flora is able to tolerate flooding over long periods. mm x 5-15 mm, rounded to obtuse to slightly Propagation and planting S. grandiflora is emarginate at the apex, slightly asymmetrical at easily propagated by direct seeding. Although the base, glabrous or sparsely pubescent on both scarification might have some positive effects the surfaces; stipels filiform, 0.75-1 mm long, pubes­ seeds usually germinate promptly without any cent, persistent; stipules broadly lanceolate, 8m m treatment. Seedlings nodulate on most soils with­ long, early deciduous. Raceme axillary, 2-4 flow­ out inoculation. Because of its rapid early growth, ered, rachis up to 65m m long; peduncle 15-35 mm it can compete very well with weeds. Vegetative long, tomentose; pedicels 15-18 mm long, pubes­ propagation by cuttings is possible but seldom cent; bracts lanceolate, 3-6 mm long, early deci­ practised. Occasionally seedlings raised in poly­ duous; bracteoles broadly lanceolate, 4-6 mm thene bags or other containers are used, to ensure long, deciduous;flowe r white, yellowish, rose-pink a better establishment. S. grandiflora is often or red; calyx 15-22 mm long, closed in young buds, planted as individual trees or in rows (spaced 1-2 splitting or breaking in various ways at anthesis, m apart) along fence lines, field borders and the the basal part persistent in fruit; standard up to bunds of rice paddies. 10.5 cm x 6 cm, no appendages at the claw; wings For wood production it can be planted very den­ up to 10.5 cm x 3 cm without a basal tooth; keel sely. Over 3000stem s per ha have been used in Aus­ up to 10.5 cm x 4.5 cm with basal tooth; staminal tralia and India. 198 FORAGES

Husbandry The foliage of S. grandiflora is gen­ Tropical Agriculture Series No 8. Westview Press, erally used in a cut-and-carry system. It is usually Boulder, Colorado, pp. 27-68. |3| Lewis, G.P., 1988. fed fresh as a supplement to other roughages, but Sesbania Adans. in the Flora Zambesiaca Region. it can also be dried. Although there are several Kirkia 13:11-51. |4|Macklin , B.& Evans , D.O. (Ed­ reports that it is browsed, the reaction to and re­ itors), 1990.Perennia l Sesbania species in agrofor- covery of bushes after direct grazing has not been estry systems. Proceedings of a NFTA/ICRAF studied. Fertilizers are rarely used and therefore workshop held in Nairobi, Kenya, March 27-31, their effects are not well known. 1989. NFTA, Waimanalo, Hawaii. 242pp . {51 Natio­ Diseases and pests Little information is avail­ nal Academy of Sciences, 1979. Tropical legumes: able on the occurrence and importance of diseases resources for the future. National Research Coun­ and pests in S. grandiflora. Eeports on the fungus cil, Washington, pp. 185-193. |6i Nguyen van Pseudocercospora sesbaniae (grey leaf-spot) are Thuân, Dy Phon, P., Niyomdham, C. & Vidal, Y., only from India, as is the occurrence of the sesba- 1987. Légumineuses - Papilionoïdées. In: Morat, nia mosaic virus. The stem-borer Azygophleps sca- Ph. (Director): Flore du Cambodge, du Laos et du laris has caused some damage in India. Larvae of Vietnam. Vol. 23. Muséum National D'Histoire the insect Bruchophagus mellipes infest and dam­ Naturelle, Laboratoire de Phanérogamie, Paris, age seeds. Susceptibility to nematodes has been pp. 56-62. reported. J.H. Heering &R.C . Gutteridge Harvesting S. grandiflora does not tolerate fre­ quent, complete defoliation; this will cause high mortality rates. Initially the sidebranche s ofa tree Sesbania sesban (L.) Merrill may be cut, leaving the main growing point untouched. After the tree has reached a height of Philipp. J. Sei., Bot. 7:23 5(1912) . 3 m or more, the leader can be cut back to heights LEGUMINOSAE above 1.5 m. 2« = 12 Yield Forage yields of S. grandiflora depend Synonyms Aeschynomene sesban L. (1753), Ses­ very much on soil fertility and the management bania aegyptiaca Poiret (1806)(a s 'Sesban aegyptia- imposed. When side branches are lopped periodi­ cus'). cally a tree can yield up to 27 kg of fresh green Vernacular names Sesban (En). Indonesia: leaves per year. jayanti (Sundanese), janti (Javanese), puri. Philip­ Wood yields of 20-25 m3/ha per year are commonly pines:katura y (Tagalog, Bikol), katodai (Ilokano). achieved in Indonesia. For green manure, yields Burma: yay-tha-kyee (Burmese), yethugyi. Cambo­ of 55 t/ha green material have been obtained in dia: snaô kôôk. Laos: sapao lom. Thailand: sami Java in 6-7 months. (central), saphaolom (northern). Vietnam: dien- Genetic resources Small germplasm collec­ dien. tions are available at the University of Hawaii Origin and geographic distribution The exact (Waimanalo, United States), ILCA (Ethiopia) and origin of S. sesban is unknown, but it is widely dis­ ATFGRC (CSIRO, Australia). tributed and cultivated throughout tropical Africa Breeding Nobreedin g work isbein g carried out. and tropical Asia. It has been introduced into trop­ Prospects This species is widely grown in the ical America. tropics and has potential for wider use, although Uses The leaves and young twigs of sesban are prospects for improvement as a livestock fodder used as fodder for ruminants. The thick branches are limited. The research needs are: to collect and stems serve as fuelwood or are used as con­ germplasm of S. grandiflora, to document diseases struction material. Sesban is frequently used to and pests, to quantify the yield of forage from improve soil fertility, either through direct N fixa­ border plantings and fences, and to study the anti- tion or through the incorporation of foliage as a nutritional factors that affect ruminants and green manure to the soil. Sesban can be grown to monogastric animals. provide shade (e.g. in coffee, cocoa, ), and Literature |1| Evans, D.O. & Macklin, B. (Edi­ as a live support (e.g. in pepper and vine) or tors), 1990. Perennial Sesbania production and windbreak for other crops (e.g. banana). Sesban is use. A manual of practical information for exten­ also used byhuman s for food from the edible leaves sion agents and development workers. NFTA, Wai­ and flowers, for fibre from the bark, for traditional manalo, Hawaii. 41 pp. |2| Evans, D.O. & Rotar, medicines from leaves and flowers, and for gum P.P., 1987. Sesbania in agriculture. Westview from the seeds and bark. SESBANIA 199

Properties The N concentration of sesban long, subglabrous or pilose, very early caducous; leaves ranges between 3.0 and 4.5% ofDM .I n vitro calyx up to 6.5 mm long, the tube glabrous, the DM digestibility often exceeds 65% because of a teeth broadly triangular with an acuminate point relatively low fibre content. Anti-nutritional fac­ 0.5-1 mm long, marginally pubescent; standard tors such as polyphenolic compounds may some­ ovate, 11-20 mm x 13-21 mm, cordate at the base, times have an adverse effect on the digestibility yellow and commonly speckled or flecked with vio­ of sesban, but their role is not clearly understood. let-purple, claw up to 4m m long, appendages with In view of reported negative effects of sesban in acuminate free tips 2-5 mm long; wings 15-19 mm monogastric animals, especially poultry, it is less x 4.5-7 mm including a claw of 4-6 mm, yellow, suitable to be used in diets for these animals. Phos­ with a broad tooth or short hook at the base; keel phorus concentrations in the edible portion are 11-21 mm x 6.5-9 mm, including a claw of 6-9 generally adequate for animal nutrition. Sesban mm, yellow or creamish, basal tooth acute at 0-20 has 55-80 seeds/g. degrees to the claw; staminal tube (9-)12-13(-17) Description Shrub or short-lived tree up to 8 m mm long; ovary glabrous or rarely somewhat tall. Stem up to 12 cm in diameter, usually pubes­ pilose; style glabrous, 5m m long. Pod subcylindri- cent, sometimes becoming glabrous. Leaves, cal, straight or slightly curved, up to 20-30 cm x including a short petiole, 2-18 cm long, pinnately 2-5 mm, straw-coloured, often with a brown blotch compound; leaflets in 6-27 pairs, linear oblong, up over each septum or reddish-brown, 10-50 seeded, to 26 mm x 5 mm, glabrous or almost so above, glabrous;sept a 4-8 mm apart. Seed subcylindrical, sometimes pubescent beneath, often pilose at the 3-4.5 mm x 2 mm x 2 mm, olive-green or brown, margins; stipules narrowly triangular, up to 7m m usually mottled. long, pubescent. Racemes 2-20 flowered, up to 20 Growth and development Sesban germinates cm long, glabrous or sparsely pilose; peduncle up and grows rapidly; plants can reach a height of to 5 cm long; pedicels 4-12 mm long, glabrous; 1.5-2 mi n 10-12 weeks after sowing under favour­ bracts and bracteoles linear lanceolate, 3-5 mm able conditions. It usually flowers and produces ripe pods within the first year after planting. The flowers are mostly visited and pollinated by members of the Hymenoptera. Other botanical information S. sesban is sub­ divided into two subspecies, ssp. sesban and ssp. punctata (DC.) J.B. Gillett, of which the latter occurs only in West Africa from Senegal to the Sudan. The main difference is the length of the sta­ minal tube: 15-17 mm in ssp. punctata and 9-13 mm in ssp. sesban. The specimens in South-East Asia belong to ssp. sesban. Based on flower colour and hairiness, ssp. sesban has been further divided into 4botanica l varieties, which are, however, without much practical value because the distinguishing characters are weak and unstable. Ecology Sesban grows in areas with a semi-arid to sub-humid climate, with a rainfall between 500 and 2000 mm per year. In East Africa it grows up to an altitude of 2300 m as it can withstand cool temperatures, but not frost. It grows in a wide range of soil types ranging from loose sandy soils to heavy clays. It tolerates saline (1.0% salt con­ centration in seedling stage to 1.4% at maturity), alkaline (pH(H20) < 10) and acidic soils as well as waterlogging and flooding. However, sesban cannot withstand waterlogging immediately after Sesbania sesban (L.) Merrill- 1,flowering branch; germination or in the early stages of seedling de­ 2,fruit; 3, seeds. velopment. Although sesban tolerates low P 200 FORAGES levels, the application of P has a positive effect on > 4 m tall may even be detrimental. Cutting fre­ its growth and nodulation. quency depends on use, soil type and climate but Propagation and planting Sesban is propa­ can be as much as 5 times per year. A cutting gated by seeds.Scarificatio n ofth e seeds,usin g hot height of 75-100 cm is best. Too low (< 50 cm) and water or acid, improves germination. The seeds are too frequent cutting decreases the lifespan of sown in a well prepared soil. The planting distance plants. In order to increase longevity, it is advis­ depends on the purpose: as a fence, planting is in able to leave some foliage on the plant when har­ rows with a spacing up to 1-2 m; as an alley crop, vesting. Although usually fed fresh as a supple­ rows (single or double) are planted 2-10 m apart ment, sesban can also be dried. with a spacing of 25-50 cm between plants; as a Yield The fodder yields of sesban depend on soil fodderbank, rows are 1-2 mapar t and plants 25-50 type, soil fertility and moisture, climate and on cm apart. Seedlings will normally form root nod­ management factors, such as planting distance, ules with native rhizobia within 3-4 weeks after cutting height and frequency. Under favourable planting. Where the effective rhizobia are absent conditions DM yields of 20t/h a per year have been and effective nodulation does not take place, inoc­ obtained. The edible fraction (leaves and young ulation with an appropriate Bradyrhizobium twigs) ranges from 30-60% depending on the strain is necessary. Sesban has rapid early growth growing conditions and cutting frequency. In a and therefore overcomes weed competition easily subtropical environment DM yields of 5t/h a over and usually requires little maintenance. If neces­ 6 months have been obtained at a cutting height sary, the selective herbicide fluazifop can be used of 100c m and cutting frequency of 8weeks . post-emergence at 2 kg of active ingredient/ha to Seed production depends on growing conditions control grass weeds.Vegetativ e propagation using but can be as much as 1-2 t/ha. stem cuttings is possible but rarely practised. Genetic resources Sesban is the most widely Husbandry Sesban is normally used by cutting collected species of the genus Sesbania Adanson, the foliage and carrying it to the animals for fodder but only few accessions are of Asian origin; most or to neighbouring fields for green manure. Natu­ were collected in Africa. Germplasm collections ral stands of sesban are known to be browsed by are available at ILCA (Ethiopia), ATFGRC ruminants. Limited experience with the effects of (CSIRO, Australia) and the University of Hawaii direct grazing by cattle have shown that stems are (Waimanalo, United States). quite brittle and are easily broken but regrowth Breeding No breeding programmes have been below the break is rapid. Direct grazing by goats, started. however, resulted in over 80% plant mortality Prospects It is only recently that sesban has because of ringbarking 8-20 cm above ground received international attention as fodder. level. Although it responds to fertilizers, espe­ Research priorities are to collect germplasm, espe­ cially P and farmyard manure, their application cially in South-East Asia, to establish proper man­ is not a common practice. agement systems for alley cropping, to investigate Diseases and pests In general the reduction in the role of anti-nutritional factors in animal feed­ yield caused by diseases and pests is not large and ing and to develop a proper feeding strategy. The very few reports are available from South-East outcome of this research will determine further Asia. Larvae of the Azygophleps Scolaris are use of sesban. reported to tunnel in the stem. Seeds can be Literature ill Evans, D.O. & Macklin, B. (Edi­ infested with larvae of Bruchophagus mellipes tors), 1990. Perennial Sesbania production and (Hymenoptera) which reduce the seed yield. Nema­ use. A manual of practical information for exten­ todes can reduce the growth of the plants. In East sion agents and development workers. NFTA, Wai­ Africa, larvae of a leaf-feeding beetle Mesoplatys manalo, Hawaii. 41 pp. ;2j Evans, D.O. & Rotar, ochroptera can completely defoliate and kill ses­ P.P., 1987. Sesbania in agriculture. Westview ban. In Australia, larvae of the moth Orgyia au- Tropical Agriculture Series No 8.Westvie w Press, stralis have caused leaf rolling and drop of young Boulder, Colorado. 192 pp. |3| Gillett, J.B., 1963. leaves. In cool moist conditions in south-eastern Sesbania in Africa (excluding Madagascar) and Queensland, lesions of the fungus Botrytis cinerea southern Arabia. Kew Bulletin 17: 91-159. ^4i girdled the stem resulting in death of leaf and stem Lewis, G.P., 1988. Sesbania Adans. in the Flora above the lesion. Zambesiaca Region. Kirkia 13:11-51. !5'' Macklin, Harvesting Sesban can be cut after it reaches B. & Evans, D.O. (Editors), 1990. Perennial Sesba­ 1-2 m height. Leaving the first cutting until it is nia species in agroforestry systems. Proceedings SETARIA 201

of a NFTA/ICEAF workshop held in Nairobi, accumulates oxalate, especially when heavily fer­ Kenya, March 27-31, 1989. NFTA, Waimanalo, tilized with N. This is normally not a problem with Hawaii. 242 pp. |6j Nguyen van Thuân, Dy Phon, ruminants, such as cattle, provided they have been P., Niyomdham, C. & Vidal, Y., 1987. Légumi­ allowed to become used to setaria, but it is a prob­ neuses - Papilionoïdées. In: Morat, Ph. (Director): lem with horses, which develop a condition known Flore du Cambodge, du Laos et du Vietnam. Vol. as big-head {Osteodystrophia fibrosa). There are 23. Muséum National D'Histoire Naturelle, Labo­ 1200-1900 seeds/g. ratoire de Phanérogamie, Paris, pp. 58-59. Description A tufted, or more rarely rhizoma- J.H. Heering &R.C . Gutteridge tous, perennial with more or less erect stems to 3 m tall. Young tillers are strongly flattened in com­ mercial cultivars, with leaf-sheath prominently Setaria sphacelata (Schumach.) Stapf & keeled and often red-pigmented, sometimes hairy. Hubbard ex M.B. Moss Culms vary in number of internodes and diameter, depending on variety and cultivar, few and narrow Kew Bull. 1929:195 (1929). in 'Nandi', up to 17 and 6-12 mm in diameter in GRAMINEAE var. splendida; nodes hairless; the culms occasio­ ïn = 18(diploid) , 36,45, 54,72 ,9 0 (decaploid) nally branching when grown in fertile soils to give Synonyms multiple inflorescences. Leaf-blade 10-70 cm x - var. sphacelata: Panicum sphacelatum Schu­ 11-12 mm in var. sericea, up to 20 mm wide in var. macher (1827); splendida, hairless or with a few long hairs close - var. sericea (Stapf) W.D. Clayton: Setaria anceps to the junction with the sheath. Inflorescence an Stapf (1930); elongated cylindrical spike-like panicle 10-50 cm - var. splendida (Stapf) W.D. Clayton: Setaria long,th e spikelets borne in groups of2 o r 3o n short splendida Stapf (1930). Vernacular names General: setaria (En). - var. splendida: splendida, broadleafed setaria (En). Malaysia: sekoi. Philippines: bunga- bunga. Thailand: ya taiwan. Vietnam: co duôi chó, co ro'm. Origin and geographic distribution Setaria originates from and is widely distributed in trop­ ical and subtropical Africa. It also occurs in Yemen. The species was first brought into cultiva­ tion as a pasture plant in Kenya and has since been widely planted throughout the tropics and subtropics, especially in Africa, Asia and Austra­ lia. It has become naturalized in many of the coun­ tries to which it was introduced. In South-East Asia it is widely planted in Malaysia and Indone­ sia but is of only minor importance in Thailand. Uses Setaria is an important forage and is used under grazing and in cut-and-carry systems. The botanical variety splendida is reported to be one of the more commonly used species for cut-and- carry systems in Indonesia and Malaysia. Properties Nitrogen concentrations vary from over 3% in very young growth to under 1% in old growth. It is difficult to compare nutrient concen­ trations in different cultivars, asthe y flower at dif­ ferent times. Setaria has a higher moisture content than some other tropical grasses. In Australia accessions have been shown to differ quite widely Setaria sphacelata (Schumach.) Stapf & Hubbard in digestibility and a range of chemical compo­ ex M.B. Moss - 1, habit; 2, ligule; 3, inflorescence; nents, especially Na. Setaria is a species which 4,bristles; 5, spikelet. 202 FORAGES

branches, each spikelet subtended by 5-15 stiff seed production, associated with higher tiller fer­ bristles varying in length and colour; spikelets tility. 'Kazungula' produces more seed than other 2.5-3 mmlong , flat onth e side ofth e lower glume, cultivars, but becomes extremely stemmy when convex on the other, the lower glume about i, the mature. Hybridization between var. splendida and upper f the length ofth e spikelet; lower floret male var. sericea and selection in Australia led to the or sterile, the upper floret bisexual, the palea and release of the seed-producing cultivar 'Splenda', lemma hardened andenclosin g thecaryopsi s when which hasyielde d well in trials throughout South- ripe. East Asia andth eSout h Pacific. Growth and development Var. splendida is Ecology In natural grasslands at moderate to near-sterile, butsee d ofothe r varieties germinates high altitudes in East Africa, setaria can consti­ reasonably readily in good conditions and estab­ tute a considerable proportion of the herbage but lishes without difficulty. If the initial stand has a is rarely dominant. Setaria grows at higher alti­ low plant density, setaria has the capacity to tudes in the tropics than other panicoid pasture thicken upi f allowed toproduc e a seed crop. Flow­ grass species, and this has been associated with ering isvariabl e within most cultivars, with plants evolutionary physiological adaptation to sub-zero and tillers within a plant poorly synchronized. night temperatures. Varieties and cultivars differ Furthermore, a single inflorescence takes from in their adaptation and yield, although all require 1-7 weeks to complete flowering. Thus the earli­ a relatively moist environment and have low est-formed spikelets may be ripe and shedding drought tolerance. In general, setaria cultivars when the latest ones have only just flowered. This have some tolerance of intermittent waterlogging. poor synchrony, together with abscission of the 'Kazungula' is considered to be one of the better spikelet soon after ripening, is a major contribu­ adapted forages forth elowlan d wettropic s receiv­ tory factor to low seed production in this species. ing at least 1200m m annual rainfall. In Malaysia In subtropical climates, setaria generally pro­ it has been commonly recommended as an duces twosee d crops a year. improved forage for sowing in smallholdings and Other botanical information Themainl y Afri­ commercial enterprises since the early 1970s. can S. sphacelata is a polyploid complex, running 'Kazungula' cantolerat e brief periods of moisture from diploid to decaploid, with thedifferen t ploidy stress and is suited to well-drained soils of lower levels crossing freely without showing clearly dif­ fertility. The variety splendida isals o well adapted ferent morphological characters. For convenience to thehumi d lowland tropics. 'Nandi', 'Narok' and only, the enormous variability ofth e complex spe­ 'Solander' are better adapted to the subtropics or cies has been divided into 5 botanical varieties elevated tropics receiving more than 1000m mo f which, however, intergrade completely andd ono t annual rainfall. represent discrete entities. Only twovarietie s are Propagation and planting Most cultivars and in commercial use: varieties are sown by seed, except for var. splen­ - var. sericea: culms4-10-noded ,3- 6m mi ndiame ­ dida, which needs tob evegetativel y planted. This ter, up to 2 m tall; basal leaf-sheaths conspi­ has to some extent limited the usefulness of var. cuously flabellate; leaf-blades 3-10 mm wide, splendida in Malaysia and Indonesia. For vegeta­ glabrous; panicle 7-25c mlong ; bristles fulvous. tive planting, plants should be topped to a height - var. splendida: culms 6-16-noded, 6-12 mm in of about 15cm ,spli t into pieces of2- 3 tillers, and diameter, very stout, up to 3 m tall; basal leaf- planted without allowing them todr yout . Success­ sheaths often flabellate; leaf-blades 10-17m m ful establishment usually exceeds 90%, providing wide, glabrous; panicle 20-50 cm long; bristles the soil remains moist following planting. Setaria fulvous. seed should be sown at a rate of at least 2 kg/ha. Cultivars of var. sericea include 'Nandi', a diploid, Establishment is most successful in a fully-culti­ and 'Narok', 'Kazungula' and 'Solander', tetrap- vated seed-bed. Setaria may besow n with a compa­ loids. 'Narok' wasselecte d for its cool-season pro­ nion legume, such as Centrosemapubescens Benth. ductivity andwinter-greennes s in frost-prone sub­ or Neonotonia wightii (Wight & Arnott) Lackey, tropical environments, but seed production is and may produce a stable pasture mixture, al­ poor, owing toth elo w percentage oftiller s bearing though var. splendida and 'Kazungula' compete inflorescences, especially in older seed production aggressively with legumes andar eles s suitable for stands. This was partially remedied with the mixed pastures. release of 'Solander', which is similar to 'Narok' Husbandry Setaria should begrow n ona relati­ in winter-greenness andyiel d butha smuc h higher vely fertile soil or fertilized, particularly with N, SORGHUM 203

if it is to yield to its potential. It responds well to Although hybrids can be produced between tetrap- applied N and yields exceeding 30 kg dry matter loids and hexaploids, and between hexaploids and per kg of N applied have been reported. It also has octaploids, they are likely to be unstable as they a high requirement for K. It is tolerant of continu­ have an odd number of genomes. ous grazing and high grazing pressures, although Prospects Setaria has been shown to be a suc­ the latter would have an adverse effect on most cessful species in Malaysia and Indonesia, and it associate legumes. In subtropical Australia, well- can be expected that its usefulness will extend to fertilized setaria pastures sustained continuous pasture developments elsewhere inth e region. The stocking rates of up to 6steer s per hectare. Similar relatively reliable establishment of setaria when stocking rates could be anticipated in tropical sown by seed is a point in its favour. The new culti- regions without a pronounced dry/cool season. var 'Splenda' is likely to take over from 'Kazun­ Diseases and pests The leaf disease caused by gula', especially in flood-prone and high rainfall Pyricularia grisea is noted in Australia but is not environments. a serious problem. Inflorescence diseases caused Literature |1|Blair , G.J., Ivory, D.A. & Evans, by Sphacelotheca sp. and Fusarium nivale can be T.R. (Editors), 1986. Forages in Southeast Asian serious in Zaire and setaria bunt, caused by Tille- and South Pacific Agriculture. ACIAR Proceed­ tia echinosperma, can devastate seed crops in ings Series No 12. ACIAR, Canberra. 202 pp. |2| Kenya. It is considered that there is little likeli­ Bogdan, A.V., 1977. Tropical pasture and fodder hood of transmitting this disease in seed, as it is plants. Longman, London, pp. 249-261. |3| Evans, spread from inflorescence to inflorescence in the T.R . &Hacker , J.B. 1992.A n evaluation ofth e pro­ field. Seed crops in Australia may also be adver­ duction potential of six tropical grasses under sely affected by caterpillars. grazing. 2. Assessment of quality using variable Harvesting Setaria isbot h grazed and fed as cut- stocking rates. Australian Journal of Experimen­ and-carry forage and can also be made into hay or tal Agriculture 32:29-37 . !4:Hacker , J.B. & Evans, silage.Thicke r stemmed cultivars, such as 'Kazun- T.R., 1992.A n evaluation of the produption poten­ gula' or 'Splenda' are less satisfactory for hay than tial of six tropical grasses under grazing. 1. Yield finer stemmed cultivars such as 'Nandi'. and yield components, growth rates and pheno­ Yield Annual DM yields of as high as 31 t/ha logy. Australian Journal of Experimental Agricul­ have been recorded for var. splendida in Indonesia ture 32:19-27. |5|Halim , R.A. (Editor), 1989.Grass ­ and 19t/h a under regular defoliation in Malaysia. lands and forage production in South-East Asia. Trials in Malaysia gave higher DM yields for Proceedings of first meeting of the regional work­ 'Kazungula' than 'Narok' or 'Nandi', and 'Nandi' ing group on grazing and feed resources in South- also yielded poorly in trials at Khon Kaen, Thai­ East Asia, 27 Feb - 3 Mar, 1989. FAO, Rome. 216 land. In Australia, dairy cows grazing var. splen­ pp. |6|Oram , R.N., 1990.Registe r ofAustralia n her­ dida have produced more milk than cows grazing bage plant cultivars. CSIRO, Australia, pp. 50-53. cultivars of other setaria varieties. In beef-produc­ |7| Skerman, P.J. & Riveros, F., 1990. Tropical tion studies var. splendida and 'Narok' produced grasses. FAO, Rome. pp. 662-669. high levels of animal weight gain, although differ­ J.B. Hacker ences between cultivars are most pronounced dur­ ing the subtropical cool season. Genetic resources Setaria is an extremely Sorghum x almum Parodi diverse species which in Africa naturally covers a wide latitudinal and altitudinal range. The exis­ Rev. Argent. Agron. 10:36 1(1943) . tence of various intra-specific taxa, at least some GRAMINEAE of which can be inter-hybridized, is an indication 2n = 40 (tetraploid) of its morphological diversity. Germplasm re­ Vernacular names Columbus grass (En). Pasto sources are held by ATFGRC (CSIRO, Australia) colon, sorgo (Sp). Philippines: batag (Taga- and a smaller collection by CIAT (Columbia). log), gau (Ifugao). Thailand: ya-sokum. Breeding Hybridization between ecotypes may Origin and geographic distribution This spe­ be carried out without difficulty. Being a cross-pol­ cies, first noted in Argentina, is considered to be linated species, combining inflorescences in a bag a natural hybrid between S. halepense (L.) Pers. when at anthesis results in a high proportion of (Johnson grass) and S. bicolor (L.) Moench (grain hybrid seed. However, the existence of an exten­ sorghum). It has been introduced into some South- sive polyploid series is a barrier to gene transfer. East Asian countries, notably Thailand, but is 204 FORAGES

only used to a very limited extent. mm long, mostly with an awn about 1 cm long; Uses Used as a forage for grazing by cattle, or glumes brown or black, completely covering the less commonly for hay or silage, Columbus grass caryopsis at maturity. has now largely been replaced by the man-made Columbus grass may be distinguished from the hybrid cultivar 'Silk'. Columbus grass is mostly widespread weed species S. halepense by its short grown in pure stands. ascending terminal rhizomes and its 5-6.5 mm Properties Herbage of Columbus grass is of rea­ long sessile spikelets. S. halepense has some rhi­ sonably high quality and N concentrations in zomes which arise from side buds and elongated leaves can be over 3% if stands are adequately fer­ and sessile spikelets 4.5-5mm long. Columbus tilized, although concentrations decrease as grass cultivars are 'Crooble' and 'Nunbank' (Aus­ plants mature. It is palatable to cattle. Young her­ tralia), 'de Soto' (United States) and 'Rietondale' bage can accumulate dangerously high levels of (South Africa). HCN which may lead to prussic acid poisoning in Ecology Columbus grass is well adapted to livestock. Unlike its putative Johnson grass par­ heavy clay soils and areas receiving 500-800 mm ent, it can be eradicated without difficulty when rainfall. It can tolerate drought and some salinity. so desired. Stands are not killed by mild frosts. It is intolerant Botany Densely tufted perennial to about 3 m of waterlogged soils. tall, producing short, thick, ascending terminal Agronomy Columbus grass is a reliable seed rhizomes. Leaf-blade flat, 30-100 cm x 15-40 mm. producer and is sown by seed, either broadcast at Inflorescence an open panicle 20-60 cm long, the 20 kg/ha, or, preferably, drilled in rows 1 m apart lowermost branches in whorls of 4-9; spikelets in at 5-7 kg/ha. Seedlings are vigorous and the spe­ short fragile racemes, paired, the lower one 5-6.5 cies normally establishes without difficulty. For­ age yields tend to be best in the season of sowing, and, although in exceptional circumstances it may persist for five years or more, mostly stands are ploughed out after the third year. Columbus grass seeds profusely and, as there is little loss from shedding, seed yields are high. Columbus grass is commonly grown on more fer­ tile soils and only low levels of N fertilization are required. On less fertile soils, the species responds well to applied N. Colombus grass is susceptible to leaf blight (Hel- minthosporium turcicum) and leaf rusts (Puccinia spp.). Columbus grass is normally grazed, and topping to a height of 30-40 cm after grazing has been rec­ ommended. Stands of Columbus grass should not be grazed within 3 months of sowing nor should young regrowth be grazed, so as to avoid the risk of prussic acid poisoning. For hay or silage, stands should be cut at early flowering stage. Dry matter yields as high as 19 t/ha per year have been recorded from two cuts. In farm practice, 4-10 t/ha per year can be expected, depending on rainfall and fertility. On an annual basis, livew- eight gains of cattle of up to 0.5 kg/head per day can be expected, while weight gains of 1.3 kg/head per day have been recorded over shorter periods. Genetic resources and breeding Due to its probable hybrid origin and the large diversity of Sorgum x almum Parodi -1, base ofplant; 2, culm- one of the parents, there is potential for extensive and leaf- part; 3, inflorescence; 4, sessile spikelet; variation within S. xalmum which may be 5,pedicelled spikelet; 6, caryopsis. exploited by the plant breeder. Columbus grass is SORGHUM 205

predominantly cross-pollinating. It may be hybri­ dized with Johnson grass, also a tetraploid, and with the diploids S. bicolor and S. x drummondii (Steud.) Millsp. &Chase . A limited germplasm col­ lection is held by ATFGRC (CSIRO, Australia). Prospects Columbus grass isuse d only to a very limited extent in South-East Asia and it isno t anti­ cipated that its usefulness will increase as better perennial are available. Literature II Bogdan, A.V., 1977. Tropical pas­ ture and fodder plants. Longman, London, pp. 264-269. .21 Oram R.N., 1990. Register of Austra­ lian herbage plant cultivars. CSIRO, Australia, p. 56.13!Silvey , M.W. &Ferraris , R., 1988.Swar d pro­ ductivity and seasonal animal production from pe­ rennial forage sorghums grazed at four stocking rates. CSIRO, Australia, Division of Tropical Crops and Pastures, Technical Memorandum No 56. 22 pp. 41Yates , J.J., Edye, L.A., Davies, J.G. & Haydock, K.P., 1964. Animal production from a Sorghum almum pasture in South-east Queens­ land. Australian Journal of Experimental Agricul­ ture and Animal Husbandry 4:326-335 . J.B. Hacker

Sorghum, cultivar 'Silk' - 1, habit; 2, ligule; 3, Sorghum, artificial perennial hybrids inflorescence; 4,pair of'spikelets.

GRAMINEAE and stems to 4 m tall, with or without a few short In = 20,4 0 rhizomes. Leaf-blade 25-40 mm wide, glabrous Vernacular names General: artificial perenni­ except for a few hairs close to the membranous al hybrids are usually known by their cultivar ligule. Inflorescence a large open pyramidal pani­ names, e.g., 'Silk', 'Krish'. Philippines: batag cle with secondary and tertiary branches, the (Tagalog), batad (Bikol), bukakau (Ilokano). Laos: branches sometimes drooping; spikelets in short, liiay, khauz f'angx. Thailand: ya khao fang. fragile racemes, paired, the lower one sessile and Origin and geographic distribution Being bisexual, with rigid shiny glumes which may be man-made hybrids, these are confined to areas straw-coloured, reddish-brown or black, depend­ where they have purposefully been sown or have ing on cultivar, and which tightly enclose the car- since become naturalized. They have only been yopsis when ripe, the upper pedicelled and male; grown to a limited extent in South-East Asia, but lemmas often awned. 'Silk' is widely grown in subtropical Australia. 'Krish' is a hybrid between S. halepense (L.) Pers. Uses Perennial sorghum hybrids are grown as and S. roxburghii Stapf; it has not been utilized short-duration perennial forages for grazing by commercially to any great extent. 'Silk' is a hybrid cattle, but can be used for cut-and-carry forage or between 'Krish' and S. arundinaceum (Desv.) for hay and silage. Stapf. In the subtropics, 'Silk' may easily be estab­ Properties Foliage is very palatable to live­ lished in spring and shows rapid early growth, stock and 'Silk' has greater in vitro digestibility early tillering and an ability to compete with than S. x almum Parodi ('Crooble'). Total free weeds. In contrast, 'Krish' has poor seedling sugars in the stem averaged about 20%. In com­ vigour and growth in spring is also slow; however, mon with other sorghum species, young herbage it outyields other sorghums provided it is not defo­ can accumulate dangerously high levels of HCN liated before 10 weeks of growth. 'Krish' flowers which may lead to prussic acid poisoning in lives­ much later than 'Silk', which is later flowering tock. There are 120-160 seeds/g. than S. x almum cultivar 'Crooble'. 'Krish' has Botany Erect perennials with numerous tillers been used as a source of resistance to sugar-cane 206 FORAGES mosaic virus in the breeding of grain sorghum (S. In Australia the difficulty in distinguishing be­ bicolor (L.) Moench). One of the merits of 'Krish' tween seeds of 'Silk' and the noxious weed S. hale- and 'Silk' is that, unlike the weed-parent S. hale- pense has led to efforts to produce a cultivar with pense, they can be eradicated without difficulty tan-coloured seeds by selection within 'Silk'. and hence dono t pose a problem to any subsequent Prospects The perennial sorghums have poten­ crop. However, this does not necessarily apply to tial in sub-humid regions where yearly cultivation other hybrid combinations. for sowing annual forages is undesirable and Ecology Perennial sorghums are well-adapted where permanent pastures are not required. In to high temperatures and are well suited to areas more humid regions, over a period ofsevera l years, receiving from 500-800 mm annual rainfall to peri­ annual forages are higher yielding when yields are ods of moisture stress. In Queensland, Australia, averaged over the lifespan of the perennial for­ 'Silk' is widely grown on clay soils, and has re­ ages. placed S. x almum as a sown forage in these situa­ Literature |1|Ferraris , R. &Silvey , M.W., 1988. tions. Sward productivity and seasonal animal produc­ Agronomy Perennial sorghum pastures may be tionfro m perennial forage sorghums grazed at four established by seed into a cultivated seed-bed, stocking rates. CSIRO, Australia. Division of either by broadcasting at 20-25 kg/ha or drilled Tropical Crops and Pastures, Technical Memoran­ in 1 m rows at 5-7 kg/ha. Perennial sorghum is dum No 56. 22 pp. |2| Oram, R.N., 1990. Register of usually sown in pure stands but can be sown with Australian herbage plant cultivars. CSIRO, Aus­ other grasses with or without legumes. Initally the tralia, pp. 66-68. mixtures grow more slowly, but they are more per­ J.B. Hacker sistent. The sorghum then provides early grazing and contributes to the suppression of weeds. Pe­ rennial sorghums are compatible with cool season Sorghum x drummondii (Steud.) Millsp. forages such as lucerne (Medicago sativa L.), annu­ & Chase al Medicago spp. or oats (Avena sativa L.) and warm season legumes such as Neonotonia wightii Publ. Field Columb. Mus. Bot. 3:2 1 (1903); Flora (Wight &Arnott ) Lackey or Lablab purpureus (L.) of Trop. E. Africa. Gram. (3):72 6(1982) . Sweet. Young plants or young regrowth should not GRAMINEAE be grazed owing to risk of prussic acid poisoning. 2n = 20 (counts of 40 are also reported, but Perennial sorghums are normally grazed, al­ require confirmation) though they are suitable for cut-and-carry systems Synonyms Sorghum sudanense (Piper) Stapf of management or for hay or silage if not allowed (1917), S. bicolor (L.) Moench ssp. drummondii to become too coarse. A recovery period in spring (Steud.) deWe t (1978). may be necessary to ensure persistence if pastures Vernacular names Sudan grass (En). Pasto have been heavily grazed during the cool season. Sudan (Sp).Philippines : batag (Tagalog), bukakau In trials in Australia which led to its release, 'Silk' (Ilokano), layagah (Sulu).Thailand : ya-sudan. consistently outyielded S. x almum 'Crooble' by Origin and geographic distribution What is 24-78%. 'Krish' outyields other forage sorghums known commercially as Sudan grass is a segregate late in the growing season. Under continuous graz­ from a natural hybrid between S. bicolor (L.) ing in sub-humid Queensland, Australia, steers Moench and S. arundinaceum (Desv.) Stapf. This grazing at stocking rates of 1.5-3 steers/ha aver­ hybrid combination is responsible for several aged liveweight gains of 150kg/hea d per year. other 'species' which are, however, unstable, and 'Krish' shows a high degree of resistance to rust revert to one or other of their parental types. They and blight, also to sugar-cane mosaic virus. have all been included in the binomial S. x drum­ Genetic resources and breeding Breeder's mondii in the Flora of Tropical East Africa. The seed ofhybri d cultivars and stocks of parental spe­ hybrid originated in the region from southern cies are held by ATPGRC (CSIRO, Australia). The Egypt to the Sudan, but has rarely been collected perennial hybrid sorghums are cross-pollinating there. It was introduced to the United States in plants which show a considerable degree of intra- 1909 and rapidly became popular as a forage, later varietal genetic diversity. Many perennial hybrid being evaluated and sown in other regions with combinations have been produced experimentally warm and dry growing season. and the wide range in variability in the genus Uses Sudan grass is used as an annual forage for allows the possibility of new improved cultivars. ruminants, but more commonly as a parent in a SORGHUM 207

wide range of inter-specific F1hybrid s in which S. bicolor, grain sorghum, is the alternative parent. Examples of such hybrids are 'Sudax', 'Zulu' and 'Bantu'. In Thailand, sorghum x sudan hybrids are utilized asfres h cut-and-carry forages or as hay for dairy cattle or water buffaloes. Properties Nitrogen concentrations can be as high as 3%, although levels fall with increasing maturity. Sudan grass is very palatable to live­ stock so the level of utilization is high and cattle are reported to consume 80% of the forage at pani­ cleemergence . In common with other sorghum spe­ cies, young herbage can accumulate quite high levels of HCN which may lead to prussic acid poi­ soning in livestock. This is rarely a serious prob­ lem, except where high levels of N fertilizer have been applied or where the grass is wilted. Prussic acid poisoning is considered to be less of a problem with Sudan grass than it is with either fodder sor­ ghum or sorghum-Sudan hybrids. There are 90-120 seeds/g. Botany Annual with erect stems to 3m tall, 3-9 mmthick . Leaf-blade lanceolate, 30-60 cm x 8-15 mm. Inflorescence an open pyramidal panicle with secondary and sometimes tertiary branches which end in short fragile racemes, which do not readily break up at maturity; spikelets paired; sessile spik- elet 6-7 mm long; glumes loosely hairy, shiny and Sorghum x drummondii (Steud.) Millsp. & Chase almost hairless when mature; upper lemma with - 1, habit flowering plant; 2, inflorescence; 3, long- an awn up to 16m m long; pedicelled spikelet about awned spikelet; 4,pedicelled and sessile spikelet; 5, as long as the sessile spikelet, but narrower. Car- caryopsis. yopsis variable, enclosed by the glumes. Seedlings emerge 5-6 days after sowing. Flower­ Ecology Sudan grass isno t adapted to the humid ing in Sudan grass is to some extent photoperiod- tropics, but is suited to warm conditions with low sensitive, but in the tropics it flowers freely. As humidity and an average annual rainfall of the racemes do not break up easily, seed produc­ 600-900 mm. It is intolerant of waterlogging but tion of Sudan grass is easier than it is with many has reasonable tolerance of salinity. tropical grasses, even though flowering within in­ Agronomy Sudan grass should be sown in a dividual plants is poorly synchronized. A number well-prepared seed-bed, either broadcast or prefer­ of 'cultivars' which are marketed as Sudan grass ably drilled in rows 25-50 cm apart (wider in drier are, in fact, hybrids with other species. There are climates) and not deeper than 2.5 cm, at seeding relatively few cultivars available within S. x rates of 8-12 kg/ha when drilled and 12-16 when drummondii as such, but considerable numbers of broadcast. Higher rates have been recommended open-pollinated and Yl hybrids have been devel­ inmor e humid areas.I t isnormall y grown as a pure oped. Examples, and their parentage, are: 'Tift' crop, although companion legumes such as Glycine (Sudan grass x 'Leoti' sweet sorghum) x Sudan max (L.) Merrill and Vigna unguiculata (L.) Walp. grass; 'Piper' ('Tift' x Sudan grass); 'Greenleaf have been used. It responds well to irrigation in ('Leoti' sweet sorghum x Sudan grass); 'Lahoma' dry climates. Relatively low levels of fertilizer are (Sudan grass x 'Leoti' sweet sorghum); 'Sucro'- normally used, but it responds to N, P and K fertil­ (perennial) (S. x almum x perennial sweet Sudan izers where these nutrients are deficient in the soil. grass); 'Sudax^FJ (male sterile grain sorghum x Cultivars differ in resistance to leaf diseases. Sudan grass); 'Zulu^Fj) (male sterile grain sor­ These diseases are of greater concern where the ghum x 'Greenleaf) (see above); 'Bantu'(Fj) (male crop is grown under conditions of higher humidity sterile grain sorghum x 'Piper') (see above). and rainfall than it is normally suited to. 208 FORAGES

Sudan grass may begrazed , chopped andfe d direct­ always in coastal areas. In cultivation it occasio­ ly, or made into hay or silage. As stems are thin, nally occurs outside its natural range. compared with the perennial sorghum species, it Uses St. Augustine grass is used for grazing in 'dries quickly, which is an advantage when hay­ open areas, but is also showing considerable prom­ making. Yields of green fodder average 20-40 t/ha isefo r use under plantation crops. It is also a popu­ but may be twice as high with optimal fertilizer lar lawn grass and is used for soil conservation. use. Milk production may be low if sulphur is Properties The quality of St. Augustine grass limited in Sudan grass forage. declines throughout the growing season, with N Genetic resources and breeding Genetic concentrations between 2.6% and 2.0% . Digestibi­ resources within S. x drummondii are limited, but lities of crude protein decline from 53% to 31% there is a considerable genetic resource associated and DM digestibilities from 60% to 50%. It is pal­ with closely related species with which it can be atable when young,bu t palatability ofol d material hybridized. Limited germplasm is maintained is low. under the binomial S. sudanense at ATFGRC Botany A stoloniferous perennial with upright (CSIRO, Australia). More extensive collections of or ascending stems, often much branched, 10-50 S. x drummondii and related hybrids are held by cm tall. Leaf-sheath tightly compressed and the USDA (Fort Collins, Colorado and Beltsville, keeled; ligule a ring of hairs, 0.5 mm high; leaf- Maryland) and ICRISAT (India). blade oblong-linear, 3-15 cm x 4-10 mm, plicate Prospects In a number of countries, commercial when young, obtuse, glabrous, slightly bluish. companies have a major interest in production of Inflorescence terminal or axillary, 5-10 cm x F: hybrid forages, and it can be anticipated that 5-10 mm, composed of 10-20 racemes each 0.5-1 there will be a continuing flow of new hybrid culti- vars with improved characteristics. As with exist­ ing cultivars, these are likely to be better adapted to the less humid parts of South-East Asia. Literature jl| Bogdan, A.V., 1977. Tropical pas­ ture and fodder plants. Longman, London, pp. 275-280. |2| Clayton, W.D. & Renvoize, S.A., 1982. Gramineae (Part 3). In: Polhill, R.M. (Editor): Flora of tropical East Africa. A.A. Balkema, Rot­ terdam, p.726 . |3|Oram , R.N., 1990.Registe r of Aus­ tralian herbage plant cultivars. CSIRO, Australia, pp. 61-66. |4| Skerman, P.J. & Riveros, F., 1990. Tropical grasses. FAO, Rome. pp. 690-694. |5| Stobbs, T.H. & Wheeler, J.L., 1977. Response by lactating cows grazing sorghum to sulphur supple­ mentation. Tropical Agriculture, Trinidad 54: 229-234. J.B. Hacker

Stenotaphrum secundatum (Walter) O. Kuntze

Revis, gen. pi. 2:79 4(1891) , ('secundum'). GRAMINEAE 2n = 18,2 7(triploid) , 36(tetraploid) , 54an d 72 Vernacular names St. Augustine grass, crab grass, buffalo grass (En).Gro s chiendent (Fr). Viet­ nam: co'quai chèo. Origin and geographic distribution Although its natural distribution is on the shores of the Stenotaphrum secundatum (Walter) 0. Kuntze - 1, Atlantic Ocean, St. Augustine grass is now found flowering plant; 2, ligule; 3, detail of inflorescence; quite extensively in Australia and the Pacific, too, 4,spikelet in two views. STYLOSANTHES 209 cmlon g and bearing 1-3 spikelets;mai n axis thick, Augustine grass yielded 5.5 t/ha of DM per year corky, flat on one surface, deeply hollowed out on in full sunlight, as measured over 6 harvests. Cor­ the other, each cavity containing a raceme, borne responding yields under 70%, 50%, 40% and 20% alternately on either side of a wavy midrib; spike- sunlight were 3.5,4.0,4.9 and 1.9 t/ha. At 40% light lets sessile, lanceolate, up to 5 mm long; lower transmission St. Augustine grass can attain its glume 1-2 mm long, nerveless; upper glume 4-5 maximum yield and can outyield signal grass (Bra- cm long, 5-7-nerved; lower floret neuter, upper chiaria decumbens Stapf). floret bisexual. Caryopsis oblongoid to obovoid, Nitrogen-fertilized pastures of St. Augustine grass about 2m m long. have produced over 1000 kg/ha of liveweight gain A form with abnormally slender inflorescences per year in the United States. St.Augustin e grass- occurs in South Africa and Brazil (called 'Natal- siratro (Macroptilium atropurpureum (DC.) Plata deme' by Sauer); a form with almost strobi- Urban) pastures beneath a sparse coconut planta­ loid inflorescences is a sterile triploid and occurs tion in Vanuatu gave annual liveweight gains of in South Africa, Australia and the Pacific islands 275-400 kg/ha. However, only 1-2 head/ha could (called 'Cape deme' by Sauer). be sustained during the growing season under a The species only flowers occasionally in the wet denser stand of coconuts at a 9m spacing. tropics, hence seed production is very poor. The Genetic resources and breeding Evaluation most popular cultivar for lawns, 'Roselawn', is of a range of accessions of St. Augustine grass is used in several countries. being carried out in South-East Asia by ACIAR. Ecology St. Augustine grass is primarily a Prospects With the recent emphasis on develop­ coastal pioneer. It occurs at altitudes from sea- ment of farm systems involving the integration of level up to 800 m, and could well grow at higher livestock and plantations in South-East Asia and elevations than this as it is one of the more eold- elsewhere in the tropics, further research on the and frost-tolerant tropical and warm temperate shade-tolerant St. Augustine grass is warranted. grasses. It grows in relatively humid areas, prefer­ Improving its productivity and seed production ring fertile soils, although it is also well-adapted would be priority objectives. to organic sandy soils in Florida (United States) Literature |1|Bogdan , A.V., 1977. Tropical pas­ and to alkaline soils. In Puerto Rico it grows well ture and fodder plants. Longman, London, pp. on soils rich in lime and on sandy soils on sloping 284-285. :2;Reynolds , S.G., 1988.Pasture s and cat­ hillsides. It responds to fertilization on poorer tle under coconuts. FAO Plant Production and soils. It tolerates short-term flooding and salt Protection Division Paper 91. FAO, Rome. pp. spray, but it does not persist where there is a pro­ 71-72. ;3| Samarakoon, S.P., Shelton, H.M. & Wil­ longed dry season and on soils with a shallow son, J.R., 1990.Voluntar y feed intake by sheep and water table. St. Augustine grass is shade-loving digestibility of shaded Stenotaphrum secundatum and produces higher yields under shade intensities and Pennisetum clandestinum herbage. The Jour­ of as low as 40% sunlight than in full sunlight. Its nal of Agricultural Science, Cambridge 114: productivity is maintained with 40% full sunlight. 143-150. j4j Sauer, J.D., 1972. Revision of Steno­ Agronomy St. Augustine grass is propagated by taphrum (Gramineae: ) with attention to planting sections of rooted stolons, preferably its historical geography. Brittonia 24: 202-222. 5; early in the wet season. The sections can be Skerman, P.J. & Riveros, F., 1990. Tropical planted about 30 cm apart in rows 60-70 cm apart, grasses. FAO, Rome. pp. 711-713. or else scattered on the soil surface and disked in. C.P. Chen One ha of grass will provide sufficient cuttings to plant 10 ha. The stolons grow quickly, although it may still take 5-6 months to form a complete Stylosanthes capitata Vogel cover. Sward formation is faster under light to moderate shade than in the open. Once estab­ Linnaea 12:7 0(1838) . lished, the dense swards resist weed invasion. LBGUMINOSAE Because of its low growth habit, St. Augustine In = 40 grass is usually grazed, although it can be cut for Origin and geographic distribution S. capi­ hay or silage. It should be closely grazed, and graz­ tata originates in South America where it occurs ing every 12-14 days to a height of 5 cm or less naturally in sub-humid and dryland areas in north­ has been suggested. When used as a lawn, it should eastern Venezuela and, mainly, in central-west, be regularly and closely mown. In one study, St. south-east and north-eastern Brazil. One cultivar 210 FORAGES and several experimental lines have spread to x 5-15 mm, apically acute, mostly densely villous other tropical regions for testing, including South- on both surfaces, with 8-12 pairs of conspicuous East Asia. veins. Inflorescence a capitate spike, terminal or Uses The main use of S. capitata is as forage in axillary, cylindrical-ovoid, upt o 7c m x 15-20 mm, permanent pastures grazed by ruminants. many-flowered; often several inflorescences in Properties S. capitata provides a palatable for­ dense clusters; bracts 1-foliate, oblong, 9-13 mm age ofmoderat e to high quality, with N concentra­ long, with 3-5 pairs of veins, and variable pilosity; tions ranging from 2-3.0%, and DM digestibility flowers papilionaceous, small, with obovoid stan­ 55-60%. Phosphorus concentrations in the her­ dard 4-6 mm long, sulphur-yellow; axis rudiment bage are moderate to low (0.09-0.18%). There are and 2 inner bracteoles present. Pod 2-articulated, 400-450 seeds/g. 5-7 mm x 2-2.5 mm, reticulately nerved; both Description A perennial herb or sub-shrub, articles usually fertile, the upper one glabrous and semi-erect to erect, with a strong taproot. Stems with a straight to uncinate beak about 1m m long. many-branched, lignified at the base; under com­ Seed colour varying from yellow, sometimes petition ascending up to 1 m; indumentum varying slightly mottled, to almost black. from glabrous to densely pilose, or with scattered Growth and development Initial growth of S. bristles. Leaves 3-foliolate; stipules oblong, 16-20 capitata is slow. Flowering and seed setting occur mm long (including teeth) and 6-8 mm wide, with mainly during the end of the rainy season and 2-3 pairs of veins; petiole 2-6 mm long, densely extend into the dry. Being a very prolific seeder, villous; rachis up to 3.5 mm long; leaflets broad- S. capitata regenerates through seedling recruit­ elliptical, oblong or sometimes obovate, 15-40 mm ment. If plants are allowed to set seed in the year of establishment, more than 11o f seeds in pods per ha can be produced. The species tends to behave like an annual and plants seldom live longer than 2-3 years. Other botanical information To date, S. capi­ tata has become a commercial pasture legume only in Colombia, South America, where a blend of five similar ecotypes was released as 'Capica'. The objective of blending was to broaden the genetic base of the cultivar and to decrease the risk of anthracnose susceptibility. Ecology S. capitata is best adapted to tropical savanna ecosystems with a sub-humid to humid cli­ mate characterized by relatively high rainfall (1000-2500 mm/year) during a growing season of 6-9 months. The species requires light-textured, well-drained, acid soils, and has a good tolerance of toxic levels of soil Al and Mn. It grows well on soils of poor fertility, including low available P. Savanna ecosystems with such soils are common in tropical America; their South-East Asian equiv­ alents would be Imperata cylindrica (L.) Raeuschel grasslands in somewhat drier areas with ultisols as the predominant soil type. Agronomy S. capitata is drilled in rows or broadcast, preferably with a grass, at a rate of 2-3 kg/ha. Germination of fresh seed is poor because of a high degree of hard-seededness; this can be overcome by mechanical or acid scarification,.or by hot-water treatment. Successful S. capitata as­ sociations have been obtained with Andropogon Stylosanthes capitata Vogel - 1, habit flowering gayanus Kunth (gamba grass), Brachiaria decum- and fruiting branch; 2,fruit with axis rudiment. bens Stapf (signal grass) and Brachiaria dictyo- STYLOSANTHES 211

neura (Fig. &D e Not.) Stapf. Although the legume Stylosanthes guianensis (Aublet) nodulates with native rhizobia, inoculation of seed Swartz with a Bradyrhizobium strain ofknow n effectiven­ ess is recommended. Svenska Vet. Akad. Handl. 10:30 1(1789) . Fertilization with P and K enhances establish­ LEGUMINOSAE ment, and despite its low soil-fertility require­ 2n = 20 ments, S. capitata responds to maintenance fertil­ Synonyms Trifoliumguianense Aublet (1775). ization with these nutrients. It persists well in Vernacular names Stylo, Brazilian lucerne association with a strongly competitive grass such (En). Luzerne du Brésil, luzerne tropicale (Fr). as Andropogon gayanus, even under fairly uncon­ Thailand: thua-satailo. trolled grazing, but the use of intermittent grazing Origin and geographic distribution The natu­ systems is suggested. ral distribution of stylo ranges from northern S. capitata has proved to be very tolerant of an- Argentina into Mexico. The variety guianensis, thracnose (Colletotrichum gloeosporioides), the which is of particular interest to South-East Asia, major disease affecting Stylosanthes spp. There has its centre of origin in Brazil and is naturally are, however, regional differences, depending on distributed to Paraguay, Bolivia, Peru, Colombia, anthracnose-strain virulence. A common pest is Venezuela, Guyana and Central America. During the pod-borer Stegasta bosqueella; it can drasti­ the 20th Century it has been introduced all over cally decrease seed yield. the tropical world and is now widely naturalized In pure legume stands, DM yields of 1-3 t/ha every in most tropical countries. 12 weeks are obtained during the rainy season. Uses Stylo isuse d as a cover crop in plantations, Dry-season yields can be very low, as plants will as a green manure crop and as a fallow crop in eventually shed their leaves completely under shifting cultivation, but it is best known as a pas­ severe drought. Nevertheless, year-round animal ture legume for humid tropical regions. production can be improved due to the presence Properties Nitrogen concentrations'range from of S. capitata in a pasture. For example, in a mix­ 1.5-3.0%. Dry matter digestibility of young plant ture with A. gayanus, liveweight gains of 150 kg/ material lies between 60% and 70%, but with steer per year have been measured compared with increasing age and lignification this may be 110kg/stee r per year in A.gayanus alone. reduced to below 40%. Because stylo is able to Genetic resources and breeding The species grow on poor soil, its P concentration is often as has been widely collected and there is a large, very low as 0.06%, but with P fertilization it can variable collection available at CIAT (Colombia). increase to over 0.30%) of DM. Stylo can grow ade­ A S. capitata breeding programme, aimed at adapt­ quately on soils with a low phosphate availability, ing 'Capica' to the higher anthracnose-stress con­ in which case animals feeding on the plant need ditions of central Brazil is now being concluded. to be supplemented with P for their normal requir­ Prospects Because of such characteristics as ements. Although stylo is readily eaten by cattle outstanding adaptation to acid and poor soils, dis­ and sheep, its palatability is not very high, which ease resistance, long-term persistence through protects it from being overgrazed. As with many seedling recruitment as a result of high seed pro­ other tropical legumes, young growth of stylo is duction, and good forage quality, S. capitata is an less palatable than young growth of grasses. important pasture legume for low-input produc­ Description A perennial herb or sub-shrub, tion systems on marginal soils in tropical savanna semi-erect to erect, with a strong taproot and small regions. Efforts to improve its dry-season perfor­ round root nodules. Stem much branched, herba­ mance seem to be justified. ceous or lignified at the base, to 1m tall, indumen­ Literature |1| Instituto Colombiano Agrope- tum varying from nearly glabrous to densely cuario, 1983.Capic a (Stylosanthes capitata Vog.). pilose, often with bristles and viscid. Leaves trifo- Boletin Técnico No 103. ICA, Bogota. 12 pp. ;2j liolate; petiole 1-12 mm, rachis 0.5-1.5 mm long; Stace, H.M. &Edye ,L .A .(Editors) , 1984.Th e biolo­ stipules 2-15 mm, adnate to the petiole, teeth 2-10 gy and agronomy ofStylosanthes . Academic Press, mm long; leaflets elliptical to lanceolate, 5-45 mm Sydney. 636 pp. |3| Thomas, D. & Grof, B., 1986. x 2-20 mm, not more than 8 times longer than Some pasture species for the tropical savannas of wide, indumentum varying as on stems. Inflores­ South America. I. Species of Stylosanthes. Her­ cence a loosely capitate spike, terminal or axilla­ bage Abstracts 56:445-454 . ry, with more than 4flowers . Flower subtended by R. Schultze-Kraft an outer bract with 3-7 mm long sheath, a 2.5-5.5 212 FORAGES

gracilis (Kunth) J. Vogel (syn. S. gracilis Kunth), but this has no agronomic value. Several cultivars of var. guianensis have been developed in Austra­ lia and South America. The cultivars released in Australia are : 'Schofield', adapted to hot humid climates; 'Cook', similar to 'Schofield' but earlier flowering, more branched and better adapted to lower temperatures; 'Endeavour' intermediate be­ tween cultivars 'Schofield' and 'Cook'; 'Graham', the earliest flowering cultivar and better adapted to subtropical conditions. 'Pucallpa', released in Peru specifically for the humid tropics and soils with a low pH. This cultivar is also widely used in tropical China as 'Pi Hua Dou 184'.Var . interme­ dia (Vogel) Hassler 'Oxley' (known as fine stem stylo) is adapted to subtropical sub-humid regions with 700-1000 mm annual rainfall and is frost-tol­ erant. The name S. guianensis is sometimes erroneously spelled S. guyanensis. Ecology Stylo is adapted to hot, humid climates, and is neither frost nor drought tolerant. It grows on all soil types, hut is particularly well adapted to poor acid soils with high Al and Mn contents. Stylosanthes guianensis (Aublet) Swartz- 1, habit Stylo is a short-day plant with a critical photoperi- flowering and fruiting branch; 2, fruit. od of between 12-14 hours, depending on cultivar. However, some cultivars have been reported to mm long outer bracteole and a 2-4.5 mm long inner require exposure to long days prior to short days bracteole; calyx tube 4-8 mm long, lobes 3-5 mm; before flowering isinitiated . Cultivars with a criti­ standard 4-8 mm x 3-5 mm, yellow to orange, cal photoperiod about 12 hours will only flower often with black stripes, wings and keel 3.5-5 mm sporadically at the equator because of daylength long. Pod usually 1-jointed, the article ovoid, 2-3 requirements. mm x 1.5-2.5 mm, glabrous or rarely with very Propagation and planting Stylo is propagated short pubescence, indistinctly veined, with a by seed. Hot water treatment (10minute s at 80°C ) minute beak, strongly inflexed. Seeds pale brown to break hardseededness improves germination or purple. rates. Seeding rate is 2-6 kg/ha. When sown Growth and development Stylo plants have a together with grasses a good seed-bed preparation juvenile phase during which floral initiation does is desirable, but when sown into an existing pas­ not take place. Stylo is a copious seed producer, ture, little or no seed-bed preparation is necessary but more than 70% may be hardseeded. Hardsee- although establishment will be slower. Stylo can dedness breaks down naturally under hot condi­ also be grown in a pure stand as a green manure, tions and can also be broken by dry or wet heat cover or fodder crop. It is suitable for small- and before sowing. large-scale agriculture. As it fixes atmospheric N, stylo contributes to Husbandry Stylo can be used by continuous or increased soil fertility, although it has been com­ rotational grazing or cutting. It responds to pared unfavourably with other legumes in this res­ improved soil fertility, particularly P, but can pect. It nodulates freely with cowpea rhizobia and grow on infertile soil. does not require inoculation. It is sometimes sown as a pioneer species in a mix­ Other botanical information In S. guianensis ture of legume species. In this role, it provides for­ 7varietie s are distinguished by morphological and age early in the life of the pasture, but is not ecological characteristics. Only var. guianensis is expected to persist in the long term. important for South-East Asia and the information Diseases and pests Stylo is susceptible to an- given here refers mainly to this variety. Another thracnose disease, caused by the fungi Colletotri­ variety adapted for humid tropical regions is var. chum gloeosporioides and C.dematium, the former STYLOSANTHES 213 being the more important one. The disease was Boulder, Colorado, United States, pp. 141-181. ;2 first reported from Brazil, but has now spread Skerman, P.J., Cameron, D.G. &Riveros , F., 1988. throughout the world bytranspor t ofinfecte d seed. Tropical forage legumes. FAO, Rome. pp. 394-407. Symptoms of the infection are black lesions on the 3. Stace, H.M. & Edye, L.A. (Editors), 1984. The leaves and stems, which eventually lead to the biology and agronomy of Stylosanthes. Academic death ofth e plants. Although the fungi can be con­ Press, Sydney, Australia. 636 pp. j4i 't Mannetje, trolled chemically, this isno t an economic proposi­ L., 1977. A revision of varieties of Stylosanthes tion, except for valuable seed crops. The approach guianensis (Aubl.) Sw.Australia n Journal of Bota­ being followed in Australia and Colombia is to ny 25: 347-362. select resistant cultivars. It has been reported by L. 't Mannetje CIAT (Colombia), that stylo is more resistant to the disease in humid than in seasonally dry trop­ ical regions, because of the presence of microor­ Stylosanthes hamata (L.) Taub. ganisms antagonistic to the fungi. Harvesting Stylo is harvested by grazing ani­ Verh. Bot. Ver. Brandenb. 32:2 2(1890) . mals or it ismow n for stall feeding or artificial dry­ LEGUMINOSAE ing. When mown, care should be taken not to cut 2n = 20, 40 woody stems too low, otherwise regrowth will be Synonyms Hedysarum hamatum L. (1759), Sty­ adversely- affected. Although usually consumed losanthes procumbens Swartz (1788). fresh, the forage can also be artificially dried. Hay Vernacular names Caribbean stylo (En). Laos: making or ensiling are not commonly practised in hnha:z liaz ngu:. Thailand: thua-hamata. the humid tropics. Origin and geographic distribution S. hamata Yield In pure stands stylo can produce DM up isindigenou s to the drier habitats ofth e Caribbean to 10 t/ha and its contribution in mixed pastures Islands, southern Florida, Central America and can amount to 2-6 t/ha, depending on soil fertility South America. In South America it is recorded and moisture level. from Colombia, Venezuela and Brazil (Ceara, Per- Cattle liveweight gains of 140 kg/animal and 400 nambuco and Bahia). Tetraploid as well as diploid kg/ha were recorded in Peninsular Malaysia in the races occur in Central and South America but third year of continuous grazing of a guinea grass tetraploids have not been recorded in the Carib­ (Panicum maximum Jacq.) pasture containing bean Islands or Florida. 18% stylo and 7% centro (Centrosema pubescens Uses The tetraploid Caribbean stylo cultivars Benth.). 'Verano' and 'Amiga' are widely used as a pasture Genetic resources Stylo is well represented in legume in northern Australia, Thailand, India, South-East Asia and seed is on sale. Germplasm Nigeria and other West African countries. It can collections are available at ATFGRC (CSIRO, also be cut and fed fresh or used to make hay. Di­ Australia) and CIAT (Colombia). ploid S. hamata accessions have so far not been Breeding Plant breeding programmes are in used as sown pasture legumes. progress in Queensland, Australia, for anthrac- Properties The Nan d P concentrations of whole nose resistance within var. intermedia. The avail­ plants of Caribbean stylo sampled at flowering able natural variation within the species allows averaged 1.8% and 0.08%, respectively. The DM selection for adapted cultivars to a wide range of digestibility and DM intake ofyoun g plant materi­ environmental conditions. Plant collections in al grown under irrigation with P fertilizer aver­ South America also offer scope for further cultivar aged 80% and 73g/k g Lw076 when fed to sheep. The development and improvement. N and P concentrations and in vitro digestibility Prospects It is unlikely that much effort will be (IVD) of leaf and stem fractions generally differ put into improving the adaptation or yield of S. from each other and decline during the growing guianensis, because there are other species of Sty­ season. In green leaves, the mean N concentration losanthes that can take its place. falls from 3.9% to 2.7%, P concentration from Literature |1| Burt, R.L., Cameron, D.G., 0.37% to 0.16% and IVD from 72% to 66%. In Cameron, D.F., 't Mannetje, L. & Lenné, J.M., stems, the corresponding changes were 2.0% to 1983. Stylosanthes. In: Burt, R.L., Rotar, P.P., 1.0% N, 0.34% to 0.06% P and 57% to 33% IVD. Walker, J.L. &Silvey , M.W. (Editors): The role of However, P levels will vary with soil P status. The Centrosema, Desmodium and Stylosanthes in IVD of inflorescence and seed is 69% and 60% re­ improving tropical pastures. Westview Press, spectively. 214 FORAGES

There are about 270 seeds/g with pods and 450 flower of the spike, 1-2 mm in the next flower, seeds/g without pods. minute or absent in upper flowers; there are two Description The tetraploid is a short-lived pe­ inner bracteoles. Fruit a loment, usually with two rennial herb, usually with an erect habit but some­ fertile articles, upper article 6-7 mm long includ­ times prostrate, 0.5 m or more tall, with much ing distally recurved beak 3-4 mm long, body with branched stems;internode s 2-8 cm long,lin e of as­ prominent nerve on each side, reticulate, minute cending whitish indumentum changing sides on hairs on edges; lower article 3.5 mm long with successive internodes, glabrous elsewhere. Leaves dense white hairs. Seeds 2.5 mm x 1.5 mm, tan to trifoliolate; leaflets narrowly elliptical-lanceo­ dark maroon, mottled. late, central leaflet 16-26 mm x 3-6 mm, acute Growth and development Embryo dormancy at base and tip, mucronate, sparsely hairy above of tetraploid S. hamata is high with up to 50% in and below, central vein prominent below, 4-6 July at the end of the wet season in Queensland, pairs lateral whitish veins,margin s entire or minu­ Australia and has disappeared by October. Hard- tely denticulate at base of whitish hairs; petioles seededness (> 97%) occurs where the mean tem­ 3-6 mm long, grooved with 2line s ofhairs ; stipules perature during seed formation is above 24° C but bidentate, adnate to the base of the petiole with is less at lower temperature (64% at 21° Can d 30% hairs on the sheath and teeth; sheath about 6 mm at 20/16°C).See d germinates freely following early long. Inflorescence axillary or terminal, indeter­ summer rain and the seedlings are able to with­ minate, with 8-14 flowers; terminal inflorescence stand considerable water stress. It flowers 9-10 with 2-3 alternate spikes; outer bracts trifoliate, weeks after germination, and plants will flower broader than stipules but venation and pilosity within 6 weeks of the start of the season. Seed similar; inner bracts unifoliate, evenly sericeous; ripens over a period of 15-16 days after flowering. corolla tube 3 mm long; standard petal 4-5 mm It largely behaves as an annual except in the sec­ wide, yellow with reddish mark; wings and keel ond year when the survival rate offirst-yea r plants petals yellow; axis rudiment 4-5 mm long in first can be as high as 93%. It avoids long dry seasons as predominantly hard seed which rapidly breaks down when the maximum diurnal soil tempera­ tures reach 50-55°C just before the onset of the wet season. The tetraploid S. hamata lines have an indetermi­ nate flowering response to short days. Other botanical information Strictly S. hamata refers to the diploid species and the tetra- ploids may be a separate species. There is genetic evidence that the tetraploids have evolved as an allotetraploid combination of S. hamata (2n = 20) and S. humilis Kunth (2n = 20); both diploid spe­ cies are sympatric or nearly so with the tetra­ ploids. Because the tetraploid S. hamata (2n = 40) is largely sympatrical with its diploid progenitors S. hamata and S. humilis, it is advantageous to dis­ tinguish between them. The tetraploid and diploid S. hamata have similarities in the presence of a fine line of indumentum on one side of each inter- node, the presence of an axis rudiment, the loment often with two fertile articles and the absence of bristles on stems. The tetraploid differs from the diploid in its short-lived perenniality, the axis ru­ diment present in the lower flowers only, the pres­ ence of a long terminal bristle on the tips of the stipules and bracts, and in its darker coloured and Stylosanthes hamata (L.) Taub. -1, habit flowering longer seed. Although the tetraploid S. hamata is and fruiting branch; 2,fruit with axis rudiment. superficially similar to S. humilis, the latter differs STYLOSANTHES 215 from the tetrapioid in having abundant bristles on trition. Alternatively, grazing animals can be di­ stems, stipules and bracts; stipules forming a fused rectly supplemented with P to meet their dietary sheath around the stem; the loment with only one requirements. fertile article; no axis rudiment and is an obligate Diseases and pests Caribbean stylo is largely annual. resistant to anthracnose disease caused by the Cultivars 'Verano' and 'Amiga' were released in fungi Colletotrichum gloeosporioides and C. dema- Queensland in 1973 and 1988 respectively. Al­ tium (see S. guianensis). There is some variation though both cultivars are morphologically simi­ in anthracnose resistance between naturally lar, they differ in that 'Amiga' gives 40-50% occurring genotypes of Caribbean stylo; 'Verano' higher seed yields and appears better adapted to and 'Amiga' show a high field resistance to anth­ high altitude (> 300 m) environments than 'Ver­ racnose. Web blight (Rhizoctonia solani) can dam­ ano'. age vegetative growth of Caribbean stylo during Ecology Tetrapioid Caribbean stylo is suited to the wet season in Queensland and head blight semi-arid to sub-humid tropical regions with short (Botrytis cinerea) can cause yield losses in seed variable growing seasons and an annual rainfall crops. The latter fungus causes extensive blossom of500-200 0 mm.I t ispoorl y adapted to subtropical blighting and apical dieback. Under wet condi­ environments. It grows on a wide variety of soil tions conducive to disease development a grey-col­ types ranging from sands to clay loams, but not on oured mould grows over the inflorescences. There heavy clays.I t isparticularl y well adapted to infer­ are no important insect pests of Caribbean stylo. tile, moderately acid and sandy surface soils with Harvesting Caribbean stylo is usually har­ very low P levels. This adaptation is enhanced by vested by grazing animals but can be cut for stall its efficiency in extracting less available forms of feeding as green feed or to make hay. Caribbean soil P. The diploid Caribbean stylo occurs on neu­ stylo paddocks that are cut may need resting from tral to slightly alkaline soils and is less adapted cutting every third or fourth year to/allow seed to acid conditions. reserves to build up. Propagation and planting Caribbean stylo is Yield In pure stands, Caribbean stylo can pro­ propagated by seed. Hard-seededness can be duce up to 10 t/ha of DM in high rainfall environ­ broken by mechanical scarification or hot water ments with adequate fertilizer. Its contribution in treatment to improve germination. Seeding rate is mixed pastures usually ranges from 1-7 t/ha usually 2-3 kg/ha. In drier regions it should be depending on soil fertility, moisture level and the sown with S. scabra Vog. cultivars 'Seca' and/or competition from grasses. Heavy grazing encour­ 'Siran' at 1 kg/ha and in the wetter areas other ages legume dominance in mixed Caribbean stylo- legumes such as Aeschynomene americana L. grass pastures. Tetrapolid S. hamata lines have a 'Glen' and Stylosanthes guianensis (Aublet) high seed potential from 1750-2000 kg/ha. Actual Swartz 'Cook' could be included in the mixture. In machine harvester yields can reach 1000 kg/ha. semi-arid environments the seed can be surface Annual liveweight gains of 140t o 160kg/hea d have sown onto existing pasture following burning. been recorded from Caribbean stylo-shrubby stylo When sown together with improved grasses, pastures on lightly fertilized poor soils in northern usually in more humid environments, a well pre­ Queensland. pared seed-bed is desirable. The tetrapioid culti­ Genetic resources Germplasm collections are vars nodulate freely with the native cowpea rhizo- maintained by ATFGKC (CSIRO, Australia) and bia and do not require inoculation. Caribbean CIAT (Colombia). stylo pastures require early grazing soon after Breeding Plant breeding programmes are in establishment to reduce the competition from progress in Queensland, Australia, to develop cul­ grasses as Caribbean stylo is less palatable than tivars with greater anthracnose resistance than the young growth of grasses. 'Verano' and 'Amiga'. Accessions with low disease Husbandry Caribbean stylo can be utilized by damage could have potential as parents for a continuous or rotational grazing or cutting. breeding programme to improve anthracnose re­ Because it can grow at very low P levels, it may sistance and in studies of the genetics of anthrac­ be necessary to apply 10 to 20 kg/ha of P at estab­ nose resistance. lishment and periodically thereafter to maintain Prospects The wide range of adaptation and soil P at 8 mg/kg so that the P concentration in productivity of Caribbean stylos will encourage the herbage reaches a level of above 0.12% during further plant improvement programmes through the growing season to insure adequate animal nu­ plant breeding and the selection of naturally 216 FORAGES

occurring genotypes. Besides resistance to an- thracnose, other desirable objectives are increased seed yield and colonizing ability. It is well suited toth e drier parts ofSouth-Eas t Asia. Literature |1| Edye, L.A.,Burt , R.L., Nicholson, C.H.L., Williams, R.J. & Williams, W.T., 1974. Classification of the Stylosanthes collection 1928-1969. CSIRO, Australia, Division of Tropical Agronomy Technical Paper No 15.2 8pp . |2| Edye, L.A., Hall, T.J., Middleton, C.H., Messer, W.B., Piggin, CM., Schlink, A.C. & Klepacki, N.M., 1991. Sward evaluation of fifteen Stylosanthes hamata accessions in twenty drytropica l environ­ ments. Tropical Grasslands 25: 1-11. |3| Oram, R.N., 1990. Register of Australian herbage plant cultivars. CSIRO,Australia , pp.256-257 . i4iStace , H.M. & Cameron, D.F., 1987. Cytogenetic review of taxa in Stylosanthes hamata sensu lato. Trop­ ical Grasslands 21:182-188. |5j Stace, H.M. &Edye , L.A. (Editors), 1984.Th e biology and agronomy of Stylosanthes. Academic Press, Sydney, Australia. 636pp . L.A. Edye &A . Topark-Ngarm

Stylosanthes humilis Kunth Stylosanthes humilis Kunth - 1,2, habit of flower­ ing and fruiting branches; 3,flower; 4, fruit. Nov. gen.sp .6 :506 ,t . 594 (1824). LEGUMINOSAE the stem and often with scattered short bristles on 2n = 20 the stem and nodes. Leaves trifoliolate; leaflets Synonyms S. figueroae Mohl. (1957). lanceolate or sometimes elliptical, acute, both sur­ Vernacular names Townsville stylo, Towns- faces nearly glabrous, with 3-6 pairs of conspicu­ ville lucerne (En).Philippines : magsaysay-lucerne ous veins; terminal leaflet up to 15m m x 3.5 mm; (Tagalog). Thailand: thua-satailo. petioles 3-5 mmlong , shortly hairy and often with Origin and geographic distribution Towns­ scattered bristles; stipules bidentate, adnate to ville stylo is widespread in Brazil and also occurs base of petiole with bristles on both sheath and in Venezuela, Colombia, Central America, Mexico teeth. Inflorescence consists of several short, and Cuba. It is adventive to Queensland (Austra­ ovoid, crowded spikes with 5-15 flowers in each lia) and is naturalized in Malaysia, Indonesia and spike;spike s hirsute without axis rudiment; flower Thailand. with oneinne r andon eoute r bracteole; calyx tube Uses Townsville stylo is a forage particularly 4-5 mmlong , lobes 1.5m mlong ; corolla bright yel­ useful in heavily grazed areas in the semi-arid to low; standard 3-4 mm x 3-4 mm. Fruit a loment, subhumid tropical and lower latitude subtropical 1.5-2.5 mm broad, reticulately nerved; only the regions with a marked dry season. upper articulation fertile, 1.5-2.5 mm long, with Properties TheN an dP concentration s of whole beak 4-10 mm long; beak 1.5-3 times the length plants range from 3.0-1.6% and 0.20-0.04% re­ of the upper articulation and strongly uncinate to spectively. The DM digestibility of young plant coiled. Seed yellowish-brown. material lies between 60% and 70%, but with Growth and development Embryo dormancy increasing agei ti sreduce d to4 0% . Seed hasa D M is high with up to 94% in July in northern Austra­ digestibility of ca. 62%.Ther e are275-30 0 seeds/g lia at the end of the wet season, but is short lived with pods and400-50 0 seeds/g without pods. and has disappeared by October. Hard-seededness Description A herbaceous, usually erect but can be as high as 100% in July but it breaks down sometimes prostrate annual, 0.5m tall, branched, rapidly to 1-6% by December when maximum usually with short white hairs along one side of diurnal soil temperatures are 50-55°C under field STYLOSANTHES 217 conditions. Hard-seed breakdown occurs to a less­ It is unsuited to heavier textured and more fertile er extent when the soil surface is moist as the tem­ soils, including cracking clays and soils derived peratures do not rise above 40°C. It germinates from basalt or other igneous rocks, and also saline freely following early summer rain and the seed­ coastal soils. lings are able to withstand considerable water Townsville stylo is sensitive to shading when stress and then grow rapidly during the vegetative grown in mixtures with taller grasses. and early reproductive stage if the soil moisture Propagation and planting Townsville stylo is is adequate. It has a short-day flowering response propagated by seed. Hard-seededness can be with a critical photoperiod of 12-14 hours.I t nodu- broken by mechanical scarification or hot water latesfreel y with the cowpea type rhizobia and does or dry heat treatment. Seeding rate is 2-3 kg/ha. not require inoculation. Once Townsville stylo In semi-arid environments, seed can be surface plants become established the major influences on sown onto existing pastures following burning just growth are water, nutrition and amount of compe­ prior to the expected start of the wet season. tition from associated grasses. Flowering is geneti­ Husbandry In Thailand, the anthracnose-resis- cally controlled with shorter critical photoperiod tant cultivar 'Khon Kaen' was released in 1984 flowering being dominant. Flowering occurs over because it showed greater productivity and persis­ 8-10 weeks and seed ripens over a similar time tence than S. hamata (L.) Taub. 'Verano' under interval. It avoids long dry seasons as predomi­ extremely heavy grazing, because of its prostrate nantly hard-seed. growth habit. It is oversown onto communal graz­ Other botanical information Collections of ing lands which are subjected to very heavy con­ Townsville stylo can be divided into two major tinuous grazing. Townsville stylo was widely used groups. One includes adventive populations from for pasture improvement in northern Australia Australia, Colombia, Dominican Republic, Ivory during 1960-1974 but, since the advent of anthrac­ Coast, Kenya, the Philippines, Tanzania, the Uni­ nose it is of little importance as it has been super­ ted States, as well as collections from Brazil of sup­ seded by the anthracnose-resistant cultivars of S. posedly indigenous material. The second group hamata, 'Verano' and 'Amiga', and of S. scabra includes collections from Venezuela, Costa Rica, Vogel 'Seca' and 'Siran'. and Mexico. The evidence suggests Diseases and pests Townsville stylo is suscepti­ that Townsville stylo is not native to Brazil and blet o anthracnose disease caused by the fungi Col­ that it first occurred around ports in north-eastern letotrichum gloeosporioides and C. dematium (see Brazil inth e late 19th Century about the same time S.guianensis). There is some variation in anthrac­ as its first recorded occurrence in Australia. nose resistance between naturally occurring geno­ The cultivars released in Queensland, Australia types ofTownsvill e stylo (e.g.'Kho n Kaen'). Blight during the late 1960's were the common natura­ (Sclerotium rolfsii) and bacterial wilt (Pseudomo­ lized type, 'Paterson' (early flowering), 'Lawson' nas solanacearum) have been recorded on Towns­ (mid season flowering) and 'Gordon' (late flower­ ville stylo in Australia. There are no known impor­ ing). All the Australian naturalized populations tant insect pests. and cultivars proved highly susceptible to the fun­ Harvesting Although usually consumed fresh gal disease anthracnose when it was accidentally by grazing animals, Townsville stylo can be cut introduced into Australia ca. 1973. and fed when grown or made into hay towards the Ecology Townsville stylo's adaptation in Aus­ end of the growing season. Prior to the advent of tralia to a wide latitudinal range (12-28°S) with anthracnose, feeding of Townsville stylo hay dur­ annual rainfall ranging from 600-1500 mm is due ing the dry season was commonly practised in to ecotypicvariatio n in critical photoperiod. Rain­ northern Australia to encourage seed dispersal by fall and latitude delineate ecotypic occurrences; cattle. ecotypes with a long critical photoperiod are re­ Yield Townsville stylo has a high seed yield po­ stricted to drier or higher latitude regions whilst tential up to 1250 kg/ha and actual machine har­ those with a shorter critical photoperiod are vested yields of 600kg/ha . Yields of dry matter can adapted to higher rainfall regions at lower lati­ range from 1-6 t/ha. tudes. Although it will persist in regions with as Genetic resources Major germplasm collec­ little as 600 mm annual rainfall, it is not suffi­ tions are held by ATFGRC (CSIRO, Australia) and ciently productive to enhance animal production CIAT (Columbia). below 800m m rainfall. It occurs on infertile sandy Breeding There are no plant breeding pro­ to clay loam soils with a pH(H20) range of 5t o 6.5. grammes in progress. 218 FORAGES

Prospects It is unlikely that any effort will be put into improving adaptation and yield of S. humilis because there are other Stylosanthes spe­ cies that can take its place. Literature |1| Edye, L.A., Burt, R.L., Nicholson, C.H.L., Williams, R.J. & Williams, W.T., 1974. Classification of the Stylosanthes collection 1928-1969. CSIRO, Australia, Division of Tropical Agronomy Technical Paper No 15. 28 pp. |2| Gil- lard, P. &Fisher , M.J., 1978.Th e ecology ofTowns - ville stylo-based pastures in northern Australia. In: Wilson, J.R. (Editor): Plant relations in pas­ tures. CSIRO, Melbourne, pp. 340-352 |3| Shaw, N.H. & 't Mannetje, L., 1970. Studies on a spear grass pasture in central coastal Queensland - the effect of fertilizer, stocking rate, and oversowing with Stylosanthes humilis on leaf production and botanical composition. Tropical Grasslands 4: 43-56. |4|Stace , H.M. &Edye , L.A. (Editors), 1984. The biology and agronomy of Stylosanthes. Aca­ demic Press, Sydney. 636pp . L.A. Edye &A . Topark-Ngarm

Stylosanthes macrocephala M.B. Ferreira &N.M . Sousa Costa

An. Soc. Bot. Brazil 28 Congr. Nac. Bot.: 87 (1978). LEGUMINOSAE Stylosanthes macrocephala M.B. Ferreira & N.M. 2re = 20? Sousa Costa - 1, habit of flowering and fruiting Origin and geographic distribution S. macro­ branch; 2,fruit with axis rudiment. cephala originates in South America; its natural distribution isrestricte d to sub-humid and dryland veins;petiol e 1-2 mm long, villous;leaflet s lanceo­ areas of central and eastern Brazil. Experimental late, 20-55 mm x 10-19 mm, pubescent or gla­ lines have spread for testing to other tropical brous, with 7-10 pairs of veins. Inflorescence a regions, including South-East Asia. capitate spike, terminal or axillary, ovoid to Uses The main use of S. macrocephala is as for­ almost spherical, 14-18 mm x 10-15 mm, with age in permanent pastures grazed by ruminants. 10-30 flowers; often several inflorescences in a Properties In many regards, S. macrocephala is cluster; bracts imbricate, 1-foliate, elliptical- similar to S. capitata Vogel. Its nutritive value, ovate, 10-12 mm x 8-9 mm,pubescent , with 11-15 however, is higher: in 6-week-old regrowth, N con­ veins, Reddish;flower s papilionaceous, small, with centrations in leaves range from 2.2%-3.5% obovoid standard 4-6 mm long, yellow, striated; (stems: 1.4-1.8%), and leaf DM digestibility from axis rudiment and 2inne r bracteoles present. Fruit 66%-75% (stems: 46-61%); P concentrations are 2-articulated, reticulately nerved; both articles moderate to low, 0.17-0.25%) in leaves and usually fertile; the upper one glabrous, 3-4 mm x 0.13-0.25% in stems. There are about 500 seeds 2.5-3 mm, with a short, straight to uncinate beak; (dehulled pods)/g. lower article villous and smaller. Seed yellow- Botany A perennial, fine-stemmed, many- brown, sometimes slightly mottled, to black. branched, and taprooted subshrub, prostrate to Like S. capitata, S. macrocephala persists through semi-erect, 20-80 cm tall; stems and branches extensive seedling recruitment. It is a prolific hairy-bristly. Leaves 3-foliolate; stipules obovate, seeder. Cultivar 'Pioneiro' has been released in 14-16 mm x 5-6 mm, with pointed teeth, pubes­ Brazil for use in the Cerrados savanna region. cent to glabrous or bristly, with 11-13 pairs of Prior to its relatively recent description, S. macro- STYLOSANTHES 219 cephala has sometimes been referred to as S. capi­ Bolivia, Colombia and Venezuela. It is also culti­ tata or S. bracteata Vogel. vated in other tropical regions besides South Ecology S. macrocephala has essentially the America, e.g. in Australia and India. same climatic and soil adaptation as S. capitata. Uses Shrubby stylo is mainly used as a pasture That is, good adaptation to savanna environments, legume for semi-arid to humid tropical regions mainly in terms oftoleranc e of the prevailing acid, with an average annual rainfall of 500-2000 mm. low-fertility oxisols and ultisols. In the State of Maharashtra, India, it is cut and Agronomy The agronomy of S. macrocephala is fed green to dairy cattle. similar to that of S. capitata. Newly harvested seed Properties The Nan d P concentrations of whole may have to be treated to reduce hard-seededness. plants sampled at flowering averaged 1.7% N Although S. macrocephala nodulates effectively (range 1.1-2.2% ) and 0.07% P (range 0.07-0.11 %). with native rhizobia, inoculation with an effective The N and P concentrations and in vitro digestibi­ strain of Bradyrhizobium is recommended. Al­ lity (IVD) of leaf and stem fractions generally though tolerant of low fertility, productivity is differ from each other and decline during the grow­ enhanced by fertilization. It is somewhat less pro­ ing season. In leaves, the mean N concentration ductive than S. capitata, during both the rainy and falls from 3% to 1.5% , P concentration from 0.3% dry seasons. Seed set and seedling recruitment are to 0.1%,an d IVD from 70% to 50%. In stems, the essential for long-term persistence. S. macroce­ corresponding changes were 1.5 % to 0.5% N,0. 3% phala is, in general, more anthracnose-tolerant to 0.03% P and 60% to 30% IVD. Shrubby stylo than any other Stylosanthes species. In Colombia, has ahig h proportion ofste m which increases from it can be seriously affected by Rhizoctonia foliar 20% in the early growth stage to 75% at the end blight. of the growing season. In grazed pastures, stem Genetic resources and breeding A large and contents of80-9 6% have been recorded. There are quite variable S. macrocephala collection is held 400-550 hulled seeds, and 600-800 delrulled seeds at CIAT (Colombia). At present there is'n o breed­ per g. ing programme for S. macrocephala. Description A perennial subshrub, erect to Prospects Unless more productive cultivars suberect to 2 m tall with a strong taproot with with better dry-season performance become avail­ small round root nodules and penetrating deeply able, the potential role of S. macrocephala for pas­ to depths up to 4m .Stem swit h variable branching, ture development in its particular ecological niche covered by short or long hairs with bristles, will most likely be limited, as S. capitata is an usually viscose, pale green to dark blue-green, equally well-adapted but more productive species. brown or red. Leaves trifoliolate; leaflets elliptical Literature |1| Schultze-Kraft, R., Costa, N.M.S. to oblong-laceolate, apex obtuse to mucronate, & Flores, A., 1984. Stylosanthes macrocephala hairy on both surfaces with 4-7 prominent veins M.B. Ferr. et S. Costa - collection and preliminary thickening terminally, pale green to dark green or evaluation of a new tropical pasture legume. Trop­ dark blue-green; terminal leaflet 20-33 mm x ical Agriculture (Trinidad) 61:230-240 . |2| Stace, 4-12 mm, length/breadth ratio 2 to 5; petiole up H.M. &Edye , L.A. (Editors), 1984.Th e biology and to 10 mm long, canaliculate above, scabrous with agronomy of Stylosanthes. Academic Press, Syd­ dense short hairs, the rachis 4-5 mm long; stipules ney, Australia. 636 pp. |3|Thomas , D. & Grof, B., obovate, 15-25 mm long including teeth, bristly- 1986. Some pasture species for the tropical savan­ hairy. Inflorescence obovoid to ellipsoid, general­ nas of South America. I. Species of Stylosanthes. ly longer than wide, 10-30 mm long, several-flow­ Herbage Abstracts 56: 445-454. ered, each flower surrounded by a bract and 3brac - R. Schultze-Kraft teoles; bracts unifoliolate, the blade about 4 mm long, the sheath 3.5-6.5 mm long with long teeth; outer bracteole 1, 2-4.5 mm long, bifid and ciliate Stylosanthes scabra Vogel at apex; axis rudiment 4-5 mm long; inner brac- teoles 2, 2-4 mm long, ciliate at apex; calyx tube Linnaea 12:6 9(1838) . 3-6.5 mm long, lobes 1.5-3.5 mm long; standard LEGUMINOSAE suborbiculate, ca. 7 mm long. Fruit a loment with 2re = 40 usually two fertile articles about 2.5 mm broad, Vernacular names Shrubby stylo (En). reticulately nerved; upper articulation 2-4 mm Origin and geographic distribution Shrubby long, shortly hairy, the lower 2-3 mm long, evenly stylo is widespread in Brazil, but also occurs in pilose throughout; beak uncinate, with short 220 FORAGES

and climatic ecotypes. In Brazil the species is very widespread and two major forms have been recog­ nized. One, often described as S. aff. scabra or S. aff. hamata by some authors, has thin stems with long, narrow leaflets, length/breadth ratio 4 to 5, with dark blue-green stems and leaves, and pro­ duces seed all year; this form mainly occurs in the coastal states of Maranhïo, Pernambuco Bahia and Rio de Janeiro. The other major form, which is more readily accepted as S. scabra, has thick woody stems and can be grouped into coastal types and continental types. The coastal types are tall robust subshrubs, leaflets elliptical and a length/ breadth ratio of 2.0 to 2.7. The continental types are shorter in growth habit and more densely branched, leaflets lanceolate or elliptical, length/ breadth ratio 2.3 to 3.9. The continental types mainly occur in the states of Goiâs, Minas Gerais, Matto Grosso, Matto Grosso do Sul, Piaui and Sào Paulo. The shrubby stylos from Colombia and Venezuela are morphologically different from the Brazilian material but more closely resemble the continen­ tal types than the coastal types of Brazil by their short growth habit and dense branching. The geo­ graphical plant groups reflect the climatic and edaphic differences between the regions. Stylosanthes scabra Vogel - 1, habit of flowering The cultivars released in Australia are: 'Seca', an and fruiting branch; 2, fruit. anthracnose-resistant cultivar which is widely adapted in Queensland; 'Fitzroy', an early flower­ hairs, 1-2 mm long, \ to \ as long as the upper ing and leafy cultivar which is very susceptible to articulation. Seed asymmetrically reniform, small anthracnose and is no longer used; and 'Siran', (< 2 mm long), pale to light brown, radical end bred to provide a multigenic resistance to anthrac­ prominent. nose. Growth and development Shrubby stylo Ecology Shrubby stylo is very drought resistant appears to have a juvenile phase during which and can survive long dry periods. It is adapted to floral initiation does not take place, followed by tropical and subtropical environments with an a short-day flowering response; the critical photo- average annual rainfall of 500-2000 mm and can period lies between 11.5-12.5 hours, depending on tolerate light frost to -3°C; heavy frosts can kill the accession. Seed yields stay at a peak for a long the crowns. It grows on a wide range of soil types, period but vary with the accession and season; but isparticularl y well adapted to infertile, moder­ 70-90 % ofth e seed may be hard-seeded. ately acid, sandy surfaced soils with very low P Other botanical information S. scabra is part levels. This adaptation is enhanced by its effi­ of a species complex in which the species cannot ciency in extracting less available forms of soil P. be separated satisfactorily. Besides S. scabra, the Shrubby stylo can grow on heavier textured soils complex comprises S. tuberculata Blake, S. ner­ than most other stylos, but is not suited to heavy vosa Macbr., S. fruticosa (Retzius) Mohlenbrock, black basaltic clays. and S. suffruticosa Mohlenbrock. If, after a badly Propagation and planting Shrubby stylo is needed taxonomie revision, the whole complex is propagated by seed. Hard-seededness can be reduced to one taxon, the correct name will be S. broken by mechanical scarification to improve fruticosa for reasons of priority. germination. Under field conditions, substantial Although shrubby stylo is very variable, most softening of hard seed occurs when the maximum plants can be readily classified into morphologi­ diurnal soil temperatures reach 50-55°C and is cal-agronomic groups that reflect geographical largelyrestricte d toth e late dry and pre-wet period STYLOSANTHES 221 in the tropics and to late spring and summer in the Yield In pure stands, shrubby stylo can produce subtropics. Seeding rate is 1-2.5 kg/ha with the DM up to 10 t/ha in areas of high rainfall, and its lower figure for drier regions. In semi-arid environ­ contribution in mixed pastures can amount to 2-7 ments, the seed can be surface sown onto existing t/ha depending on soil fertility and moisture pasture following a burn; in humid environments levels. Annual liveweight gains of cattle of when sown together with improved grasses, a well 140-160 kg/head have been recorded from shrubby prepared seed-bed is desirable. Shrubby stylo stylo pastures on lightly fertilized poor soils in nodulates freely with cowpea type rhizobia and northern Queensland (Australia). does not require inoculation. In drier regions it Seed yields of up to 620kg/h a have been recorded. should be sown with S. hamata (L.)Taub , cultivars Genetic resources Germplasm collections are 'Verano' or 'Amiga'. In wetter areas it should be maintained at ATFGRC (CSIRO, Australia) and sown with other legumes such as Aeschynomene CIAT (Colombia). americana L. cultivars 'Glenn' or 'Lee' and S. guia- Breeding Plant breeding programmes are in nensis (Aublet) Swartz cultivar 'Cook' could be progress in Queensland, Australia. Cultivar included in the mixture. 'Siran', a composite of 3 lines, was released in Husbandry Shrubby stylo isuse d by continuous August 1990; it incorporates a multigenic resis­ or rotational grazing or cutting. Because it can tance to anthracnose derived from crosses and grow at very low P levels, it may be necessary to selections of the anthracnose-resistant accessions apply 10-20 kg/ha of P as superphosphate or alter­ Q 10042 x CPI 93116 (Line 1), an early flowering natively, directly supplement the grazing animals selection of 'Seca' (Line 4), and CPI 55860 x Q with P to meet their dietary requirements. Al­ 10042 (Line 8). Lines 1, 4 and 8 have been named though it is readily eaten by cattle and sheep, it 'Jecuipe', 'Recife' and 'Feira' respectively for plant is not very palatable. As with most Stylosanthes cultivar rights registration. Current research has species, young growth of shrubby stylo is less pal­ shown that partial resistance to anthracnose is atable than young growth of grasses. This assists stable, heritable and improvable through breed­ in seedling establishment and regeneration of ing. Future plant breeding objectives will aim at shrubby stylo and enables long-term persistence. combining both major gene and partial resistance Diseases and pests Shrubby stylo is susceptible genes against anthracnose in new cultivars. to anthracnose disease caused by the fungi Colleto­ Prospects The extreme drought resistance and trichum gloeosporioides and C.dematium; the culti­ wide range of adaptation of shrubby stylo will en­ vars 'Seca' and 'Siran' are resistant to anthrac­ courage further plant improvement programmes nose. Blight (Sclerotium rolfsii) has been recorded through plant breeding and the selection of natu­ on shrubby stylo inThailand . Two important stem- rally occurring genotypes. Besides resistance to borers damage shrubby stylo: Caloptilia sp.i n Bra­ anthracnose, other desirable objectives for plant zil and Colombia, and Platyomopsis pedicornis in improvement include increased frost resistance Queensland, Australia. Calotilia sp. is the major and higher nutritive value. pest of Stylosanthes in South America. The larvae Literature |ll Edye, L.A., Burt, R.L., Nicholson, tunnel through lower stems, considerably weaken­ C.H.L., Williams, R.J. & Williams, W.T., 1974. ing susceptible accessions which usually die Classification of the Stylosanthes collection within 2 years. Field screening has identified 1928-1969. CSIRO,Australia , Division of Tropical accessions that are resistant to stem-borer. Iso­ Agronomy Technical Paper No 15.2 8pp . |2| Mass, lated occurrences of Platyomopsis have been B.L., 1989. Die tropische Weideleguminose Stylo­ recorded in Queensland but it is not a serious pest; santhes scabra Vog. - Variabilität, Leistungs­ the larvae hollowed the taproot and lower main stand und Möglichkeiten züchterischer Verbes­ stem and killed the infested plants. have serung [The tropical pasture legume Stylosanthes been recorded on shrubby stylo in Queensland and scabra Vog. - variability, productivity and im­ they can kill older plants. provement possibilities through breeding]. Land- Harvesting Shrubby stylo is harvested by graz­ bauforschung Völkenrode, Sonderheft 97. Institut ing animals or it is cut for stall feeding. When für Grünland-und Futterpflanzenforschung, mown, care should be taken not to cut the woody Braunschweig, Deutschland. 191 pp. j3| Stace, stems too low, otherwise regrowth will be adver­ H.M. &Edye , L.A. (Editors), 1984.Th e biology and sely affected and some plants will die. In semi-arid agronomy of Stylosanthes. Academic Press, Syd­ environments, forage can be made into hay to­ ney, Australia. 636pp . wards the end of the growing season. L.A. Edye &A . Topark-Ngarm 222 FORAGES

Themeda triandra Forssk.

Fl. Aegypt.-Arab.: 178(1775) . GRAMINBAB 2M = 20, 30,40 ,50 ,60 , 70, 80 Synonyms Themeda imberbis (Retzius) Cooke (1908), T. australis (R. Br.) Stapf (in part) (1919). Vernacular names Red oat grass, kangaroo grass (En).Indonesia : merakan lanang (Javanese). Philippines: bagokbok (Tagalog), samsamon (Ilo- kano), ipatpatey (Bontok). Cambodia: sbö':w. Laos: hnaaz fèèk. Thailand: ya-faek. Vietnam: co' bông cao ru'ng khôp, co'tam hung. Origin and geographic distribution The natu­ ral distribution of red oat grass is pantropical/sub- tropical with extension into the temperate zone as a summer growing grass. It is most likely of Gond- wanan origin with a predominant distribution in Australia, South-East Asia, India and Africa. Uses Red oat grass is used as part of the natural­ ly occurring savanna pastures for domestic live­ stock production and wildlife. Properties In native pastures the N concentra­ tion ofre d oat grass ranges from 2.7% in green ma­ terial from the early flush of the growing season to 0.5% in dry forage by the end of the dry season. Digestibility varies from about 60% to 35% . Botany A tufted leafy perennial, growing up to Themeda triandra Forssk. - 1, flowering plant; 2, 1.5 m tall, erect, culms usually unbranched but ligule; 3,group ofspikelets; 4,two spikelets. quite variable in morphology, smooth, often becoming golden at maturity, nodes hairless. 1 in the involucral and pedicellate spikelets which Leaves both basal and on the culms; leaf-sheath are male, sterile or reduced to a lemma; fertile spik­ keeled, coarsely striate, hairless below but becom­ elet sessile and awned, about 8m m long including ing quite short-hairy towards the ligule, the mar­ the dark brown, sharply pointed, densely and gins translucent; ligule short and membranous, stiffly hairy barbed callus; awn up to 7 cm long, later splitting into a broad rim of short hairs with puberulous, twisted and geniculate, hygroscopi- some longer, marginal hairs; leaf-blade 15-50 cm cally active so that the seed can bore itself into x 2-5 mm, green or sometimes bluish, folded with the ground. Caryopsis lanceolate, channelled on a more or less hairy clasping base, becoming more one side. or less flat towards the acutely pointed tip, flex- The fruits ('seeds') ofre d oat grass fall individually uous, smooth or slightly rough down the margins. to the ground. They probably have similar seed Inflorescence a loose, narrow spathate usually dormancy and germination control mechanisms to reddish-coloured panicle, the flexuous peduncled speargrass Heteropogon contortus (L.) Beauv. ex racemes slightly nodding; panicle branches Roemer & Schultes and other Andropogoneae. slender and smooth, often quite widely separated Flower initiation is obligately to facultatively along the axis and bearing clusters of one to sever­ short-day in the tropics, short-day to day-neutral al racemes (spikelet clusters), each at first in the subtropics and long-day in the temperate enclosed by, and later subtended by a strongly zone. There is some evidence of a cold stratifica­ keeled, sheath-like spathe; spikelet clusters are tion response in materials occurring in subalpine composed of a sessile, fertile awned spikelet sur­ areas, these forms also being polyploid. The repro­ rounded by an involucre of 2 pairs of sessile and ductive system is also likely to be obligately to 1 pair of pedicellate male or sterile (often reduced) facultatively apomictic as in the majority of spikelets; florets 2i n the fertile spikelet, the lower Andropogoneae. sterile and reduced to a lemma, the upper bisexual, The species is very polymorphic, and based on the THEMEDA 223 hairiness of the inflorescence parts, has been sub­ defoliation with the different pattern of synchrony divided repeatedly in the literature. Without clear of growing points in these two species at the start correlations with geographic distribution, these of the growing season. subdivisions are of little value. No diseases and pests of any consequence have Ecology Red oat grass is a typical andropogo- been recorded. noid grass ofth e tropical and subtropical savannas The annual yield depends very much on the which are characterized by a marked seasonality amount and favourable distribution of rainfall of warm wet, and cooler dry seasons. It is a occurring during the warm season. The normal savanna grass in seasonally wet/dry climates of savanna environment means that growth and moderate rainfall. However, it also occurs in quite quality are seasonal so that there is a fluctuating arid environments along drainage lines and run-on amount of biomass and level of feed quality on areas. It is relatively drought-tolerant, with its offer for livestock grazing. Production is likely to main range of distribution being within 500-1200 range between 0.5-5 t/ha ofDM ,dependin g on con­ mm of annual rainfall. It has been recorded at ditions. above 3000m i n the Himalayas. However, an inter­ Genetic resources and breeding It is unlikely esting departure from this is its adaptation to that substantial germplasm collections are being growing during the warm season of the temperate maintained. zone provided there ismoistur e at that time. In the Prospects In production systems where a stable subtropics red oat grass will produce green growth alternative can be established, either naturally as in the cool season if there is some out-of-season with replacement by Heteropogon contortus, or by rainfall, which is atypical of andropogonoid conversion to planted pastures, there could be lit­ grasses. However, it is sensitive to continued tle disadvantage in losing red oat grass. However, intensive grazing and this unseasonal growth where there are no viable and acceptable alterna­ places it at greater risk from overgrazing when tives or where there is the need to preserve its par­ almost everything else is senesced and dormant. ticular type of ecosystem, as in some national park It has been shown to be particularly sensitive to systems, it may be important to maintain it, which heavy grazing when the growing points are elon­ requires careful management of livestock or wild­ gating late in the growing season. Like Heteropo­ life. gon contortus, red oat grass responds to dry-season Literature 1| Bogdan, A.V., 1977. Tropical pas­ burning, but being more sensitive to grazing, it ture and fodder plants. Longman, London, pp. will predominate in areas which are burnt and not 286-290. |2| Bor, N.L., 1960. The grasses of Burma, significantly grazed, e.g. some road and railway Ceylon, India and Pakistan. Pergamon Press, exclosures. It grows on a wide range of soil types Oxford, p. 254. |3| Lazarides, M., 1980.Th e tropical except heavy clays. grasses of southeast Asia. Phanerogamarum Agronomy The wide variety of growth forms Monographieae Tomus XII, J. Cramer, Vaduz, pp. and habitats of red oat grass makes generalizing 76-78.14! Mott, J.J., Ludlow, M.M., Richards, J.H. somewhat hazardous. Its intolerance of prolonged & Parsons, A.D., 1992. Effects of moisture supply heavy grazing is sometimes belied by areas which in the dry season and subsequent defoliation on have survived many years of close grazing or mow­ the persistence of the savanna grasses Themeda ing where the species has developed a prostrate triandra, Heteropogon contortus and Panicum growth form. Heteropogon contortus often replaces maximum. Australian Journal of Agricultural red oat grass following sustained grazing and dry- Research 43: 241-260. |5| Skerman, P.J. &Riveros , season burning. F., 1990. Tropical grasses. FAO, Rome. pp. As red oat grass is susceptible to prolonged heavy 721-724. |6| Tothill, J.C. & Hacker J.B., 1983. The grazing pressure, it requires very careful control grasses of southern Queensland. University of of grazing when used in conjunction with manage­ Queensland Press, Brisbane [as T. australis]. pp. ment practices such as oversowing legumes or pro­ 402-404. vision of feed supplements such as mineral licks, J.C . Tothill urea/molasses, and other externally supplied feeds. These practices usually lead to increased grazing pressure, especially early in the growing season when red oat grass is more susceptible to defoliation than is Heteropogon contortus. This appears to be becasue of the interactive effect of 224 FORAGES

Thysanolaena latifolia (Roxb. ex Hörnern.) Honda

Journ. Fac. Sei. Tokyo, Sect. 3.Bot . 3:31 2(1930) . GRAMINBAB 2n = 24 Synonyms Melica latifolia Roxb. ex Hörnern. (1819), maxima Roxb. (1820), Thysano­ laena maxima (Roxb.) Kuntze (1891). Vernacular names Tiger grass (En). Indonesia: awis (Sundanese), menjalin wuwu (Javanese), lan- tebung (Makasar). Malaysia: buloh teberau, rum- put buloh. Philippines: tambu (Tagalog), gatbo (Bikol), buybuy (Ilokano). Laos: kh'èèm kh'ôông. Thailand: tongkong, laolaeng (northern), ya-yung (southern). Vietnam: dot, dông trung hôa tha'o. Origin and geographic distribution Tiger grass occurs from India to Indo-China and China and throughout Malesia. It is also occasionally cultivated outside this region. Uses Young leaves and stem tips are used to feed cattle and buffaloes. Its large inflorescences are used in making brooms. The grass is occasionally planted for ornamental purposes and as a hedge. Properties One leaf sample in Thailand had a N concentration of 1.2 %. The in vitro DM digesti­ bility of leaves ranged from 40% to 60% . Botany A strongly tufted, very robust perennial Thysanolaena latifolia (Roxb. ex Hörnern.) Honda grass with erect or slightly spreading solid bam­ -1, flowering culm; 2, ligule; 3,part of inflorescence boo-like culms up to 3.5 m tall. Leaf-sheath hairy branch; 4, spikelet. along outer margin; ligule a scarious membrane, 1-2 mm long; leaf-blade lanceolate-acuminate, ing is required at the early stage of establishment. 30-65 cm x 3-7.5 cm, base broad and rounded or Genetic resources and breeding It is unlikely subcordate, margins scaberulous, conspicuously that substantial germplasm collections are being glaucous beneath. Inflorescence a terminal huge maintained. and drooping panicle, up to 140 cm long, well Prospects More attention should be given to exserted, branches divided and subdivided into this species as a source of feed for cattle in high­ many branchlets; spikelets awnless, short pedi- land areas. celled, falling with part of the pedicel, often in Literature jl| Falvey, F., Pongpiachan, P. & pairs on a common peduncle, 2-flowered; lower Hengmichai, P., 1981. The quality of native pas­ glumes clasping. Caryopsis subglobose to ovoid, tures grazed by highland cattle. Thai Journal of 0.6m m long, reddish-brown. Agricultural Science 14:97-105. |2|Gilliland , H.B., In light shade seedlings grow slowly at first, but Holttum, R.E. & Bor, N.L., 1971. Grasses of are then able to compete with other low-growing Malaya. In: Burkill, H.M. (Editor): Flora of plants.I t flowers throughout the year at lower alti­ Malaya. Vol. 3. Government Printing Office, tudes. Singapore, p. 45. |3| Holm, J., 1971.Feedin g tables. Ecology Tiger grass grows from 150-2000 m alti­ Nutrition Laboratory of Thai-German Dairy Pro­ tude in valleys and on lightly shaded slopes, in ject, Livestock Breeding Station, Chiangmai, ravines and onrive r banks.I t tends to grow in asso­ Thailand, p. 27. |4j Mehra, K.L. & Fachrurozi, Z., ciation with trees (often bamboo forests), solitarily 1985.Indonesia n economic plant resources: forage or in small groups, and not in full sunlight. crops. Lembaga Biologi Nasional - LIPI, Bogor, Agronomy Tiger grass can be propagated by No 31. p. 50. |5| Santos, J.V., 1986. Grasses. In: rhizomes,roote d culms or seeds. It is cultivated for Umali, R.M., Zamora, P.M., Gotera, R.R. & Jara, broom making, but not primarily for forage. Weed- R.S. (Editors):Guid e to Philippine flora and fauna. TRIFOLIUM 225

Vol. 4. Natural Resources Management Center and University ofth e Philippines. Manila, p. 127. C. Manidool

Trifolium repens L.

Sp. PL: 767(1753) . LEGUMINOSAE 2re = 32 (tetraploid), but 16,28 ,4 8an d 64 occur as well Vernacular names White clover, Dutch clover (En). Trèfle blanc (Fr). Indonesia: semanggi landa (Javanese). Thailand: thua clover. Origin and geographic distribution The like­ ly centre of origin isth eMediterranea n region but white clover isindigenou s toth ewhol e of Europe, Central Asia west of , and to small areas in Morocco andTunisia . It is naturalized in many countries, particularly in North America, China, Australia and in higher latitudes of South America. Although of most importance in mild temperate and Mediterranean climates, white clover isnaturalize d inth e higher rainfall subtrop- ics and in the elevated tropical areas such as the highlands ofPapu a New Guinea. Uses In the tropics and subtropics, white clover is used as a forage legume in grazed pastures. It Trifolium repens L. 1, flowering plant part; 2, is also an important species for pasture hay and flower; 3, fruit. silage in temperate regions. The flowers are con­ sidered asa n important source of honey. cm in diameter, with up to 40 white or light pink Properties White clover ison eo f the best quali­ flowers, reflexed after fertilization; calyx 3-5mm , ty temperate legumes andi sbette r in quality than corolla 4-13 mm long. Pod linear-oblongoid, 4-5 any tropical legume. Nitrogen percentages in mm long, with 3-4 seeds. Seed ovoid to reniform, foliage range from 3.5-5% and DM digestibilities ca. 1.5m mlong , usually pale yellow, occasionally range from 60-80%. Cultivars adapted to warmer reddish-brown. climates tend to have higher glycoside levels, but White clover isa ver y variable species, and mainly not high enough to affect animal production. based on flower colour andsiz e offlower s and leaf­ There are 1400-1800 seeds/g. lets, numerous botanical varieties have been dis­ Botany A perennial herb which, particularly in tinguished. No satisfying classification exists. subtropical areas, can also persist as an annual. There are numerous cultivars of white clover, for The primary taproot seldom persists for more than description purposes roughly classified based on one year (subtropics) or two years (temperate size into small, intermediate and large types.Tw o areas). The prostrate succulent stolons which de­ widely used ones in the subtropics are 'Haifa' and velop shallow adventitious roots are short-lived, 'Louisiana SI', both being heavier seeders in the usually for less than oneyear . Leaves trifoliolate, subtropics than 'Ladino' or 'Grassland Huia'. long-petioled (1-30 cm);leaflet s elliptical toobo - In temperate areas white clover grows actively vate to heart-shaped, 0.5-4 cm x 1-1.5 cm, margin during summer. In contrast, in the subtropics it denticulate, glabrous, frequently with white or grows in the cooler months with a peak late in light green markings that are often crescent spring if there is adequate moisture. There is con­ shaped; petiole length and leaflet size vary mark­ siderable stolon death over late summer and early edly with cultivar and decrease with increasing autumn. This death is associated with high grazing pressure. Peduncle 2-30 cm long, gla­ minimum and maximum temperatures and also brous; inflorescence umbellate, globose, 1.5-3.5 with moisture stress during dry periods, with rot- 226 FORAGES ting of roots and stolons in wet conditions, and repens is an amphidiploid and its related diploic! with competition from warm season grasses. species are thought to be T. nigrescens Viv., a self- Growth from late autumn onwards is then from incompatible Mediterranean annual, and T. occi­ surviving stolons and from seedlings which dentale Coombe, a self-compatible perennial indig­ emerge until early spring. Soil seed reserves can enous to southern England, south-western France reach 10000seeds/m 2 in good white clover pastures and Spain. in the subtropics, whereas poor pastures may have Prospects While breeding and selection may less than 1000 seeds/m2. improve the persistence and productivity of white Ecology White clover does not grow well on clover in tropical highlands and subtropical areas drought-susceptible soils with low soil moisture where it is adapted, it isunlikel y that breeding will storage or on waterlogged soils, particularly substantially increase its area of adaption. where waterlogging coincides with high tempera­ Literature ;1| Baker, M.J. & Williams, W.M., tures. White clover requires high levels of fertility, 1987. White clover. CAB International, Wall- with a pH(H20) of over 5.0 and preferably 5.5., and ingford, UK. 534 pp. |2;Evan s A.M., 1976. Clovers. available soil P levels ofove r 20mg/kg .I t is suscep­ In: Simmonds, N.W. (Editor): Evolution of crop tible to high levels of available soil Mn and Al. plants. Longman, London, pp. 175-179. 3! Gibson, Agronomy Although hand-harvested seed of P.B. & Cope, W.A., 1985. White clover. In: Taylor, white clover is hard-seeded, commercial seed N.L. (Editor): Clover science and technology. No usually has low levels of hard seed. Under favour­ 25. American Society of Agronomy, Madison, Uni­ able conditions it can be established by broadcast­ ted States, pp.471-490 . \4\Jones , R.M., 1984. White ing into closely grazed pastures, but there is more clover (Trifolium repens) in subtropical southeast prospect ofsucces s with sowing intoroug h cultiva­ Queensland, III.Increasin g clover and animal pro­ tion or a fully prepared seed-bed. Seed should be duction by use of lime and flexible stocking rates. inoculated with an appropriate Rhizobium strain Tropical Grasslands 18: 186-192. 5j Oram, R.N., before sowing if it is sown in an area where it has 1990. Register of Australian herbage plant culti- not been grown before. White clover based pas­ vars. CSIRO,Australia , pp. 111-117. |6| Ostrowski, tures should be well grazed, especially during the H., 1972.Whit e clover (Trifolium repens) in subtro­ summer period when the companion grasses are pical Australia - a review. Tropical Grasslands 6: growing quickly and the clover is growing under 97-106. i7iZohary , M. &Heller , D.,1984 .Th e genus adverse temperature conditions. Bloat (tympani­ Trifolium. The Israel Academy of Sciences and tis) can be a problem in spring when clover growth Humanities, Ahva Printing Press, Jerusalem, pp. is vigorous and there is little grass in the pasture. 167-171. Probably the most important diseases are those R.M. Jones &S.M.M . Kersten associated with weak pathogens such as Pythium middletonii which attack stolons and roots during wet and hot conditions during summer. Rust pus­ Trifolium semipilosum Fresen. tules on leaves caused by Uromyces spp. can be more conspicuous but are much less serious. Pests Flora 22: 52(1839) . of white clover in the subtropics are usually of mi­ LEGUMINOSAE nor importance. Total annual DM yields range 2n = 16 from < 1 to > 7 t/ha. Peak DM yields on offer in Vernacular names Kenya white clover, Kenya well grazed pastures seldom exceed 1t/h a and are clover (En). more often under 0.5 t/ha. However, as white Origin and geographic distribution T. semipi­ clover is selectively grazed such yields substan­ losum occurs naturally in the humid and equatori­ tially improve animal production. Good clover al high altitude areas of Kenya, Uganda, Tanza­ growth results in increased soil nitrogen status. nia, Yemen and Ethiopia. Now it is cultivated Genetic resources and breeding Germplasm elsewhere in the high altitude tropics and in the collections are held in many countries. An impor­ subtropics and sometimes it is naturalized locally. tant collection is held by DSIR Grasslands Divi­ Uses Kenya white clover ismainl y used for graz­ sion, Palmerston North, New Zealand. The main ing ruminants, although it is alsofe d to pigs, chick­ breeding programme to improve white clover for ens and ducks in Papua New Guinea. use in the subtropics has been carried out in Flor­ Properties The N concentration and digestibi­ ida, United States, leading to the release of culti- lity of Kenya white clover leaves and stems are var 'Osceola'. There is some evidence that T. only slightly lower than white clover {Trifolium TRIFOLIUM 227

repens L.) and higher than for almost all tropical continuing development of nodal roots. legumes. Nitrogen concentrations range from Two varieties are distinguished: 2-5% and P concentrations from 0.2-0.5%. Sodi­ - var. semipilosum (synonyms var. microphyllum um percentages are often below 0.1%.Ther e are Chiov. and var. kilimanjaricum Baker), with 700-900 seeds/g. leaflets twice as long as broad and silky pilose Botany A perennial herb, initially with a crown underneath; pods pilose at least at apex. and taproot, with prostrate pilose stems radiating - var. glabrescens Gillett, with leaflets about as from this, which root at the nodes ifi n contact with long as broad, glabrescent underneath and gla­ moist soil. Leaf trifoliolate with pilose petiole brous pods. much longer than the leaflets; leaflets euneate- Var. glabrescens naturally occurs in rather more obovate to elliptical or orbicular, 0.4-2 cm x 0.4 humid areas than var. semipilosum, often in associ­ -1.1 cm, glabrous above and pilose on the margins, ation with Pennisetum clandestinum Höchst, ex midrib and usually on the lower half of the two lat­ Chiov. eral leaflets; petiole length varies with grazing Cultivar 'Safari' is described as being var. glabres­ pressure, from 2-20 cm. Inflorescence globose, cens. 5-40 flowered, about 2 cm across; flowers white to Ecology Kenya white clover grows in the low pale pink; calyx 4-6 mm long, corolla 8-10 mm altitude subtropics and elevated tropics long. Pods oblongoid, 5-6 mm long with 2-6 seeds. (1000-3000 m). In its natural habitat in East Seed irregularly discoid, ca. 1.5 mm across, dull Africa, annual rainfall is 550-1400 mm. Tempera­ yellow, light brown or olive green to black. ture requirements for growth are intermediate be­ The primary taproot usually only lives for 1-2 tween those of most tropical legumes and white years and the prostrate stems then persist through clover. It grows very well in Papua New Guinea at sites with a diurnal temperature range of 10-20°C and at elevations as low as 1300 m, pro­ vided the soil is fertile. It can flower and set seed on the equator and also at latitudes in excess of 30°. In the subtropics, flowering occurs mainly during the cooler months. Plants of 'Safari' can resist moderate frosting but top growth is killed by heavy frosts. It grows at pH range of 5-7.5 with greater tolerance of low Ca and high Al and Mn than white clover. It requires levels of available soil P > 20 mg/kg. It cannot tolerate very dry or waterlogged sites. In Papua New Guinea it grows very well in abandoned gardens and along road­ sides. Agronomy Seed of 'Safari' can be very hard- seeded, particularly when hand-harvested, and scarification is usually required. Seed should be inoculated with the appropriate rhizobium inocu­ lum (e.g. CB782). Although it can be established by surface broadcasting into undisturbed swards, it isbes t established into a seed-bed.I n the subtrop­ ics, it is best to sow in autumn when temperatures are falling yet soil moisture is often favourable. First year pastures often go through a period when growth is poor and leaves pucker and turn yellow, orange or red. The reasons for this are not clear, although sometimes ascribed to 'rugose leaf curl', but the plants often or usually grow out ofth e con­ dition. However, new plants arising from seedling recruitment in an established sward can also be Trifolium semipilosum Fresen. -1, flowering plant; affected. 2,flower; 3, fruit. The main disease described for 'Safari' has been 228 FORAGES rugose leaf curl. This was initially ascribed to a Agronomy of Kenya white clover cultivar Safari. virus and then to a Rickettsia-like organism. How­ CSIRO, Australia, Division of Tropical Crops and ever, the symptoms on young stands may not nec­ Pastures. Information service. Sheet 41-3.4 pp . ,4 essarily be caused by that disease. Slugs can sever­ Oram, R.N., 1990. Register of Australian herbage ely defoliate swards of 'Safari' during sustained plant cultivars. CSIRO, Australia, pp. 163-164. |5j periods of wet weather, and weevils can damage Skerman, P.J., Cameron, D.G. &Riveros' , F., 1988. swards by damaging or severing larger roots. Rot­ Tropical forage legumes. FAO, Rome. pp. 456-462. ting of roots and stolons by Pythium spp. has been :6|' t Mannetje, L., 1966.A punche d card key to spe­ recorded during hot, moist conditions. cies of Trifolium L. in Africa, south of the Sahara, 'Safari' combines well with a widerang e of tussock excluding Ethiopia. East African Agriculture and and stoloniferous grasses in mixed pastures. Al­ Forestry Journal 31: 261-270. |7j Zohary, M. & though 'Safari' stems can grow upwards from the Heller, D., 1984. The genus Trifolium. The Israel soil surface, 'Safari' pastures should be well Academy of Sciences and Humanities, Ahva Print­ grazed, particularly when the grasses are growing ing Press, Jerusalem, pp. 176-177. actively. This ensures that stolons remain close to R.M. Jones &A.K . Benjamin the ground and continue to develop new nodal roots to make up for the death of older roots. 'Safari' swards can develop high reserves of soil Tripsacum andersonii J.R. Gray seed, of over 10000 seeds/m2, but recruitment of new seedlings is not reliable. 'Safari' can spread Phytologia 33: 204(1976) . through seed dispersed in faeces. Given levels of GRAMINEAE above 10-20% legume in the sward, 'Safari' pas­ In = 64 tures can give high production of milk or live- Vernacular names Guatemala grass (En). weight gain. Malaysia: rumput jelai, rumput jagung. Thailand: Genetic resources and breeding Germplasm ya-kuatemala. collections are held by ATFGRC (CSIRO, Austra­ Origin and geographic distribution Guate­ lia) and ILCA (Ethiopia).Attempt s to cross 'Safari' mala grass isnativ e to Central America and north­ with T. repens have been unsuccessful. Some ern South America (from Mexico to Peru).I t is now plants in Ethiopia and Kenya look like intermedi­ widely cultivated throughout the tropics includ­ ates between T. semipilosum and T. burchellianum ing South-East Asia. Ser., and var.glabrescens might be originally a nat­ Uses Green foliage of Guatemala grass is widely ural hybrid of these 2 species. used as cut fodder. In Sri Lanka, it has been used Prospects It is highly unlikely that the zone of as a soil cover in tea plantations. In Malaysia, it adaptation will be extended beyond the wetter has been planted extensively in rubber and oil subtropics and elevated tropics. Even within this palm estates for control of erosion, for mulching zone, the main limitation of'Safari' isit s unreliabi­ and as a soil conditioner in drained swamps. It can lity of successful establishment and persistence, be a weed in cultivated areas. which is partly related to the condition described Properties Regrowth harvested at 4, 6, 10 and as rugose leaf curl. Although it is showing consid­ 12 weeks had N concentrations of 2.2, 2.1,1. 3 and erable promise in Papua New Guinea highlands, 0.8%. Digestible crude protein percentages at the this unreliability will continue to limit its use in same growth stages were 9.8%, 9.1%, 5.0% and many areas unless the problems can be defined and 2.2% as assessed by using cattle. Six-week-old solutions found. regrowth from plants grown on 'acid sulphate' soil Literature ill Cook, B.G., Mulder, J.C. & Pow­ contained 1.5% N, 0.14% P,2.6% oK , 0.13% Ca and ell, B., 1985.A survey to assess the influence of en­ 0.11% Mg. vironment and management on frequency, vigour Botany Robust perennial grass, forming large and chemical composition of Trifolium semipilo­ mats up to 5 m across of tangled stolons and rhi­ sum in south eastern Queensland. Tropical Grass­ zomes, from which erect to decumbent flowering lands 19: 49-59. |2| Gillett, J.B., 1971. Trifolieae. culms are produced; stolons and culms up to 5 cm In: Milne-Redhead, E. & Polhill, R.M. (Editors): in diameter, flowering culms 2-3 m tall. Lower Flora of tropical East Africa. Leguminosae 4 - leaf-sheaths shortly pubescent near the base, Papilionoideae 2. Crown Agents for Oversea Gov­ upper ones glabrous; ligule a membraneous ridge, ernments and Administrations, London, pp. slightly hairy behind; leaf-blade sessile, linear-lan­ 1027-1028. |3| Jones, R.M. & Cook, B.G., 1981. ceolate,u pt o 120c m x 5-10 cm,shortl y tomentose TRIPSACUM 229

in suitable sites at higher altitudes (about 1800 m) has also been observed. Good growth can also be maintained when planted in plantations with about 50% shade. It is adapted to a wide range of soil types including ultisols, oxisols, peats, acid sulphate soils and Bris soils (coastal marine sands) with a pH(H20) range of 2.7-4.5. Guatemala grass is adapted to inundated and marshy conditions. It has remarkable tolerance to shallow water tables, but poor tolerance to soil moisture stress. Agronomy Guatemala grass isplante d vegetati- vely, either through splitting the rooted tillers from clumps, byrhizome s or through stem cuttings containing 3 nodes. There are no known commer­ cial suppliers of seed in South-East Asia. Green foliage is cut by hand for pen feeding rather than being conserved for hay. Thevigorou s growth of Guatemala grass has been demonstrated in its annual DM yields, at a 6-week cutting interval with maintenance fertilization of 200 kg/ha of N, 40 kg/ha of P and 100 kg/ha of K per year, of 23 t/ha on a sedimentary soil, 21 t on a peat soil, 24 t on an acid sulphate soil and 2 t on a Bris soil. The low production on the Bris soil is attributed to water stress and high soil temperatures. It outyielded 20 other grass species on acid sulphate soil. While only 30% of plants died over a 3-year Tripsacum andersonii J.R. Gray - 1, habit; 2, cutting experiment, mortality might have been ligule; 3, inflorescence; 4, male spikelet; 5, female higher if plants had been subjected to more fre­ spikelet. quent defoliation. Adequate soil moisture (either from rainfall or irrigation) and fertilizer are the above, glabrous below. Inflorescences terminal two important factors involved in maintaining and axillary, composed of 3-7 branches (terminal) productive Guatemala grass. In view of its declin­ and 1-4 branches (axillary); branches subdigita- ing quality with increasing age of regrowth, sup­ tely arranged on a short primary axis and each plementation with a leguminous shrub such as raceme consists of several basal fruitcases (pistil­ Leucaena leucocephala (Lamk) de Wit may best late spikelets) with numerous staminate spikelets exploit its high yield potential to optimize animal on a continuous rachis above them; pistillate spik­ production. No serious disease or pest problems elets 6-10 mm long, sunken in cavities ofth e indur­ have been noted so far. ated rachis and covered by the outer glume, disar­ Genetic resources and breeding It is unlikely ticulating individually at maturity; staminate that any substantial germplasm collections are spikelets paired at each rachis node (one sessile, being maintained and there are no known breed­ one shortly pedicelled), 6-10 mm long, with coria­ ing programmes. T. andersonii behaves cytologi- ceous 10-15-nerved glumes. cally as if its genome consists of a triploid Tripsa­ In Central America T. andersonii is often confused cum L. genome (54 chromosomes) and a haploid with T. latifolium Hitchcock (2n = 36), T. laxum Zea L. genome (10 chromosomes). Cytogenetic Nash (2n = 36), and T. maizar Hernandez & Ran­ studies indicate that T. andersonii originated as dolph (2n = 36, 72). T. laxum and T. maizar bear a hybrid between a species of Tripsacum (2n = 36) 10-50 racemes per terminal inflorescence and the and a species oîZea (2n = 20).Th e Tripsacum par­ male section of each raceme is typically pendent. ent probably was T. latifolium of Central America T. latifolium has sessile, 4-6 mm long, staminate and the Zea parent either Z. mays L. ssp. mays spikelets. T. andersonii is very sterile. (domesticated maize) or Z. mays L. ssp. mexicana Ecology Although known for its excellent per­ (Schrad.) Iltis (annual teosinte). formance in lowland conditions, vigorous growth T. andersonii is remarkably uniform morphologi- 230 FORAGES cally, indicating that the species had a single Uses Sabi grass is mainly used as a forage for hybrid origin. Its wide distribution in Central and ruminants but can also be used for reseeding South America suggests a rather ancient origin. denuded areas. Prospects Under high rainfall tropical environ­ Properties Chemical composition and nutritive ments, Guatemala grass isa proven species for ero­ value of Sabi grass depend on soil fertility but sion control, soil conditioning and mulching, and young green leaves typically contain up to 2.5% it is also a high-yielding forage species. If com­ N, 0.2% P and are 65-70% digestible. In the late bined with legumes in a feeding system, its utiliza­ wet season equivalent values are 1.2 %, 0.15% and tion for animal production could be further 55-60%i respectively. Dry leaves and stems are improved. much lower in quality and typical values are 0.5% Literature 1 Aminah, A. & Wong, C. C, 1991. N and 0.05% P. Sabi grass is palatable. There are Dry matter productivity and chemical composi­ 600-1000 seeds/g. tion of some promising grasses grown on acid sul­ Botany A creeping, perennial grass of variable phate soil.In :Recen t innovation in the animal and size and growth habit usually with short stolons animal products industry. 14th Malaysian Society or often tufted. Culms smooth, more or less erect, for Animal Production (MSAP) Conference, Gen- up to 150 cm tall, 5 mm in diameter, often branch­ ting Highlands, Malaysia, May 8-9, 1991. MSAP ing at the nodes, and sometimes rooting at lower and The Federation of Livestock Farmers Associa­ nodes; nodes prominent, densely covered with tion of Malaysia (FLFAM), Kuala Lumpur, pp. short, silky hairs. Leaf-sheaths close, covered with 92-96. |2| Devendra, C, 1979. Malaysian feed- erect, white, tubercle-based hairs particularly in ingstuffs. MARDI, Kuala Lumpur, Malaysia. 145 their upper part, shorter than the internodes; pp. |3!d e Wet, J.M.J., Fletcher, G.B., Hilu, K.W. ligule a rim of short (1 mm) hairs. Leaf-blade &Harlan , J.R., 1983.Origi n of Tripsacum anderso- broadly linear to narrowly lanceolate, 2-30 cm x nii (Gramineae). American Journal of Botany 70: 706-711. 4; de Wet, J.M.J., Gray, J.R. & Harlan, J.R., 1976.Systematic s of Tripsacum (Gramineae). Phytologia 33: 203-227. |5| Wong, C.C., Chen, C.P. & Ajit, S.S., 1982. A report on pasture and fodder introduction in MARDI. MARDI Report No 76. MARDI, Kuala Lumpur. 35pp . |6iWycherley , P.R. & Ahmad Azli, M.Y., 1974. Grasses in Malaysian plantations. Rubber Research Institute of Malay­ sia. Rajiv Printers, Kuala Lumpur, pp. 58-59. C.P. Chen

Urochloa mosambicensis (Hack.) Dandy

J. Bot. 69:5 4(1931) . GRAMINEAE 2n = 28, 30, 42 Synonyms Panicum mosambicense Hack. (1888). Vernacular names Sabi grass (En). Thailand: ya sabee. Origin and geographic distribution Sabi grass originated in the drier (400-800 mm annual rainfall), frost-free regions ofsouther n and eastern Africa. It has been introduced into many countries including Australia, India, Sri Lanka, Burma, Thailand, Indonesia, Fiji and Hawaii. It is used commercially and is now naturalized in northern Urochloa mosambicensis (Hack.) Dandy - 1, habit; Australia. 2, inflorescence; 3, spikelets. UROCHLOA 231

3-20 mm, pale to bright green, hirsute on both sur­ There are no important diseases or pests of Sabi faces; midrib prominent, often purplish, apex grass. tapering to a fine point. Inflorescence composed Sabi grass can be used by continuous or rotational of 3-15 racemes (3-90 mm long) on a simple com­ grazing, or by cutting for feeding as fresh material mon axis up to 15c mlon g but often much less; spik- or for making hay. It will stand close defoliation. elets are borne on the lower side ofth e rachis only, Sabi grass responds to N and P and is aggressive 3-5 mm x 1.5-3 mm; lower glume ovate-lanceo­ at high fertility. It combines well with legumes and late, 3-nerved, with a single bristle 1-2 mm long; is a common associate grass for Stylosanthes spp. lower lemma with a conspicuous fringe of bristles Sabi grass produces 1-8 t/ha of DM per year 1-1.5 mm long. Caryopsis light buff or cream. depending on seasonal conditions, soil fertility Seeds of Sabi grass germinate early in the wet sea­ and associate species. son and vegetative growth continues until soil Genetic resources and breeding There is wide water is exhausted. Flowering commences 3-4 variation in Sabi grass for morphological and weeks after the start of the wet season and conti­ agronomic characteristics. The collection held at nues until growth ceases. Seed matures in 3-4 ATFGRC (CSIRO, Australia) mainly came from weeks. Leaves live for 5-25 weeks depending South Africa. mainly on water supply. Plants are often short­ There are no active breeding programmes and im­ lived (3-4 years) but new plants are formed from provement will depend on selection from natural seedling recruitment. Cultivar 'Nixon' has been populations. Important objectives to date have developed in Australia from material introduced been adaptability to a range ofenvironments , yield from Zimbabwe. and quality. Sabi grass is closely related to and much resembles Prospects Sabi grass is an effective colonizing, Urochloa oligotricha (Fig. & De Not.) Henrard palatable, and grazing-tolerant forage of reason­ which has a 5-nerved lower glume, but intermedi­ able quality suited for sowing in the drier regions ates occur. Both species also resemble the South of South-East Asia. It is recommended for use in African Urochloa stolonifera (Goossens) Chippend, the 'Three Strata Forage System' in drier areas of which is smaller and with its spikelets untidily Indonesia. arranged. Literature 111 Bogdan, A.V., 1977. Tropical pas­ Ecology Sabi grass is commonly grown in areas ture and fodder plants. Longman, New York. pp. with 500-1200 mm annual rainfall with a pro­ 295-296. |2| Burt, R.L., Williams, W.T., Gillard, P. nounced warm season and a dry season of 5 to 9 & Pengelly, B.C., 1980. Variation within and be­ months. It has poor frost tolerance and rapidly hays tween some perennial Urochloa species. Austra­ off when soil moisture is exhausted. In its natural lian Journal of Botany 28: 343-356. 13! Clayton, habitat in Africa of wooded grassland and deci­ W.D. &Renvoize, S.A., 1982.Urochloa . In: Polhill, duous bushland, it occurs up to 1400 m altitude. R.M. (Editor): Flora oftropica l East Africa. Grami- Flower initiation occurs over a range of daylengths neae. A.A. Balkema, Rotterdam. Part 3. pp. but it is earlier at 12hour s than 9o r 15hours . Sabi 600-607. j4| Devendra, C. (Editor), 1990. Shrubs grass is adapted to a wide range of soils but will and tree fodders for farm animals. Proceedings of not tolerate permanently flooded or waterlogged a workshop held in Denpasar, Indonesia, 24-29 conditions. In Africa it is a common roadside weed July 1989. IDRC, Ottawa, Canada. 349 pp. |5| Mc- and often grows in disturbed or overgrazed areas. Ivor, J.G., 1985. The growth of Urochloa acces­ Agronomy Although Sabi grass can be propa­ sions in grazed swards with Stylosanthes near gated vegetatively it is normally established from Townsville, north Queensland. Australian Jour­ seed. Freshly harvested seed is dormant but dor­ nal of Experimental Agriculture 25: 61-69. 161 mancy breaks down after 9 to 12 months storage. Oram, R.N., 1990. Register of Australian herbage Hammer milling to destroy the hard lemma can plant cultivars. CSIRO, Australia, pp. 87-88. 17! increase germination. Sabi grass can be readily es­ Whiteman, P.C. & Gillard, P., 1971. Species of tablished under a variety ofsowin g conditions, but Urochloa as pasture plants. Herbage Abstracts41 : is best established by sowing on or near the soil 351-357. surface of a well cultivated seed-bed during the J.G. Mclvor early wet season. Oversowing into undisturbed pasture isofte n a complete failure, or takes several years to develop into a pasture. Seeding rates range from 1-5 kg/ha. 232 FORAGES

Vigna parkeri Baker

Journ. Bot. 20:6 9(1882) . LBGUMINOSAE 2re = 22 Synonyms Dolichos maranguënsis Taub. (1892), Vigna maranguënsis (Taub.) Harms (1915), V. gra­ cilis sensu auctt., non (Guill. &Perr. ) Hook.f. Vernacular names Creeping vigna, (Papua New Guinea) (En).Philippines :balaton g (Bontok), bulligan (Ifugao). Origin and geographic distribution Creeping vigna isnativ e to tropical East and Central Africa, but has been recorded most frequently in Kenya, Uganda and Tanzania. It is being sown in the humid subtropics in Australia and Florida (United States) and has become naturalized in the high­ lands of Papua New Guinea. Uses While the main use of creeping vigna is as a pasture legume, it is also proving useful in Papua New Guinea as a protein supplement for pigs, and as a green manure and cover crop in shifting culti­ vation. Properties Creeping vigna provides quality for­ age. In one report, leaves comprised 65% ofth e top growth and had a N concentration of4. 1% and DM digestibility of 61%, while stems had a N concen­ tration of 2% and DM digestibility of 55%. It is generally readily eaten by cattle but there is some Vigna parkeri Baker - 1, flowering and fruiting doubt about its acceptance by goats. plant part; 2,leaf. Botany A perennial twining or prostrate stolo- niferous herb. Stolons remain appressed to the soil parkeri (flowers blue or rose, leaflets ovate, only surface, while the primary axis, and laterals devel­ on Madagascar); ssp. acutifolia Verde, (flowers oping from stolon nodes, have a twining habit. mainly yellow, leaflets elliptical or ovate, in Taproot slender, freely nodulating; nodal roots at Kenya, Tanzania, ); and ssp. maran­ first fibrous becoming small taproots as new guënsis (Taub.) Verde, (flowers mainly blue or pur­ crowns develop. Stems slender, sparsely to densely ple, leaflets round, in East and Central Africa). covered with mostly spreading hairs. Leaves trifo- The occurrence ofman y intermediate forms makes liolate; leaflets round, ovate to ovate-lanceolate, the subdivision unpractical. 1-9 cm x 1-5 cm, rounded to acuminate at the One introduction of creeping vigna which became apex and rounded to subacute at the base, pubes­ naturalized in parts of south-east Queensland has cent on both faces, margins entire and densely been released as cultivar 'Shaw' and is now being ciliate; petiole 1-8.5 cm long; stipules lanceolate, planted in other parts of the humid subtropics and up to 8 mm long, bilobed at the base, persistent. the tropical highlands. It appears that only one Inflorescence axillary, with 2-5(-10) flowers per form (of ssp. maranguënsis) was introduced from raceme; peduncle 2-13 cm long; flowers blue, yel­ Africa into Papua New Guinea at Aiyura in the low or white, occurring in alternate pairs inserted Eastern Highlands about 1958, from where it has on either side of a glandular node;standar d oblate, spread to the Southern Highlands, Western High­ 5-8(-12) mm x 5-8(-10) mm, glabrous. Pod lands and Enga Provinces. linear-oblong, compressed, mostly l-2(-3) cm x The ecotype from Papua New Guinea and 'Shaw' 4.5-5.5 mm, 2-5-seeded. Seed oblong-ovoid, 3-4 are distinguishable by the more acuminate, often mm x 2-3 mm x 2 mm, grey-to-brown/black mot­ paler leaflets and the stronger twining tendency tled. in the former. Both have blue flowers and a pale Three subspecies have beeen distinguished: ssp. crescent on the leaflets of some plants. In south- VlGNA 233

eastern Queensland, latitude 26°S , some types swards in Queensland, but had no lasting effect. flower throughout the growing season, while Two other fungi, Rhizoctonia solani and Colletotri­ others, including the Papua New Guinea ecotype chum truncatum have been isolated from leaf and and 'Shaw' do not flower until about May. The stem lesions in seed crops of 'Shaw', but can be Papua New Guinea ecotype flowers throughout largely controlled by avoiding accumulation of a the year in its naturalized habitat in the highlands bulk ofmatur e top growth. between 5°an d 7°15'S. Creeping vigna is usually grazed since its pro­ Ecology In general, creeping vigna has similar strate, low twining growth habit does not readily environmental requirements as greenleaf desmo- lend itself to a cut-and-carry system. High quality dium (Desmodium intortum (Miller) Urban). The hay can be made from creeping vigna, but weather Papua New Guinea ecotype and 'Shaw', both of conditions in its preferred habitats are rarely ideal which have uncertain origins, are adapted to moist for hay-making. Improved utilization by cattle of soils in the high altitude tropics and low altitude grasses such as Pennisetum clandestinum, Axono- subtropics where annual rainfall exceeds 1100 mm pus affinis A. Chase and even Imperata cylindrica per year. They appear to grow best at cooler tem­ (L.) Raeuschel has been observed where they are peratures, and when cut by frost, mostly recover associated with this legume. from crowns. They are tolerant of poorly fertile, No yield data are available from South-East Asia, acid soils (down to pH(H20) 5.0), and although but in Queensland, autumn presentation DM both are also tolerant of low available soil P, have yields of 1650 kg/ha of 'Shaw' were obtained from responded to applied P. They cannot withstand a grass/legume pasture stocked at 1.5 beast/ha. prolonged dry periods, but in the event of stand Genetic resources and breeding A limited loss through drought, they can regenerate from range of germplasm material is held at ATFGRC the often high levels of soil seed, measured at 50 (CSIRO, Australia). The species is only adapted to 2 2 seeds/m under close grazing to over 1000 seeds/m humid highland areas in South-East Asia and has under lenient grazing. Both possess moderate not attracted and does not warrant breeding work, shade tolerance. It can spread through dense at least not until wild types have been fully grasses such as Pennisetum clandestinum Höchst, exploited. ex Chiov. by stolons, sometimes at over 1 m/year. Prospects Creeping vigna is a useful species, It may be spread over considerable distances especially in acid soil areas where sward-forming through ingested seed being voided in the dung. grasses are common and in which pastures are sub­ Agronomy High levels of hard seed have been jected to heavy grazing. At this stage, its potential recorded and, where immediate germination is has only been partly realized. Evaluation of exist­ required, seed may need to be scarified. This can ing collections in tropical highland and subtropi­ be achieved using mild abrasion or hot water (pre­ cal lowland areas in South-East Asia is required. liminary results suggest 10minute s at 70°C). Seed Literature il! Cook, B.G. & Jones, R.M., 1987. is best sown at 2 kg/ha into a clean, well prepared Persistent new legumes for intensive grazing. 1. seed-bed. It may be sown on the surface or at shal­ Shaw creeping vigna. Queensland Agricultural low depth. Subsequent rolling improves establish­ Journal 113: 89-91. 2 Gillett, J.B., Polhill, R.M. ment. Native cowpea rhizobia give effective nodu- &Verdcourt , B.1971 .Papilionoidea e (2).In : Milne- lation. Renovation of old stands can be achieved Redhead, E. & Polhill, R.M. (Editors): Flora of through limited cultivation, using soil seed tropical East Africa. Leguminosae 4- Papilionoi­ reserves. Where seed is unavailable or more rapid deae 2. Crown Agents for Oversea Governments establishment is required, creeping vigna may be and Administrations, London, pp. 635-637. 3 established from rooted stolons. Jones, R.M. &Clements , R.J. 1987.Persistenc e and Creeping vigna is adapted to both lenient and productivity of Centrosema virginianum and heavy grazing. However, to achieve maximum Vigna parkeri cv. Shaw under grazing on the benefit as a forage and for N fixation, stands coastal lowlands of South-East Queensland. Trop­ should bemaintaine d between 20an d 50cm .I n low- ical Grasslands 21: 55-64. 4 Oram, R.N., 1990.Reg ­ fertility soils, nutrient deficiencies should be cor­ ister of Australian herbage plant cultivars. rected, paying special attention to P and Mo. CSIRO, Australia, pp. 269-270. 5! Pentney, C.J., In Papua New Guinea, no major disease or pest Whiteman, P.C., & Sivasupiramaniam, S. 1984. problems have been recorded. A fungus (Sclero­ Studies on the germination, phenology and Rhizo- tium rolfsii) and a root-knot nematode (Meloido- bium requirements of Vigna parkeri. Tropical gyne javanica) have caused some damage in 'Shaw' Grasslands 18: 66-74. |6| Verdcourt, B. 1979. A 234 FORAGES manual of New Guinea legumes. Botany Bulletin Number 11.Offic e of Forests, Division of Botany, Lae, Papua New Guinea, pp. 522-523. B.G. Cook &A.K . Benjamin

Zoysia matrella (L.) Merrill

Philip. J. Sc. 7:23 0 (1912). GRAMINEAE 2n = 20, 40 Synonyms Agrostis matrella L. (1771), Zoysia pungens Willd. (1801),Z. tenuifolia Willd. ex Thiele (1834). Vernacular names Manila grass, Siglap grass, Korean grass (En). Mascarene grass (Am). Chien­ dent gazon (Fr).Indonesia : jukut kakawatan hija u (Sundanese), rebha sekem-sekeman (Madura), bu- rikit (Ceram). Malaysia: rumput siglap. Philip­ pines: damong-alat, barit-baritan (Tagalog), malakuwerdas (Pangasinan). Singapore: Siglap grass. Thailand: ya-nuannoi. Vietnam: cong cong. Origin and geographic distribution Manila grass occurs naturally along the coasts ofth e Indi­ an Ocean, the Chinese Sea to the Ryukyu Islands, and particularly in all South-East Asian countries. Now it is extensively cultivated (also inland) pan- tropically. Uses Manila grass is used as forage on sandy Zoysia matrella (L.) Merrill - 1, habit flowering soils in coastal areas where other grasses are not plant; 2, ligule; 3,part of inflorescence. adapted. It is a good sand-binder and is cultivated for lawns. tinctly exserted above the foliage, up to 4.2 cm Botany A mat-forming perennial, often stoloni- long; distribution as above. ferous, rarely rhizomatous, up to 35 cm high. Sto­ - var. pacifica Goudswaard: leaf-blade 0.7-1 mm lons to 45c m long. Leaf-sheath hairy at the throat; wide, usually erect; peduncle, even in fruit, not ligule membranous and finely hairy, ca. 0.25 mm or hardly exserted from the uppermost sheath, long; leaf-blade lanceolate, up to 8 cm x 3 mm the raceme therefore not or little exserted above when expanded, erectopatent to patent, base more the foliage, up to 1.5 cm long; distribution: or less cordately contracted into a short pseudo- Taiwan to the Ryukyu Islands and Loo Choo petiole. Inflorescence a terminal, spike-like Island, Marianes, Solomons and in the Moluccas. raceme, up to 4 cm long, rachis somewhat wavy; Ecology Z. matrella can grow up to 300 m alti­ pedicels 0-3 mm long; spikelets ovoid-oblongoid, tude, but is primarily adapted to very sandy soils 2-3.8 mm x 1 mm; lower glume usually absent, on coastal areas. It is often found covering the upper glume 5-nerved; palea absent. Caryopsis ground under coconut plantations on sandy coast­ with a straight truncate base. al soils. Seedling growth is slow but after 5-8 weeks strong Agronomy It is easily established by rhizomes. new shoots send out tough rhizomes leading to the Frequent grazing is required to stimulate the formation of a turf mat. There is considerable vari­ growth of young leaves, which are palatable to ation in growth habit, rhizome development and stock. It is low yielding and not suited to cutting the shape of spikelets. for forage. When used as a lawn grass it is fre­ Two varieties are distinquished: quently mown to keep it weed-free and green. - var. matrella: leaf-blade 1.5-2.6 mm wide, more Genetic resources and breeding It is unlikely or less spreading; peduncle exserted from the that substantial germplasm collections are being uppermost sheath at anthesis, the raceme dis- maintained. ZoYsiA 235

Prospects Theforag e yield from this species will inevitably be very low and thus research is not warranted. Literature jl! Gilliland, H.B., Holttum, E.E. & Bor, N.L., 1971. Grasses of Malaya. In: Burkill, H.M. (Editor): Flora of Malaya. Vol. 3. Govern­ ment Printing Office, Singapore, pp. 109-111. |2j Goudswaard, P.C., 1980.Th e genus Zoysia (Grami- neae) in Malesia. Blumea 26:169-175. 13, Häfliger, E. & Scholz, H., 1981. Grass weeds 2. Ciba-Geigy Ltd., Basel, Switzerland, p. 137. '4| Mehra, K.L. & Fachrurozi, Z., 1985. Indonesian economic plant resources: forage crops. Lembaga Biologi Nasio- nal - LIPI, Bogor, No 31. p. 51. C. Manidool 3 Minor forages

In this tentative list the species name is followed by its botanical family name, and synonyms are given in the following, indented lines. The list contains species which the major historical handbooks on economically useful plants of South-East Asia have noted as primarily being used for forage. In the Prosea books species of minor importance within a commodity group are normally described briefly. In this volume it was decided not to dothi s because theoretically all non-poisonous plant speciesmigh t beconsidere d tob eforages . Nevertheless, itwa sthough t useful to present this list of 'minor forage species' as compiled in the Prosea documenta­ tion system, realizing that the list in no way claims to be complete.

Acroceras crassiapiculatum (Merrill) Alston - Gramineae Panicum crassiapiculatum Merrill rigidula Steudel var. rigidula - Gramineae Agrostis infirma Buse Acroceras tonkinense (Belanse) Bor - Gramineae Acroceras sparsum Stapf Acroceras zizanioides (Kunth) Dandy - Gramineae Panicum oryzoides Swartz Acroceras oryzoides (Swartz) Stapf ciminica (L.) Stapf- Gramineae Coridochloa cimicina Nees (R. Br.) Hitchc. - Gramineae Coridochloa semialata Nees Alysicarpus rugosus (Willd.)DC . -Leguminosae Hedysarum rugosum Willd. Alysicarpus wallichii Wight & Arnott Apluda mutica L.- Gramineae Apluda varia Hackel Arthraxon castratus Bor- Gramineae Arthraxon pilipes Backer Arthraxon hispidus (Thunberg ex Murray) Makino - Gramineae Arthraxon hispidus (Thunberg ex Murray) Makino var. robustior Weizen - Gra­ mineae Arthraxon typicus Koord. Arthraxon microphyllus Höchst. - Gramineae Arundinella holcoides (Kunth) Trinius - Gramineae Arundinella agrostoides Trinius Arundinella nepalensis Trinius - Gramineae Arundinellapumila (Höchst.) Steudel - Gramineae Arundinella tenella Nees & Wight MINOR FORAGES 237

Arundinella setosa Trinius - Gramineae Asthenochloa tenera Buse - Gramineae Avena fatua L.- Gramineae Avena sativa L.- Gramineae Avena sterilis L.- Gramineae Bauhinia hirsuta Weinmann - Leguminosae Bothriochloa bladhii (Retzius) S.T. Blake - Gramineae Amphilophis glabra Stapf Andropogon intermedins R. Br. Amphilophis intermedia Stapf Bothriochloa modesta (Backer) Backer &Henrar d - Gramineae Andropogon modestus Backer Brachiaria eruciformis (J.E. Smith) Griseb. - Gramineae Panicum eruciforme J.E. Smith Brachiaria ramosa (L.) Stapf- Gramineae Panicum ramosum L. Brachiaria reptans (L.) C.Gardne r &C.E . Hubbard - Gramineae Panicum reptans L. Urochloa reptans (L.) Stapf Brachiaria villosa (Lamk) A. Camus - Gramineae Panicum coccospermum Steudel sylvaticum P. Beauv. - Gramineae Briza minor L.- Gramineae Bromus catharticus Vahl - Gramineae Bromus unioloides Kunth Bromus insignis Buse - Gramineae Calamagrostis australis Buse - Gramineae Capillipedium assimile (Steudel) Camus - Gramineae Andropogonparviflorus Backer, non Roxburgh Carex baccans Nees - Carex remota L. ssp. alta (Boott) Kuek. - Cyperaceae Cenchrus inflexus R. Br.- Gramineae Ceratonia siliqua L.- Leguminosae Chloris inflata Link - Gramineae Chloris barbata auct., non Swartz Chloris pumilio R. Br.- Gramineae Chloris divaricata R. Br. (misapplied to Chloris pumilio R. Br.) Chloris truncata R. Br. - Gramineae Chloris virgata Swartz - Gramineae Chrysopogon subtilis (Steudel) Miquel - Gramineae Andropogon subtilis Steudel Coelachne infirma Buse - Gramineae Coelachnepulchella auct., non R. Br. Commelina diffusa Burm.f. - Commelinaceae Commelina nudiflora L. Cyperus brevifolius (Rottb.) Hassk. - Cyperaceae Kyllinga brevifolia Rottb. Cyperus castaneus Willd. - Cyperaceae Cyperus compressus L.- Cyperaceae 238 FORAGES

Cyperus haspan L.- Cyperaceae Cyperus iria L. Cyperus kyllingia Endl. - Cyperaceae Kyllinga monocephala Rottb. Cytisuspalmensis (Christ) Hutch. - Leguminosae Dactylis glomerata L.- Gramineae Dactyloctenium verticillatum Backer - Gramineae Dichanthium caricosum (L.) Camus - Gramineae Andropogon caricosus L. Diectomis fastigiata (Swartz) Kunth - Gramineae Andropogon fastigiatus Swartz Digitaria abludens (Roemer &Schultes ) Veldk.- Gramineae Digitaria pedicellaris Prain Digitaria eminens (Steudel) Backer - Gramineae Digitaria junghuhniana (Steudel) Henrard - Gramineae Digitaria perrottetii Backer Digitaria longiflora (Retzius)Pers.-Gramineae Digitaria ternata (A.Richard ) Stapf-Gramineae Digitaria ropalotricha Buse Digitaria violascens Link - Gramineae Digitaria pertenuis Buse retroflexa (Vahl) Panz. - Gramineae Dinebra arabica Jacq. Drymaria cordata (L.)Willd . ex Roemer &Schulte s - Caryophyllaceae Drymaria hirsuta Bartling - Caryophyllaceae Echinochloa stagnina (Retzius) Beauv. - Gramineae Panicum stagninum Retzius Ectrosia leporina R. Br. - Gramineae Enteropogon dolichostachyus (Lagasca) Keng - Gramineae Chloris digitata Steudel Epipremnum pinnatum (L.) Engl.- Araceae Rhaphidophora merrillii Engl. Eragrostis atrovirens (Desf.) Trinius ex Steudel - Gramineae Eragrostis elegantula Steudel Eragrostis bahiensis Schultes - Gramineae Eragrostis elongata (Willd.) Jacq. - Gramineae Eragrostis brownii Nees ex Hook. & Arnott Eragrostis interrupta (R. Br.) Beauv. - Gramineae Eragrostis japonica (Thunberg ex Murray) Trinius - Gramineae Eragrostis (Koeler) Link - Gramineae Eragrostis cilianensis (All.) Link ex Vign. Eragrostis nigra Nees ex Steudel - Gramineae Eragrostispilosa (L.) Beauv. - Gramineae Eremochloa ciliaris (L.)Merril l - Gramineae Eremochloa horneri Buse Eremochloa malayana Ridley Eriachne obtusa R. Br.- Gramineae Eriachnepallescens R. Br. - Gramineae Eriachne tincta Nees - Gramineae MINOR FORAGES 239

Eriachne triseta Nees ex Steudel - Gramineae nubica (Steudel) Hackel &Stap f ex Thell. - Gramineae Eriochloa decumbens F.M. Bailey Eriochloaprocera (Retzius) C.E. Hubbard - Gramineae Eriochloa ramosa (Retzius) 0. Kuntze Eriochloa subglabra (Nash) Hitchc. - Gramineae Euchlaena mexicana Schrader - Gramineae Eulalia fimbriata (Hackel) O. Kuntze - Gramineae Pollinia fimbriata Hackel Eulalia fulva (R. Br.) O. Kuntze - Gramineae Pollinia fulva (R. Br.) Benth. Eulalia quadrinervis (Hackel) 0. Kuntze - Gramineae Pollinia quadrinervis Hackel Eulalia trispicata (Schultes) Henrard - Gramineae Pollinia argentea (Brongn.) Trinius Eustachys tenera (Presl) Camus - Gramineae Chloris tenera Scribner Festuca leptopogon Stapf- Gramineae Festuca nubigena Jungh. - Gramineae Ficus globosa Blume - Moraceae Ficus hirta Vahl - Moraceae Fimbristylis annua (All.) Roemer &Schulte s - Cyperaceae Fimbristylis miliacea (L.)Vah l - Cyperaceae stricta Brongn. - Gramineae Hackechloa granulans (L.) 0. Kuntze - Gramineae Manisuris granulans L.f. Helictotrichon virescens (Steudel) Henrard - Gramineae Avena junghuhnii Buse Hemarthria vaginata Buse - Gramineae Rottboellia vaginata (Buse) Backer Heteropogon triticeus (R. Br.) Stapf-Gramineae Andropogon triticeus R. Br. horsfieldii (Kunth) Maxim. - Gramineae Holcus lanatus L.- Gramineae Homalomena aromatica (Roxburgh) Schott - Araceae Homalomena cordata Schott - Araceae Homalomenaphilippinensis Engl. Hydrilla verticillata (L.f.) Royle- Hydrocharitaceae Hymenachne amplexicaulis (Rudge) Nees - Gramineae Hymenachne myurus Beauv. Hymenachne aurita (Nees) Backer - Gramineae Panicum auritum Presl Hymenachne interrupta (Willd.) Buse - Gramineae Hyparrhenia filipendula (Höchst.) Stapf-Gramineae Andropogon filipendulus Höchst. paliens (Swartz) Munro - Gramineae Ichnanthus vicinus (F.M. Bailey) Merrill (misapplied to Ichnanthuspallens (Swartz) Munro) Indigofera cordifolia K. Heyne ex Roth - Leguminosae 240 FORAGES

Indigofera glandulosa Wendl. - Leguminosae Psoralea leichhardtii F. v. Muell. Isachne albens Trinius - Gramineae Isachne beneckei Hackel - Gramineae Isachne globosa (Thunberg ex Murray) O. Kuntze - Gramineae Isachne miliacea Roth ex Roemer &Schulte s - Gramineae Isachne myosotis Nées - Gramineae Isachne pangerangensis Zollinger &Moritz i - Gramineae Isachne rigida Nees ex Miquel Ischaemum arundinaceum F.v.Mueller ex Benth. - Gramineae Ischaemum digitatum Brongn. - Gramineae Ischaemum laxum R. Br. - Gramineae Leersia hexandra Swartz - Gramineae Lemna minor L.- Lemnaceae Leptaspis banksii R. Br. - Gramineae Leptaspis cochleata Thwaites - Gramineae Leptochloa filiformis (Lamk) Beauv. - Gramineae Leptochloa polystachya (R. Br.) Benth. (misapplied to Leptochloa filiformis (Lamk) Beauv.) Leptochloa malabarica (L.) Veldk.- Gramineae Diplachnepolystachya (Forsskal) Backer fusca (L.) Beauv. ex Roemer & Schultes Panicum malabaricum (L.) Merrill Leptochloa tectoneticola (Backer) Jansen - Gramineae Diplachne tectoneticola Backer Lepturus repens (Forster) R. Br. - Gramineae Monerma repens Beauv. Lespedeza thunbergii (DC.) Nakai - Leguminosae Lespedeza sieboldi Miquel Ligustrum lucidum Aiton - Oleaceae Lindernia ciliata (Colsm.) Pennell - Scrophulariaceae Bonnaya ciliata Burkil l Bonnaya serrata Burkill Lolium multiflorum Lamk - Gramineae Lolium italicum A. Braun Loliumperenne L.- Gramineae Lolium temulentum L.- Gramineae Lophatherum gracile Brongn. - Gramineae Macaranga hosei King ex Hook.f.- Euphorbiaceae Mapania kurzii Clarke - Cyperaceae Melinis minutiflora Beauv. - Gramineae Microchloa indica (L.f.) Beauv. - Gramineae Microlaena stipoides (Labill.) R. Br. - Gramineae Mnesithea glandulosa (Trinius) Koning &Sosef - Gramineae Coelorrhachis glandulosa Stapf Rottboellia glandulosa Trinius Mnesithea laevis (Retzius) Kunth - Gramineae Mnesithea mollicoma (Hance) Camus - Gramineae Mnesithea pubescens Ridley MINOR FORAGES 241

Mnesithea rottboellioides (R. Br.) Koning &Sosef - Gramineae Rottboellia ophiuroides Benth. Morus alba L.- Moraceae Muehlenbergia huegelii Trinius - Gramineae Myriostachya wightiana (Nees) Hook.f.- Gramineae Najas indica (Willd.) Chamisso - Najadaceae Najas falciculata A. Braun Najas minor Allioni - Najadaceae Nertera depressa Banks &Soland . - Rubiaceae Ophiurus exaltatus (L.) O. Kuntze - Gramineae Oplismenus burmanni (Retzius) Beauv. - Gramineae Hoplismenus burmani P. Beauv. Oplismenus compositus (L.) Beauv. - Gramineae Hoplismenus compositus P. Beauv. Oplismenus undulatifolius (Ard.) Beauv. - Gramineae Hoplismenus undulatifolius P. Beauv. Opuntia cochenillifera Miller- Cactaceae Nopalea coccinellifera (Miller) Salm-Dyck Opuntia monacantha Haw.- Cactaceae Oryza fatua Koenig ex Trinius - Gramineae Oryza granulata Nees &Arnot t ex Steudel - Gramineae Oryza meyeriana (Zollinger &Moritzi ) Bâillon- Gramineae ' Oryza minuta Presl - Gramineae Oryza latifolia Desv. Oryza ridleyi Hook.f. - Gramineae Panicum ambiguum Trinius - Gramineae Urochloa paspaloides Presl Panicum brevifolium L.- Gramineae Panicum caudiglume Hackel - Gramineae Panicum luzonense Presl - Gramineae Panicum tuberculatum Presl (misapplied to Panicum luzonense Presl) Panicum notatum Retzius - Gramineae Panicum miliare Lamk Panicum montanum Roxburgh Panicumpaludosum Roxburgh- Gramineae Panicum sarmentosum Roxburgh - Gramineae Panicum trichoides Swartz - Gramineae Panicum trypheron Schultes - Gramineae Panicum walense Mez- Gramineae Panicum humile Nees ex Steudel Paspalidium flavidum (Retzius) Camus - Gramineae Panicum flavidum Retzius Paspalidium geminatum (Forsskal) Stapf- Gramineae Panicum geminatum Forsskal Paspalidium punctatum (Burm.f.) Camus - Gramineae Panicum mucronatum Roth ex Roemer & Schultes Paspalum orbiculare Forster f. - Gramineae Paspalum vaginatum Swartz - Gramineae Pennisetum alopecuroides (L.) Sprengel - Gramineae 242 FORAGES

Pennisetum compressum R. Br. Perotis indica (L.) Kuntze - Gramineae Phalaris arundinacea L.- Gramineae Pisonia umbellifera (J. &G . Forster) Seemann - Nyctaginaceae Pisonia excelsa Blume Pistia stratiotes L.- Araceae Planchonella longepetiolata (King &Prain ) H.J. Lam - Sapotaceae Planchonella pohlmanniana Burkill - Sapotaceae Pluchea lanceolata (DC.) C.B. Clarke - Compositae Poa annua L.- Gramineae Poa trivialis L.- Gramineae Pollinia clavigera Backer ex Heyne - Gramineae Pollinia dispar Nees ex Steudel - Gramineae Pollinia grata Hackel - Gramineae Pollinia nuda Trinius - Gramineae Pollinia spectabilis Trinius - Gramineae Polypogon monspeliensis (L.) Desv.- Gramineae Polytoca bracteata R. Br. - Gramineae Polytoca macrophylla Benth. - Gramineae Polytrias amaura (Buse) Kuntze - Gramineae Eulalia praemorsa (Nees) Hackel (misapplied to Polytrias amaura (Buse) Kuntze) Pseudochinolaena polystachya (Kunth) Stapf-Gramineae Panicum uncinatum Raddi Pseudopogonatherum contortum (Brongn.) Camus - Gramineae Pollinia contorta Backer Pseudoraphis spinescens (R. Br.) Vickery - Gramineae Chamaeraphis aspera Nees Pseudosorghum zollingeri (Steudel) Camus - Gramineae Andropogon zollingeri Steudel Rhagodia hastata R. Br. - Chenopodiaceae Rhagodiaparabolica R. Br. - Chenopodiaceae Richardsonia brasiliensis Gomez —Rubiacea e Rottboellia exaltata L.f. - Gramineae Rytidospermapilosa (R. Br.) Connor &Edga r - Gramineae Danthonia pilosa R. Br. Saccolepis angusta (Trinius) Stapf-Gramineae Saccolepis indica (L.)A . Chase - Gramineae Hymenachne indica Buse Saccolepis myosuroides (R. Br.) Chase ex Camus - Gramineae Saraca dives Pierre - Leguminosae Schismatoglottis rupestris Zollinger &Moritz i - Araceae Schismatoglottis latifolia Miquel Schizachyrium brevifolium (Swartz) Nees ex Buse - Gramineae Andropogon brevifolius Swartz Schizachyrium sanguineum (Retzius) Alston - Gramineae Andropogon sanguineus Merrill Schizachyrium semiberbe Nees - Gramineae Scirpus erectus Poiret - Cyperaceae MINOR FORAGES 243

Sclerachne punctata R. Br. - Gramineae Scrotochloa urceolata (Roxburgh) Judziewicz - Gramineae Leptaspis urceolata R. Br. Semecarpus kurzii Engl. - Anacardiaceae Setaria barbata (Lamk) Kunth - Gramineae Panicum barbatum Lamk Setaria clivalis (Ridley) Veldk.- Gramineae Panicum chamaeraphoides Hackel Setaria geniculata (Lamk) Beauv. - Gramineae Setaria glauca (L.) Beauv. - Gramineae Panicum lutescens Weigel Setaria palmifolia (Koenig) Stapf-Gramineae Panicum palmifolium Koenig Setaria verticillata (L.) Beauv. - Gramineae Panicum verticillatum L. Setaria viridis (L.) Beauv. - Gramineae Panicum viride L. Sorghum affine Camus - Gramineae Sorghum halepense (L.) Pers.- Gramineae Andropogon halepensis Brot. Andropogon halepensis Brot. var. propinquus (Kunth) Merrill (misapplied to Sorghum halepense (L.) Pers.) • Sorghum nitidum (Vahl) Pers. - Gramineae Andropogon amboinicus Merrill Sorghumplumosum (R. Br.) Beauv. - Gramineae Andropogonplumosus Backer, non Kunth Sorghum sudanense (Piper) Stapf- Gramineae Spergula arvensis L.- Caryophyllaceae diandrus (Retzius) Beauv. -Gramineae Sporobolus humilus Presl - Gramineae Sporobolus tremulus (Willd.) Kunth Sporoboluspulchellus R. Br. - Gramineae Sporobolus virginicus (L.) Kunth - Gramineae Streblochaete longiaristum (Richard) Pilger-Gramineae Koordersiochloa javanica Merrill Symphytum asperum Lepechin - Boraginaceae Symphytum asperrimum J. Donn Thelepogon elegans Roth ex Roemer &Schulte s - Gramineae Themeda arguens (L.) Hackel - Gramineae Themeda gigantea (Cav.) Hackel - Gramineae Themeda villosa Durand &Jackso n (misapplied to Themeda gigantea (Cav.) Hackel) Thuarea involuta (Forster) R. Br. ex Roemer &Schulte s - Gramineae Thuarea sarmentosa Pers. Trema aspera Blume - Ulmaceae rosea Nees - Gramineae Trifolium incarnatum L.- Leguminosae Tripogon exiguus Buse - Gramineae Tripsacum latifolium Hitchc. - Gramineae 244 FORAGES

Vigna trilobata (L.) Verde. - Leguminosae Dolichos trilobatus L. Phaseolus trilobatus (L.) Schreb. Wallichia densiflora (Martius) Martius - Palmae 4 Forages with other primary use

In this tentative list the commodity group ofprimar y use is given in parentheses and synonyms are given in the following, indented lines. The list contains spe­ cies which have been noted in the major historical handbooks on economically useful plants of South-East Asia as having a secundary use as forage. It is not a list of all species that are used as forage.

Abelmoschus esculentus (L.) Moench (vegetables) Hibiscus esculentus L. Acacia farnesiana (L.)Willd . (essentail-oil plants) Acacia mearnsii De Wild, (dye and tannin-producing plants) Acacia decurrens auct. non Willd. Acacia nilotica (L.)Willd . ex Del. (dye and tannin-producing plants) Acacia arabica (Lamk) Willd. / Acacia tomentosa (Roxburgh) Willd. (auxiliary plants in agriculture and forestry) Acanthus ilicifolius L. (medicinal and poisonous plants) Acanthus volubilis Wallich Adenostemma lavenia (L.) Kuntze (medicinal and poisonous plants) Aganope heptaphylla (L.) Polhill (vegetables) Derris heptaphylla Merrill Ailanthus altissima (Miller) Swingle (medicinal and poisonous plants) Ailanthus glandulosa Desf. Ailanthus vilmoriniana Dode Albiziaprocera (Roxburgh) Benth. (timber trees) Alphitonia excelsa Reisseck ex Endlicher (timber trees) Amaranthus spinosus L. (medicinal and poisonous plants) Amaranthus viridis L. (vegetables) Amaranthus gracilis Desf. Amorphophalluspaeoniifolius (Dennst.) Nicolson (plants mainly producing car­ bohydrates) Amorphophallus campanulatus Blume Amorphophallus variabilis Blume (plants mainly producing carbohydrates) Andropogon nardus L. (essentail-oil plants) Cymbopogon citratus Stapf (misapplied to Andropogon nardus L.) Andropogon nardus L. var. tortilis (Presl) Merrill (misapplied to Andropo­ gon nardus L.) Annona muricata L. (edible fruits and nuts) Annona squamosa L. (edible fruits and nuts) Anthocephalus chinensis (Lamk) A.Rich , ex Walp. (timber trees) Anthocephalus cadamba (Roxburgh) Miquel 246 FORAGES

Anthocephalus indicus Richard Neolamarckia cadamba (Roxburgh) Bosser Antidesma bunius (L.) Sprengel (edible fruits and nuts) Antidesma dallachyanum Bâillon Antidesma rumphii Tulasne Apium graveolens L. (vegetables) Arachis hypogaea L. (pulses) Areca triandra Roxburgh (stimulants) Areca borneensis Beccari Artabotrys suaveolens Blume (ornamental plants) Artocarpus altilis (Parkinson) Fosberg (edible fruits and nuts) Artocarpus communis J.R. &G . Forster Artocarpus camansi Blanco Artocarpus integer (Thunberg) Merrill (edible fruits and nuts) Artocarpus champeden (Loureiro) Stokes Azadirachta indica Adr. Jussieu (medicinal and poisonous plants) Melia indica Brandis Azollapinnata R. Br. (lower plants) Bambusa bambos Backer (bamboos) Beta vulgaris L. (vegetables) Bidens pilosa L. var. radiata Schultz-Bip. (medicinal and poisonous plants) Bidenspilosus L. var. albus Schultz-Bip. Boerhavia diffusa L. (medicinal and poisonous plants) Bombax ceiba L. (timber trees) Gossampinus heptaphylla Bakh. Bombax malabaricum DC. Bombax valetonii Hochr. (timber trees) Bombax insigne Wallich Bombax larutense Ridley Borassus flabellifer L. (plants mainly producing carbohydrates) Brassica juncea (L.) Czernova (vegetables) Brassica rugosa Prain Brassica integrifolia (West) Rupr. Bridelia retusa (L.)Sprenge l (timber trees) Brugmansia Candida (Pers.) Safford (ornamental plants) Pseudodatura arborea van Zyp Brugmansia suaveolens Humb. &Bonpl . (ornamental plants) Pseudodatura suaveolens van Zyp Datura suaveolens Humb. & Bonpl. Bruguiera gymnorhiza (L.) Savigny (dye and tannin-producing plants) Bruguiera conjugata Merrill Butea monosperma (Lamk) Taubert (dye and tannin-producing plants) Butea superba Roxburgh (medicinal and poisonous plants) Caesalpinia crista L. (medicinal and poisonous plants) Caesalpinia nuga (L.)W.T . Aiton Caesalpinia digyna Rottler (dye and tannin-producing plants) Cajanus cajan (L.) Millsp. (pulses) Cytisus cajan L. Cajanus indicus Spreng. FORAGES WITH OTHER PRIMARY USE 247

Canarium sylvestre Gaertner (plants producing exudates) Canavalia maritima (Aublet) du Petit-Thouars (auxiliary plants in agriculture and forestry) Canavalia rosea (Swartz) DC. Canna edulis Ker (plants mainly producing carbohydrates) Cannabis sativa L. (medicinal and poisonous plants) Careya arborea Roxburgh (timber trees) Carthamus tinctorius L. (vegetable oils and fats) Carum carvi L. (spices and condiments) Cassia mimosoides L. (auxiliary plants in agriculture and forestry) Cassia siamea Lamk (timber trees) Ceibapentandra (L.) Gaertner (fibre plants) Cicer arietinum L. (pulses) Citrullus lanatus (Thunberg) Matsum. &Naka i ssp. vulgaris (Schrader) Fursa (vegetables) Citrullus vulgaris Schrader ex Ecklon & Zeyher Citrus limon (L.)Burm.f . (edible fruits and nuts) Claoxylon indicum (Reinw. ex Blume) Hassk. (spices and condiments) Claoxylon polot Merrill Cleome gynandra L. (vegetables) Gynandropsis gynandra (L.) Merrill Cocos nucifera L. (vegetable oils and fats) / Coix lachryma-jobi L. (cereals) Colocasia esculenta (L.)Schot t (plants mainly producing carbohydrates) Combretum quadrangulare Kurz (medicinal and poisonous plants) Commelina benghalensis L. (vegetables) Corchorus olitorius L. (fibre plants) Cordia dichotoma Forster f. (medicinal and poisonous plants) Cordia obliqua Willd. Coriandrum sativum L. (spices and condiments) Corypha utan Lamk (fibre plants) Corypha elata Roxburgh Crotalaria alata Buch.-Ham. &Roxburg h ex D.Do n (auxiliary plants in agricul­ ture and forestry) Crotalaria anagyroides Kunth (auxiliary plants in agriculture and forestry) Crotalaria ferruginea Graham ex Benth. (auxiliary plants in agriculture and forestry) Crotalaria zanzibarica Benth. (auxiliary plants in agriculture and forestry) Crotalaria usaramoensis Baker f. Cucurbita pepo L. (vegetables) Cyamopsis tetragonoloba (L.) Taubert (auxiliary plants in agriculture and forestry) Cyamopsis psoralioides DC. Cyanotis cristata (L.) D.Do n (vegetables) Cyathea amboinensis (v.A.v.R.) Merrill (lower plants) Alsophila amboinensis v.A.v.R. Cyperus cyperoides (L.) 0. Kuntze (medicinal and poisonous plants) Mariscus sieberianus Nees ex Clarke Cyperus elatus L. (fibre plants) 248 FORAGES

Cyperuspilosus Vahl (auxiliary plants in agriculture and forestry) Cyperus rotundus L. (plants mainly producing carbohydrates) Cyrtococcum accrescens (Trinius) Stapf (auxiliary plants in agriculture and forestry) Cyrtococcum oxyphyllum (Steudel) Stapf (auxiliary plants in agriculture and forestry) Panicum pilipes Nees &Arnot t ex Buse Cyrtococcum trigonum (Retzius) Camus (auxiliary plants in agriculture and forestry) Panicum trigonum Retzius Desmodium tortuosum (Swartz)DC . (auxiliary plants in agriculture and forestry) Desmodium purpureum (Miller) Fawc. & Rendle Digitaria didactyla Willd. (ornamental plants) Dimeria ornithopoda Trinius (ornamental plants) Echinochloa frumentacea (Roxburgh) Link (cereals) Eichhornia crassipes (Martius) Solms (auxiliary plants in agriculture and forestry) Pontederia crassipes Mart. Eichhornia speciosa Kunth Elaeis guineensis N.J. Jacquin (vegetable oils and fats) Elephantopus scaber L. (medicinal and poisonous plants) Eleusine indica (L.)Gaertne r f. coracana (L.) Hook.f. ex Backer (cereals) Eleusine coracana Gaertner Enhalus acoroides (L.f.) L.C. Richard ex Steudel (fibre plants) Eragrostis te/(Zuccagni) Trotter (cereals) Erianthus arundinaceus (Retzius) Jeswiet (fibre plants) Saccharum arundinaceum Retzius Erythrina subumbrans (Hassk.) Merrill (auxiliary plants in agriculture and forestry) Erythrina lithosperma Miquel Fagopyrum esculentum Moench (cereals) Ficus altissima Blume (fibre plants) Ficus benghalensis L. (timber trees) Ficus indica L. Ficus grossularioides Burm.f. (vegetables) Ficus alba Reinw. ex Blume Ficus hispida L. (medicinal and poisonous plants) Ficus padana Burm.f. (plants producing exudates) Ficus toxicaria L. Ficus religiosa L. (medicinal and poisonous plants) Ficus superba (Miquel) Miquel (vegetables) Ficus variegata Blume (plants producing exudates) Fimbristylis dichotoma (L.)Vah l (auxiliary plants in agriculture and forestry) Fimbristylis diphylla Vahl Firmiana colorata (Roxburgh) R. Br. (fibre plants) Erythropsis colorata (Roxburgh) Burkill Galinsogaparviflora Cav. (vegetables) Garuga floribunda Decaisne (timber trees) Garuga abilo Merrill FORAGES WITH OTHER PRIMARY USE 249

Glycine max (L.) Merrill (pulses) Gossypium arboreum L. var. obtusifolium (Roxburgh) Roberty (fibre plants) Gossypium nangking Meyen Gossypium obtusifolium Roxburgh Gossypium hirsutum L. (fibre plants) Guazuma ulmifolia Lamk var. tomentosa K. Schumann (fibre plants) Guizotia abyssinica (L.f.) Cass, (vegetable oils and fats) Helianthus annuus L. (vegetable oils and fats) Hevea brasiliensis (Willd. ex A. Jussieu) Muell. Arg. (plants producing exudates) Hibiscus cannabinus L. (fibre plants) Hibiscus sabdariffa L. (vegetables) Hibiscus tiliaceus L. (fibre plants) Homalomena alba Hassk. (medicinal and poisonous plants) Homalomena javanica Alderw. Hydnocarpus castanea Hook.f. &Thomso n (timber trees) Hydnocarpus heterophylla Blume (vegetable oils and fats) Taraktogenos heterophylla Slooten Hydnocarpus kurzii (King) Warb, (vegetable oils and fats) Taraktogenos kurzii King Hydnocarpuspolypetala (Slooten) Sleumer (vegetable oils and fats) Taraktogenos polypetala Slooten / Hygrorhiza aristata (Retzius) Nees ex Wight &Arnot t (cereals) Hymenodictyon orixense (Roxburgh) Mabberly (timber trees) Hymenodictyon excelsum Wallich Hymenodictyon timoriense Klotsch ex Walp. Imperata conferta (Presl) Ohwi (fibre plants) Imperata contracta Hitchc. Imperata exaltata Brongn. (misapplied to Imperata conferta (Presl) Ohwi) Indigofera spicata Forssk. var. spicata (auxiliary plants in agriculture and forestry) Indigofera hendecaphy HaJacq . Inocarpus fagiferus (Parkinson) Fosberg (ornamental plants) Inocarpus edulis J.R. &G . Forster Ipomoea aquatica Forssk. (vegetables) Ipomoea reptans Poiret Ipomoea batatas (L.)Lam k (plants mainly producing carbohydrates) Ipomoeapes-caprae (L.) Sweet (medicinal and poisonous plants) Justicia procumbens L. (medicinal and poisonous plants) Rostellaria procumbens (L.) Nees Kydia calycina Roxburgh (fibre plants) Lablabpurpureus (L.) Sweet (pulses) Dolichos lablab L. Lablab niger Medikus Lactuca indica L. (vegetables) Lagerstroemia indica L. (ornamental plants) Lannea coromandelica (Houtt.) Merrill (ornamental plants) Lannea grandis (Dennst.) Engl. Lantana camara L. (medicinal and poisonous plants) 250 FORAGES

Lantana aculeata L. Leucas lavandulifolia J.E. Smith (medicinal and poisonous plants) Leucas linifolia (Roth) Sprengel Limnocharis flava (L.) Buchenau (vegetables) Litchi chinensis Sonn, (edible fruits and nuts) Nephelium litchi Cambess. Litchi philippinensis Radlk. Euphoria didyma Blanco Litsea garciae Vidal (edible fruits and nuts) Litsea sebifera (Blume) Blume Litsea glutinosa (Loureiro) C.B.Robinso n (timber trees) Litsea chinensis Lamk Lycopersicon esculentum Miller (vegetables) Solanum lycopersicum L. Macaranga gigantea (H.G. Reichenbach &Zollinger ) Muell. Arg. (timber trees) Macaranga incisa Gage Madura cochinchinensis (Loureiro) Corner (dye and tannin-producing plants) Cudrania javanensis Trécul Cudrania pubescens Trécul Macrotyloma uniflorum (Lamk) Verde, (pulses) Dolichos uniflorus Lamk Dolichos biflorus auct. mult., non L. Mangifera indica L. (edible fruits and nuts) Manihot esculenta Crantz (plants mainly producing carbohydrates) Manihot utilissima Pohl Melastoma malabathricum L. (medicinal and poisonous plants) Melastomapolyanthum Blume (medicinal and poisonous plants) Melastoma malabathricum L.f .polyanthum L. Melastoma sanguineum Sims (medicinal and poisonous plants) Melastoma decemfidum Roxburgh Merremia hederacea (Burm.f.) H. Hallier (medicinal and poisonous plants) Metroxylon sagu Rottboel (plants mainly producing carbohydrates) Metroxylon rumphii (Willdenow) Martius Metroxylon squarrosum Beccari Mimosa invisa Martius ex Colla (auxiliary plants in agriculture and forestry) Miscanthus japonicus Andersson (fibre plants) Mollugopentaphylla L. (medicinal and poisonous plants) Monochoria hastata (L.)Solm s (vegetables) Monochoria vaginalis (Burm.f.) Presl (vegetables) Moringa oleifera Lamk (spices and condiments) Mucunapruriens (L.) DC. cv. group Utilis (auxiliary plants in agriculture and forestry) Mucuna aterrima (Piper &Tracy ) Merrill Mucuna cochinchinensis (Loureiro) A. Chev. Mucuna deeringiana (Bort.) Merrill Murdannia nudiflora (L.) Brenan (vegetables) Aneilema nudiflorum R. Br. Musa xparadisiaca L. (Musa AAB group) (edible fruits and nuts) Musa paradisiaca L. var. sapientum Kuntze FORAGES WITH OTHER PRIMARY USE 251

Musa sapientum L. var.paradisiaca Baker Neyraudia reynaudiana (Kunth) Keng ex Hitchc. (ornamental plants) Neyraudia madagascariensis Hook.f. var. zollingeri Hook.f. Triraphis madagascariensis Hook.f. ex Prain Nymphoides spp. (vegetables) Limnanthemum spp. Oldenlandia corymbosa L. (medicinal and poisonous plants) Opuntia elatior Miller (ornamental plants) Opuntia nigricans Haw. Oryza sativa L. (cereals) Pandanus affinis Kurz (medicinal and poisonous plants) Pandanus aurantiacus Ridley Pandanus helicopus Kurz (fibre plants) Pandanus johorensis Martelli Panicum miliaceum L. (cereals) Paraserianthes falcataria (L.) Nielsen (auxiliary plants in agriculture and forestry) Albizia falcata sensu Backer Parkinsonia aculeata L. (ornamental plants) Passiflora foetida L. (auxiliary plants in agriculture and forestry) Peltophorum pterocarpum (DC.) Backer ex Heyne (dye and tannin-producing plants) Pennisetum americanum (L.) K. Schum. ex Leeke (cereals) / Pennisetum glaucum R. Br. Pennisetum macrostachyum (Brongn.) Trinius (ornamental plants) Pennisetum villosum R. Br. ex Fresen. (ornamental plants) Perilla frutescens (L.) Britton (vegetable oils and fats) Perilla ocimoides L. Phaseolus coccineus L. (pulses) Phaseolus multiflorus Lamk Phaseolus lunatus L. (pulses) Phragmites australis (Cav.) Steudel (fibre plants) Phragmites vulgaris (Lamk) Crepin Phragmites karka (Retzius) Trinius ex Steudel (fibre plants) Phragmites communis Trinius (misappplied to Phragmites karka (Retzius) Trinius ex Steudel) Phyllodiumpulchellum (L.) Desv. (medicinal and poisonous plants) Desmodium pulchellum (L.) Benth. Phytelephas macrocarpa Ruiz &Pavo n (ornamental plants) Pimpinella anisum L. (medicinal and poisonous plants) Plantago major L. (medicinal and poisonous plants) Pogonatherum crinitum (Thunberg ex Murray) Trinius ex Kunth (auxiliary plants in agriculture and forestry) Pogonatherum paniceum (Lamk) Hackel Pongamia pinnata (L.) Pierre (medicinal and poisonous plants) Portulaca oleracea L. (vegetables) Premna serratifolia L. (timber trees) Premna integrifolia L. Premna foetida Reinw. ex Blume 252 FORAGES

Prosopis juliflora (Swartz) DC. (auxiliary plants in agriculture and forestry) Prosopis spicigera L. (auxiliary plants in agriculture and forestry) Pseuderanthemum racemosum (Roxburgh) Radlk. (vegetables) Pteridium aquilinum Kuhn (lower plants) Pueraria lobata (Willd.) Ohwi (plants mainly producing carbohydrates) Pueraria triloba (Loureiro) Backer Pueraria hirsuta (Thunberg) Schneider Putranjiva roxburghii Wallich (medicinal and poisonous plants) Quercus roburL. (timber trees) Ricinus communis L. (vegetable oils and fats) Rivina humilis L. (medicinal and poisonous plants) Saccharum officinarum L. (plants mainly producing carbohydrates) Saccharum ravennae (L.) Murray (fibre plants) Sagittaria sagittifolia L. (plants mainly producing carbohydrates) Sagittaria sagittaefolia L. Samanea saman (Jacq.) Merrill (ornamental plants) Enterolobium saman (Jacq.) Prain Santalum acuminatum (R. Br.) A. DC. (essential-oil plants) Eucarya acuminata (R. Br.) Sprague & Summerhayes Sapium discolor (Champ, ex Benth.) Muell.Arg. (timber trees) Schismatoglottis calyptrata (Roxburgh) Zollinger &Moritz i (vegetables) Schleichera oleosa (Loureiro) Oken (vegetable oils and fats) Scoparia dulcis L. (medicinal and poisonous plants) Sechium edule (Jacq.) Swartz (vegetables) Sesamum orientale L. (vegetable oils and fats) Sesamum indicum L. Sesamum radiatum Schumacher (vegetable oils and fats) Sesbania bispinosa (Jacq.) W.F. Wight (fibre plants) Sesbania aculeata (Willd.) Pers. Sesbania javanica Miquel (ornamental plants) Sesbania roxburghii Merrill Sesuviumportulacastrum (L.)L . (vegetables) Setaria italica (L.) Beauv. (cereals) Panicum viride L. var. italica L. Shorea roxburghii G. Don (timber trees) Shorea cochinchinensis Pierre Shorea siamensis Miquel (timber trees) Pentacme siamensis (Miquel) Kurz Sida cordifolia L. (medicinal and poisonous plants) Sida rhombifolia L. (medicinal and poisonous plants) Smithia sensitiva Aiton (medicinal and poisonous plants) Sonchus oleraceus L. (vegetables) Sonchus wightianus DC. (vegetables) Sonchus arvensis L. (misapplied to Sonchus wightianus DC.) Sophora tomentosa L. (medicinal and poisonous plants) Sorghum bicolor (L.) Moench (cereals) Andropogon sorghum Brot. Sorghum vulgare Pers. Sparganophorus sparganophora (L.) C. Jeffrey (auxiliary plants in agriculture FORAGES WITH OTHER PRIMARY USE 253

and forestry) Sparganophorus vaillantii Gaertner Sphaeranthus africanus L. (medicinal and poisonous plants) Sporobolus indicus (L.)R . Br. (fibre plants) Sporobolus berteroanus Hitchc. & Chase Stachytarphetajamaicensis (L.)Vah l (medicinal and poisonous plants) Streblus asper (Retzius) Loureiro (medicinal and poisonous plants) Syagrus oleracea (Martius) Beccari (timber trees) Tamarindus indica L. (edible fruits and nuts) Tephrosiapurpurea (L.)Pers . (auxiliary plants in agriculture and forestry) Theobroma cacao L. (vegetable oils and fats) Theobroma pentagona Bernoulli Tragus racemosus Aiton exHook.f . (auxiliary plants in agriculture and forestry) Trema orientalis (L.) Blume (auxiliary plants in agriculture and forestry) Tridax procumbens L. (auxiliary plants in agriculture and forestry) Trigonella foenum-graecumh. (spices and condiments) Triumfetta bartramia L. (fibre plants) Triumfetta rhomboidea Jacq. Turnera ulmifolia L. (medicinal and poisonous plants) Typhonium trilobatum (L.) Schott (plants mainly producing carbohydrates) Urena lobata L. (fibre plants) Vallisneria spiralis L. (vegetables) Vernonia cinerea (L) Less, (medicinal and poisonous plants) ' Vetiveria zizanoides (L.) Nash (essentail-oil plants) Andropogon zizanioides Urban Vigna aconitifolia (Jacq.) Maréchal (pulses) Phaseolus aconitifolius Jacq. Vigna radiata (L.)Wilcze k (pulses) Phaseolus radiatus L. Phaseolus aureus Roxburgh Vigna umbellata (Thunberg) Ohwi &Ohash i (pulses) Phaseolus calcaratus Roxburgh Vigna unguiculata (L.)Walp . (pulses) Dolichos unguiculatus L. Dolichos sinensis L. Vigna sinensis (L.) Hassk. Wollastonia biflora (L.)DC . (medicinal and poisonous plants) Wedelia biflora DC. Wollastonia moluccana (Blume) DC. (medicinal and poisonous plants) Wedelia moluccana Boerl. Zea mays L. (cereals) Ziziphus mauritiana Lamk (edible fruits and nuts) Ziziphus jujuba (L.)Gaertne r (non Miller; often cited as Lamk) Ziziphus xylopyrus (Retzius) Willd. (edible fruits and nuts) Literature

Ahmed Tajuddin, Z., 1991.Proceeding s of the international livestock-tree work­ shop, 5-9th December 1988,Serdang , Malaysia. MARDI/FAO, Rome. 133pp . Blair, G.J., Ivory, D.A.& Evans ,T.R . (Editors), 1986.Forage s in Southeast Asian and South Pacific agriculture. ACIAR Proceedings No 12.ACIAR , Canberra. 202pp . Bogdan, A.V., 1977.Tropica l pasture and fodder plants. Longman, London. 474 pp. Burt, R.L., Rotar, P.P., Walker, J.L. & Silvey, M.W., 1983. The role of Centro- sema, Desmodium and Stylosanthes in improving tropical pastures. Westview Tropical Agriculture Series No 6.Westvie w Press, Boulder, Colorado, United States. 292pp . Cameron, D.G., 1987.Bette r pastures for the tropics. 2nd ed. Yates Seeds, Toow- oomba, Australia. 75pp . Chin, F.Y., 1989. Justification, government policy, measures and strategy for the development of grazing reserves in Peninsular Malaysia. In: Halim, R.A. (Editor):Proceeding s workshop on grasslands and forage production in South East Asia, 27 February - 3 March 1989, Serdang, Malaysia. FAO, Rome. pp. 77-82. Cooper, J.P., 1970. Potential production and energy conversion in temperate and tropical grass. Herbage Abstracts 40:1-15. Crowder, L.V. &Chhedda , H.R., 1982.Tropica l grassland husbandry. Longman, London.562 pp. Devendra, C. (Editor), 1990. Shrubs and fodders for farm animals. Proceedings ofa workshop in Denpasar, Indonesia, 24-29 July 1989.IDRC , Ottawa. 349pp . Falvey, J.L. &Hengmichai , P., 1979.Invasio n ofImperat a cylindrica (L.) Beauv. by Eupatorium species. Journal of Range Management 32: 340-344. Food and Agriculture Organization, 1989. Production yearbook. Vol. 43. FAO, Rome. Food and Fertilizer Technology Center for the Asian and Pacific Region, 1984. Asian pastures: recent developments in pasture research and development in southeast Asia. FFTC, Taipei, Taiwan. 220pp . Ghani, A.N.A. &Wang , K.A., 1990.Developmen t and evaluation of agroforestry systems for fodder production. In: Devendra, C. (Editor): Shrubs and fodders for farm animals. IDRC, Ottawa, pp. 319-330. Halim, R.A. (Editor), 1989.Grassland s and forage production in south-east Asia. Proceedings regional working group on grazing and feed resources of south­ east Asia. FAO, Rome. 216pp . Hopkinson J.M., 1986.See d production in tropical species. In: Blair, G.J., Ivory, D.A. & Evans, T.R. (Editors): Forages in Southeast Asia and South Pacific Agriculture. ACIAR Proceedings No 12.ACIAR , Canberra, pp. 188-192. LITERATURE 255

Humphreys, L.R., 1980.A guid e tobette r pastures for the tropics and sub-tropics. 4th ed. Wright Stephenson &Co. , Ermington, Sydney. 96pp . Humphreys, L.R., 1981. Environmental adaptation of tropical pasture plants. MacMillan, London. 261pp . Humphreys, L.R., 1991. Tropical pasture utilisation. Cambridge University Press, Cambridge. 206pp . Johnson, R.W. & Tothill, J.C., 1985. Definition and broad geographic outline of savanna lands. In: Tothill, J.C. &Mott , J.J. (Editors): Ecology and manage­ ment of the world's savannas. Australian Academy of Science, Canberra & Commonwealth Agricultural Bureaux, Farnham Royal, England, pp. 1-13. Jones, R.M., Tothill, J.C. &Jones , R.J. , 1984.Pasture s and pasture management in the tropics and subtropics. Occasional Publication No 1. Tropical Grass­ land Society of Australia, Brisbane. 74pp . Kerati-Kasikorn, P., 1984.Soi l nutrient deficiencies affecting pasture production in northeast and northern Thailand. In: Asian pastures: recent advances in pasture research and development in southeast Asia. Food and Fertilizer Tech­ nology Center for the Asian and Pacific Region, Taipei, Taiwan, pp. 159-172. Kerridge, P.C., Edwards, D.G. & Sale, P.W.G., 1986. Soil fertility constraints - amelioration and plant adaptation. In: Blair, G.J. , Ivory, D.A. &Evans , T.R. (Editors): Forages in Southeast Asia and South Pacific agriculture. ACIAR Proceedings No 12.ACIAR , Canberra, pp. 179-187. Little, D.A., Kompiang, S. &Petheram , R.J., 1989.Minera l composition of Indo­ nesian ruminant forages. Tropical Agriculture (Trinidad) 66:33-37 . Lopez, F.S.S. &Vergara , B.S.,1988 .Herbag e production from a rice crop. Philip­ pine Journal of Crop Science 13:15-19. Manidool, C, 1990. Pasture development and seed production in Thailand. ACIAR Forage Newsletter, Canberra. No 13.pp . 11-12. Manidool, C. & Chantkam, S., 1986. Feed resources for small holder dairy pro­ duction. ASPAC Food and Fertilizer Technology Center. Extension Bulletin No 236.16 pp. McWilliam, J.R., 1978. Response of pasture plants to temperature. In: Wilson, J.R. (Editor): Plant relations in pastures. CSIRO, Melbourne, pp. 17-34. Mehra, K.L. & Fachrurozi, Z., 1985.Indonesia n economic plant resources: for­ age crops. LBN 31, Seri Sumber Daya Alam 117. Lembaga Biologi Nasional -LIPI, Bogor, Indonesia. 61pp . Moog, F.A., 1986. Forages in integrated food cropping systems. In: Blair, G.J., Ivory, D.A. & Evans, T.R. (Editors): Forages in Southeast Asian and South Pacific agriculture. ACIAR Proceedings No 12. ACIAR, Canberra, pp. 152-156. National Academy of Sciences, 1979.Tropica l legumes: resources for the future. Washington, D.C., United States. 330pp . Niewolt, S., 1982. Climate and agricultural planning in Peninsular Malaysia. MARDI, Serdang, Malaysia. 139pp . Nitis, I.M., Lana, K., Sukanten, W. Suarna, M. & Putra, S., 1990. The concept and development of the three strata forage system. In: Devendra, C. (Editor): Shrubs and fodders for farm animals. IDRC, Ottawa, pp. 92-102. Nurjaya, O., Nitis, I.M. &Britten , E.J., 1983.Evaluatio n of annual and perenni­ al temperate pasture legumes at medium elevation in the tropics at Bali, Indo­ nesia: a preliminary investigation. Tropical Grasslands 17:122-128. 256 FORAGES

Oram, R.N., 1990.Registe r ofAustralia n herbage plant cultivars. 3rd ed. CSIRO, Melbourne. 304pp . Perkins, J., Petheram, R.J., Rachman, R. & Semali, A., 1986. Introduction and management prospects for forages in Southeast Asia and the south Pacific. In:Blair , G.J. , Ivory, D.A. &Evans ,T.R . (Editors): Forages in Southeast Asian and South Pacific agriculture. ACIAR Proceedings No 12. ACIAR, Canberra. pp. 15-23. Raghavan, C.V., 1990. Availability and use of shrubs and tree fodders in India. In:Devendra , C.(Editor) :Shrub s and fodders for farm animals. IDRC, Ottawa, pp. 196-210. Rangkuti, M., Siregar, M.E. &Roesyat , A., 1990. Availability and use of shrubs and tree fodders in Indonesia. In: Devendra, C. (Editor): Shrubs and fodders for farm animals. IDRC, Ottawa, pp. 266-278. Ranjhan, S.K., 1986.Source s offee d for ruminant production in Southeast Asia. In: Blair G.J., Ivory, D.A. &Evans , T.R. (Editors): Forages in Southeast Asian and South Pacific agriculture. ACIAR Proceedings No 12.ACIAR , Canberra, pp. 24-28. Remenyi, J.V. & McWilliam, J.R., 1986.Ruminan t production trends in South­ eastern Asia and the South Pacific, and the need for forages. In: Blair, G.J, Ivory, D.A. & Evans, T.R. (Editors): Forages in Southeast Asian and South Pacific agriculture. ACIAR Proceedings No 12.ACIAR , Canberra, pp. 1-6. Reynolds, S.G., 1988. Some factors of importance in the integration of pastures and cattle with coconuts. Journal of Biogeography 15: 31-39. Schultze-Kraft, R. & Clements, R.J. (Editors), 1990. Centrosema: biology, agro­ nomy and utilization. CIAT, Cali, Columbia. 667pp . Shaw, N.H. & Bryan, W.W., 1976. Tropical pasture research. Principles and methods. Commonwealth Agricultural Bureaux, Farnham Royal, Bucks, England. 454pp . Shelton, H.M. & Stür, W.W., 1991. Forages for plantation crops. ACIAR Pro­ ceedings No 32.ACIAR , Canberra. 162pp . Skerman, P.C., Cameron, D.G. &Riveros , F., 1988.Tropica l forage legumes. 2nd ed. FAO Plant Production and Protection Series No 2.FAO , Rome. 692pp . Skerman, P.J. &Riveros , F., 1990.Tropica l forage grasses. FAO, Rome. 832pp . Soedomo,R. , Ginting, P.G. &Blair , G.J. , 1986.Non-leguminou s trees and shrubs as forage for ruminants. In: Blair, G.J., Ivory, D.A. & Evans, T.R. (Editors): Forages in Southeast Asia and South Pacific agriculture. ACIAR Proceed­ ings No 12.ACIAR , Canberra, pp. 51-54. Soerjani, M., 1970. Alang-alang (Imperata cylindrica (L.) Beauv.) Pattern of growth as related to its problem of control. BIOTROP Bulletin 1.8 7pp . Stace, H.M. & Edye, L.A. (Editors), 1984. The biology and agronomy of Stylo- santhes. Academic Press, Sydney. 636pp . Staples,J.B. , Colman, P.A. &Crowther , D.E., 1983.Productivit y and persistence of horse gram (Macrotyloma uniflorum) under dry season grazing. Tropical Grasslands 17:152-155. Tajuddin, Z.A., 1991.Proceeding s of the international livestock-tree cropping workshop, 5-9 December 1988,Serdang , Malaysia. MARDI/FAO, Kuala Lum­ pur. 133pp . Thahar, A. & Petheram, R.J., 1983. Ruminant feeding systems in West Java, Indonesia. Agricultural Systems 10:87-97 . LITERATURE 257

Tinnimit, P. 1985. Rubber seed meal and oilpalm meal for livestock feeding in Thailand. In: Wanapat, M. & Devendra, C. (Editors): Relevance of cropping residues as animal feeds in developing countries. Proceedings of an interna­ tional workshop held in Khon Kaen, Thailand. November 29 - December 2, 1984. Funny Press, Bangkok, pp. 368-381. Topark-Ngarm, A., 1990. Shrubs and fodders in farming systems in Asia. In: Devendra, C. (Editor): Shrubs and fodders for farm animals. IDRC, Ottawa, pp. 12-21. Tothill, J.C. &Hacker , J.B., 1983.Th e grasses of southern Queensland. Univer­ sity of Queensland Press, St. Lucia, Brisbane. 475pp . Troll, C, 1966. Seasonal climates of the earth. The seasonal course of natural phenomena in the different climatic zones of the earth. In: Rodenwaldt, E. &Justaz , H.J. (Editors): World maps of climatology. Springer Verlag, Berlin, pp. 19-28. Trung, L.T., 1990. Availability and use of shrubs and tree fodders in the Philip­ pines. In: Devendra, C. (Editor): Shrubs and fodders for farm animals. IDRC, Ottawa, pp. 279-294. United Nations Educational, Sciencific and Cultural Organization, 1979.Trop ­ ical grazing land ecosystems. A state-of-the-art report. UNESCO, Paris. 655 pp. Verdcourt, B., 1979.A manual of New Guinea legumes. Botany Bulletin No 11. Office of Forests, Division of Botany, Lae. 645pp . / Wanapat, M. & Devendra, C. (Editors), 1985. Relevance of cropping residues as animal feeds in developing countries. Proceedings of an international workshop held in Khon Kaen, Thailand. November 29 - December 2, 1984. Funny Press, Bangkok. 459pp . Whiteman, P.C., 1980. Tropical pasture science. Oxford University Press, Oxford. 392pp . Whyte, R.O., 1974.Tropica l grazing lands:communitie s and constituent species. Dr. W. Junk NV, The Hague. 222pp . Wiersema, J.H., Kirkbride, J.H. &Gunn, CR., 1990.Legum e (Fabaceae) nomen­ clature in the USDA germplasm system. United States Department of Agri­ culture, Technical Bulletin No 1757.57 2pp . Wilaipon, B.& Pongskul , V., 1984.Pastur e establishment in the farming systems ofnorther n Thailand. In: Asian pastures: recent advances in pasture research and development in southeast Asia. Food and Fertilizer Technology Center for the Asian and Pacific region, Taipei, Taiwan, pp. 65-85. Wong, C.C., Mohd. Sharudin, M.A. &Rahim , H., 1985.Shad e tolerance of some tropical forages for integration with plantations. 2. Legumes. MARDI Research Bulletin 13: 249-269. Wong, C.C.,Rahim , H. &Mohd . Sharudin, M.A. , 1985.Shad e tolerance potential of some tropical forages for integration with plantations. 1.Grasses . MARDI Research Bulletin 13: 225-247. Acknowledgments

Our thanks are due to - the Executive Board of the Wageningen Agricultural University for support­ ing the Prosea Programme; - the Netherlands Ministry of Education &Science , for financial support; - the Netherlands Ministry of Agriculture, Nature Management & Fisheries, for financial support; - the Netherlands Ministry ofForeig n Affairs, Directorate-General for Interna­ tional Cooperation, for financial support of the South-East Asian Network of the Prosea Programme; - the coordinating institutions of the Prosea Programme for providing facili­ ties for the Prosea staff; - the Departments of Tropical Crop Science and Plant Taxonomy of Wage­ ningen Agricultural University, for providing facilities for the Prosea Publi­ cation Office; - the Centre for Research &Developmen t in Biology, Bogor, Indonesia, for pro­ viding facilities for the Prosea Network Office; - the Department of Field Crops and Grassland Science, Wageningen Agricul­ tural University, for awarding Mr R.M. Jones a senior research fellowship to allow him to work on the editing ofthi s volume; - the Centre for Agricultural Publishing & Documentation (Pudoc), Wage­ ningen, the Netherlands, in particular Drs J.M. Schippers, Director, for his support, and his staff for cooperation and for documentation facilities; - Dr R.J. Clements, Chief ofth e CSIRODivisio n ofTropica l Crops and Pastures for allowing Mr R.M. Jones to work on this volume for much of his time in 1991; - Mr R.J. Williams of the CSIRO Division of Tropical Crops and Pastures for checking distribution records of many ofth e species; - Dr J.B. Hacker of the CSIRO Division of Tropical Crops and Pastures for checking the illustrations; - Ms S.M.M. Kersten of the Department of Field Crops and Grassland Science, Wageningen Agricultural University, for general editorial assistance; - the Prosea Country Offices in South-East Asia, for providing vernacular names and access to literature which is not readily available elsewhere, through records entered in a special data base; - Dr J.E. Vidal and Mrs Dy Phon of the Laboratoire de Phanérogamie of the Muséum National d'Histoire Naturelle, Paris, for assistance with vernacular names for Laos and Cambodia; - Mrs A. van der Lande, Bennekom, the Netherlands, for assistance with typ­ ing; - all persons, institutions, publishers and authors mentioned in the list 'Sources of illustrations', for authorization to use these illustrations. Acronyms of organizations

- ACIAR: Australian Centre for International Agricultural Research (Can­ berra, Australia) - ARFSN: Asian Rice Farming Systems Network (IRRI, Los Bafios, the Philip­ pines) - ATFGRC: Australian Tropical Forage Genetic Resource Centre (CSIRO, Brisbane, Australia) - BIOTROP: South-East Asian Regional Centre for Tropical Biology (Bogor, Indonesia) - CATIE: Centro Agronómico Tropical de Investigación y Ensenanza [Centre for Tropical Agricultural Research and Training] (Turrialba, Costa Rica). - CENARGEN: Centro Nacional de Recursos Genéticos [National Centre for Genetic Resources] (Brasilia, Brazil) - CIAT: Centro Internacional de Agricultura Tropical [International Center for Tropical Agriculture] (Cali, Columbia) - CNPGC: Centro Nacional de Pesquisa de Gado de Corte [National Centre for Beef Cattle Research] (Campo Grande, Brazil) - CSIRO: Commonwealth Scientific and Industrial Research Organization (Canberra, Australia) - DSIR: Department of Scientific and Industrial Research (Palmerston-North, New Zealand) - EMBRAPA: Empresa Brasileira de Pesquisa Agropecuâria [Brazilian Centre for Animal Production Research] (Brasilia, Brazil) - FAO:Foo d and Agriculture Organization ofth e United Nations (Rome, Italy) - FFTC: Food and Fertilizer Technology Center for the Asian and Pacific Region (Taipei, Taiwan) - FLFAM: Federation of Livestock Farmers Association of Malaysia (Kuala Lumpur, Malaysia) - HITAHR: Hawaiian Institute of Tropical Agriculture and Human Resources (University of Hawaii, Honolulu, United States) - IBPGR: International Board of Plant Genetic Resources (Rome, Italy) - ICA: Instituto Columbiano Agropecuario [Colombian Institute for Animal Production Research] (Bogota, Columbia) - ICARDA: International Center for Agricultural Research in the Dry Areas (Aleppo, Syria) - ICRAF: International Council for Research in Agroforestry (Nairobi, Kenya) - ICRISAT: International Crops Research Institute for the Semi-Arid Tropics (Hyderabad, India) - IDRC: International Development Research Centre (Ottawa, Canada) - ILCA: International Livestock Center for Africa (Addis Ababa, Ethiopia) - IPB: Institute of Plant Breeding (University ofth e Philippines, Los Bahos) 260 FORAGES

IRRI:Internationa l Rice Research Institute (Los Banos, the Philippines) LBN:Lembag a Biologi Nasional [National Biological Institute] (Bogor, Indo­ nesia) LIPI: Lembaga Ilmu Pengetahuan Indonesia [Indonesian Institute of Sciences] (Jakarta, Indonesia) MARDI: Malaysian Agricultural and Development Institute (Serdang, Malaysia) MSAP: Malaysian Society for Animal Production (Kuala Lumpur, Malaysia) NFTA: Nitrogen Fixing Tree Association (Waimanolo, Hawaii, United States) PORIM: Palm Oil Research Institute of Malaysia (Kuala Lumpur, Malaysia) QDPI: Queensland Department of Primary Industries (Australia) UNESCO: United Nations Educational, Scientific, and Cultural Organiza­ tion (Paris, France) USDA: United States Department of Agriculture (Washington, D.C., United States) Glossary

abaxial: on the side facing away from the axis or along the stem, as distinct from opposite or stem (dorsal) whorled abscission: the natural detachment of leaves, amphicarpic: amphicarpous; posessing two kinds branches, flowers or fruits offruit , differing in character or time of ripening accession: in germplasm collections: plant materi­ amplexicaul: stem-clasping, when the petiole-leaf, al of a particular collection, usually indicated or stipule, is dilated at the base, and embraces with a number the stem accessory buds: those additional to the axillary and anastomosis: cross connection ofbranches ; in bota­ normal buds; more than one bud in an axil ny: union of one vein with another, the connec­ accrescent: increasing in size with age tion forming a reticulation achene: a small dry indéhiscent one-seeded fruit androecium: the male element; the stamens as a that does not split open unit of the flower actinomorphic: radially symmetrical; applied to androgynophore: a column on which stamens and flowers that can be bisected in more than one carpels are borne / vertical plane annual: a plant that completes its life cycle in one aculeate: furnished with prickles; prickly year acuminate: ending in a narrowed, tapering point annulate: ring-shaped with concave sides annulus: a ring or a ring-like part acute: sharp; ending in a point with straight or anther: the part of the stamen containing the pol­ slightly convex sides len adaxial: on the side facing the axis (ventral) anthesis: the time the flower is expanded, or, more adnate: united with another part; with unlike parts strictly, the time when pollination takes place fused, e.g., ovary and calyx tube antrorse: directed upwards (opposed to retrorse) adpressed (appressed): lying flat for the whole apetalous: without petals or with a single perianth length of the organ apex: the growing point of a stem or root adventitious: not in the usual place, e.g. roots on apical: at the point of any structure stems, or buds produced in other than terminal apices: plural of apex or axillary positions on stems apiculate: ending abruptly in a short point aerenchyma: a spongy tissue having large thin- apomict: an organism reproducing by apomixis walled cells and large intercellular spaces, serv­ apomixis: reproduction by seed formed without ing for aeration or floating tissue sexual fusion (apomictic) agosporic: a type of apomixis in which a diploid aposporic: with suppression of spore-formation, gametophyte is produced from the sporophyte the prothallus developing direct from the asex­ without spore formation ual generation agrostology: that part of botany dealing with appendage: a part added to another; attached sec­ grasses; graminology ondary or subsidiary part, sometimes projecting albumen: the nutritive material stored within the or hanging seed, and in many cases surrounding the embryo applanate: flattened out or horizontally expanded (endosperm) appressed (adpressed): lying flat for the whole allopolyploid (alloploid): a polyploid with more length of the organ than two sets of chromosomes, derived from dif­ arachnoid: like a cobweb ferent species; allotriploid with three sets, allo- aréole: irregular squares or angular spaces marked tetraploid with four sets, etc. out on a surface, e.g. of a fruit; a small cell or alternate: leaves, etc., inserted at different levels cavity 262 FORAGES aril: an expansion of the funicle enveloping the callus: small hard outgrowth at the base of spike- seed, arising from the placenta; sometimes lets in some grasses occurring as a pulpy cover (arillus) calyx: the outer envelope of the flower, consisting arillode: a false aril, a coat of the seed not arising of sepals, free or united from the placenta campanulate: bell-shaped arilloid: like an aril canaliculate: channelled, with a longitudinal article: a segment of a constricted pod or fruit, as groove in Desmodium canopy: the uppermost leafy layer of a tree or a for­ articulate: jointed, or with places where separation est takes place naturally capitate: headed, like the head of a pin in some stig­ articulation: a joint, popularly applied to nodes of mas, or collected into compact head-like clusters grasses as in some inflorescences ascending: curving or sloping upwards capitellate: diminutive of capitate asynchronous: not synchronous, not existing or capsule: a dry dehiscent fruit composed of two or occurring at the same time more carpels and either splitting when ripe into attenuate: gradually tapering valves, or opening by slits or pores auricle: a small lobe or ear, an appendage to the carinal: relating to the keel in aestivation (the leaf manner in which the parts of a flower are folded awn: a bristle-like appendage, especially occur­ up before expansion) when the carina (keel) ring on the glumes of grasses includes the other parts of the flower axil: the upper angle between the leaf and the stem carpel: one of the foliar units of a compound pistil axillary: arising from the axil or ovary; a simple pistil has only one carpel axis: the main or central line of development of a . cartilaginous: hard and tough plant or organ caruncle: an outgrowth of a seed near the hilum barb: a hooked hair at the base of spikelets in some caryopsis: the fruit of a grass, in which the outer grasses layer (testa) of the seed proper is fused to the bark: the tissue external to the vascular cambium ovary wall collectively, being the secondary phloem, cortex cauline: belonging to the stem or arising from it and periderm chalaza: basal part of the ovule or seed where it beaked: used offruit s which end in a long point is attached to the funiculus and the point at bearded: awned; having tufts of hairs which vascular tissues enter and spread into the bidentate: having two teeth; doubly dentate, as ovule when the marginal teeth are also toothed chalazal: pertaining to the chalaza bidenticulate: minutely bidentate chartaceous: papery biennial: a plant which flowers, fruits and dies in ciliate: with a fringe of hairs along the edge its second year or season clavate: club-shaped or thickened towards the end bifid: cleft into two parts at the tip claw: the narrow part of a petal or sepal biotype: a population or race in which all the indi­ cleft: cut halfway down viduals have the same genetic constitution cleistogamous: pollination and fertilization taking binate: consisting of two members place within the unopened flowers bipinnate: when the primary divisions (pinnae) of clone: a group of plants originating by vegetative a pinnate leaf are themselves pinnate propagation from a single plant and therefore of bisexual: having both sexes present and functional the same genotype in the same flower coenocarpium: the collective fruit of an entire blade: the expanded part of a leaf or petal inflorescence, e.g. as a pineapple or fig bole:th e main trunk of a tree, with a distinct stem compatibility: in floral biology: capable of cross- or bract: a reduced leaf subtending a flower, flower self-fertilization; in plant propagation: stock- stalk or (a part of an) inflorescence scion combinations resulting in a lasting union bracteole: a secondary bract on the pedicel or close compound: of two or more similar parts in one under the flower organ, as in a compound leaf or compound fruit bristle: a stiff hair or a hair-like stiff slender body concave: hollow bullate: surface much blistered or puckered confluent: blended into one, passing by degrees one bunch grass: grass growing in clusters into the other caducous: falling off early conical: having the shape of a cone (cone-shaped) GLOSSARY 263 conjugate: coupled cystolith: mineral concretions, usually of calcium connate: united or joined carbonate ona cellulose stalk connivent: having a gradually inward direction, as deciduous: finally falling off; a perennial plant in many petals (convergent) that sheds its leaves more or less simultaneous­ convex: having amor e or less rounded surface ly,bein g leafless for a while cordate: heart-shaped, as seen at the base of a decumbent: reclining or lying on the ground, but deeply-notched leaf,etc . with thesummi t ascending cordiform: heart-shaped decurrent: extending down and adnate toth e stem, coriaceous: ofleather y texture as occurs insom e leaves corolla: the inner envelope ofth e flower of free or decussate (of leaves): arranged inopposit e pairs on united petals the stem, with each pair perpendicular to the corymb: indeterminate flat- or convex-topped preceding pair inflorescence in which the branches or pedicels dehiscent: opening spontaneously when ripe, e.g., start from different points, but attain approxi­ capsules, legume pods deltoid: shaped like an mately the same level, with the outer flowers equilateral triangle opening first (amodifie d panicle) dentate: margin prominently toothed with the corymbose: flowers arranged to resemble a corymb pointed teeth directed outwards cotyledon: seed leaf, the primary leaf. Dicotylous denticulate: minutely toothed embryos have twocotyledon s and monocotylous determinate: of inflorescences: when the terminal embryos have one or central flower of an inflorescence opens first cover crop: a crop planted to prevent soil erosion and the prolongation of the axis is arrested. Of and toprovid e humus and/or fodder shoot growth: when extension growth takes the crenate: themargi n notched with blunt or rounded form of a flush, i.e. only the previously formed teeth leaf primordia unfold crenulate: crenate (scalloped), but the teeth them­ dicotyledon: angiosperm with two cotyledons or selves small seed-leaves cross-pollination: placement of pollen from one digestibility: the percentage of a foodstuff taken flower onth e stigma of a flower ofanothe r plant into the digestive tract that is absorbed into the which isno to f the same clone body cross-protection: protection of crop plants against digitate: a compound leaf whose leaflets diverge infection by a virulent form of a virus by prior from thesam e point like thefinger s ofa hand inoculation with an innocuous form of that dimorphic: of two forms, as may occur with virus branches, etc. crown: corona; a short rootstock with leaves; the dioecious: with unisexual flowers andwit h thesta - base ofa tufted, herbaceous, perennial grass;th e minate and pistillate flowers on different plants aerial expanse ofa tree, not including the trunk (dioecy) culm: theste m of grasses and sedges diploid: with two sets (genomes) of chromosomes, cultivar (cv., cvs): an agricultural or horticultural as occurs in somatic or body cells; usually writ­ variety that has originated and persisted under ten 2n, having twice the basic chromosome cultivation, asdistinc t from a botanical variety. number ofth e haploid germ cells A cultivar name should always be written with discoid: resembling a disk or discus, being flat and an initial capital letter and given single quo­ circular tation marks (e.g. sabi grass 'Nixon') disk: aflesh y orelevate d development ofth e recep­ cuneate: wedge-shaped; triangular, with the nar­ tacle within the calyx, or corolla or stamens, row enda t the point of attachment, asth e bases often lobed and nectariferous of leaves or petals distal: situated farthest from the place of attach­ cusp: a sharp, rigid point ment cuspidate: abruptly tipped with a sharp rigid point distichous: regularly arranged in two opposite cyme: a determinate inflorescence, often flat- rows on either side ofth e stem topped, inwhic h thecentra l flowers open first DM: dry matter cymose: flowers in a cyme-like inflorescence that dormancy: a term used to denote the inability of a may orma yno tb ea true cyme resting plant orplan t part (e.g.th eseed , ori n tree cymule: a diminutive, usually few-flowered cyme cropsusuall y thebuds )t ogro w ort olea f out even or portion of one under favourable environmental conditions 264 FORAGES

dorsal: back; referring to the back or outer surface floccose: covered with dense hairs that fall away of a part or organ (abaxial) in tufts, locks or flocci ear:th e spike of a grass floret: a small flower, one of a cluster as in grasses eccentric: one-sided, out of the centre or Compositae ecotype: a biotype resulting from selection in a par­ flush: a brief period of rapid shoot growth, with ticular habitat unfolding of the leaf primordia which had accu­ ellipsoid(al): a solid object which is elliptical in mulated during the previous quiescent period section fodder: something fed to domesticated animals, elliptic(al): shaped like an ellipse especially coarse, dried food from plants (hay, emarginate: having a notch at the end; indented straw, leaves) with an acute sinus foliolate (2-,3-, 4- etc.): with 2-,3- ,4 - leaflets emasculate: to remove the anthers from a bud or follicle: a dry, unicarpellate fruit, dehiscing by the flower before the pollen is shed ventral suture to which the seeds are attached embryo: the rudimentary plant within a seed, de­ forage: grassland and fodder plants suitable as feed veloped from a zygote (sexual) or from other nu­ for herbivores, usually with lower nutrient con­ clei in the embryo sac or cells of the nucellus centration and digestibility than concentrates or integuments (apomictic) such as grain endocarp: the innermost layer of the pericarp or foveola: a small pit fruit wall fruit: the ripened ovary with adnate parts entire (botany): with an even margin fulvous: yellow, tawny epicalyx: an involucre of bracts below the flower, funicle (funiculus): the little cord which attaches resembling an extra calyx the ovule or seed to the placenta epicotyl: the young stem above the cotyledons funnelform: salver-shaped, as the corolla of Asysta- epigeal: above ground; in epigeal germination the sia gangetica (L.) T. Anderson cotyledons are raised above the ground fusiform: spindle-shaped; tapering towards each erect: directed towards summit, not decumbent end from a swollen middle erectopatent: between spreading and erect gene: the unit of inheritance located on the chro­ evergreen: bearing foliage all year long; a plant mosome that changes its leaves gradually geniculate: abruptly bent so as to resemble the exsert, exserted: protruded beyond, as stamens knee-joint beyond the tube of the corolla genome: a set of chromosomes as contained within extra-axillary: beyond or outside the axil the gamete and corresponding to the haploid extrorse: an anther which dehisces outwardly to­ chromosome number of the species wards the perianth genotype: the genetic make-up of an organism com­ falcate: sickle-shaped prising the sum total of its genes, both dominant fascicle: a cluster of flowers, leaves, etc., arising and recessive; a group of organisms with the from the same point same genetic make-up ferruginous: rust-coloured germplasm: the genetic material that provides the fertile (botany): bearing pollen which fecundates physical basis of heredity; also a collection of the ovules; said of pollen-bearing anthers or of genotypes of an organism seed-bearing fruits glabrate: destitute ofpubescenc e or any roughness fertilization: union of the gametes (egg and sperm) glabrescent: becoming glabrous or nearly so to form a zygote glabrous: devoid of hairs fibrous: having much woody fibre glandular: having or bearing secreting organs or fig: the fleshy multiple fruit derived from the glands inflorescence of Ficus spp. (syconium) glaucous: pale bluish-green, or with a whitish filament: thread; the stalk supporting the anther bloom which rubs off filiform: slender; threadlike globose: spherical or nearly so fimbriate: fringed glumes: the chaffy or membranous two-ranked flabellate: fan-shaped, dilated in a wedge shape, members of the inflorescence of grasses and sim­ sometimes plaited (folded) ilar plants; lower glume and upper glume, two flaky: lamelliform, in the shape of a plate or scale sterile bracts at the base of a grass spikelet flexuose, flexuous: zigzag;ben t alternately in oppo­ granulöse (granular): composed of or covered with site directions grain-like tiny particles GLOSSARY 265 gynoecium: the female part or pistil of a flower, hypanthium: a cup-like receptacle usually derived consisting, when complete, of one or more ovar­ from the fusion of the floral envelopes and ies with their styles and stigmas androecium on which are seemingly borne the gynophore: a stalk supporting the gynoecium calyx, corolla and stamens formed by elongation of the receptacle hypocotyl: the young stem below the cotyledons habit: external appearance or way of growth of a hypogeal: below ground; in hypogeal germination plant the cotyledons remain below ground within the habitat: the kind of locality in which a plant testa grows imbricate: overlapping like tiles; in a flower bud haploid: having a single set (genome) of chromo­ when one sepal or petal is wholly external and somes in a cell or an individual, corresponding one wholly internal and the others overlapping to the chromosome number (n) in a gamete at the edges only hard-seededness: impermeability of the seed-coat, imparipinnate: pinnate with an odd terminal leaf­ which avoids quick germination let harvest index: the total harvestable produce as a impressed: marked with slight depressions fraction of the total biomass produced by the incised: cut deeply crop in a given year incompatibility: in floral biology: not capable of herb: any which is not woody cross- or self-fertilization; in plant propagation: herbaceous: not woody no stock-scion combinations resulting in a last­ heritability: the proportion of variability that ing union results from genetic causes; also that proportion indéhiscent: not opening when ripe of the variation of a population that is transmit­ indented: forced inward to form a depression ted to progeny indeterminate: of inflorescences: a sequence in hermaphrodite: bisexual; in flowers, with stamens which the terminal flowers are the last to open, and pistil in the same flower so that the floral axis may be prolonged indefini­ heterogamous: with two or more kinds or forms of tely by the terminal meristem. Of shoot growth: flowers when the shoot apex forms and unfolds leaves heterozygote: an organism with different genes at during extension growth, so that shoot growth corresponding loci of homologous chromo­ can continue indefinitely somes; consequently producing unlike gametes indumentum: any covering, as hairiness heterozygous: the condition in which homologous inferior ovary: situated below the sepals,petal s and chromosomes of an individual possess different stamens alleles at corresponding loci inflexed: turned abruptly or bent inward hilum: the scar left on a seed indicating its point inflorescence: the arrangement and mode of devel­ of attachment opment of the flowers on the floral axis hirsute: hairy, with long, tolerably distinct hairs infructescence: a ripened inflorescence in the fruit­ hispid: covered with long rigid hairs or bristles ing stage hispidulous: minutely hispid inoculum: material used for inoculation, e.g. rhizo- homomorphous: uniform in shape bia in soil to promote the growth of certain Legu- homozygote: an individual whose homologous minosae chromosomes carry identical genes at corres­ internode: the portion of the stem between two ponding loci nodes homozygous: possessing identical genes at corres­ in vitro: outside the living body and in an artificial ponding loci on homologous chromosomes environment hull: see husk involucral: belonging to an involucre husk: the outer covering of certain fruits or seeds involucre: a ring of bracts surrounding several hyaline: almost transparant flowers or their supports, as in the heads of Com- hybrid: the first generation offspring of a cross be­ positae or the umbels in Umbelliferae tween two individuals differing in one or more involute: having the edges of the leaves rolled genes inwards hybridization: the crossing ofindividual s of unlike isozymes: multiple distinct molecular forms of an genetic constitution enzyme that differ in net electrical charge; hygroscopic: susceptible to extending or shrinking important to the investigation of the molecular on application or removal of water or vapour basis for cellular differentiation and morphoge- 266 FORAGES

nesis, and increasingly used to clarify genotypic lobe:an y division of an organ or specially rounded relationships division isthmus: anarrowe d connection between two parts lobed:o f leaves: divided, but not into separate leaf­ IVD: in vitro digestibility lets IVOMD: in vitro organic matter digestibility locular: divided by internal partitions into com­ jugate: connected or yoked together; e.g. in leaves partments as in anthers and ovaries 1-n jugate: with 1-n pairs of leaflets lament: lomentum; a legume or pod which is con­ juvenile phase (stage): the period between germina­ tracted between the seeds, falling apart at the tion and the first signs of flowering, during constrictions into one-seededj oint swhe n mature which vegetative processes preclude flower ini­ Malesia: the bio-geographical region including tiation even under the most favourable condi­ Malaysia, Indonesia, the Philippines, tions Singapore, Brunei and Papua New Guinea keel (carina): a ridge like the keel of a boat; the margin: the edge or boundary line of a body two anterior and united petals of a papiliona­ marginate: furnished with a margin of distinct ceous corolla; the principal nerve of a sepal or character glume mass selection: a system of breeding in which seed keeled (carinate): having a keel or carina from individuals selected on the basis of pheno- kernel: the nucellus of an ovule or of a seed, that type is used to grow the next generation is, the whole body within the coats membranous: thin and semi-transparant, like a knee: an abrupt bend in a stem or tree-trunk fine membrane labyrinthine seed: seed with the testa filling the meristem: undifferentiated tissue of the growing crevices between the transverse lobes and folds point whose cells are capable of dividing and de­ of the cotyledons which closely adhere together veloping into various organs and tissues lacerate(d): torn, or irregularly cleft merous: (4-,5-,etc.) with 4,5 ,etc .part s or numbers lamina: see blade of sepals,petal s etc. lanceolate: lance-shaped; much longer than broad, midrib: the main vein of a leaf which is a continua­ being widest at the base and tapering to the apex tion of the petiole lateral: fixed on or near the side of an organ : angiosperms having a single coty­ latex: a juice, usually white and sometimes sticky, ledon or seed-leaf which exudes from broken surfaces of some monoecious: with unisexual flowers, but male and plants female flowers borne on the same plant lax: loose, distant monogastric: having a stomach with only one com­ leaflet: one part of a compound leaf partment (e.g. swine, chicks) lemma: the lower of the two membranous bracts monopodial: of a primary axis which continues its enclosing the flower in grasses; the lower of the original line of growth from the same apical mer­ two glumes which surround each floret in the istem to produce successive lateral branches spikelet of grasses morphotype: a special form of a plant species, lenticel: lenticular masses ofloos e cells protruding usually only slightly differing from the normal through fissures in the periderm on stems, fruits habit and roots; they usually arise beneath individual mucro: a sharp terminal point stomata and their main function is gaseous mucronate: ending abruptly in a short stiff point exchange mucronulate: diminutive of mucronate lenticular: shaped like a doubly convex lens muricate: rough, with short and hard tubercular lignification: the thickening or hardening of the excrescences cell-wall by secondary deposits mycorrhiza: a symbiotic association of roots with ligulate: with or possessing a ligule a fungal mycelium which may form a layer out­ ligule: a membranous outgrowth on the upper sur­ side the root (ectotrophic) or within the outer face of a grass leaf at the junction of the sheath root tissue (endotrophic) and the blade. It may be presented by a ridge or nectar: a sweet fluid exuded from various parts of by a line of hairs; an elongated flattened strap- the plant (e.g.fro m the flower to attract pollina­ shaped structure tors) limb: the expanded part of a tubular corolla, as dis­ nematode: small elongated cylindrical worm-like tinct from the tube or throat; the lamina of a leaf micro-organism, free-living in soil or water, or or of a petal parasitic in animals or plants GLOSSARY 267

nerve: a strand of strengthening and/or conducting paripinnate: a pinnate leaf without the odd termi­ tissue running through a leaf, which starts from nal leaflet the midrib and diverges or branches throughout parthenocarpy: the production offrui t without true the blade fertilization neuter: sexless, neither male nor female; having patent: spreading neither functional stamens nor pistils pedicel: stalk of each individual flower of an inflor­ node: the point on the stem or branch at which a escence leaf or lateral is borne pedicellate: borne on a pedicel nodulation: formation of root-nodules peduncle: the stalk of an inflorescence or partial nucellus: the nutritive tissue in an ovule inflorescence -: the inverse or opposite condition (obtriangu- pellucid: translucent lar, obcordate, etc.) peltate: shield-shaped, as a leaf attached by its oblanceolate: reverse of lanceolate lower surface to a stalk instead of by its mar­ oblate: flattened at the poles gin obligate: necessary, essential; the reverse of facul­ pendent: hanging down from its support tative pendulous: drooping; hanging down oblique: slanting; of unequal sides pentamerous: having five parts in a flower-whorl oblong: longer than broad, with the sides parallel perennial: a plant living for several years and or almost so usually flowering each year oblongoid: a solid object which is oblong in section perfect flower: a flower possessing both male and obovate: reverse of ovate female organs obovoid: a solid object which is obovate in section perianth: the floral leaves as a whole, including obtuse: blunt or rounded at the end sepals and petals if both are present opaque: neither shining nor transparent petal: a member of the inner series of perianth seg­ opposite: of leaves and branches when two are ments which are often brightly coloured borne at the same node on opposite sides of the petiolar: borne on, or pertaining to a a petiole stem petiolate: having a petiole orbicular: flat with a more or less circular outline petiole: the stalk of a leaf orthotropic: having a more or less vertical direc­ petiolule: the stalk of a leaflet tion of growth phenology: the complex annual course of flushing, ovary: that part of the pistil, usually the enlarged quiescence, flowering, fruiting and leaf fall in a base, which contains the ovules and eventually given environment becomes the fruit phenotype: the physical or external appearance of ovate: egg-shaped; a flat surface which is scarcely an organism as distinguished from its genetic twice as long as broad with the widest portion constitution (genotype); a group of organisms below the middle with similar physical or external make-up ovoid: a solid object which is egg-shaped (ovate) photoperiod: length of day favouring optimum in section functioning of an organism ovule: the immature seedsi n the ovary before fertil­ phyllotaxy: the arrangement of leaves or floral ization parts on their axis palea: the upper oftw o membranous bracts enclos­ physiological varieties (races): pathogens of the ing the flower in grasses same species which are structurally similar, but palmate: lobed or divided like the palm of the hand which differ in physiological and pathological panicle: an indeterminate branched racemose characteristics inflorescence pilose: hairy with rather long soft hairs paniculate: resembling a panicle pilosity: hairiness papilionaceous: a butterfly-shaped corolla, as in pinna (plural pinnae): a primary division or leaflet Papilionaceae of a pinnate leaf papillae: soft superficial glands or protuberances pinnate: a compound leaf with the leaflets papillose: covered with minute nipple-like protu­ arranged along each side of a common rachis berances pinnatifid: with the margin pinnately cleft pappus: the various tufts of hairs on achenes or pistil: the female part of a flower (gynoecium) of fruits; the limb ofth e calyx of Compositae florets one or more carpels, consisting, when complete, papyraceous: papery, like paper of one or more ovaries, styles and stigmas 268 FORAGES pistillate: a unisexual flower with pistil, but no sta­ raceme: an unbranched elongated indeterminate mens inflorescence with stalked flowers opening from pistillode: a sterile, often reduced pistil the base upwards placenta: the part of the ovary to which the ovules racemose: raceme-like are attached rachis (plural: rachides): the principal axis of an plano-convex: flat on one side and convex on the inflorescence or a compound leaf other radical: arising from the root, or its crown plagiotropic: having an oblique or horizontal direc­ radicle: the first root of an embryo or germinating tion of growth seed plumose: featherlike with fine hairs, as on the sides receptacle: the flat, concave or convex part of the of some bristles stem from which the parts of the flower arise plumule: the primary bud of an embryo or germi­ recurved: bent or curved downward or backward nating seed reflexed: abruptly bent or turned downward or pod: a dry and many-seeded dehiscent fruit, a backward legume or silique reniform: kidney-shaped pollen: spores or grains borne by the anthers con­ resupinate: upside down, or apparently so taining the male element (gametophyte) reticulate: netted, as when the smallest veins of a pollination: the transfer of pollen from the dehisc­ leaf are connected together ing anther to the receptive stigma retrorse: turned or directed backward or downward polyembryony: the production of two or more (opposed to antrorse) embryos within an ovule rhizobia: the organisms forming root-nodules in polygamous: with unisexual and bisexual flowers Leguminosae in the same plant rhizoid: root-like polymorphic: polymorphous, with several or var­ rhizome: an underground stem which is distin­ ious forms; variable as to habit guished from a root by the presence of nodes polyphenol: a polyhydroxy phenol with buds and leaves or scales posterior: next to or towards the main axis rhombic: shaped like an equilateral oblique-angled precocious: exceptionally early in development; figure flowering and fruiting at an early age rhomboid (botany):quadrangular , with the lateral prickle: a sharp, relatively stout outgrowth from angles obtuse the outer layers root-nodules: small swellings on roots of legumi­ primordial: first in order of appearance nous and other plants, containing nitrogen-fix­ procumbent: lying along the ground ing bacteria (rhizobia) prolific: fruitful, producing offspring rootstock: see rhizome prop roots: aerial roots rosette: a cluster of leaves or other organs in a cir­ prostrate: lying flat on the ground cular form protandrous: stamens shedding pollen before the rotund: rounded in outline, somewhat orbicular, stigma is receptive but slightly inclined towards oblong proximal: the part nearest the axis (as opposed to rudimentary: of organs which are imperfectly de­ distal) veloped and nonfunctional pseudogamy: parthenogenetic fruiting, as pollina­ rugose: wrinkled tion without impregnation of ovules rugulose: somewhat wrinkled pseudopetiole: a structure resembling a petiole, but ruminant: an animal that chews again what has not being one been swallowed (e.g. sheep, cows, camels, goats) puberulent: covered with soft fine hairs runner: a specialized stem that develops from a leaf puberulous: minutely pubescent axil at the crown of a plant, grows horizontally pubescent: covered with soft short hairs along the ground, and forms a new plant at one pulvinule: the swollen base of a petiolule of the nodes, usually at or near the tip (as in pulvinus: the swollen base of the petiole ) punctate: marked with dots or translucent glands scabrid: rough to the touch p.v.: see physiological varieties (races) scabridulous: slightly rough pyriform: resembling a pear in shape scabrous: see scabrid quadrangular: four-cornered scalariform: having markings suggestive of a lad­ quadrate: approximately square or cubical der GLOSSARY 269

scale: reduced leaf, usually sessile, thin and dry, spikelet: a secondary spike, one of the units com­ and seldom green prising the inflorescence in grasses, consisting scandent: climbing of one or more florets on a thin axis, subtended scarification (seed): scarifying, to cut or soften the by a common pair of glumes wall ofa hard seed to hasten germination spine: a short stiff straight sharp-pointed hard scrub: vegetation whose growth isstunte d because structure of lack of water coupled with strong transpira­ spinescent: ending in a spine or sharp point tion spinose: having spines (spinous) scurfy: bearing small scales on the surface (lepi- spinulose: with small spines dote; scaly) stamen: one of the male reproductive organs of a seed:th ereproductiv e unit formed from a fertilized flower; a unit ofth e androecium ovule, consisting of embryo and seed-coat, and, staminate: a flower bearing stamens butn o pistil in some cases, also endosperm staminode: an abortive or rudimentary stamen self-sterile: failure to complete fertilization and without a perfect anther obtain seed after self-pollination standard (flower part): the fifth or posterior petal self-pollination: pollination with pollen from the of a papilionaceous corolla same flower or from other flowers of plants of stellate: star-shaped, as of hairs with radiating the same clone branches self-compatible: see self-fertile sterile: failing to complete fertilization and pro­ self-fertile: capable offertilizatio n andsettin g seed duce seed as a result of defective pollen or after self-pollination ovules; not producing seed capable of germina­ semi-: half, incomplete (e.g. semi-inferior ovary) tion; lacking functional sexual organs (sterili- sepal: a member ofth e outer series of perianth seg­ ty) ments stigma: theportio n of the pistil which receives the septate: divided bya partition pollen septum (plural septa): a partition or cross-wall stilt roots: the oblique adventitious roots of the seriate: serial, disposed in series of rows mangrove andsimila r forms sericeous: silky stipe: thestal k supporting a carpel or gynoecium serrate: toothed like a saw, with regular pointed stipel (stipella): a minute stipule on a partial teeth pointing forwards petiole ofcompoun d leaves sessile: without a stalk stipitate: having a stipe or special stalk seta(e): a bristle-like body stipulate: with orbearin g stipules setose: bristle-pointed; terminating gradually in a stipule: a scale-like or leaf-like appendage at the fine, sharp point base ofa leaf petiole setulose: resembling a fine bristle stolon: a trailing stem usually above the ground sheath: a tubular structure surrounding an organ which is capable of producing roots and shoots or part, asth elowe r part ofth e leaf clasping the at its nodes stem in grasses stoloniferous: bearing a stolon or stolons shoot: a young growing branch or twig straggling: divaricate, extremely divergent shrub: a woody plant which branches from the striate: marked with fine longitudinal parallel base, allbranche s being equivalent lines, as grooves or ridges silique: a dry and many-seeded dehiscent fruit style: the part of the pistil connecting the ovary splitting into 2valve s with a false partition with the stigma simple (botany): not compound, as in leaves with sub-: somewhat or slightly (e.g. subacute) a single blade subglobose: nearly globular sod (turf): the upper stratum ofsoi l bound by grass subulate: awl-shaped and plant roots into a thick mat subverticillate: inimperfec t or irregular whorls spathate: furnished with a spathe succulent: juicy, fleshy spathe: a large bract enclosing a flower cluster sucker: a shoot, usually originating from adventi­ spathulate: spoon-shaped tious buds on the roots or basal stem parts, spicate: spike-like which does not fit in the architectural model, spiciform: spike-like but is capable of repeating the model spike: a simple indeterminate inflorescence with suffrutescent: obscurely shrubby sessile flowers along a single axis sulcate: grooved or furrowed 270 FORAGES

superior (ovary): an ovary with the perianth tufted: growing in tufts (caespitose) inserted below or around its base, the ovary tunicated: provided with a dry papery covering being attached at its base only round a bulb or corm suturai: relating to a suture tussock: a tuft of grass or grass-like plants suture: the line of junction of two carpels; the line umbilicus: see hilum or mark of splitting open unarmed: destitue of prickles or other armature; sympodial: of a stem in which the growing point pointless either terminates in an inflorescence or dies, uncinate: hooked growth being continued by a subtending lateral uncinulate: diminutive of uncinate growing point undershrub: any low shrub; partially herbaceous syncarpous: of an ovary composed of two or more shrub, the ends ofth e branches perishing during united carpels the winter synchronous: happening, existing or arising at the undulate: wavy, said for instance of a leaf margin same time if the waves run in a plane at right angles to the taproot: the primary descending root, forming a plane of the leaf blade direct continuation ofth e radicle unifoliolate: with one leaflet only, but in origin a taxon (plural taxa): a term applied to any taxono­ compound leaf mie unit irrespective ofit s classification level unisexual: of one sex, having stamens or pistils TDN: total digestible nutrients only tendril: a thread-like climbing organ formed from valve: a piece into which a capsule naturally sepa­ the whole or part of a stem, leaf or petiole rates at maturity tepal: a segment of a perianth, sepal or petal variety: botanical variety which is a subdivision of terete: cylindrical; circular in transverse section a species; an agricultural or horticultural vari­ terminal: borne at the end or apex ety is referred to as a cultivar terrestrial: on or in the ground vein: a strand of vascular tissue in a flat organ, testa: the outer coat of the seed such as a leaf tetrafoliolate: with four leaflets velutinous: see velvety tetraploid: having four times the basic number of velvety: with a coating of fine soft hairs; the same chromosomes, usually written An as tomentose but denser so that the surface thorn: a woody sharp-pointed structure formed resembles (and feels like) velvet from a modified branch venation: the arrangement ofth e veins in a leaf thyrse: a compound inflorescence composed of a ventral: faces central axis (adaxial), opposed to panicle (indeterminate axis) with the secondary dorsal and ultimate axes cymose (determinate) versatile (botany): turning freely on its support, as thyrsiform: shaped like a thyrse many anthers on their filaments thyrsoid: like a thyrse villose (villous): with long weak hairs tomentose: densely covered with short soft hairs vine: a plant having a stem that is too slender to torus: receptacle hold itself erect and therefore supports itself by trailing: prostrate, but not rooting climbing over an object tree: a perennial woody plant with an evident per­ viny: trailing or climbing sistent trunk violaceous: violet-coloured trifoliate: three-leaved viscid: sticky trifoliolate: with three leaflets viviparous: germinating or sprouting from seed or triploid: having three times the basic number of bud while attached to the parent plant chromosomes, usually written 3re warty: covered with firm roundish excrescences tripping (flowers): to help in pollinating whorl: more than two organs ofth e same kind aris­ triquetrous: three-edged, with three salient angles ing at the same level truncate: cut off more or less squarely at the end wing: any membraneous expansion attached to an trunk: the main stem of a tree (as opposed to the organ; a lateral petal ofa papilionaceous corolla roots and branches) xerophytic: relating to a plant structurally adapted tubercle: a small tuberlike excrescence for life and growth with a limited water supply tuberculate: covered with warty protuberances tuberiform: resembling a tuber tuberous: producing tubers or resembling a tuber Sources of illustrations

All illustrations have been redrawn and adapted Axonopus compressus: Häfliger, E. & Scholz, H., by Mrs, P. Verheij-Hayes, using the following 1980. Grass weeds. Documenta, Ciba-Geigy, sources: Basel. Vol. 1.Aquarell e (without page number). Bothriochloa pertusa: Matthew, K.M., 1982. Illus­ Aeschynomene americana: Skerman, P.J., trations on the flora of the Tamilnadu Carnatic. Cameron, D.G. &Riveros , F., 1988.Tropica l for­ The Rapinat Herbarium, Tiruchirapalli, India. age legumes. FAO, United Nations, Rome, Italy, PI. 816 (flowering plant, ligule); Skerman, P.J. p. 207, Fig. 35 (leafy branch); Koorders, S.H., &Riveros , F., 1990.Tropica l grasses. FAO, Uni­ 1926. Exkursionsflora von Java. Band 4, Atlas. ted Nations, Rome, Italy, p. 231, Fig. 15.14 Gustav Fischer Verlag, Jena, Deutschland. Lie­ (inflorescence, sessile and pedicelled spikelet). ferung 7(1).p . 960,Fig . 1252 (flowering and fruit­ Brachiaria brizantha: Clayton, W.D., 1972. Grami- ing branch). neae. In: Hepper, F.N. (Editor): Flora of West Aeschynomene falcata: original drawing by P. Ver­ Tropical Africa. 2nd ed. Crown Agents for Over­ heij-Hayes, Prosea, Wageningen. sea Governments and Administration's. London. Albizia lebbeck: Townsend, C.C., 1974. Legumi- Vol. 3(2). p. 441, Fig. 442. nales. In: Townsend, C.C. & Evan Guest (Edi­ Brachiaria decumbens: leafy branches: drawing tors): Flora of Iraq. Vol. 3. Ministry of Agricul­ provided by CIAT; Ibrahim, K.M. & Kabuya, ture & Agrarian Reform, Baghdad, Iraq. p. 33, C.M.S., 1987. An illustrated manual of Kenya PI. 5. grasses. FAO, United Nations, Rome, Italy, p. Alysicarpus vaginalis: Ohashi, H., 1977.Legumino - 330, PI. 47 (inflorescence). sae. In: Hui-lin Li et al. (Editors): Flora of Brachiaria dictyoneura: leafy branch: drawing pro­ Taiwan. Epoch Publishing Company, Taipei. vided by CIAT; Ibrahim, K.M. & Kabuya, Vol. 3. p. 171, PI. 516. C.M.S., 1987. An illustrated manual of Kenya Andropogon gayanus: Bowden, B.N., 1964.A n out­ grasses. FAO, United Nations, Rome, Italy, p. line of the morphology and anatomy of Andropo­ 332, PI. 49 (inflorescence). gon gayanus var. bisquamulatus (Höchst.) Brachiaria distachya: Gilliland, H.B., Holttum, Hack. The Journal of the Linnean Society of R.E., Bor, N.L., 1971.Grasse s ofMalaya . In: Bur- London, Botany 58:510 ,Text-fig . 1. kill, H.M. (Editor): Flora of Malaya. Govern­ Arachis glabrata: drawing provided by the Queens­ ment Printing Office, Singapore. Vol. 3. p. 177, land Herbarium, Meiers Road, Indooroopilly, Q Fig. 36 (leafy branch); Häfliger, E. &Scholz , H., 4068, Australia. 1980. Grass weeds. Documenta, Ciba-Geigy, Arachis pintoi: drawing provided by the Queens­ Basel. Vol. 1.p . 14(inflorescence , spikelet). land Herbarium, Meiers Road, Indooroopilly, Q Brachiaria humidicola: leafy plant part: drawing 4068, Australia. provided by CIAT; Ibrahim, K.M. & Kabuya, Arundinaria pusilla: Camus, E.G. & A., 1923. In: C.M.S., 1987. An illustrated manual of Kenya Lecomte, M.H. &Gagnepain, F. (Editors): Flore grasses. FAO, United Nations, Rome, Italy, p. générale de lTndo-Chine. Vol. 7. p. 586, Fig. 45 334, PL 51 (inflorescence). (leafy twig, apex of inflorescence); habit of Brachiaria mutica: Soerjani, M., Kostermans, plant with rhizome: drawing provided by the A.J.G.H., Tjitrosoepomo, G. (Editors), 1987. author. Weeds of rice in Indonesia. Balai Pustaka, Ja­ Asystasia gangetica: Engler, A. & Prantl, K. (Edi­ karta, Indonesia, p. 393,Fig . 4.180. tors), 1895. Die natürlichen Pflanzenfamilien. Brachiaria ruziziensis: habit: drawing provided by W. Engelmann Verlag, Leipzig. Vol. 4 (3b). p. CIAT; original drawing by P. Verheij-Hayes, 326, Fig. 131. after Reekmans 11040 (WAG) (inflorescence). 272 FORAGES

Brachiaria subquadripara: drawing provided by Chloris gayana: Häfliger, E. & Scholz, H., 1981. CIAT. Grass weeds. Documenta, Ciba-Geigy, Basel. Calliandra calothyrsus: Westphal, E. & Jansen, Vol. 2. p. 48 (ligule, spikelets); aquarelle (with­ P.C.M. (Editors), 1989. Plant Resources of out page number) (leafy plant parts); original South-East Asia. A selection. Pudoc, Wage- drawing by P. Verheij-Hayes after Pawek 7199 ningen. p.70 . (WAG) (inflorescence). Calopogonium caeruleum: D'Arcy, W.G., 1980. Chrysopogon aciculatus: Stanfield, D.P., 1970. List Calopogonium. In: Dwyer, J.D. et al. (Editors): of illustrations. Supplement to grasses of the Flora ofPanama . Annals ofth e Missouri Botani­ flora of Nigeria. Ibadan University Press, cal Garden 67: 559, Fig. 8 (flowering branch, Nigeria. PI.4 3(flowerin g plant); Gilliland, H.B., flower); de Martius, C.F.P. (Editor), 1859. Flora Holttum, R.E., Bor, N.L., 1971. Grasses of Brasiliensis. Vol. 15(1),T . 38 (fruits). Malaya. In: Burkill, H.M. (Editor): Flora of Calopogonium mucunoides: Verdcourt, B., 1979. A Malaya. Government Printing Office, manual of New Guinea legumes. Botany Bulle­ Singapore. Vol. 3.p . 235, Fig. 52 (triad of spike- tin No 11. Office of Forests, Division of Botany, lets). Lae, Papua New Guinea, p.482 ,Fig . 114 (flower­ Chrysopogon orientalis: Matthew, K.M., 1988. Fur­ ing branch); Purseglove, J.W., 1968. Tropical ther illustrations on the flora of the Tamilnadu Crops. Dicotyledons. Longmans, London. Vol. 1. Carnatic. The Rapinat Herbarium, Tiruchira- p. 219,Fig . 32 (fruiting branch). palli, India. PI. 736. Canavalia ensiformis: Botton, H. &Halle , N, 1958. Clitoria ternatea: Milne-Redhead, E. & Polhill, Les plantes de couverture. Guide pratique de R.M. (Editors), 1971. Flora of Tropical East reconnaissance et d'utilisation des Légumi­ Africa - Leguminosae 4 - Papilionoideae 2. neuses en Côte d'Ivoire 2.Journa l d'Agriculture Crown Agents for Oversea Governments and Ad­ tropicale et de botanique appliquée 5:99 ,PI . 27. ministrations, p. 516, Fig. 75 (flowering branch, Cenchrus ciliaris: Humphreys, L.R., 1980. A guide fruit); Botton, H. & Halle, N, 1958. Les plantes tobette r pastures for the tropics and sub-tropics. de couverture. Guide pratique de reconnais­ 4th ed. Wright Stephenson & Co., Ermington, sance et d'utilisation des Légumineuses en Côte Sydney, p. 25 (habit); Häfliger, E. & Scholz, H., d'Ivoire 2.Journa l d'Agriculture tropicale et de 1980. Grass weeds. Documenta, Ciba-Geigy, botanique appliquée: 133,P L 33 (flower). Basel. Vol. 1. p. 20 (ligule); Tothill, J.C. & Codariocalyx gyroides: Backer, C.A. & van Sloo- Hacker, J.B., 1983. The grasses of southern ten, D.F., 1924. Geïllustreerd handbook der Queensland. University of Queensland Press, Javaansche theeonkruiden en hunne beteekenis Brisbane, Australia, p.14 8(inflorescence) ; Sker- voor de cultuur [Illustrated handbook of the man, P.J. & Riveros, F., 1990. Tropical grasses. Javanese weeds in tea and their agricultural sig­ FAO, United Nations, Rome, Italy, p. 267, Fig. nificance]. Algemeen proefstation voor thee, 15.22 (spikelet with involucre). Batavia, p. 140, Fig. 140 (flowering and fruiting Centotheca latifolia: Bor, N.L., 1960. The grasses branch); Ohashi, H., 1973. The Asiatic species of of Burma, Ceylon, India and Pakistan. Perga- Desmodium and its allied genera (Legumino­ mon Press, Oxford, U.K. p.458 ,Fig . 50. sae). Ginkgoana 1:42 , Fig. 8 (fruit). Centrosema acutifolium: drawing provided by Crotalaria juncea: Botton, H. &Halle , N, 1958. Les CIAT. plantes de couverture. Guide pratique de recon­ Centrosema macrocarpum: drawing provided by naissance et d'utilisation des Légumineuses en CIAT. Côte d'Ivoire 2. Journal d'Agriculture tropicale Centrosema pascuorum: drawing provided by the et de botanique appliquée: 49,P L 16. author. Cynodon dactylon: Häfliger, E. &Scholz , H., 1981. Centrosemapubescens: Botton, H. &Halle , N, 1958. Grass weeds. Documenta, Ciba-Geigy, Basel. Les plantes de couverture. Guide pratique de Vol.2 .Aquarell e (without page number) (flower­ reconnaissance et d'utilisation des Légumineuses ing plant); p. 53(ligule , spikelet). en Côted'Ivoir e 2.Journa l d'Agriculture tropicale Cynodon nlemfuensis: original drawing by P. Ver­ et debotaniqu e appliquée 5.p . 127,PI . 31. heij-Hayes, Prosea, Wageningen, from Tanner Chamaecrista rotundifolia: Adams, G.T. (Editor), 5417 (WAG) (flowering plant); Ibrahim, K.M. & 1985.Annua l report 1984-1985. CSIRO, Division Kabuya, C.M.S., 1987. An illustrated manual of of Tropical Crops &Pastures , Brisbane, Austra­ Kenya grasses. FAO, United Nations, Rome, lia, (front cover). Italy, p. 388,Fig . 105 (spikelet). SOURCES OF ILLUSTRATIONS 273

Dactyloctenium aegyptium: Häfliger, E. & Scholz, Dichanthium annulatum: Cope, T.A., 1982. Poa- H., 1981.Gras s weeds. Documenta, Ciba-Geigy, ceae. In: Nasir, E. &Ali , S.I. (Editors): Flora of Basel. Vol. 2. Aquarelle (without page number) Pakistan. No 143. Department of Botany, Uni­ (flowering plant);p . 56(ligule , spikelet). versity of Karachi, p. 281, Fig. 30 (flowering Desmanthus virgatus: Verdcourt, B.,1979 .A manu­ plant, spikelet, caryopsis); Häfliger, E. &Scholz , al of New Guinea legumes. Botany Bulletin No H., 1980. Grass weeds. Documenta, Ciba-Geigy, 11. Office of Forests, Division of Botany. Lae, Basel. Vol. 1.p . 39 (ligule). Papua New Guinea, p. 144,Fig . 36. Digitaria ciliaris: Soerjani, M., Kostermans, Desmodium heterocarpon: Backer, C.A. & van A.J.G.H., Tjitrosoepomo, G. (Editors), 1987. Slooten, D.F., 1924. Geïllustreerd handbook der Weeds of rice in Indonesia. Balai Pustaka, Ja­ Javaansche theeonkruiden en hunne beteekenis karta, Indonesia, p.403 ,Fig .4.185 . voor de cultuur [Illustrated handbook of the Digitaria eriantha: Leistner, O.A. (Editor), 1990. Javanese weeds in tea and their agricultural sig­ Grasses of southern Africa. Memoirs ofth e bota­ nificance]. Algemeen proefstation voor thee, nical survey of South Africa No 58. p. 106, Fig. Batavia, p. 141,Fig . 141 (flowering and fruiting 63. branch); Ohashi, H., 1973.Th e Asiatic species of Digitaria milanjiana: Skerman, P.J. &Riveros , F., Desmodium and its allied genera (Legumino- 1990. Tropical grasses. FAO, United Nations, sae). Ginkgoana 1: 199, Fig. 62(7) (fruit), p.201 , Rome, Italy, p. 361, Fig. 15.50(leaf y plant, inflor­ Fig. 64(3) (seeds). escence); Hacker, J.B., 1983.Inheritanc e of sto­ Desmodium hetercarpon ssp. ovalifolium: original lon development, rhizome development and seti- drawing by P. Verheij-Hayes, Prosea, Wage­ gerous lemmas in the Digitaria milanjiana ningen, from living material. complex, and its taxonomie significance. Aus­ Desmodium heterophyllum: Backer, C.A. & van tralian Journal of Botany 31:358 , Fig. 1 (spike- Slooten, D.F., 1924. Geïllustreerd handbook der lets). Javaansche theeonkruiden en hunne beteekenis Echinochloa colona: Soerjani, M., Kostermans, voor de cultuur [Illustrated handbook of the A.J.G.H., Tjitrosoepomo, G. (Editors), 1987. Javanese weeds in tea and their agricultural sig­ Weeds of rice in Indonesia. Balai Pustaka, Ja­ nificance]. Algemeen proefstation voor thee, karta, Indonesia, p. 417,Fig . 4.192. Batavia, p. 142, Fig. 142 (flowering and fruiting Echinochloa crus-galli: Soerjani, M., Kostermans, branch); Ohashi, H., 1973.Th e Asiatic species of A.J.G.H., Tjitrosoepomo, G. (Editors), 1987. Desmodium and its allied genera (Legumino- Weeds of rice in Indonesia. Balai Pustaka, Ja­ sae). Ginkgoana 1:231 ,Fig . 67(6) (fruit, seeds). karta, Indonesia, p.419 , Fig.4.193 . Desmodium incanum: Hosaka, E.Y. & Ripperton, Eragrostis tenella: Soerjani, M., Kostermans, J.C., 1944.Legume s ofHawaiia n Ranges. Hawaii A.J.G.H., Tjitrosoepomo, G. (Editors), 1987. Agricultural Experiment Station Bulletin No Weeds of rice in Indonesia. Balai Pustaka, Ja­ 93. University of Hawaii, Honolulu, Hawaii, p. karta, Indonesia, p.425 ,Fig . 4.196. 31, Fig. 11 (vegetative plant part, flower, fruit, Eragrostis unioloides: Soerjani, M., Kostermans, seed); Wagner, W.L., Herbal, D.R. & Sohmer, A.J.G.H., Tjitrosoepomo, G. (Editors), 1987. S.H., 1990. Manual of the flowering plants of Weeds of rice in Indonesia. Balai Pustaka, Ja­ Hawaii. Bishop Museum Special Publication 83, karta, Indonesia, p.427 ,Fig . 4.197. Honolulu. Vol 1. p. 668, PI. 90 (flowering plant Ficus subcordata: drawing (fruiting branch) and part). photograph (habit) provided by the author. Desmodium intortum: Legel, S., 1983.Tropica l for­ Flemingia macrophylla: Westphal, E. & Jansen, age legumes and grasses. Part 1.Legumes , p.25 . P.C.M. (Editors), 1989. Plant Resources of Desmodium triflorum: Matthew, K.M., 1988. Fur­ South-East Asia. A selection. Pudoc, Wage- ther illustrations on the flora of the Tamilnadu ningen: p.129 . Carnatic. The Rapinat Herbarium, Tiruchira- Gliricidia sepium: Westphal, E. & Jansen, P.C.M. palli, India, p. 158, PI. 158 (habit, enlarged twig, (Editors), 1989. Plant Resources of South-East flowers); Ohashi, H., 1973. The Asiatic species Asia. A selection. Pudoc, Wageningen: p.137 . of Desmodium and its allied genera (Legumino- Heteropogon contortus: Bor, N.L., 1960.Th e grasses sae). Ginkgoana 1:231 ,Fig . 67(7) (fruit, seed). of Burma, Ceylon, India and Pakistan. Perga- Desmodium uncinatum: Legel, S., 1983. Tropical mon Press, Oxford, U.K. p.164 ,Fig .6 (habi t flow­ forage legumes and grasses. Part 1. Legumes, p. ering plant); Häfliger, E. & Scholz, H., 1980. 29. Grass weeds. Documenta, Ciba-Geigy, Basel. 274 FORAGES

Vol. 1. p. 61 (ligule); Skerman, P.J. & Riveros, Leucaena leucocephala: Westphal, E. & Jansen, F., 1990.Tropica l grasses. FAO, United Nations, P.C.M. (Editors), 1989. Plant Resources of Rome, Italy, p.447 ,Fig . 15.78 (two pairs of heter- South-East Asia. A selection. Pudoc, Wage­ ogamous spikelets). ningen, p.173 . Hymenachne acutigluma: Soerjani, M., Koster- Lotononis bainesii: Verdcourt, B., 1979. A manual mans, A.J.G.H., Tjitrosoepomo, G. (Editors), of New Guinea legumes. Botany Bulletin No11 . 1987.Weed s of rice in Indonesia. Balai Pustaka, Office ofForests ,Divisio n ofBotany , Lae, Papua Jakarta, Indonesia, p. 433,Fig . 4.200. New Guinea, p. 586,Fig . 148. Imperata cylindrical Soerjani, M., Kostermans, Macroptilium atropurpureum: drawing provided A.J.G.H., Tjitrosoepomo, G. (Editors), 1987. by the Department ofFiel d Crops and Grassland Weeds of rice in Indonesia. Balai Pustaka, Ja­ Science, Wageningen Agricultural University. karta, Indonesia, p.435 ,Fig . 4.201(habit , seeds); Macroptilium lathyroides: Verdcourt, B., 1979. A Häfliger, E. & Scholz, H., 1980. Grass weeds. manual of New Guinea legumes. Botany Bulle­ Documenta, Ciba-Geigy, Basel. Vol. 1. p. 63 tin No 11. Office of Forests, Division of Botany, (ligule, inflorescence, spikelet). Lae, Papua New Guinea, p. 506,Fig . 126. Ischaemum ciliare: Chien-Chang Hsu, 1978.Grami - Macroptilium longepedunculatum: drawing pro­ neae. In: Hui-lin Li et al. (Editors): Flora of vided by the Department of Field Crops and Taiwan. Epoch Publishing Company, Taipei. Grassland Science, Wageningen Agricultural Vol. 5. p. 669, PI. 1472 (habit flowering plant); University. Gilliland, H.B., Holttum, R.E., Bor, N.L., 1971. Macrotyloma axillare: Verdcourt, B., 1979.A man­ Grasses of Malaya. In: Burkill, H.M. (Editor): ual of New Guinea legumes. Botany Bulletin No Flora of Malaya. Government Printing Office, 11. Office of Forests, Division of Botany, Lae, Singapore. Vol. 3. Opposite p. 263, PL 20 (habit Papua New Guinea, p. 531,Fig . 129. stoloniferous part); Häfliger, E. & Scholz, H., Medicago sativa: Verdcourt, B., 1979. A manual of 1980. Grass weeds. Documenta, Ciba-Geigy, New Guinea legumes. Botany Bulletin No 11. Basel. Vol. 1.p . 64 (ligule, spikelet). Office ofForests ,Divisio n ofBotany , Lae, Papua Ischaemum magnum: Wycherley, P.R. & Ahmad New Guinea, p. 566,Fig .143 . Azli, M.Y., 1974. Grasses of Malayan planta­ Microstegium ciliatum: Gilliland, H.B., Holttum, tions, p. 40,PI . 8. R.E., Bor, N.L., 1971.Grasse s of Malaya. In: Bur- Ischaemum muticum: Gilliland, H.B., Holttum, kill, H.M. (Editor): Flora of Malaya. Govern­ R.E., Bor, N.L., 1971.Grasse s of Malaya. In: Bur- ment Printing Office, Singapore. Vol. 3. p. 248, kill, H.M. (Editor): Flora of Malaya. Govern­ Fig. 54. ment Printing Office, Singapore. Vol. 3. p. 256, Mikania cordata: Holm, L.G., Plucknett, D.L., Pan- Fig. 56 (leafy branch); Chien-Chang Hsu, 1974. cho, J.V. & Herberger, J.P., 1977. The world's Taiwan grasses. Taiwan Provincial Education worst weeds:distributio n and biology. The East- Association, Taipei, Taiwan, p. 716, Fig. 253 West Center, University of Hawaii Press, Hono­ (flowering branch); Häfliger, E. & Scholz, H., lulu, Hawaii, United States, p. 321,Fig .133 . 1980. Grass weeds. Documenta, Ciba-Geigy, Mimosa pudica: Soerjani, M., Kostermans, Basel. Vol. 1.p . 65 (ligule, pair of spikelets). A.J.G.H., Tjitrosoepomo, G. (Editors), 1987. Ischaemum rugosum: Soerjani, M., Kostermans, Weeds of rice in Indonesia. Balai Pustaka, Ja­ A.J.G.H., Tjitrosoepomo, G. (Editors), 1987. karta, Indonesia, p. 331,Fig . 4.150. Weeds of rice in Indonesia. Balai Pustaka, Ja­ Neonotonia wightii: Legel, S., 1983.Tropica l forage karta, Indonesia, p. 441, Fig. 4.204. legumes and grasses. Part 1. Legumes, p. 35 Ischaemum timorense: Soerjani, M., Kostermans, (flowering and fruiting plant); Milne-Redhead, A.J.G.H. ,Tjitrosoepomo , G.(Editors) , 1987.Weed s E. &Polhill , R.M. (Editors), 1971.Flor a of Trop­ ofric e in Indonesia. Balai Pustaka, Jakarta, Indo­ ical East Africa - Leguminosae 4 - Papilionoi- nesia, p. 443,Fig . 4.205 (flowering plant part, pair deae 2. Crown Agents for Oversea Governments of spikelets, caryopsis in two views); Häfliger, E. and Administrations, p. 531, Fig. 79 (flower, &Scholz , H.,1980 . Grass weeds.Documenta , Ciba- fruit, seeds). Geigy, Basel. Vol. 1. p.6 7(ligule) . Ottochloa nodosa: Chien-Chang Hsu, 1974. Taiwan Leptochloa chinensis: Häfliger, E. & Scholz, H., grasses. Taiwan Provincial Education Associa­ 1981. Grass weeds. Documenta, Ciba-Geigy, tion, Taipei, Taiwan, p. 548, Fig. 168 (flowering Basel. Vol. 2. Aquarelle (without page number) branch, branchlet with two spikelets); Chien- (flowering plants), p. 94(ligule , spikelet). Chang Hsu, 1978. Gramineae. In: Hui-lin Li et- SOURCES OF ILLUSTRATIONS 275

al. (Editors): Flora ofTaiwan . Epoch Publishing Pennisetum polystachion: Soerjani, M., Koster­ Company, Taipei. Vol. 5.p . 567,P L 1434 (ligule). mans, A.J.G.H., Tjitrosoepomo, G. (Editors), Panicum maximum: Bor, N.L., 1960. The grasses 1987. Weeds of rice in Indonesia. Balai Pustaka, of Burma, Ceylon, India and Pakistan. Perga- Jakarta, Indonesia, p. 467, Fig. 4.217. mon Press, Oxford, U.K. p. 328,Fig . 37. Pennisetum purpureum: Westphal, E. & Jansen, Panicum maximum var. trichoglume: Humphreys, P.C.M. (Editors), 1989. Plant Resources of L.R.,1980 .A guid et obette r pastures for the trop­ South-East Asia. A selection. Pudoc, Wage­ ics and sub-tropics. 4th ed. Wright Stephenson ningen, p. 221. &Co. , Ermington, Sydney, p.36 . Puerariaphaseoloides: Botton, H. &Halle , N,1958 . Panicum repens: Holm, L.G., Plucknett, D.L., Pan- Les plantes de couverture. Guide pratique de cho, J.V. & Herberger, J.P., 1977. The world's reconnaissance et d'utilisation des Légumi­ worst weeds:distributio n and biology. The East- neuses en Côte d'Ivoire 2.Journa l d'Agriculture West Center, University of Hawaii Press, Hono­ tropicale et de botanique appliquée 5: 105, Fig. lulu, Hawaii, United States, p. 354, Fig. 148. 28. Paspalum conjugatum: Hitchcock, A.S., 1950. Saccharum spontaneum: Gilliland, H.B., Holttum, Manual of the grasses of the United States. 2nd R.E., Bor, N.L., 1971.Grasse s of Malaya. In: Bur- ed. revised by A. Chase. United States Depart­ kill, H.M. (Editor): Flora of Malaya. Govern­ ment ofAgriculture , Miscellaneous Publication ment Printing Office, Singapore. Vol. 3. p. 225, No 200, Washington, p. 614, Fig. 892 (flowering Fig.4 9(bas e ofplant , flowering culm, culm node, plant); Häfliger, E. & Scholz, H., 1980. Grass panicle, branch with spikelets); Matthew, K.M., weeds. Documenta, Ciba-Geigy, Basel. Vol. 1. p. 1982.Illustration s on the flora of the Tamilnadu 95(ligule , spikelet). Carnatic. The Rapinat Herbarium, Tiruchira- Paspalum dilatatum: Holm, L.G., Plucknett, D.L., palli, India. PI. 928(ligule) . Pancho, J.V. &Herberger , J.P., 1977.Th e world's Sesbania grandiflora: Sachet, M.H., 1987. The lit­ worst weeds: distribution and biology. The East- toral species of Sesbania (Leguminosae) in the West Center, University of Hawaii Press, Hono­ South Pacific islands and its relations. Bulletin lulu, Hawaii, United States, p.359 ,Fig . 151 (plant du Muséum National D'Histoire Naturelle, 4esé ­ habit, ligule); Häfliger, E. & Scholz, H., 1981. rie, section B,Adansonia . Vol. 9.p . 11, PL 3. Grass weeds.Documenta , Ciba-Geigy, Basel. Vol. Sesbania sesban: Havard-Duclos, B., 1967. Les 2.p .9 6(inflorescence , spikelet). plantes fourragères tropicales. Techniques Paspalum distichum: Soerjani, M., Kostermans, Agricoles et Productions Tropicales, Collection A.J.G.H., Tjitrosoepomo, G. (Editors), 1987. dirigée par René Coste, 10. G.-P. Maisonneuve Weeds of rice in Indonesia. Balai Pustaka, Ja­ & Larose, Paris, p. 185, Fig. 70 (flowering karta, Indonesia, p.461 , Fig. 4.214. branch); Busson, F., 1965. Plantes alimentaires Paspalum notatum: Häfliger, E. &Scholz , H., 1980. de l'Ouest africain. Ministère de la Coopération Grass weeds. Documenta, Ciba-Geigy, Basel. et al., Paris, p. 244, Fig. 95(fruit , seeds). Vol. 1. Aquarelle (without numbered page) Setaria sphacelata: drawing provided by the au­ (flowering plant); p. 102(ligule , spikelet). thor (leafy plant part); Häfliger, E. &Scholz , H., Paspalumplicatulum: Skerman, P.J. &Riveros, F., 1980. Grass weeds. Documenta, Ciba-Geigy, 1990. Tropical grasses. FAO, United Nations, Basel. Vol. 1. p. 131 (ligule, spikelet, bristle); Rome, Italy, p. 580, Fig. 15.121 (habit); Häfliger, Skerman, P.J. & Riveros, F., 1990. Tropical E. & Scholz, H., 1980. Grass weeds. Documenta, grasses. FAO, United Nations, Rome, Italy, p. Ciba-Geigy, Basel. Vol. 1.p . 104(ligule , inflores­ 663. Fig. 15.140 (inflorescence). cence, spikelets). Sorghum x almum: Skerman, P.J. & Riveros, F., Paspalum scrobiculatum: Soerjani, M., Koster­ 1990. Tropical grasses. FAO, United Nations, mans, A.J.G.H., Tjitrosoepomo, G. (Editors), Rome, Italy, p.671 ,Fig . 15.142. 1987. Weeds of rice in Indonesia. Balai Pustaka, Sorghum, artificial perennial hybrids: drawing Jakarta, Indonesia, p.457 ,Fig . 4.212. provided by the Department of Field Crops and Pennisetum clandestinum: Skerman, P.J. & Rive- Grassland Science, Wageningen Agricultural ros, F., 1990. Tropical grasses. FAO, United University. Nations, Rome, Italy, p. 605, Fig. 15.126 (habit); Sorghum x drummondii: Skerman, P.J. &Riveros , Häfliger, E. & Scholz, H„ 1980. Grass weeds. F., 1990.Tropica l grasses. FAO, United Nations, Documenta, Ciba-Geigy, Basel. Vol. 1. p. 112 Rome, Italy, p. 691, Fig. 15.146; inflorescence: (ligule, inflorescence, spikelet). diapositive ofth e editors. 276 FORAGES

Stenotaphrum secundatum; Skerman, P.J. &River - (Editor): Flora of Malaya. Government Printing os, F., 1990. Tropical grasses. FAO, United Office, Singapore. Vol. 3.p .46 , Fig. 1(spikelet) . Nations, Rome, Italy, p. 712, Fig. 15.152 (flower­ Trifolium repens: Zohary, M. & Heller, D., 1984. ing plant, portion of spike); Häfliger, E. & The genus Trifolium. The Israel Academy of Scholz, H., 1980.Gras s weeds. Documenta, Ciba- Sciences and Humanities, p. 169,PI . 52. Geigy, Basel. Vol. 1.p . 137(ligule , spikelets). Trifolium semipilosum: Zohary, M. & Heller, D., Stylosanthes capitata: original drawing by P. Ver- 1984. The genus Trifolium. The Israel Academy heij-Hayes , Prosea, Wageningen, from Irwin et of Sciences and Humanities, p. 178,PI . 54. al. 14908 (WAG) (habit); Mohlenbrock, R.H., Tripsacum andersonii: Wycherley, P.R. & Ahmad 1963. Further considerations in Stylosanthes Azli, N.Y., 1974. Grasses of Malayan planta­ (Leguminosae). Rhodora 65: 250,Fig . 1 (fruit). tions, (plant habit); Matthew, K.M., 1982. Illus­ Stylosanthes guianensis: Westphal, E. & Jansen, trations on the flora of the Tamilnadu Carnatic. P.C.M. (Editors), 1989. Plant Resources of The Rapinat Herbarium, Tiruchirapalli, India, South-East Asia. A selection. Pudoc, Wage­ p. 952,PI .95 2(ligule , inflorescence); Häfliger, E. ningen, p.25 6(flowerin g branch); Mohlenbrock, & Scholz, H., 1980. Grass weeds. Documenta, R.H., 1963. Further considerations in Stylo­ Ciba-Geigy, Basel. Vol. 1.p . 141 (spikelets). santhes (Leguminosae). Rhodora 65:254 , Fig. 19 Urochloa mosambicensis: Chippendall, L.K.A., (fruit). 1955. A guide to the identification of grasses in Stylosanthes hamata: original drawing by P. Ver- South Africa. In: Meredith, D. (Editor): The heij-Hayes, Prosea, Wageningen, from Burt s.n. grasses and pastures of South Africa. Part 1. (habit); Mohlenbrock, R.H., 1963.Furthe r consi­ Central News Agency, p. 383, Fig. 327 (leafy derations in Stylosanthes (Leguminosae). Rho­ plant part, inflorescence); Tothill, J.C. & dora 65: 254,Fig . 16 (fruit). Hacker, J.B., 1983. The grasses of southern Stylosanthes humilis: Verdcourt, B.,1979 .A manu­ Queensland. University of Queensland Press, al of New Guinea legumes. Botany Bulletin No Brisbane, Australia, p. 426 (spikelets). 11. Office of Forests, Division of Botany, Lae, Vigna parkeri: drawing provided by the Depart­ Papua New Guinea, p. 374, Fig. 87 (flowering ment of Field Crops and Grassland Science, branches, flower); Mohlenbrock, R.H., 1963. Wageningen Agricultural University. Further considerations in Stylosanthes (Legu­ Zoysia matrella: Chien-Chang Hsu, 1978. Grami­ minosae). Rhodora 65:257 ,Fig . 30 (fruit). neae. In: Hui-lin Li et al. (Editors): Flora of Stylosanthes macrocephala: original drawing by P. Taiwan. Epoch Publishing Company, Taipei. Verheij-Hayes, Prosea, Wageningen, from Irwin Vol. 5. p. 509, PI. 1414 (flowering plant part); et al. 31823 (WAG) (habit); Sousa Costa, N.M. Chien-Chang Hsu, 1974. Taiwan grasses. & Ferreira, M.B., 1984. Species of Stylosanthes. Taiwan Provincial Education Association, Tai­ In:Stace , H.M. &Edye , L.A. (Editors):Th e biolo­ pei,Taiwan , p.450 ,Fig . 120(ligule , part of inflor­ gy and agronomy of Stylosanthes. Academic escence). Press, Sydney, p. 31, Fig. 3 (fruit). Stylosanthes scabra: original drawing by P. Ver- heij-Hayes, Prosea, Wageningen, from Belem 3886 (WAG) (habit); Mohlenbrock, R.H., 1963. Further considerations in Stylosanthes (Legu­ minosae). Rhodora 65: 251,Fig . 6 (fruit). Themeda triandra: Bor, N.L., 1960. The grasses of Burma, Ceylon, India and Pakistan. Pergamon Press, Oxford, U.K. p. 253,Fig . 27. Thysanolaena latifolia: Chien-Chang Hsu, 1978. Gramineae. In: Hui-lin Li et al. (Editors): Flora of Taiwan. Epoch Publishing Company, Taipei. Vol. 5.p . 396,PI . 1374(flowerin g branch); Chien- Chang Hsu, 1974. Taiwan grasses. Taiwan Pro­ vincial Education Association, Taipei, Taiwan, p. 238, Fig. 13 (ligule, part of inflorescence branch); Gilliland, H.B., Holttum, R.E., Bor, N.L., 1971. Grasses of Malaya. In: Burkill, H.M. Index ofscientifi c plant names

Page numbers printed inbol d refer to main treatment.

Abelmoschus esculentus (L.)Moenc h 245 Alloteropsis ciminica (L.)Stap f 236 Acacia spp.17 3 Alloteropsis semialata (R. Br.)Hitchc .23 6 Acacia arabica (Lamk) Willd.24 5 Alphitonia excelsa Reisseck ex Endlicher 245 Acacia decurrens auct. nonWilld .24 5 Alsophila amboinensis v.A.v.R. 247 Acacia farnesiana (L.)Willd .24 5 Alysicarpus nummularifolius (Willd.) DC.4 2 Acacia mearnsii DeWild .24 5 Alysicarpus ovalifolius (Schum.) Leonard 43 Acacia nilotica (L.)Willd . exDel .24 5 Alysicarpus rugosus (Willd.) DC.23 6 Acacia tomentosa (Roxburgh) Willd. 245 Alysicarpus vaginalis (L.) DC. 29, 42 Acanthus ilicifolius L.24 5 Alysicarpus wallichii Wight &Arnot t 236 Acanthus volubilis Wallich 245 Amaranthus gracilis Desf.24 5 Acroceras crassiapiculatum (Merrill) Alston 236 Amaranthus spinosus L.24 5 Acroceras oryzoides (Swartz) Stapf 236 Amaranthus viridis L.24 5 / Acroceras rigidula Steudel Amorphophallus campanulatus Blume24 5 - var.rigidul a 236 Amorphophallus paeoniifolius (Dennst.) Nicolson Acroceras sparsum Stapf 236 245 Acroceras tonkinense (Belanse) Bor23 6 Amorphophallus variabilis Blume24 5 Acroceras zizanioides (Kunth) Dandy 236 Amphilophis glabra Stapf 237 Adenostemma lavenia (L.)Kuntz e 245 Amphilophis intermedia Stapf 237 Aeschynomene americana L. 29,37 ,65 ,215 , 221 Amphilophis pertusa (L.)Nas h exStap f 54 - cv.'Glenn ' 38, 215, 221 Anastrophus compressus Schlechtendal ex Doell. - cv. 'Lee' 38, 221 53 Aeschynomene apoloana Rusby 39 Andropogon L.1 9 Aeschynomene falcata (Poiret) DC. 29, 39 Andropogon aciculatus Retzius 92 - cv.'Bargoo ' 39, 40 Andropogon amboinicus Merrill 243 Aeschynomene grandiflora (L.) L.19 6 Andropogon annulatus Forssk. 118 Aeschynomene javanica Miquel 37 Andropogon bisquamulatus Höchst.4 4 Aeschynomene sesban L.19 8 Andropogon brevifolius Swartz 242 Aeschynomene spicata Poiret 112 Andropogon caricosus L.23 8 Aeschynomene villosa Poiret 37,3 8 Andropogon contortus L.13 7 Aganope heptaphylla (L.)Polhil l 245 Andropogon fastigiatus Swartz 238 Agati grandiflora (L.)Desv .19 6 Andropogon filipendulus Höchst.23 9 Agrostis infirma Buse 236 Andropogon gayanus Kunth 28,44 , 81, 82,83 , 85, Agrostis matrella L. 234 194,210 , 211 Agrostis maxima Roxb.22 4 - cv.'Kent ' 45 Ailanthus altissima (Miller) Swingle 245 - var.bisquamulatu s (Höchst.) Hack. 44,45 , 46 Ailanthus glandulosa Desf.24 5 - var. tridentatus (Höchst.) Hack. 44 Ailanthus vilmoriniana Dode24 5 Andropogon halepensis Brot. 243 Albizia Durazz. 41,4 2 - var.propinquu s (Kunth) Merrill 243 Albizia falcata sensu Backer 251 Andropogon intermedius R.Br .23 7 Albizia latifolia Boivin 40 Andropogon modestus Backer 237 Albizia lebbeck (L.)Benth . 29,4 0 Andropogon nardus L.24 5 Albizia procera (Roxb.) Benth. 42,24 5 - var. tortilis (Presl) Merrill 245 278 FORAGES

Andropogon parviflorus Backer, non Roxburgh - var. robustior Weizen 236 237 Arthraxon microphyllus Höchst. 236 Andropogon pertusus (L.)Willd . 54 Arthraxon pilipes Backer 236 Andropogon plumosus Backer, non Kunth 243 Arthraxon typicus Koord. 236 Andropogon repens (Roxb.) Steud. 145 Artocarpus altilis (Parkinson) Fosberg 246 Andropogon sanguineus Merrill 242 Artocarpus camansi Blanco 246 Andropogon sorghum Brot. 252 Artocarpus champeden (Loureiro) Stokes 246 Andropogon squamulatus Höchst. 44 Artocarpus communis J.R. &G . Forster 246 Andropogon subtilis Steudel 237 Artocarpus integer (Thunberg) Merrill 246 Andropogon tridentatus Höchst. 44 Arundinaria ciliata A. Camus 50 Andropogon triticeus R. Br. 239 Arundinaria pusilla A. Chevalier &A . Camus 27,5 0 Andropogon wightianus Nees ex Steudel 93 Arundinella Raddi 19 Andropogon zizanioides Urban 253 Arundinella agrostoides Trinius 236 Andropogon zollingeri Steudel 242 Arundinella holcoides (Kunth) Trinius 236 Aneilema nudiflorum R. Br. 250 Arundinella nepalensis Trinius 236 Annona muricata L. 245 Arundinella pumila (Höchst.) Steudel 236 Annona squamosa L. 245 Arundinella setosa Trinius 237 Anthocephalus cadamba (Roxburgh) Miquel 245 Arundinella tenella Nees &Wigh t 236 Anthocephalus chinensis (Lamk) A. Rich, ex Arundo L. 19 Walp. 245 Asthenochloa tenera Buse 237 Anthocephalus indicus Richard 246 Asystasia coromandeliana Nees 51 Antidesma bunius (L.) Sprengel 246 Asystasia gangetica (L.) T. Anderson 30, 51 Antidesma dallachyanum Bâillon 246 Avena fatua L. 237 Antidesma rumphii Tulasne 246 Avena junghuhnii Buse 239 Apium graveolens L. 246 Avena sativa L. 206,23 7 Apluda mutica L. 236 Avena sterilis L. 237 Apluda varia Hackel 236 Axonopus Beauv. 19 Arachis L. 47,49 , 50,183 Axonopus affinis A. Chase 47,179, 233 - section Arachis 48 Axonopus compressus (Swartz) P. Beauv. 24, 25, - section Caulorhizae Krap. &Greg. , (nom. nud.) 27,53 49 Azadirachta indica Adr. Jussieu 246 - section Erectoides Krap. &Greg . (nom. nud.) 48 Azolla pinnata R. Br. 246 - section Rhizomatosae Krap. &Greg . (nom. Bambusa bambos Backer 246 nud.) 47 Bauhinia hirsuta Weinmann 237 - series Eurhizomatosae Krap. &Greg . (nom. Beta vulgaris L. 246 nud.) 47 Bidens pilosa L. - series Prorhizomatosae Krap. &Greg . (nom. - var. radiata Schultz-Bip. 246 nud.) 47 Bidens pilosus L. Arachis glabrata Benth. 29,4 6 - var. albus Schultz-Bip. 246 - cv. 'Arb' 47 Boerhavia diffusa L. 246 - cv. 'Arblick' 47 Bombax ceiba L. 246 - cv. 'Arbrook' 47 Bombax insigne Wallich 246 - cv. 'Florigraze' 47,48 Bombax larutense Ridley 246 Arachis hypogaea L. 49,24 6 Bombax malabaricum DC. 246 Arachis pintoi Krap. &Greg. , nom. nud. 29,48 ,59 , Bombax valetonii Hochr. 246 63 Bonnaya ciliata Burkill 240 - cv. 'Amarillo' 49,5 0 Bonnaya serrata Burkill 240 Arachis prostrata Benth. 46 Borassus flabellifer L. 246 Areca borneensis Beccari 246 Bothriochloa bladhii (Retzius) S.T. Blake 237 Areca triandra Roxburgh 246 Bothriochloa decipiens (Hack.) C.E. Hubbard 55 Artabotrys suaveolens Blume 246 Bothriochloa insculpta (A.Rich. ) A. Camus 55 Arthraxon castratus Bor 236 Bothriochloa longifolia (Hack.) Bor 55 Arthraxon hispidus (Thunberg ex Murray) Ma- Bothriochloa modesta (Backer) Backer &Hen - kino 236 rard 237 INDEX OF SCIENTIFIC PLANT NAMES 279

Bothriochloa pertusa (L.)A . Camus 28,5 4 Caesalpinia nuga (L.) W.T. Aiton 246 - cv. 'Dawson' 55 Cajanus cajan (L.) Millsp. 20,29 ,32 ,24 6 - cv. 'Medway' 55 Cajanus indicus Spreng. 246 Bothriochloa panormitana (Pari.) Pilger 55 Calamagrostis australis Buse 237 Bothriochloa radicans (Lehm.) A. Camus 55 Calliandra Benth. 69 Brachiaria spp. 30,46 ,59 ,6 3 Calliandra calothyrsus Meissn. 20,29 ,6 8 Brachiaria arrecta (Th. Dur. &Schinz ) Stent 64 Calliandra confusa Sprague &Rile y 68 Brachiaria bequaertii Robyns 58 Calliandra tetragona Benth. 69 Brachiaria brizantha (A.Eich. ) Stapf 28, 56,58 ,59 , Calopogonium caeruleum (Benth.) Sauv. 29, 71, 66 73, 74,110 - cv. 'La Libertad' 57 Calopogonium mucunoides Desv. 29,71 , 72,86 , - cv. 'Marandü' 57,5 8 110,193 Brachiaria decumbens Stapf 28, 56,57 ,58 ,61 , 66, Canarium sylvestre Gaertner 247 112,145,153,169, 209,21 0 Canavalia Adans. 76 - cv. 'Basilisk' 59 Canavalia ensiformis (L.) DC. 29,7 4 Brachiaria dictyoneura (Fig. &D e Not.) Stapf 28, Canavalia gladiata (Jacq.) DC. 76 50,59,60,62,81,83,210 - var. ensiformis (L.) Benth. 74 - cv. 'Llanero' 60, 61 Canavalia maritima (Aublet) du Petit-Thouars Brachiaria distachya (L.) Stapf 27, 61 247 Brachiaria eruciformis (J.E. Smith) Griseb. 237 Canavalia rosea (Swartz) DC. 247 Brachiaria humidicola (Rendle) Schweick. 28,59 , Canavalia virosa (Roxb.) Wight &Arnot t 76 60, 62, 65,109,110 Canna edulis Ker 247 - cv. 'INIAP-701' 62 Cannabis sativa L. 99,24 7 / - cv. Tully' 62 Capillipedium Stapf 19 Brachiaria keniensis Henr. 60 Capillipedium assimile (Steudel) Camus 237 Brachiaria miliiformis (J. &C .Presl ) Chase 67 Carex baccans Nees 237 Brachiaria mutica (Forssk.) Stapf 28, 64, 97,140 Carex remota L. Brachiaria obvoluta Stapf 60 - ssp. alta (Boott) Kuek. 237 Brachiaria purpurascens (Raddi) Henr. 64 Careya arborea Roxburgh 247 Brachiaria ramosa (L.) Stapf 237 Carthamus tinctorius L. 247 Brachiaria reptans (L.) C.Gardne r &CE . Hub­ Carum carvi L. 247 bard 237 Cassia spp. 88 Brachiaria ruziziensis Germain &Evrar d 28,50 , Cassia bauhiniifolia Kunth 89 65 Cassia bifoliolata DC. ex Colladon 89 - cv. 'Kennedy' 66 Cassia fabaginifolia Kunth 89 Brachiaria subquadripara (Trin.) Hitchc. 28,6 7 Cassia mimosoides L. 247 Brachiaria villosa (Lamk)A . Camus 237 Cassia monophylla Vellozo 89 Brachypodium sylvaticum P. Beauv. 237 Cassia rotundifolia Persoon 88 Brassica integrifolia (West) Rupr. 246 Cassia siamea Lamk 247 Brassica juncea (L.) Czernova 246 Ceiba pentandra (L.) Gaertner 247 Brassica rugosa Prain 246 Cenehrus ciliaris L. 28,77 ,85,131,13 6 Bridelia retusa (L.) Sprengel 246 - cv. 'American' 78 Briza minor L. 237 - cv. 'Biloela' 78 Bromus catharticus Vahl 237 - cv. 'Boorarra' 78 Bromus insignis Buse 237 - cv. 'Gayndah' 78 Bromus unioloides Kunth 237 - cv. 'Higgins grass' 78 Brugmansia Candida (Pers.) Safford 246 - cv. 'Lawes' 78 Brugmansia suaveolens Humb. &Bonpl . 246 - cv. 'Molopo' 78 Bruguiera conjugata Merrill 246 - cv. 'Nunbank' 78 Bruguiera gymnorhiza (L.) Savigny 246 - cv. 'Tarewinnebar' 78 Butea monosperma (Lamk) Taubert 246 - cv. 'West Australian' 78 Butea superba Roxburgh 246 Cenehrus inflexus R. Br. 237 Caesalpinia crista L. 246 Cenehruslappaceus L. 79 Caesalpinia digyna Rottler 246 Centotheca lappacea (L.) Desv. 79 280 FORAGES

Centotheca latifolia (Osbeck) Trinius 27,7 9 Citrullus vulgaris Schrader ex Ecklon & Zeyher Centrosema spp.45 ,81 , 83 247 Centrosema acutifolium Benth. 29,8 0 Citrus limon (L.) Burm.f. 247 - cv. 'Vichada' 81 Claoxylon indicum (Reinw. ex Blume) Hassk. 247 - var. matogrossense nom. nud. 81 Claoxylon polot Merrill 247 - var. orinocense nom. nud. 81 Cleome gynandra L. 247 Centrosema grandiflorum Benth. 83 Clitoria mearnsii De Wild. 94 Centrosema lisboae Ducke 82 Clitoria schiedeana Schlecht. 87 Centrosema macrocarpum Benth. 29,8 2 Clitoria tanganicensis Micheli 94 Centrosema molle Martius ex Benth. 86 Clitoria ternatea L. 29,9 4 Centrosema pascuorum Martius ex Benth. 29,8 4 - cv. 'Milgarra' 95 - cv. 'Bundey' 85 Clitoria zanzibarensis Vatke 94 - cv. 'Cavalcade' 85 Cocos nucifera L. 247 Centrosema pubescens Benth. 29,57 ,59 ,60 ,63 , 65, Codariocalyx gyroides (Roxb. ex Link) Hassk. 29, 66, 67,71 , 72,73 ,74 ,80 , 81, 82,83 ,86 , 97,109,131, 97 145,162,170,175,190,192,193, 202, 213 Codariocalyx motorius (Houtt.) Ohashi 98 - cv. 'Belalto' 87 Coelachne infirma Buse 237 Centrosema schiedeanum (comb, ined.) 87 Coelachne pulchella auct., non R. Br. 237 Ceratonia siliqua L. 237 Coelorrhachis glandulosa Stapf 240 Chamaecrista rotundifolia (Persoon) Greene 29, Coix L. 19 88 Coix lachryma-jobi L. 247 - cv. 'Wynn' 88,8 9 Colocasia esculenta (L.) Schott 247 - var. grandiflora (Bentham) Irwin &Barneb y 89 Combretum quadrangulare Kurz 247 - var. rotundifolia 89 Commelina benghalensis L. 247 Chamaeraphis aspera Nees 242 Commelina diffusa Burm.f. 237 Chloris Swartz 19 Commelina nudiflora L. 237 Chloris abyssinica A.Richar d 90 Corchorus olitorius L. 247 Chloris barbata auct., non Swartz 237 Cordia dichotoma Forster f. 247 Chloris digitata Steudel 238 Cordia obliqua Willd. 247 Chloris divaricata R. Br. 237 Coriandrum sativum L. 247 Chloris gayana Kunth 28,74 ,78 ,90,153,175,17 9 Coridochloa cimicina Nees 236 - cv. 'Callide' 90 Coridochloa semialata Nees 236 - cv. 'Karpedo' 91 Corypha elata Roxburgh 247 - cv. 'Katambora' 90,9 1 Corypha utan Lamk 247 - cv. 'Masaba' 91 Crotalaria L. 99 - cv. 'Nzoia' 91 Crotalaria alata Buch.-Ham. & Roxburgh ex D. - cv. 'Pioneer' 91 Don 247 - cv. 'Pokot' 91 Crotalaria anagyroides Kunth 247 - cv. 'Samford' 91 Crotalaria ferruginea Graham ex Benth. 247 Chloris glabrata Anderss. 90 Crotalaria juncea L. 29,9 8 Chloris inflata Link 237 - cv. 'Kanpur 14' 100 Chloris pumilio R. Br. 237 - cv. 'T6' 100 Chloris tenera Scribner 239 - cv. 'Tropic Sun' 100 Chloris truncata R. Br. 237 Crotalaria usaramoensis Baker f. 247 Chloris virgata Swartz 237 Crotalaria zanzibarica Benth. 247 Chrysopogon Trin. 19 Cucurbita pepo L. 247 Chrysopogon aciculatus (Retzius) Trinius 27,9 2 Cudrania javanensis Trécul 250 Chrysopogon orientalis (Desv.) A. Camus 27,9 3 Cudrania pubescens Trécul 250 Chrysopogon subtilis (Steudel) Miquel 237 Cuscuta australis R. Br. 167 Chrysopogon wightianus (Nees ex Steudel) Cuscuta chinensis Lamk 167 Thwaites 93 Cyamopsis psoralioides DC. 247 Cicer arietinum L. 247 Cyamopsis tetragonoloba (L.) Taubert 247 Citrullus lanatus (Thunberg) Matsum. & Nakai Cyanotis cristata (L.) D. Don 247 - ssp.vulgari s (Schrader) Fursa 247 Cyathea amboinensis (v.A.v.R.) Merrill 247 INDEX OF SCIENTIFIC PLANT NAMES 281

Cymbopogon Sprengel 19 - var. virgatus 106 Cymbopogon citratus Stapf 245 Desmodium Desv. 107 Cynodon spp. 183 Desmodium aparines (Link) DC. 114 Cynodon dactylon (L.) Pers. 28,47, 100,102 Desmodium canum Schinz &Thellun g 112 - cv. 'Brazos' 101 Desmodium gyroides (Eoxb. ex Link) DC. 97 - cv. 'Coastal' 101 Desmodium heterocarpon (L.) DC. 29,106,110, 111 - cv. 'Coastcross-1' 101 - cv. 'Florida' 106,108 - cv. 'Hardie' 101 - ssp. angustifolium Ohashi 107 - cv. 'Oklan' 101 - ssp.heterocarpo n 107 - cv. 'Tifton 68' 101 - ssp.heterocarpo n var. ovalifolium (Wallich ex - cv. 'Tifton 78' 101 Prain) Eugayah 73,108 - var. dactylon 101 - ssp. ovalifolium (Prain) Ohashi 29,57 ,59 ,60 , 63, Cynodon glabratus Steudel 100 72,107,108,145,190,194 Cynodon lemfuensis 102 - cv. 'Itabela' 109 Cynodon nlemfuensis Vanderyst 28,102 Desmodium heterophyllum (Willd.) DC. 19,29 ,59 , - cv.'Costa Eica'103 63, 72,110,190 - cv. 'Florico' 103 - cv. 'Johnstone' 111 - cv. 'Florona' 103 Desmodium incanum DC. 29,112 - cv. 'Ona' 103 Desmodium intortum (Miller) Urban 29, 66,113, - var. nlemfuensis 102 114,117,175,184, 233 - var. robustus Clayton &Harla n 102 - cv. 'Greenleaf' 115 Cynodon polevansii Stent 100 - cv. 'Tengeru' 115 Cynosurus aegyptius L. 104 Desmodium ovalifolium Wallich ex Merrill 108, Cyperus brevifolius (Eottb.) Hassk. 237 109 Cyperus castaneus Willd. 237 Desmodium papuanum C.T. White 97 Cyperus compressus L. 237 Desmodium parvifolium Blanco 116 Cyperus cyperoides (L.) O. Kuntze 247 Desmodium polycarpum DC. 108 Cyperus elatus L. 247 - var. ovalifolium Wallich ex Prain 108 Cyperus haspan L. 238 Desmodium pulchellum (L.) Benth. 251 Cyperus iria L. 238 Desmodium purpureum (Miller) Fawc. & Eendie Cyperus kyllingia Endl. 238 248 Cyperus pilosus Vahl 248 Desmodium reticulatum Champ, ex Benth. 107 Cyperus rotundus L. 248 Desmodium sandwicense E. Meyer 115 Cyrtococcum Stapf 19 Desmodium tortuosum (Swartz) DC. 248 Cyrtococcum accrescens (Trinius) Stapf 248 Desmodium triflorum (L.) DC. 19,29, 116 Cyrtococcum oxyphyllum (Steudel) Stapf 248 Desmodium uncinatum (Jacq.) DC. 29,113,114, Cyrtococcum trigonum (Eetzius) Camus 248 116,175 Cytisus cajan L. 246 - cv. 'Silverleaf 117,118 Cytisus palmensis (Christ) Hutch. 238 Dichanthium annulatum (Forssk.) Stapf 27,118 Dactylis glomerata L. 238 Dichanthium caricosum (L.) A. Camus 119,23 8 Dactyloctenium Willd. 19 Dichanthium mucronulatum Jansen 119 Dactyloctenium aegyptium (L.)Willd . 27,104 Diectomis fastigiata (Swartz) Kunth 238 Dactyloctenium verticillatum Backer 238 Digitaria Haller 19,122 Danthonia pilosa E. Br. 242 Digitaria abludens (Eoemer &Schultes ) Veldk. 238 Datura suaveolens Humb. &Bonpl . 246 Digitaria adscendens Henrard 119 Derris heptaphylla Merrill 245 Digitaria ciliaris (Eetzius) Koeler 28,119 Desmanthus virgatus (L.) Willd. 29,105 Digitaria decumbens Stent 121 - cv. 'Bayamo' 106 Digitaria didactyla Willd. 248 - cv. 'Mare' 106 Digitaria dilatata (Poiret) Coste 178 - cv. 'Yuman' 106 Digitaria distachya (L.) Pers. 61 - var. acuminatus (Benth.) Isely 106 Digitaria eminens (Steudel) Backer 238 - var. depressus (Humb. &Bonpl . ex Willd.) B. Digitaria endlichii Mez 123 Turner 106 Digitaria eriantha Steudel 28,47 , 74, 97,111,112, - var. glandulosus Turner 106 121,124,153 282 FORAGES

- ssp. eriantha sensu P.D.F. Kok 121,122 Echinochloa crus-galli (L.) P. Beauv. 27,125,126 - cv. 'Advance' 122 - ssp. colona (L.) Honda 125 - cv. 'Premier' 121,122 Echinochloa frumentacea (Roxburgh) Link 248 - ssp.pentzi i (Stent) P.D.F. Kok 121,122 Echinochloa stagnina (Retzius) Beauv. 238 - cv. 'Slenderstem' 122 Ectrosia leporina R. Br. 238 - cv. 'Taiwan' 122 Eichhornia crassipes (Martius) Solms 248 • - cv. 'Transvala' 122 Eichhornia speciosa Kunth 248 - ssp. stolonifera (Stapf) P.D.F. Kok 121 Elaeis guineensis N.J. Jacquin 248 - ssp.transvaalensi s P.D.F. Kok 121 Elephantopus scaber L. 248 Digitaria junghuhniana (Steudel) Henrard 238 Eleusine Gaertner 19,104 Digitaria longiflora (Retzius) Pers. 238 Eleusine aegyptia (L.)Desf . 104 Digitaria marginata Link 119 Eleusine coracana Gaertner 248 Digitaria milanjiana (Rendle) Stapf 28,122,123 Eleusine indica (L.) Gaertner - cv. 'Jarra' 124 - f. coracana (L.) Hook.f. ex Back 248 - cv. 'Mardi' 123,124 Enhalus acoroides (L.f.) L.C. Richard ex Steudel Digitaria mombasana CE. Hubbard 123 248 Digitaria pedicellaris Prain 238 Enterolobium saman (Jacq.) Prain 252 Digitaria pentzii Stent 121 Enteropogon dolichostachyus (Lagasca) Keng 238 - var. stolonifera (Stapf) Henrard 121 Epipremnum pinnatum (L.) Engl. 238 Digitaria perrottetii Backer 238 Eragrostis v. Wolf. 19 Digitaria pertenuis Buse 238 Eragrostis amabilis (L.) Hook. &Arnot t 127 Digitaria polevansii Stent Eragrostis atrovirens (Desf.) Trinius ex Steudel - ssp.peteran a Henrard 123 238 Digitaria ropalotricha Buse 238 Eragrostis bahiensis Schultes 238 Digitaria sanguinalis auct., non Scop. 119 Eragrostis brownii Nees ex Hook. &Arnot t 238 Digitaria setivalva Stent 123,124 Eragrostis cilianensis (All.) Link ex Vign. 238 Digitaria smutsii Stent 121 Eragrostis elegantula Steudel 238 Digitaria swynnertonii Rendle 123 Eragrostis elongata (Willd.) Jacq. 238 Digitaria ternata (A.Richard ) Stapf 238 Eragrostis interrupta (R. Br.) Beauv. 238 Digitaria valida Stent 121 Eragrostis japonica (Thunberg ex Murray) Tri­ Digitaria violascens Link 238 nius 238 Dimeria ornithopoda Trinius 248 Eragrostis megastachya (Koeler) Link 238 Dinebra arabica Jacq. 238 Eragrostis nigra Nees ex Steudel 238 Dinebra retroflexa (Vahl)Panz . 238 Eragrostis pilosa (L.) Beauv. 238 (L.) Beauv. ex Roemer & Schultes Eragrostis tef (Zuccagni) Trotter 248 240 Eragrostis tenella (L.) P. Beauv. ex Roemer & Diplachne polystachya (Forsskal) Backer 240 Schultes 27,127 Diplachne tectoneticola Backer 240 Eragrostis unioloides (Retzius) Nees ex Steudel Dipterocarpus spp. 51 27,128 Dolichos axillaris E. Meyer 161 Eremochloa ciliaris (L.) Merrill 238 Dolichos biflorus auct. muit., non L. 250 Eremochloa horneri Buse 238 Dolichos ensiformis L. 74 Eremochloa malayana Ridley 238 Dolichos lablab L. 249 Eriachne obtusa R. Br. 238 Dolichos maranguënsis Taub. 232 Eriachne pallescens R. Br. 238 Dolichos phaseoloides Roxb. 192 Eriachne tincta Nees 238 Dolichos sinensis L. 253 Eriachne triseta Nees ex Steudel 239 Dolichos trilobatus L. 244 Erianthus arundinaceus (Retzius) Jeswiet 248 Dolichos unguiculatus L. 253 Eriochloa decumbens F.M. Bailey 239 Dolichos uniflorus Lamk 250 Eriochloa nubica (Steudel) Hackel &Stap f ex Drymaria cordata (L.)Willd . ex Roemer & Thell. 239 Schultes 238 Eriochloa procera (Retzius) CE. Hubbard 239 Drymaria hirsuta Bartling 238 Eriochloa ramosa (Retzius) O. Kuntze 239 Echinochloa Beauv. 19 Eriochloa subglabra (Nash) Hitchc. 239 Echinochloa colona (L.) Link 27,125 Erythrina lithosperma Miquel 248 INDEX OF SCIENTIFIC PLANT NAMES 283

Erythrina subumbrans (Hassk.) Merrill 248 Glycine javanica auct. mult., non L. 169 Erythropsis colorata (Roxburgh) Burkill 248 Glycine max (L.)Merril l 207,24 9 Eucarya acuminata (R. Br.) Sprague & Summer- Glycyne wightii (Wight &Arnott ) Verde. 169 hayes 252 Gossampinus heptaphylla Bakh. 246 Euchlaena mexicana Schrader 239 Gossypium arboreum L. Eulalia fimbriata (Hackel) 0. Kuntze 239 - var. obtusifolium (Roxburgh) Roberty 249 Eulalia fulva (R. Br.) 0. Kuntze 239 Gossypium hirsutum L. 249 Eulalia praemorsa (Nees) Hackel 242 Gossypium nangking Meyen 249 Eulalia quadrinervis (Hackel) 0. Kuntze 239 Gossypium obtusifolium Roxburgh 249 Eulalia trispicata (Schultes) Henrard 239 Guazuma ulmifolia Lamk Eupatorium cordatum Burm.f. 166 - var. tomentosa K. Schumann 249 Euphoria didyma Blanco 250 Guizotia abyssinica (L.f.) Cass. 249 Eustachys tenera (Presl) Camus 239 Gynandropsis gynandra (L.) Merrill 247 Fagopyrum esculentum Moench 248 Hackechloa granularis (L.) O. Kuntze 239 Festuca leptopogon Stapf 239 Hedysarum diffusum Vellozo 39 Festuca nubigena Jungh. 239 Hedysarum falcatum Poiret 39 Ficus spp. 131 Hedysarum gyroides Roxb. ex Link 97 Ficus alba Reinw. ex Blume 248 Hedysarum hamatum L. 213 Ficus altissima Blume 248 Hedysarum heterocarpon L. 106 Ficus benghalensis L. 248 Hedysarum heterophyllum Willd. 110 Ficus calophylloides Elmer 129 Hedysarum intortum Miller 114 Ficus drupacea Thunb. 130 Hedysarum racemosum Aublet 112 Ficus elastica Roxb. ex Hörnern. 130 Hedysarum rugosum Willd. 236 Ficus fairchildii Backer 129 Hedysarum triflorum L. 116 Ficus garciniifolia Miquel 129 Hedysarum uncinatum Jacq. 116 ' Ficus globosa Blume 239 Hedysarum vaginale L. 42 Ficus grossularioides Burm.f. 248 Helianthus annuus L. 249 Ficus hirta Vahl 239 Helictotrichon virescens (Steudel) Henrard 239 Ficus hispida L. 248 Hemarthria vaginata Buse 239 Ficus indica L. 248 Hemigymnia fusca Ridley 171 Ficus padana Burm.f. 248 Hemigymnia multinodis Stapf 171 Ficus religiosa L. 248 HeteropogonPers. 19 Ficus stricta Miquel 130 Heteropogon contortus (L.)P . Beauv. ex Roemer Ficus subcordata Blume 30,129,136 &Schulte s 27,137, 222,22 3 - var. malayana Corner 130 Heteropogon triticeus (R. Br.) Stapf 239 - var. subcordata 130 Hevea brasiliensis (Willd. exA .Jussieu ) Muell. Ficus superba (Miquel) Miquel 248 Arg. 249 Ficus toxicaria L. 248 Hibiscus cannabinus L. 249 Ficus variegata Blume 248 Hibiscus esculentus L. 245 Fimbristylis annua (All.) Roemer &Schulte s 239 Hibiscus sabdariffa L. 249 Fimbristylis dichotoma (L.) Vahl 248 Hibiscus tiliaceus L. 131,136, 249 Fimbristylis diphylla Vahl 248 Hierochloe horsfieldii (Kunth) Maxim. 239 Fimbristylis miliacea (L.) Vahl 239 Holcus lanatus L. 239 Firmiana colorata (Roxburgh) R. Br. 248 Holcus latifolius Osbeck 79 Flemingia congesta Roxb. exAit . f. 131 Holcus pertusus L. 54 Flemingia latifolia Benth. 131 Homalomena alba Hassk. 249 Flemingia macrophylla (Willd.)Merr . 20,29, 131 Homalomena aromatica (Roxburgh) Schott 239 Galinsoga parviflora Cav. 248 Homalomena cordata Schott 239 Garnotia stricta Brongn. 239 Homalomena javanica Alderw. 249 Garuga abilo Merrill 248 Homalomena philippinensis Engl. 239 Garuga floribunda Decaisne 248 Hoplismenus burmani P. Beauv. 241 Gliricidia maculata (Kunth) Kunth ex Walp. 133,135 Hoplismenus compositus P. Beauv. 241 Gliricidia sepium (Jacq.) Kunth ex Walp. 20,29 , Hoplismenus undulatifolius P. Beauv. 241 70,131,133 Hydnocarpus castanea Hook.f. &Thomso n 249 284 FORAGES

Hydnocarpus heterophylla Blume 249 Ischaemum aristatum L. 143 Hydnocarpus kurzii (King) Warb. 249 Ischaemum arundinaceum F.v.Mueller ex Benth. Hydnocarpus polypetala (Slooten) Sleumer 249 240 Hydrilla verticillata (L.f.) Royle 239 Ischaemum ciliare Retzius 27,142 Hygrorhiza aristata (Retzius) Nees exWigh t & Ischaemum colladoa Sprengel 146 Arnott 249 Ischaemum digitatum Brongn. 240 Hymenachne acutigluma (Steudel) Gilliland 27, Ischaemum glabratum Presl 145 139 Ischaemum indicum (Houtt.) Merrill 143 - cv. 'Olive' 140 Ischaemum indicum auct. (non (Houtt.) Merrill Hymenachne amplexicaulis (Rudge) Nees 140, 239 142 Hymenachne amplexicaulis sensu Backer 139 Ischaemum laeve Ridley 144 Hymenachne aurita (Nees) Backer 239 Ischaemum laxum R. Br. 240 Hymenachne indica Buse 242 Ischaemum macrurum Stapf ex Ridley 148 Hymenachne interrupta (Willd.) Buse 239 Ischaemum magnum Rendle 28,144 Hymenachne myurus Beauv. 239 Ischaemum muticum L. 28,145 Hymenachne myurus sensu Burkill 139 Ischaemum repens Roxb. 145 Hymenachne pseudointerrupta C. Mueller 139 Ischaemum rugosum Salisb. 28,146 Hymenodictyon excelsum Wallich 249 - var. rugosum 147 Hymenodictyon orixense (Roxburgh) Mabberly - var. segetum Hackel 147 249 Ischaemum segetum Trinius 146 Hymenodictyon timoriense Klotsch ex Walp. 249 Ischaemum timorense Kunth 28,148 Hyparrhenia Andersson 19 Justicia gangetica L. 51 Hyparrhenia filipendula (Höchst.) Stapf 239 Justicia procumbens L. 249 Ichnanthus pallens (Swartz) Munro 239 Koordersiochloa javanica Merrill 243 Ichnanthus vicinus (P.M. Bailey) Merrill 239 Kydia calycina Roxburgh 249 Imperata Cyr. 19 Kyllinga brevifolia Rottb. 237 Imperata arundinacea Cyr. 140 Kyllinga monocephala Rottb. 238 Imperata conferta (Presl) Ohwi 249 Lablab niger Medikus 249 Imperata contracta Hitchc. 249 Lablab purpureus (L.) Sweet 20,29 ,32 ,96 ,206 ,24 9 Imperata cylindrica (L.)Raeusche l 18,24 ,25 , 27, Lactuca indica L. 249 51, 71,98,132,134,137,140,145,165,178, 210,23 3 Lagerstroemia indica L. 249 - var. major (Nees) C.E. Hubbard 140 Lagurus cylindricus L. 140 Imperata exaltata Brongn. 249 Lannea coromandelica (Houtt.) Merrill 131,136, Indigofera spp. 19 249 Indigofera cordifolia K. Heyne ex Roth 239 Lannea grandis (Dennst.) Engl. 249 Indigofera glandulosa Wendl. 240 Lantana aculeata L. 250 Indigofera hendecaphylla Jacq. 249 Lantana camara L. 249 Indigofera spicata Forsskal Leersia hexandra Swartz 240 - var. spicata 249 Lemna minor L. 240 Inocarpus edulis J.R. &G . Forster 249 Leptaspis banksii R. Br. 240 Inocarpus fagiferus (Parkinson) Fosberg 249 Leptaspis cochleata Thwaites 240 Ipomoea aquatica Forsskal 249 Leptaspis urceolata R. Br. 243 Ipomoea batatas (L.)Lam k 20, 249 Leptochloa chinensis (L.) Nees 28,149 Ipomoea pes-caprae (L.) Sweet 249 Leptochloa filiformis (Lamk) P. Beauv. 150,24 0 Ipomoea reptans Poiret 249 Leptochloa malabarica (L.)Veldk . 240 Isachne albens Trinius 240 Leptochloa panicea (Retzius) Ohwi 150 Isachne beneckei Hackel 240 Leptochloa polystachya (R. Br.) Benth. 240 Isachne globosa (Thunberg ex Murray) O. Kuntze Leptochloa polystachya Benth. sensu Burkill 150 240 Leptochloa tectoneticola (Backer) Jansen 240 Isachne miliacea Roth exRoeme r &Schulte s 240 Lepturus repens (Forster) R. Br. 240 Isachne myosotis Nees 240 Lespedeza sieboldi Miquel 240 Isachne pangerangensis Zollinger &Moritz i 240 Lespedeza thunbergii (DC.) Nakai 240 Isachne rigida Nees ex Miquel 240 Leucaena spp. 153,154 Ischaemum aristatum auct., non L. 142 Leucaena diversifolia (Schlecht.) Benth. 152,154 INDEX OF SCIENTIFIC PLANT NAMES 285

Leucaena glauca (Willd.) Benth. 150 - cv. 'Maldonado' 160 Leucaena latisiliqua (L.) Gillis 150 Macroptilium purpureum 79 Leucaena leucocephala (Lamk) de Wit 20,29 ,34 , Macrotyloma axillare (E. Meyer) Verde. 30,161 53, 57,59 ,60 ,65 ,66 ,70 , 97,124,131,132,133,135, - cv. 'Archer' 161,162 142,150,173,175,192, 229 - var. axillare 161 - cv. 'Cunningham' 152 - var. glabrum (E. Meyer) Verde. 161,162 - cv. 'K8' 152 - var. macranthum 161 - cv. 'K29' 152 - var. macranthum (Brenan) Verde. 161 - cv. 'K67' 152 Macrotyloma uniflorum (Lamk) Verde. 30,32 , 33, - cv. 'K156' 152 163, 250 - cv. 'K636' 152,154 Mangifera indica L. 250 - cv. 'KX1'152,153,15 4 Manihot esculenta Crantz 20,30 ,32 ,33 ,25 0 - cv. 'KX2' 152,153,154 Manihot utilissima Pohl 250 - cv. 'KX3' 152,153,154 Manisuris granularis L.f. 239 - var. glabrata Rose 152 Mapania kurzii Clarke 240 - var. leucocephala 152 Mariscus sieberianus Nees ex Clarke 247 Leucaena pallida Britton &Ros e 154 Medicago spp. 175,20 6 Leucas lavandulifolia J.E. Smith 250 Medicago falcata L. 164 Leucas linifolia (Roth) Sprengel 250 Medicago glomerata Balbis 164 Ligustrum lucidum Aiton 240 Medicago glutinosa M. v. Bieb. 164 Limnanthemum spp. 251 Medicago sativa L. 30,91,136, 163,175, 206 Limnocharis flava (L.) Buchenau 250 - cv. 'Trifecta' 164 Lindernia ciliata (Colsm.) Pennell 240 Meibomia heterophylla (Willd.) Kuntze 110 Litchi chinensis Sonn. 250 Meibomia triflora (L.) Kuntze 116 / Litchi philippinensis Radlk. 250 Meibomia uncinata (Jacq.) Kuntze 116 Litsea chinensis Lamk 250 Melastoma decemfidum Roxburgh 250 Litsea garciae Vidal 250 Melastoma malabathricum L. 250 Litsea glutinosa (Loureiro) C.B. Robinson 250 - f. polyanthum L. 250 Litsea sebifera (Blume) Blume 250 Melastoma polyanthum Blume 250 Lolium italicum A. Braun 240 Melastoma sanguineum Sims 250 Lolium multiflorum Lamk 240 Melia indica Brandis 246 Lolium perenne L. 240 Melica latifolia Roxb. ex Hörnern. 224 Lolium temulentum L. 240 Melinis minutiflora Beauv. 74,24 0 Lophatherum gracile Brongn. 240 Merremia hederacea (Burm.f.) H. Hallier 250 Lotononis bainesii Baker 29, 91,154 Metroxylon rumphii (Willdenow) Martius 250 - cv. 'Miles' 154,155 Metroxylon sagu Rottboel 250 Lycopersicon esculentum Miller 250 Metroxylon squarrosum Beccari 250 Macaranga gigantea (H.G. Reichenbach &Zoll ­ Microchloa indica (L.f.) Beauv. 240 inger) Muell. Arg. 250 Microlaena stipoides (Labill.) R. Br. 240 Macaranga hosei King ex Hook.f. 240 Microstegium ciliatum (Trinius) A. Camus 28,165 Macaranga incisa Gage 250 Microstegium montanum (Nees) Henrard 165 Maclura cochinchinensis (Loureiro) Corner 250 Microstegium rufispicum (Steud.) A. Camus 165 Macroptilium atropurpureum (DC.) Urban 29,72 , Mikania cordata (Burm.f.) B.L. Robinson 30,166 78, 91, 97,138,145,155,160,170,175,184,190, 209 Mikania micrantha Kunth 167 - cv. 'Siratro' 155 Mikania scandens (L.)Willd . 166,167 Macroptilium domingense Urban 160 Mikania scandens Hook.f. 166 Macroptilium gracile (Poepp.) Urban 160 Mikania volubilis (Vahl) Willd. 166 Macroptilium lathyroides (L.) Urban 29,65 , 91, Milium compressum Swartz 53 158 Mimosa invisa Martius ex Colla 19,25 0 - cv. 'Murray' 158 Mimosa lebbeck L. 40 - var. lathyroides 158 Mimosa pigra L. 51 - var. semierectum (L.)Urba n 158 Mimosa pudica L. 19,24 ,25 ,30, 167 Macroptilium longepedunculatum (Benth.) - var. hispida Brenan 168 Urban 30,159 - var. pudica 168 286 FORAGES

- var. tetrandra (Humb. &Bonpl . ex Willd.) DC. Nopalea coccinellifera (Miller) Salm-Dyck 241 167,168 Notonia wightii Wight &Arnot t 169 - var. unijuga (Duchass. &Walp. )Griseb . 167,168 Nymphoides spp. 251 Mimosa sirissa Eoxb. 40 Oldenlandia corymbosa L. 251 Mimosa tetrandra Humb. &Bonpl . ex Willd. 167 Ophiurus exaltatus (L.) O. Kuntze 241 Mimosa unijuga Duchass. &Walp . 167 Oplismenus burmanni (Retzius) Beauv.241 ' Mimosa virgata L. 105 Oplismenus compositus (L.) Beauv. 241 Miscanthus japonicus Andersson 250 Oplismenus undulatifolius (Ard.) Beauv. 241 Mnesithea glandulosa (Trinius) Koning &Sose f Opuntia cochenillifera Miller 241 240 Opuntia elatior Miller 251 Mnesithea laevis (Retzius) Kunth 240 Opuntia monacantha Haw. 241 Mnesithea mollicoma (Hance) Camus 240 Opuntia nigricans Haw. 251 Mnesithea pubescens Ridley 240 Oryza fatua Koenig ex Trinius 241 Mnesithea rottboellioides (R. Br.) Koning &Sose f Oryza granulata Nees &Arnot t ex Steudel 241 241 Oryza latifolia Desv. 241 Moghania macrophylla (Willd.) Kuntze 131 Oryza meyeriana (Zollinger &Moritzi ) Bâillon Mollugo pentaphylla L. 250 241 Monerma repens Beauv. 240 Oryza minuta Presl 241 Monochoria hastata (L.)Solm s 250 Oryza ridleyi Hook.f. 241 Monochoria vaginalis (Burm.f.) Presl 250 Oryza sativa L. 28,32 ,33 ,25 1 Moringa oleifera Lamk 250 Ottochloa Dandy 19 Morus alba L. 241 Ottochloa nodosa (Kunth) Dandy 28,171 Mucuna aterrima (Piper &Tracy ) Merrill 250 Pandanus affinis Kurz 251 Mucuna cochinchinensis (Loureiro) A. Chev. 250 Pandanus aurantiacus Ridley 251 Mucuna deeringiana (Bort.) Merrill 250 Pandanus helicopus Kurz 251 Mucuna pruriens (L.)DC . Pandanus johorensis Martelli 251 - cv. group Utilis 250 Panicum spp. 136,170 Muehlenbergia huegelii Trinius 241 Panicum acutigluma Steudel 139 Murdannia nudiflora (L.) Brenan 250 Panicum ambiguum Trinius 241 Musa x paradisiaca L. (Musa AAB group) 250 Panicum auritum Presl 239 Musa paradisiaca L. Panicum barbatum Lamk 243 - var. sapientum Kuntze 250 Panicum brevifolium L. 241 Musa sapientum L. Panicum brizanthum Höchst, ex A. Rich. 56 - var. paradisiaca Baker 251 Panicum caudiglume Hackel 241 Myriostachya wightiana (Nees) Hook.f. 241 Panicum chamaeraphoides Hackel 243 Najas falciculata A. Braun 241 Panicum coccospermum Steudel 237 Najas indica (Willd.) Chamisso 241 Panicum colonum L. 125 Najas minor Allioni 241 Panicum convolutum P. Beauv. ex Sprengel 176 Neolamarckia cadamba (Roxburgh) Bosser 246 Panicum crassiapiculatum Merrill 236 Neonotonia wightii (Wight &Arnott ) Lackey 30, Panicum crus-galli L. 126 169,173,175, 202, 206 Panicum dactylon L. 100 - cv. 'Clarence' 170 Panicum dictyoneurum Fig. &D e Not. 60 - cv. 'Cooper' 170 Panicum distachyon L. 61 - cv. 'Malawi' 170 Panicum eruciforme J.E. Smith 237 - cv. 'Tinaroo' 170,175 Panicum flavidum Retzius 241 - ssp.petitian a (A.Rich. ) Lackey 170 Panicum geminatum Forsskal 241 - ssp.pseudojavanic a (Taubert) Lackey 170 Panicum humidicola Rendle 62 - ssp.wighti i 170 Panicum humile Nees ex Steudel 241 Nephelium litchi Cambess. 250 Panicum lutescens Weigel 243 Nertera depressa Banks &Soland . 241 Panicum luzonense Presl 241 Neyraudia madagascariensis Hook.f. Panicum malabaricum (L.)Merril l 240 - var. zollingeri Hook.f. 251 Panicum maximum Jacq. 20,28 ,30 ,70 , 81, 83, 86, Neyraudia reynaudiana (Kunth) Keng ex Hitchc. 87,112,131,153,172,174,194, 213 251 - cv. 'Gatton' 173 INDEX OF SCIENTIFIC PLANT NAMES 287

- cv. 'Hamil' 173 - cv. 'Charu' 179 - cv. 'Makueni' 172,173 - cv. 'Natsugumo' 179 - cv. 'Natsukaze' 174 - cv. 'Raki' 179 - cv. 'Riversdale' 70 - var. dilatatum 179 - cv. 'Vencedor' 174 - var. pauciciliatum Parodi 179 - var. trichoglume Eobijns 28,78 , 91,136,162,173, Paspalum distichum L. 28,180 174 Paspalum horneri Henrard 185 - cv. 'Pétrie' 174 Paspalum lamprocaryon K. Schum. 185 Panicum miliaceum L. 251 Paspalum notatum Flueggé 28,47, 181 Panicum miliare Lamk 241 - cv. 'Pensacola' 182 Panicum miliiforme J. &C .Pres l 67 - cv. 'Tifthi-1' 182 Panicum montanum Roxburgh 241 - var. latiflorum Doell. 182 Panicum mosambicense Hack. 230 - var. notatum 182 Panicum mucronatum Roth ex Roemer & Schultes - var. saureae 182 241 - var. saureae Parodi 182 Panicum muticum Forssk. 64 Paspalum orbiculare Forster f. 186, 241 Panicum nodosum Kunth 171 Paspalum ovatum Trinius 178 Panicum notatum Retzius 241 Paspalum paspalodes Scribner 180 Panicum ofyzoides Swartz 236 Paspalum platense Sprengel 178 Panicum palmifolium Koenig 243 Paspalum platicaulon Poiret 53 Panicum paludosum Roxburgh 241 Paspalum plicatulum Michaux 28,183 Panicum pilipes Nees &Arnot t ex Buse 248 - cv. 'Bryan' 184 Panicum plicatulum (Michaux) Kuntze 183 - cv. 'Hartley' 184 Panicum polygamum Swartz 172 - cv. 'Rodds Bay' 184 Panicum polystachion L. 189 Paspalum scrobiculatum L. 28,185 Panicum purpurascens Raddi 64 - cv. 'Paltridge' 186 Panicum ramosum L. 237 - var. auriculatum (Presl) Merrill 185 Panicum repens L. 28,176 - var. bispicatum Hackel 185,186 Panicum reptans L. 237 - var. horneri (Henrard) Koning &Sose f 185,186 Panicum sarmentosum Roxburgh 241 - var. lanceolatum Koning &Sose f 185,186 Panicum sphacelatum Schumacher 201 - var. scrobiculatum 185 Panicum stagninum Retzius 238 Paspalum undulatum Poiret 183 Panicum subquadriparum Trin. 67 Paspalum vaginatum Swartz 181,186, 241 Panicum trichoides Swartz 241 Paspalum zollingeri Steudel 185 Panicum trigonum Retzius 248 Passiflora foetida L. 251 Panicum trypheron Schultes 241 Peltophorum pterocarpum (DC.) Backer ex Heyne Panicum tuberculatum Presl 241 251 Panicum uncinatum Raddi 242 Pennisetum Rich. 19 Panicum verticillatum L. 243 Pennisetum alopecuroides (L.) Sprengel 241 Panicum viride L. 243 Pennisetum americanum (L.) K. Schum. ex Leeke - var. italica L. 252 20, 28,32 , 33,119,192, 251 Panicum walense Mez 241 Pennisetum atrichum Stapf &Hubbar d 189 Paraserianthes falcataria (L.) I. Nielsen 41, 251 Pennisetum cenchroides Rich. 77 Parkinsonia aculeata L. 251 Pennisetum ciliare (L.) Link 77 Paspalidium Stapf 19 Pennisetum clandestinum Höchst, ex Chiov. 28, Paspalidium flavidum (Retzius) Camus 241 187, 227, 233 Paspalidium geminatum (Forsskal) Stapf 241 - cv. 'Breakwell' 188 Paspalidium punctatum (Burm.f.) Camus 241 - cv. 'Crofts' 188 Paspalum L. 19,187 - cv. 'Noonan' 188 Paspalum auriculatum Presl 185 - cv. 'Whittet' 188 Paspalum cartilagineum Presl 185 Pennisetum compressum R. Br. 242 Paspalum commersonii Lamk 185 Pennisetum glaucum R. Br. 251 Paspalum conjugatum Bergius 28,177 Pennisetum macrostachyum (Brongn.) Trinius Paspalum dilatatum Poiret 28,178 251 288 FORAGES

Pennisetum pedicellatum Trin. 190 Poa tenella L. 127 Pennisetum polystachion (L.) Schultes 28,189 Poa trivialis L. 242 - ssp. atrichum (Stapf &Hubbard ) Brunken 190 Poa unioloides Retzius 128 - ssp.polystachio n 190 Pogonatherum crinitum (Thunberg ex Murray) - ssp.setosu m (Swartz) Brunken 190 Trinius ex Kunth 251 Pennisetum purpureum x Pennisetum america- Pogonatherum paniceum (Lamk) Hackel 251 num20 Pollinia argentea (Brongn.) Trinius 239 Pennisetum purpureum Schumach. 19,20 ,21 , 28, Pollinia ciliata Trin. 165 30,190,191 Pollinia clavigera Backer ex Heyne 242 - cv. 'Mott' 192 Pollinia contorta Backer 242 Pennisetum setosum (Swartz) L. Rich. 189 Pollinia dispar Nees ex Steudel 242 Pennisetum subangustum (Schum.) Stapf &Hub ­ Pollinia fimbriata Hackel 239 bard 189 Pollinia fulva (R. Br.) Benth. 239 Pennisetum villosum R. Br. ex Fresen. 251 Pollinia grata Hackel 242 Pentacme siamensis (Miquel) Kurz 252 Pollinia nuda Trinius 242 Perilla frutescens (L.) Britton 251 Pollinia quadrinervis Hackel 239 Perilla ocimoides L. 251 Pollinia spectabilis Trinius 242 Perotis indica (L.) Kuntze 242 Polypogon monspeliensis (L.) Desv. 242 Phalaris arundinacea L. 242 Polytoca bracteata R. Br. 242 Phaseolus spp. 157 Polytoca macrophylla Benth. 242 Phaseolus aconitifolius Jacq. 253 Polytrias amaura (Buse) Kuntze 242 Phaseolus atropurpureus DC. 155 Polytrias indica (Houtt.) Veldk. 143 Phaseolus aureus Roxburgh 253 Pongamia pinnata (L.) Pierre 251 Phaseolus calcaratus Roxburgh 253 Pontederia crassipes Mart. 248 Phaseolus campestris Bentham 159 Portulaca oleracea L. 251 Phaseolus coccineus L. 251 Premna foetida Reinw. ex Blume 251 Phaseolus lathyroides L. 158 Premna integrifolia L. 251 Phaseolus longepedunculatus Bentham 159 Premna serratifolia L. 251 Phaseolus lunatus L. 251 Prosopis juliflora (Swartz) DC. 252 Phaseolus multiflorus Lamk 251 Prosopis spicigera L. 252 Phaseolus radiatus L. 253 Pseuderanthemum racemosum (Roxburgh) Radlk. Phaseolus semierectus L. 158 252 Phaseolus trilobatus (L.) Schreb. 244 Pseudochinolaena polystachya (Kunth) Stapf 242 Phleum indicum Houtt. 143 Pseudodatura arborea van Zyp 246 Phragmites Adanson 19 Pseudodatura suaveolens van Zyp 246 Phragmites australis (Cav.) Steudel 251 Pseudopogonatherum contortum (Brongn.) Phragmites communis Trinius 251 Camus 242 Phragmites karka (Retzius) Trinius ex Steudel Pseudoraphis spinescens (R. Br.) Vickery 242 251 Pseudosorghum zollingeri (Steudel) Camus 242 Phragmites vulgaris (Lamk) Crepin 251 Psoralea leichhardtii F.v . Muell. 240 Phyllodium pulchellum (L.) Desv. 251 Pteridium aquilinum Kuhn 252 Phytelephas macrocarpa Ruiz &Pavo n 251 Pueraria hirsuta (Thunberg) Schneider 252 Pimpinella anisum L. 251 Pueraria javanica (Benth.) Benth. 192 Pisonia excelsa Blume 242 Pueraria lobata (Willd.) Ohwi 252 Pisonia umbellifera (J. &G . Forster) Seemann 242 Pueraria phaseoloides (Roxb.) Benth. 30,45 ,57 , Pistia stratiotes L. 242 59, 60,63 ,65 ,67 ,71 , 73,74 , 86,110,145,166,192 Planchonella longepetiolata (King &Prain ) H.J. - var. javanica (Benth.) Baker 192,193 Lam 242 - var. phaseoloides 193 Planchonella pohlmanniana Burkill 242 Pueraria triloba (Loureiro) Backer 252 Plantago major L. 251 Putranjiva roxburghii Wallich 252 Pluchea lanceolata (DC.) C.B. Clarke 242 Quercus robur L. 252 Poa amabilis L. 127 Rhagodia hastata R. Br. 242 Poa annua L. 242 Rhagodia parabolica R. Br. 242 Poa chinensis L. 149 Rhaphidophora merrillii Engl. 238 INDEX OF SCIENTIFIC PLANT NAMES 289

Khaphis orientalis Desv. 93 Sesbania formosa (F. Mueller) N. Burbridge 197 Richardsonia brasiliensis Gomez 242 Sesbania grandiflora (L.) Poiret 20, 29,196 Ricinus communis L. 252 Sesbania javanica Miquel 252 Rivina humilis L. 252 Sesbania roxburghii Merrill 252 Robinia grandiflora L. 196 Sesbania sesban (L.) Merrill 20, 29,198 Rostellaria procumbens (L.) Nees 249 - ssp.punctat a (DC.) J.B. Gillett 199 Rottboellia exaltata L.f. 242 - ssp.sesba n 199 Rottboellia glandulosa Trinius 240 Sesuvium portulacastrum (L.)L . 252 Rottboellia ophiuroides Benth. 241 Setaria anceps Stapf 201 Rottboellia vaginata (Buse) Backer 239 Setaria barbata (Lamk) Kunth 243 Rytidosperma pilosa (R. Br.) Connor &Edga r 242 Setaria clivalis (Ridley) Veldk. 243 Saccharum L. 19 Setaria geniculata (Lamk) Beauv. 243 Saccharum arundinaceum Retzius 248 Setaria glauca (L.) Beauv. 243 Saccharum officinarum L. 20,28 ,32 ,33 ,25 2 Setaria italica (L.) Beauv. 252 Saccharum ravennae (L.) Murray 252 Setaria palmifolia (Koenig) Stapf 243 Saccharum sinense Roxb. 196 Setaria sphacelata (Schumach.) Stapf & Hubbard Saccharum spontaneum L. 28,195 ex M.B. Moss 28,74,115,117,153,170 , 201 - ssp. aegyptiacum (Willd.) Hack. 195 - cv. 'Splenda' 202, 203 - ssp.spontaneu m 195 - var. sericea (Stapf) W.D. Clayton 201, 202 Saccolepis angusta (Trinius) Stapf 242 - cv. 'Kazungula' 202, 203 Saccolepis indica (L.) A. Chase 242 - cv. 'Nandi' 201, 202, 203 Saccolepis myosuroides (R. Br.) Chase ex Camus - cv. 'Narok' 202,20 3 242 - cv. 'Solander' 202 Sagittaria sagittaefolia L. 252 - var. sphacelata 201 Sagittaria sagittifolia L. 252 - var. splendida (Stapf) W.D. Clayton 201, 202, Samanea saman (Jacq.) Merrill 252 203 Santalum acuminatum (R. Br.) A. DC. 252 Setaria splendida Stapf 201 Sapium discolor (Champ, ex Benth.) Muell.Arg. Setaria verticillata (L.) Beauv. 243 252 Setaria viridis (L.) Beauv. 243 Saraca dives Pierre 242 Shorea cochinchinensis Pierre 252 Schismatoglottis calyptrata (Roxburgh) Zollinger Shorea roxburghii G. Don 252 &Moritzi252 Shorea siamensis Miquel 252 Schismatoglottis latifolia Miquel 242 Sida cordifolia L. 252 Schismatoglottis rupestris Zollinger & Moritzi Sida rhombifolia L. 252 242 Smithia sensitiva Aiton 252 Schizachyrium brevifolium (Swartz) Nees ex Buse Solanum lycopersicum L. 250 242 Sonchus arvensis L. 252 Schizachyrium sanguineum (Retzius) Alston 242 Sonchus oleraceus L. 252 Schizachyrium semiberbe Nees 242 Sonchus wightianus DC. 252 Schleichera oleosa (Loureiro) Oken 252 Sophora tomentosa L. 252 Scirpus erectus Poiret 242 Sorghum x almum Parodi 28, 203,205 ,206 , 207 Sclerachne punctata R. Br. 243 - cv. 'Crooble' 204,205 ,20 6 Scoparia dulcis L. 252 - cv. 'de Soto' 204 Scrotochloa urceolata (Roxburgh) Judziewicz 243 - cv. 'Rietondale' 204 Sechium edule (Jacq.) Swartz 252 Sorghum, artificial perennial hybrids 28,20 5 Semecarpus kurzii Engl. 243 - cv. 'Krish' 205,20 6 Sesamum indicum L. 252 - cv. 'Silk' 204,205 ,20 6 Sesamum orientale L. 252 Sorghum xdrummondi i (Steud.) Millsp. & Chase Sesamum radiatum Schumacher 252 29,119, 205,20 6 Sesban aegyptiacus Poiret 198 - cv.'Bantu '20 7 Sesbania Adanson 200 -cv .'Greenlea f20 7 Sesbania aculeata (Willd.) Pers. 252 -cv .'Lahoma '20 7 Sesbania aegyptiaca Poiret 198 -cv .'Piper '20 7 Sesbania bispinosa (Jacq.) W.F. Wight 252 -cv .'Sucro '20 7 290 FORAGES

- cv. 'Sudax' 207 - var. intermedia (Vogel) Hassler 212, 213 - cv. 'Tift' 207 - cv. 'Oxley' 212 - cv. 'Zulu' 207 Stylosanthes guyanensis 212 Sorghum affine Camus 243 Stylosanthes hamata (L.)Taub . 27,30 ,56 , 78,119, Sorghum arundinaceum (Desv.) Stapf 205, 206 131,136, 213,217 , 220, 221 Sorghum bicolor (L.) Moench 203, 205,206 ,207 , - cv. 'Amiga' 213,215 ,217 ,220 , 221 252 - cv. 'Verano' 50,213 ,215 ,217 ,220 , 221 - ssp. drummondii (Steud.) de Wet 206 Stylosanthes humilis Kunth 30,214 ,21 6 Sorghum halepense (L.) Pers. 203,204 ,205 ,206 , - cv. 'Gordon' 217 243 - cv. 'Khon Kaen' 217 Sorghum nitidum (Vahl) Pers. 243 - cv. 'Lawson' 217 Sorghum plumosum (E. Br.) Beauv. 243 - cv. 'Paterson' 217 Sorghum roxburghii Stapf 205 Stylosanthes macrocephala M.B. Ferreira &N.M . Sorghum sudanense (Piper) Stapf 206, 208,24 3 Sousa Costa 30,21 8 Sorghum vulgare Pers. 252 - cv. 'Pioneiro' 218 Sparganophorus sparganophora (L.) C. Jeffrey Stylosanthes nervosa Macbr. 220 252 Stylosanthes procumbens Swartz 213 Sparganophorus vaillantii Gaertner 253 Stylosanthes scabra Vogel 30,89,131,136 , 215, Spergula arvensis L. 243 217, 219 Sphaeranthus africanus L. 253 - cv. 'Feira' 221 Sporobolus berteroanus Hitchc. &Chas e 253 - cv. 'Fitzroy' 220 Sporobolus diandrus (Retzius) Beauv. 243 - cv. 'Jecuipe' 221 Sporobolus humilus Presl 243 - cv. 'Recife' 221 Sporobolus indicus (L.) R. Br. 253 - cv. 'Seca' 215,217 ,220 ,22 1 Sporobolus pulchellus R. Br. 243 - cv. 'Siran' 215,217 ,220 , 221 Sporobolus tremulus (Willd.) Kunth 243 Stylosanthes suffruticosa Mohlenbrock 220 Sporobolus virginicus (L.) Kunth 243 Stylosanthes tuberculata Blake 220 Stachytarpheta jamaicensis (L.)Vah l 253 Syagrus oleracea (Martius) Beccari 253 Stenolobium caeruleum Benth. 71 Symphytum asperrimum J. Donn 243 Stenotaphrum secundatum (Walter) O. Kuntze 29, Symphytum asperum Lepechin 243 208 Tamarindus indica L. 253 - cv. 'Roselawn' 209 Taraktogenos heterophylla Slooten 249 Stenotaphrum secundum 208 Taraktogenos kurzii King 249 Streblochaete longiaristum (Richard) Pilger 243 Taraktogenos polypetala Slooten 249 Streblus asper (Retzius) Loureiro 253 Tephrosia purpurea (L.) Pers. 253 Stylosanthes spp. 26,34 , 45,136,138,156,175,184, Thelepogon elegans Roth exRoeme r & Schultes 211,213,219,221,231 243 Stylosanthes bracteata Vogel 219 Themeda Forsskal 19 Stylosanthes capitata Vogel 30,209 ,218 ,21 9 Themeda arguens (L.) Hackel 243 - cv. 'Capica' 210, 211 Themeda australis (R. Br.) Stapf 222 Stylosanthes figueroae Mohl. 216 Themeda gigantea (Cav.) Hackel 243 Stylosanthes fruticosa (Retzius) Mohlenbrock 220 Themeda imberbis (Retzius) Cooke 222 Stylosanthes gracilis Kunth 212 Themeda triandra Forssk. 24,28 ,22 2 Stylosanthes guianensis (Aublet) Swartz 20,30 , Themeda villosa Durand &Jackso n 243 57, 59,60 ,66 ,72 ,87 , 91,142,143,145,190, 211, Theobroma cacao L. 253 215, 221 Theobroma pentagona Bernoulli 253 - var. gracilis (Kunth) J. Vogel 212 Thuarea involuta (Forster) R. Br. ex Roemer & - var. guianensis 211, 212 Schultes 243 - cv. 'Cook' 212,215 ,22 1 Thuarea sarmentosa Pers. 243 - cv. 'Endeavour' 212 Thysanolaena latifolia (Roxb. ex Hörnern.) Honda - cv. 'Graham' 212 28, 224 - cv. 'Pi Hua Dou 184' 212 Thysanolaena maxima (Roxb.) Kuntze 224 - cv. 'Pucalpa' 212 Tragus racemosus Aiton ex Hook.f. 253 - cv. 'Schofield' 212 Trema aspera Blume 243 INDEX OF SCIENTIFIC PLANT NAMES 291

Trema orientalis (L.) Blume 253 - ssp.maranguënsi s 232 Tricholaena rosea Nees 243 - ssp.maranguënsi s (Taub.) Verde. 232 Tridax procumbens L. 253 - ssp.parker i 232 Trifolium burchellianum Ser. 228 - var. matrella 234 Trifolium guianense Aublet 211 Vigna radiata (L.)Wilcze k 96,25 3 Trifolium incarnatum L. 243 Vigna sinensis (L.) Hassk. 253 Trifolium nigrescens Viv. 226 Vigna trilobata (L.)Verde . 244 Trifolium occidentale Coombe 226 Vigna umbellata (Thunberg) Ohwi &Ohash i 253 Trifolium repens L. 30,180,183,184, 225,226 ,22 8 Vigna unguiculata (L.)Walp . 20,30 ,32 ,33 ,91 , 207, - cv. 'Grassland Huia' 225 253 - cv. 'Haifa' 225 Wallichia densiflora (Martius) Martius 244 - cv. 'Ladino' 225 Wedelia biflora DC. 253 - cv.'Louisiana SI'225 Wedelia moluccana Boerl. 253 - cv. 'Osceola' 225 Wollastonia biflora (L.) DC. 253 Trifolium semipilosum Fresen. 30,180, 226, 228 Wollastonia moluccana (Blume) DC. 253 - cv. 'Safari' 227, 228 Zea L. 229 - var. glabrescens Gillett 227, 228 Zea mays L. 20,29 ,32 ,33 ,25 3 - var. kilimanjaricum Baker 227 - ssp.may s 229 - var. microphyllum Chiov. 227 - ssp.mexican a (Schrad.) Iltis 229 - var. semipilosum 227 Ziziphus jujuba (L.)Gaertne r (non Miller; often Trigonella foenum-graecum L. 253 cited as Lamk) 253 Tripogon exiguus Buse 243 Ziziphus mauritiana Lamk 253 Tripsacum L. 229 Ziziphus xylopyrus (Retzius) Willd. 253 Tripsacum andersonii J.R. Gray 20,29 ,22 8 Zoysia matrella (L.) Merrill 28,23 4 Tripsacum latifolium Hitchcock 229,24 3 - var. matrella 234 / Tripsacum laxum Nash 229 - var. pacifica Goudswaard 234 Tripsacum maizar Hernandez &Randolp h 229 Zoysia pungens Willd. 234 Triraphis madagascariensis Hook.f. ex Prain 251 Willd. exThiel e 234 Triumfetta bartramia L. 253 Triumfetta rhomboidea Jacq. 253 Turnera ulmifolia L. 253 Typhonium trilobatum (L.) Schott 253 Urena lobata L. 253 Urochloa distachya (L.) Nguyen 61 Urochloa maxima (Jacq.) R. Webster - var. trichoglume (Robijns) R. Webster 174 Urochloa mosambicensis (Hack.) Dandy 29,85 , 131,136,153, 230 - cv. 'Nixon' 231 Urochloa oligotricha (Fig. &D e Not.) Henrard 231 Urochloa paspaloides Presl 241 Urochloa reptans (L.) Stapf 237 Urochloa stolonifera (Goossens) Chippend. 231 Vallisneria spiralis L. 253 Vernonia cinerea (L)Less . 253 Vetiveria zizanoides (L.) Nash 253 Vigna aconitifolia (Jacq.) Maréchal 253 Vigna gracilis sensu auctt., non (Guill. & Perr.) Hook.f. 232 Vigna maranguënsis (Taub.) Harms 232 Vigna parkeri Baker 30,180,183, 232 - cv. 'Shaw' 232, 233 - ssp. acutifolia Verde. 232 Index of vernacular plant names

Page numbers printed in bold refer to main treatment.

acacia courant 105 bagokbok 222 blem bem 139 caker ayam 119 acacia savane 105 bahay193 blembem 61,142 calliandra 68 African foxtail 77 Bahia grass 47,181 blue pea 94 calopo 72, 86,110,193, African stargrass 102 bakbaba180 blukar 166 194 agati sesbania 196 bakbaka100 bo chét150 carabao grass 177 akar entimor 106 balatong 232 bô kêt tây 40 Caribbean stylo 213 akar lupang 166 balete 129 Brazilian lucerne 211 carpon desmodium 106 akar seleguri 43 balog-balog 94 broadleaf carpet grass cassava 20,25 ,33 , 76, alabang-grass 118 balok-balok 133 53 192 alang-alang 140 baludyangan 119 broadleafed setaria 201 cây dau ma 132 alfalfa 163 bamboo grass 50,119 brobos 42 cây dâu ra 75 alyce clover 42 banana 20,132,167, brodjo lego 166 cây diên keo 105 American jointvetc h 192,198 brown seed paspalum cây me dât 43 37,65 bang s'an dam 94 183 cây the the 43 amorseco 92 banig-usa 43,171 brown top paspalum centipede grass 148 amorseko 92 bânkrap168 183 centro 59,65 ,66 ,67 , 71, 'âm'pëu préi 195 banuko 90 buffalo clover 42 73, 80, 86, 87, 97, anchan 94 banyonan tembaga165 buffalo grass 172,177, 109,162,170,193, andudukot aridekdiket bariri magwakat 165 208 194,213 79 bariri-magwakat 171 buffelgrass 77,136 centurion 84 'ang s'an dam 94 barit-baritan 234 bug's egg grass 127 ceroma 166 'ângkiëdèi 196 barnyard millet 126 bujangkagetl68 chaax kh'aa 171 anhdào gi'al33 batad44,205 bukakau 44,205 ,20 6 chamchuri 40 aninapla 40 batag 44, 203,205 ,20 6 bulak-bulak 51 Chanvre indien 98 anjang37 batiki blue grass 142 bulligan 232 chhlong 176 apa apa 131 baya 51 buloh teberau 224 chicken-panic grass apatot133 baylu-patong 79 bulum 140 126 asem-aseman 37 bayokibok 126 bunch speargrass 137 chiendent dactyle 100 Asian pigeon-wings 94 bebontengan 149 bunga biru 94 chiendent gazon 234 asinan 180 bejeng-benjeng 137 bunga-bunga 201 chreh 40 asistasiya 51 bendalaut 176 buntot-pusa 191 co chi104 atro 155 beringan 131 buntut meyong sisir co duôi chó 201 Australian grass 178 berita 172 106 coduó i phung149 awis 224 Bermuda grass 47,100 bunut bingin 130 coro' m 201 awnless barnyard betel vine 198 bunut bulu 130 co san cäp 177 grass 125 bias-biasin 185 bunut lengis 129 co san dâû 181 axillaris 161 bikas 166 bunut lulub 130 co' bông cao ru'ng bâcnhonl39 blabakan 61 burikit 234 khôp 222 baay dâm'nnaëp 106 black pepper 134,151, butterfly pea 86,9 4 co' chân gà 104 babain 168 186,196,198 buybuy 224 c6' chi' 100 babaka-nalabaga 64 black speargrass 137 caeruleum calopo 71 co' cu'a gà 176 badang-badang 75 blady grass 140 cai duoi chon 132 co' duôi voi 191 INDEX OF VERNACULAR PLANT NAMES 293

co' duôi voi nho' 189 dâu dai 193 gogon 140 Indian hemp 98 co' kê to 172 dâu ma 193 gome-gome 61 ipatpatey 222 co' lach 195 dâu ngua 75 grain sorghum 203, 206, ipil-ipil 150 co' long tây 64 dâu ra 75 207 jack bean 74 co' long vu'c 125 dâu rùng 193 grape 101 jajagoanl26 cô' longvu e 126 dâu tây 75 green panic 162,174 jampang jemprak 119 co' mât 61 daua 146 green panic grass 172 jampang pahit 177 co' may dông 93 daun mules 116 green summer grass 61 jangkiri 149 co' móe 79 day trung châu long 86 greenleaf desmodium jant i 198 co' ông 176 delilan 116 66,114, 233 Japanese love-grass co' san dep 178 desmodium 108,194 gros chiendent 208 127 co' san dôi 180 detidid 166 groundnut 49,136 jaringan 185 co' san tru'ng 185 didok-chi 104 Guatemala grass 228 jawa n 126 co' tru'ng 185 dien-dien 198 gudé oyod 43 jayanti 198 co'bông 92 digit grass 121,123 guinea grass 81,83 , jelamparam 119 co'may 92 dilang-butiki 86 112,153,172,174, 213 Johnson grass 203,204 , co'ông 100 dodder 167 guinga 125 205 co'quai chèo 208 dông trung höa tha'o gulong lapas 146 jointvetch 39,6 5 co'tam hung 222 224 gumadep 116 jujuket 139 co'tranh i40 dót 224 habas 75 jukut canklor 165 coast button grass 104 drought grass 145 hahapaan131 jukut domdoman 92 cock's foot 126 durian 52 halanaw 178 jukut inggris 64 cocoa 51, 97,132,134, Dutch clover 225 hah the 110 jukut jampang mang- 151,167,198 East Indian walnut 40 handalawi 191 gung148 coconut 19,48 ,67 ,68 , elena 150 hemp 99 jukut jarem-116 94,135,167,173,177, elephant grass 21,191 herbe à cheval 183 jukut kakawatan100 183,194, 209 enieo 117 herbe d'éléphant 191 jukut kakawatan hijau codarrio 97 fagotier 196 herbe de Guinée 172 234 coffee 97,132,134,151, false tamarind 105 herbe de para 64 jukut karukuan 127 167,198 f a:z langab 168 herbe de Rhode 90 jukut kidang 79 cogongrass 140 feather pennisetum 189 herbe polisson 137 jukut pahit 53 Columbus grass 203 fève Jacques 75 hetero 59, 63,110 jukut randan 146 common finger grass fine-leaved guinea heuheulangan 110 julukan 97 121 grass 174 hila hitu lama 166 jungle rice 125 cong cong 234 finger grass 123 himbis-puyo 116 jute 99 Congo grass 65 foea banga 192 hindi grass 118 ka-du-ai-nu 146 Congo signal grass 65 gagayut180 hnaaz fèèk 222 ka-gyi-the-myet 146 Cordofan pea 94 galeria 163 hnha:z kh'a: 140 ka-thum' théét 150 cori grass 67 galud-galud 102 hnha:z khwa:k 93 kacang asu 72 cotton 101 gamal 133 hnha:z liaz ngu: 213 kacang batang 158 couch grass 100 gamba grass 44, 81,83 , hnha:z ph'è:d 100 kacang kayu betina cowpea 33 210 hnhaaz khachoon189 106 crab grass 119,20 8 gatbo 224 hnhaaz nhung 127,165 kacang mekah 75 creeping beggar weed gau 203 hom sam muang 132 kacang meongan 37 112 gauai-gauai 196 hông mai 133 kacang parang 75 creeping vigna 232 gewawini 132 horse bean 75 kacang parang putih crête-de-coq 126 giant ipil-ipil 150 horse gram 33 75 crowfoot grass 104 giant rhodes grass hung th'ao 44 kacang telang 94 dakayang 125 gigirinling 100 huyêt tha'o lô 55 kacang turi 196 dal grass 139 ginger grass 180 huyo-huyop 106 kaci 106 Dallis grass 178 giting princesa 94 ikug-kuting 189 kadalan 61 damong-alat 234 glagah 195 ilalang 140 kadatuwa 97 damong-balang 104 gliricidia 133 Indian bluegrass 54 kaimi 112 dâu biê'c 94 glycine 169,173 Indian couch grass 54 kaimi clover 112 294 FORAGES kâk'tung 98 koda grass 185 luh-buluhan 127 oats 206 kaliskis-dalag 116 kodo millet 185 luy-a-luy-a 176 oil palm 19,20 , 51, 52, kalo-kawayan 177 kodra millet 185 luya-luyahan 176 53, 54, 71, 72,74,120 , kalopogonium 72 koer 117 luzerne 163 167,171,186, 228 kâm'phé:m 97 kogon 140 luzerne du Brésil 211 one-leaf clover 42 kambengan 140 koko 40 luzerne tropicale 211 orok-orok lembut 98 kampu 40 kolatay 102 Madras hemp 98 padang togu 128 kangaroo grass 222 kolokanting 94 magsaysay-lucerne 216 padi burung 125,126 kapot-kapot 100 kom ka 165 mahae-nok 132 padông mod liinz 110 kara bedog 75 Kordofan pea 94 maiyarep 168 paillotte 140 karaparak 37 Korean grass 234 maiwa peanut 47 paitan 177 karaparapak sara naw koro-korosan 90 maize 20,33 , 76,101, pala belang 185 72 koronivia grass 60,62 , 229 palay-maya 149 karay-kagay 98 109 makahiya 168 palisade grass 56 karetet lempad 79 krandang 192 makahiyang-lalaki 37 palisade signal grass 56 karikut-ritkut 114 krathin 150 malabalatong 132 palo-maria 150 kariskis 150 krus-krusan 104 malakuwerdas 234 pan 196 karukauâyan 149 kuang 79 malit-kalabaw 180 pangola grass 47,74 , kathin 150 kucingan 168 malu-malu 168 97, 111, 121,122,153 katodai 198 kudzu tropika 192 mangkit-parang 106 pangrot 92 katuday 196 kukuwatit 133 mani-mani 106 panic élevé 172 katuray 198 kulape 53,177 mani-manian 43 panic rampant 176 kauat-kauat 177 kumpai batu 185 Manila grass 234 papahitan 53 kaw kua chai 52 kunai 140,150 mara-kawayan 64 para grass 64, 97,140 kawad-kawad 100 kyet-mei 140 marabotum 129 paspalum 178 kawakawayanan 171 lablab 32 maralaya 176 paspalum dilaté 178 kawit-kawitan 77 laclay-guinan 132 marapagay126 passion fruit 151 kembang telang 94 lagaylay 75 marisekos 92 pasto colon 203 kemuchut 92 lagoli 191 Mascarene grass 234 pasto Sudan 206 kemunchup 92 lagtom 126,139 Mauritius grass 64 pattes de poule 126 Kenya clover 226 lalang 140 memalu 168 pazun-sa-myet 125 Kenya white clover 226 lamhani 180 menjalin wuwu 224 pearl millet 33,119 keo dâu 150 lamtoro 150 merakan 137 pega-pega 112,114,117 kerunong padi 176 lamtoro gung 150 merakan lanang 222 Pensacola Bahia grass kh'ê: kha:w 196 langil 40 metubong 176 181 kh'è: nooyz 133 lantebung 224 Milanje finger grass perennial horse gram kh'èèm kh'ôông 224 lao 195 123 161 kh'oonz koong khaaw laolaeng 224 mile-a-minute 166 perennial sweet Sudan 150 lau 195 molasses grass 74 grass 207 khae196 lawa-lawa 178 môm timor 148 petai belalang 150,196 khae-ban 196 layagah 206 môm trui 145 petai cina 150 khae-farang 133 le 50 môm u 146 ph'ek 50 khamin ling 132 lesu-kesu 86 mother of cocoa 133 ph'ông 195 khamin naang 132 leucaena 34,97 , 98,135, mungbean 96 phasey bean 65,158 khauz f angx 205 142,150,173,174 muôn'g la tron 88 phong195 khauz nôk 125 lim xanh 40 mye-sa myet 104 pied 193 khê fàlangx 133 liriksidia 133 mye-sa-myet 100 pigeon pea 32 khikaiyan 166 llanos macro 159 myet-ihi 126 pineapple 52 khonthi din 108 lotononis 154 myet-kha 176 pinto peanut 48, 59,6 3 kikuyu grass 187 love grass 92 myet-le-gra 104 pit-pit 195 kitoke 40 lüay 205 myet-thi 125 plicatulum 183 klamaran 177 lue lac 98 narrow leaf carpet po:th'üan g 98 klandingan 150 lucerne 136,163, 206 grass 47 po-thuang 98 knot grass 180 lucuntu grass 148 napier grass 191 pok kepokan 131 INDEX OF VERNACULAR PLANT NAMES 295

pongola grass 121 rumput embun 55 rumput udang 128 siratro 97,155,170, 209 pötuk 37 rumput gajah 191 rüssèi préïch 50 siris 40 puero 73,86, 192 rumput gerek telinga ruzi grass 65 sisek tenggiling 116 pulang-puwit 125 142 Sabi grass 153, 230 sisik betok 110 puri 198 rumput hantu 51 sabung-sabungan 185 slender guinea grass purple bean 155 rumput italia 180 sagisi 77 174 putok-putukan 98 rumput jagung 228 sai129 slender panic grass 171 putri malu 168 rumput ja jagoa n kecil saka-saka 119 smao anchien 100 quick stick 133 125 sakata 178 smao bek kbol 126 ramput kolam padang rumput jelai 228 salay 55 smao srauv 146 128 rumput kekusa basar salay-parang 55 smau 'âm'bôk 110 ramput kumaranting 126 salohot 92 smau 'ânnchien 185 176 rumput kemarau 145 salsapot 93 smau bânla 168 rebha kartuut 104 rumput kerbau 177 salt-water couch grass smau kaè lolook 116 rebha las-alasan 55 rumput kerbau dera- 180 smau kântraëy 92 rebha luh-buluhan 172 pah 106 sami 198 smau kôô 64 rebha sekem-sekeman rumput kerupet 145 samsamon 222 smau phluk 176 234 rumput ketih belalang sanagori 97 smutgrass 142 rebu bawang 185 185 sano khon 37 snaô kôôk 198 red oat grass 222 rumput kinangan 185 sano-bok 37 so düa 196 rhizoma peanut 46 rumput kuda 172 sano-don 37 soja perene 169 Rhodes grass 74,90 , rumput kumpai 139 santing 72 song chong 126 153,175 rumput kusa-kusa 125 sapao lorn 198 songtha'oto 118 rice 20,25 , 33,100,120, rumput lilit kain 79 saphaolom 198 sorghum 20,75 ,203 ,20 6 125,127,136,146,168, rumput malela 64 saramattagrass 146 sorgo negro 203 171,177,178,179,180, rumput melayu 144 sat thu 133 sour paspalum 177 183,186 rumput mele 146 satintail 140 Spanish clover 110 rokdönao 37 rumput melela 64 sbö':w 140,22 2 speargrass137, 222 round-leafed cassia 88 rumput meiera minyak scrobic 185 splendida 201 roundleaf cassia 88 61 seashore centipede St. Augustine grass 208 rubber 19,20 ,43 , 48, 51, rumput minyak 61,100 grass 145 starfruit 52 53, 71, 72,74 , 94,120, rumput miuyak 104 sekoi 201 stargrass 102 132,149,165,166,167, rumput nyonya 51 selaput tunggul 166 stylo 59,66 ,87,142 , 211 171,173,174,186, 228 rumput padang 142 semanggi 117 s'iiak pied 193 ruiköjing 37 rumput pahit 53 semanggi landa 225 subabul150 rukut dukut 93 rumput pait 171 sensitive 168 sue sat 98 rukut-dukut 102 rumput para 64 sensitive plant 168 Sudan grass 119,20 6 rum'choan94 rumput parit 53 sentro 86 sugar cane 20, 33,101, rumput ala negri 177 rumput pencasilan 181 serengan jantan 131 167,196 rumput apet 148 rumput rawa 171 sesban 196,198 suket balungen 176 rumput australi 178 rumput sarang buaya setaria 74,153,170, 201 suket blembem 146 rumput banggala 172 148,171 sheda grass 118 suket ceker ayam 119 rumput barek sisek pu- rumput siglap 234 shrubby stylo 89,21 9 suket dringoan 104 tih 116 rumput signal 58 sibat-sibatan 137 suket emprit-empritan rumput bebek 125 rumput sisek betok 110 sidda 195 127 rumput benggala 172 rumput sisek naga 110 Siglap grass 234 suket krisik 185 rumput berus kucing rumput telinga tikus 110 signal grass 56,57 ,58 , suket londo 172 189 rumput temaga 79 61,153,169, 209,21 0 suket lorodan 79 rumput buloh 224 rumput tembaga jan- silverleaf desmodium suket pring-pringan rumput bunga putih 51 tan 145 116 171 rumput colok cinia 146 rumput tembaga kasar silverleaf Spanish suket putihan 54 rumput dara 79 144 clover 117 suket resap 145 rumput ekor chawi 146 rumput tulo santadok singkamasaso 193 sukit grinting 100 rumput ekor kucing 189 185 sipadi 129 sukut jareman 110 296 FORAGES sukut kolonjono 64 tomato 166 ya khon 64 ya-phraek 100 summer grass 119 tongkong 224 ya khon moi maemai 79 ya-phungchu 93 sunn hemp 98 top mo' lato 132 ya kom ka 165 ya-plongkhaonok 119 sweet pitted grass 54 torog-torog 39,168 ya kumba 44 ya-plonglaman 126 sword bean 76 toroy196 ya mardi digit 123 ya-rot 90 talahib 195 torpedo grass 176 ya noksichomphu 125 ya-sakatnamkhem 180 tambu 224 Townsville lucerne 216 ya plong 139 ya-sakhorbik 185 tanageb 61 Townsville stylo 216 ya ruzi 65 ya-sata 102 tanglehead 137 trang qu'a ba hoa 116 ya sabee 230 ya-siknaentonnon 58 tanner grass 64 tràng qu'a di diêp 110 ya signal luey 60 ya-sokum 203 tapak jalak 104 trahg qu'a di qu'a 106 ya siknaentontang 56 ya-sudan 206 tarisi 40 trang qu'a xoan 108 ya taiwan 201 ya-surinam 58 tat-awwa 185 trat4 0 ya waen 118 ya-tanhoi 116 tatebu 195 trèfle blanc 225 ya wai 142 ya-tansaill6 tea 134,167, 228 trinh nü' 168 ya-bahia 181 ya-waitham 145 tebalan 43 trôm' préi 110 ya-bupfen 77 ya-yonhu 142,149 tekik 40 tropical finger grass ya-chanakat 176 ya-yung 224 telur ikan 176 119 ya-chaochu 92 yah-tin-nok 119 temuchut 92 tropical kudzu 59, 65, ya-daenlit 178 yam 134 th'üb nhub 168 67, 71,109,166,192 ya-dokkhao 149 yateenka 61 thoua huat 132 truc th'ao 50 ya-enieo 79 yay-tha-kyee 198 three-flowered beggar- tûchihhleo 119 ya-faek 222 yaya 51 weed 116 tucgiéân 90 ya-hangma 55 yethugyi 198 thua clover 225 tugot-manok 104 ya-hep 177 thua desmanthus 105 turi 196 ya-hom 55 thua desmodium 97 turi rawa 39 ya-kha 140 thua karopo 72 turmeric 198 ya-khachonchop 189 thua khaek 75 tuton 125 ya-khaemman 176 thua kleen leap 114 tuwi 196 ya-khaipu 128 thua lisong tao 48 ulam 52 ya-khaonok 126 thua lisongna 43 uoko166 ya-khikhrok 92 thua peelenian soibean utod 50 ya-khikhuyu 187 169 vaay 193 ya-khitroei 92 thua sealulium 71 vanilla 134,196 ya-khuiphaikhon 171 thua sian paa 193 verano stylo 119 ya-kinni 172 thua-anfanfa 163 water couch grass 180 ya-klethoi 116 thua-hamata 213 West-Indian pea 196 ya-kuatemala 228 thua-lai 86 white clover 183, 225, ya-laman 171 thua-lotononit 154 226, 227 ya-lem 137 thua-phi 158 wick grass 139 ya-maengmi 110 thua-satailo211,216 wild pea bean 158 ya-malaysia 53 ya-metnga 149 thua-sentro 86 woolly finger grass 123 ya-nepia 191 thua-sirato 155 wrinkle duck-beak 146 ya-ngap 168 thwax chu:b 98 wrinkled grass 146 ya-nokkhao 125 thwâx fak ph'aaz 75 wunut 129 ya-noksichomphu 146 thwàx h'èè h'üad 132 xâû hô' 168 ya-nomnon 177 tibi 129 xuân tha'o do' 128 ya-nuannoi 234 tiehs 75 xuân tha'o min 127 ya-nuatrusi 137 tigbau 195 ya baimaln 53 ya-onoi 176 tiger grass 224 ya daeng 146 ya-pakkhwai 104 timunan 149 ya harng krarork 127 ya-phek 50 tinitrigo 146 ya hindi 118 ya-phet 50 tinloy 93 ya humidicola 62 ya-phlikhathiulum 183 to-bao 150 ya khao fang 205 The Prosea Foundation (Plant Resources of South-East Asia)

Name, location, legal status, and structure

- Prosea is a Foundation under Indonesian law, with an international charter, domiciled in Bogor. It is an autonomous, non-profit, international agency, governed by a Board of Trustees. It seeks linkages with existing regional and international organizations; - Prosea is an international programme focusing on the documentation of information on plant resources of South-East Asia; - Prosea consists of a Network Office at Bogor (Indonesia) coordinating 6 Country Offices in South-East Asia, and a Publication Office in Wageningen (the Netherlands).

Participating institutions /

- Forest Research Institute of Malaysia (FRIM), Karung Berkunci 201, Jalan FRI Kepong, 52109 Kuala Lumpur, Malaysia; - Indonesian Institute of Sciences (LIPI), Widya Graha, Jalan Gatot Subroto 10,Jakart a 12710, Indonesia; - Institute of Ecology & Biological Resources (IEBR), Nghia Do, Tu Liem, Hanoi, Vietnam; - Papua New Guinea University ofTechnolog y (UNITECH), Private Mail Bag, Lae, Papua New Guinea; - Philippine Council for Agriculture, Forestry and Natural Resources Research &Developmen t (PCARRD), Los Bahos, Laguna, the Philippines; - Thailand Institute of Scientific &Technologica l Research (TISTR), 196 Pha- honyothin Road, Bang Khen, Bangkok 10900, Thailand; - Wageningen Agricultural University (WAU), Costerweg 50, 6701 BH Wage­ ningen, the Netherlands.

Objectives

- to document and make available the existing wealth of information on the plant resources of South-East Asia for education, extension work, research and industry; - to make operational a computerized data bank on the plant resources of South-East Asia; - to publish the results in the form of an illustrated, multi-volume handbook in English; - to promote the dissemination of the information gathered. 298 FORAGES

Target groups

- those professionally concerned with plant resources in South-East Asia and working in education, extension work, research and commercial production (direct users); - those in South-East Asia depending directly on plant resources, obtaining relevant information through extension (indirect users).

Activities

- the establishment and operation of data bases; - the publication of books; - the sponsorship, support and organization of training courses; - research into topics relevant to Prosea's purpose; - the publication and disseminaton of reports and the research results.

Implementation

The programme period has been tentatively divided into 3phases : - preliminary phase (1985-1986):publicatio n of 'Plant Resources of South-East Asia, Proposal for a Handbook' (1986); - preparatory phase (1987-1990): establishing cooperation with South-East Asia through internationalization, documentation, consultation and publica­ tion;reachin g agreement onth e scientific, organizational and financial struc­ ture of Prosea; - implementation phase (1991-1995): compiling, editing and publishing of the handbook; making operational the computerized data bank with the texts and additional information; promoting the dissemination of the information obtained.

Documentation

A documentation system has been developed for information storage and retrie­ val called SAPRIS (South-East Asian Plant Resources Information System). It consists of 6dat a bases: - BASELIST: primarily a checklist of more than 6200plan t species; - CATALOG: references to general literature on plant resources; - PREPHASE: references to literature from South-East Asia; - ORGANYM:reference s to institutions and their research activities; - PERSONYM: references to specialists; - TEXTFILE: the published handbook volumes and additional information.

Publication

The handbook in blue cover (hardbound) is distributed by Pudoc, the low-price edition in green cover (paperback) by Prosea only in the developing countries of South-East Asia and the Pacific, the bibliographies by Prosea and the miscel­ laneous publications by Pudoc. THE PROSEA FOUNDATION 299

The handbook - No 1. Pulses. L.J.G. van der Maesen and Sadikin Somaatmadja (Editors). Pudoc, Wageningen. 1989/ESCAP CGPRT Centre, Bogor. 1990 (out of print)/ Prosea, Bogor. 1992. - No 2. Edible fruits and nuts. E.W.M. Verheij and R.E. Coronel (Editors). Pudoc, Wageningen. 1991/Prosea, Bogor. 1992. - No 3. Dye and tannin-producing plants. R.H.M.J. Lemmens and N. Wulijarni- Soetjipto (Editors). Pudoc, Wageningen. 1991/Prosea, Bogor. 1992. - No 4. Forages. L. 't Mannetje and R.M. Jones (Editors). Pudoc, Wageningen. 1992/Prosea, Bogor. 1992. - No 5 (1). Timber trees. Major commercial timbers. I. Soerianegara and R. H. M. J. Lemmens (Editors), (expected publication date 1993). - No 6. Rattans. J. Dransfield and N. Manokaran (Editors), (expected publica­ tion date 1993). - No 7. Bamboos. S. Dransfield and E. A. Widjaja (Editors), (expected publica­ tion date 1993). - No.8 .Vegetables . J. S.Siemonsm a and K. Piluek (Editors), (expected publica­ tion date 1992/1993).

Bibliographies - Bibliography 1:Pulses . Edition 1.N . Wulijarni-Soetjipto and J.S. Siemonsma (Editors). Prosea, Bogor. 1990. - Bibliography 2: Edible fruits and nuts. Edition 1. N. Wulijarni-Soetjipto and L. S. Anema (Editors). Prosea, Bogor/Pudoc, Wageningen. 1992. - Bibliography 3:Dy e and tannin-producing plants. Edition 1.N . Wulijarni-Soet­ jipto and J.S. Siemonsma (Editors). Prosea, Bogor/Pudoc, Wageningen. 1991.

Miscellaneous - A Selection. E. Westphal and P.CM. Jansen (Editors). Pudoc, Wageningen. 1989. - Basic list of species and commodity grouping. Version 1.R.H.M.J . Lemmens, P.C.M. Jansen, J.S. Siemonsma, F.M. Stavast (Editors). Prosea Project, Wageningen. 1989.(ou t of print). - Basic list of species and commodity grouping. Final version. P.C.M. Jansen, R.H.M.J. Lemmens, L.P.A. Oyen, J.S. Siemonsma, F.M. Stavast and J.L.C.H. van Valkenburg (Editors). Pudoc, Wageningen. 1991. - Proceedings of the First PROSEA International Symposium, May 22-25, 1989, Jakarta, Indonesia. J.S. Siemonsma and N. Wulijarni-Soetjipto (Edi­ tors). Pudoc, Wageningen. 1989.(ou t of print).

In brief, Prosea is

- an international programme, focused on plant resources of South-East Asia; - interdisciplinary, covering the fields of agriculture, forestry, horticulture and botany; - a research programme, making knowledge available for education and exten­ sion; - ecologically focused on promoting plant resources for sustainable tropical land-use systems; 300 FORAGES

- committed to conservation of biodiversity; - committed to rural development through diversification of resources and application of farmers' knowledge.

Prosea Network Office

Centre for Research &Developmen t in Biology Jalan Raya Juanda 22-24 P.O. Box 234 Bogor 16122, Indonesia tel.:(0251)322859 telex: c/o 48304 BPPHBG fax: c/o (62)(251) 324869/326425

Prosea Publication Office

Wageningen Agricultural University P.O. Box 341 6700A H Wageningen, the Netherlands tel.:(08370)84587 telex: 45917 BURLU fax: (31X8370) 82206 CHINA ß----'"r TAIWAN ;W^- Keyo fisland s (/), states (s), regions (r) and provinces (p). BURMA x"! MALAYSIA Northern r3 0 Mo rota i /7 3 East Malaysia r 13-14 Palawan /1 5 Nias i3 9 ') LAOS Johor s1 2 Panay i2 3 North Sulawesi p6 2 Kedah s3 Samar /2 5 North Sumatra p3 8 16 «elantan s6 Southern Tagalog r2 0 Obi / 75 \ Langkawi /'2 Southern Mindanao r3 1 Riau p 40 K /,r' '^-' { , ( Melaka s1 1 Sulu Archipelago /3 5 Riau Archipelago /4 3 i.V' Negeri Sembilan s1 0 Western Mindanao r3 3 Seram / 78 J i >' 18) Pahang s8 Siberut /4 2 x tA Y J Peninsular Malaysia f\° - ' - y ' INDONESIA Simeuluë /3 7 k.uv y THAILAND r (West Malaysia) r1-1 2 Aceh p3 6 Singkep /' 45 Perak s5 Ambon /7 9 South-East Sulawesi p6 5 \ 19 !" „. Perlis s1 Aru Islands /8 2 South Kalimantan p5 9 J »l ' " 21 Pinang s4 Bali /6 7 South Sulawesi p6 4 VIETNAM ( 20 \ '• PHIL) Sabah s1 4 Bangka /4 9 South Sumatra p4 8 Sarawak s1 3 Belitung /5 0 ç •- Sula Islands /7 6 CAMBODIA 22 Selangor s 9 Bengkulu p4 7 Sumba /7 0 „ Terengganu s7 Buru i7 7 Sumbawa /6 9 &' " 25 Butung /6 6 J ' , ',2Y4 ^.v Talaud Islands /6 1 1/ .s i\ j23 PHILIPPINES Central Java p5 3 Tanimbar Islands /8 1 26 Central Kalimantan p5 8 À?J° L ", ' ,2s", > Babuyan Islands /1 6 Timor /7 2 15 £J i i , > Basilan /' 34 Central Sulawesi p6 3 West Daya Islands /' 80 y -•'' 27/^29 ' Bicol r2 1 East Java p5 5 West Java p5 2 Bohol /2 9 East Kalimantan p6 0 West Kalimantan p5 7 / _/ Flores /7 1 i .'. 30 i o Cagayan Valley r1 8 West Sumatra p4 1 WH 3 Cebu i2 8 Halmahera i7 4 Yogyakarta p5 4 ( ' .33.i \ fl l Mindanao Central Mindanao r3 2 Irian Jaya p8 4 ) Jambi p4 6 ^ * X/\ o i_j- \7-—-S ^ Lombok ;6 8 5 Mindoro /' 22 New Britain /8 6 \ 36 \> ! s Madura /5 6 o, \ 61 i' /2 7 Papua r8 5 \9.v i \ H' V/10 ,\.„ ƒ " 13 37^ -.R38 *f/73 2 ; Borneo ', '^N ^ SINGAPORE , « ;r*w 60 ,^.,..> ƒ./' r • /-" • ik '74 39^-» V--JV ^JÄ43^ Je! 61 / V 2?< \ Sumatra p. ^ C 57 CsV« \

Mentöwa/Xv x ^ 6 ,-\.7^\1 Q 4 4 58 % \ { '^^J' Islands '\. kX-- f^ S '"•3 Sulawesi ""-•>-- *l "-^ 76 "~\ Moluccas l x< "% \ \47\ 4848 "V^P Bismarck NN^ a r^( 50 1 i ^ 'j) 84 Archipelago ^s' INDONESIA New Guinea ]. 87 <ÏS Ù/p*i VH; w^->l-^v^ ! s .—v^ - - -N^ J36,,.^/ 56 fi / 85 «^ 52 V---W rx.--r~. vr~_? 82(^» 80 ^ \ 53 v' « Lesser * PAPUÂ NEW GUINEA

Java 5"4-^V,~__ •*•—••^JîCN J /81 ^jQ^k^iskr J'J :*-• ^ ^—-—-• ^ y ^ L~ét \ 67 "68'~~'69 < 71 s T 72 X 70 tit. 89 MAP OF SOUTH-EAST ASIA FOR PROSEA Names ofcountrie s incapita l letters and islands inlowe r case; numbers refer toth e key. C AUSTRALIA f& 1986 fB