Revision of the Genus Pueraria Dc with Some Notes on Teyleria Backer (Leguminosae)
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AGRICULTURAL UNIVERSITY WAGENINGEN PAPERS 85-1 (1985) REVISION OF THE GENUS PUERARIA DC WITH SOME NOTES ON TEYLERIA BACKER (LEGUMINOSAE) L.J .G .VA NDE R MAESEN Genetic Resources Unit ICRISAT P.O. Patancheru 502324 Andhra Pradesh, India CropEvolution Laboratory University of Illinois Urbana,Illinois 61801, U.S.A. Present address: Department of Plant Taxonomy Agricultural University POB8010 6700ED Wageningen, The Netherlands Received: 22-XI-1984 Dateo fpublicatio n 31-V-198 5 (LOn Agricultural University Wageningen The Netherlands 1985 CIP ISBN90-6754-066- 8 ISSN0169-345X © Agricultural University,Wageningen ,th eNetherlands ,1985 . Nopar t ofthi spublication , apart from bibliographic data and brief quotations embodied in critical reviews,ma y be reproduced, recorder or published in any form including print, photocopy, microform, elektronic or elektromagnetic re cord without written permission from the publisher Agricultural University, P.O.Bo x 1901,670 0H BWageningen ,th eNetherlands . Printedi nth eNetherland sb yDrukkeri jVeenma nb.v. ,Wageningen . CONTENTS 1. INTRODUCTION 5 2. HISTORYO FTH EGENU SPUERARI AD C 5 3. RELATIONSHIPSO FPUERARI AI NTH ETRIB EPHASEOLEA E 6 4. MORPHOLOGY OFPUERARI A 7 5. GEOGRAPHICAL DISTRIBUTION ANDECOLOG Y 7 6. BIOSYSTEMATICSO FPUERARI A 8 7. DESCRIPTION OFTH EGENU S 9 8. SECTIONALARRANGEMENT S 11 8.1. SectionPuerari a 12 8.1.1. SubsectionPuerari a 12 8.1.2. SubsectionNonnudiflora eva nde rMaese nsubsect .no v 12 8.1.3. SubsectionPulcherrim ava nde rMaese nsubsect .no v 12 8.2. SectionSchizophyllo nBake r 13 8.3. SectionBreviramula eva nde rMaese nsect ,no v 13 9. KEYT OTH E SPECIESO FPUERARI A 13 10. ALPHABETICAL TREATMENTO FTH ESPECIE S 15 1.Puerari aalopecuroide sCrai b 15 2.Puerari abellaPrai n 18 3. Puerariacalycin aFranche t 20 4. Puerariacandolle iGrah .e xBent h 23 5.PuerariaedulisPam p 27 6.Puerari aimbricat ava nde rMaese nsp .no v 32 7.Puerari alace iCrai b 35 8.Puerarialobat a(Willd. )Ohw i 37 9.Puerariamirific aA .Shaw .&Suva t 63 10.Puerari apeduculari s(Grah .e xBenth. )Bent h 64 11.Puerari aphaseoloide s(Roxb. )Bent h 71 12.Puerari apulcherrim a(Kds)Mer r 88 13.Puerariarigen sCrai b 93 14.Puerari asikkimensi sPrai nCrai b . 96 15.Puerari astrict aKur z 100 16.Puerari atuberos a(Roxb .ex .Willd. )D C 105 17.Puerari awallichi iD C 109 Agric. Univ. Wageningen Papers 85-1 (1985) 3 11. MATERIALO FUNCERTAI N DISPOSITION 112 12. EXCLUDED SPECIESAND REJECTE DNAMES ,WIT H NOTESO NTEYLERI A .... 117 13. ACKNOWLEDGEMENTS 124 SUMMARY 125 RESUME 126 SAMENVATTING 127 REFERENCES 128 INDEXT OPLAN TNAME S 131 Agric. Univ. Wageningen Papers 85-1 (1985) 1. INTRODUCTION The genus Pueraria has not been the subject of a monograph since Bentham's revision of 1867. Since then, Pueraria has served more or less as a receptacle for speciesno t easilyclassifie d elsewhere.Som e speciesof Pueraria are important as fodder, green manure crops, or weeds; some produce edible tubers; some havemedicina l or poisonous properties, but many are rare or only locally abun dant, andpoorl y collected. Becauseo fit sclos eaffinit y to Glycine, LACKEY(1977 , 1981)inspecte d the genus in some detail in the course of his study of the Phaseo- leae. Based on the holdings at Kew, he tentatively reorganized the genus and his infra-generic classification formed a solid base for the present revision. He excluded many species obviously belonging to other genera and simplified the synonymy. The present author prepared this monograph while on sabbatical leave from ICRISAT, India, at the Crop Evolution Laboratory, University of Illinois, Ur- bana, Illinois, USA, upon the suggestion of Drs. J. R. HARLAN and T. HYMO- WITZ. At Urbana the long-standing agricultural research program, is a major factor inth eenormou s increasei n soybean, {Glycinemax (L.) Merr.) production in the USA with the State of Illinois as the largest producer. Since live material of only a few species of Pueraria was available, this study was first tackled as one of classical taxonomy. If and when more species become available,furthe r biosystematic research should help to elucidate the infraspecif- icrelationship s of Pueraria spp. and those between Glycine, Pueraria, and other genera in Glycininae. When introgression of the related genera into Glycine proves possible, another germplasm pool can be created by tapping Pueraria spp. whichmigh tcontribut e valuable genest oth e soybean. Some ethnobotanical aspects are touched upon in the notes, but for a detailed survey I refer to the paper by BODNER & HYMOWITZ. The expertise available at the Department of Plant Taxonomy, Agricultural University, Wageningen, the Netherlands, was an important factor to getting this account published. 2. HISTORY OF THE GENUS PUERARIA DC. DE CANDOLLEestablishe d thegenu sPueraria in 1825fo r two species, Pueraria tuberosa (Roxb. ex Willd.)DC. and P. wallichiiDC , the former is generally ac cepted as the type species (HUTCHINSON, 1964). The genus was named after M. N. PUERARI, a fellow Swissan d friend of DE CANDOLLE, aProfesso r at Copenha gen and a pupil and friend of the botanist VAHL. In 1831,Wallich' s Catalogue ofth eEas t Indian Herbarium listed somespecie snames ,whic h weno w consider asnomina nuda, but which wereuse d by BENTHAMan d others. In 1852, BENTHAM Agric. Univ. WageningenPapers 85-1 (1985) 5 established thegenu sNeustanthus to accommodate Dolichosphaseoloides Roxb. and took in P. wallichiiDC , since he thought the pod of P. tuberosa was articu late(se e§4) .Upo n further investigation (BENTHAM, 1867)o fth e supposed articu lations,th epod s appeared tob econtracte d wherea n ovuleha d failed to develop, whichi sfrequentl y so;therefore , BENTHAMmerge d hisNeusthanthus into Puerar iawit h nine speciesa t that time. Many more names have been proposed by independent workers since Ben- tham's time but most become synonymous when more adequate material repre senting a wider range of diversity is studied. LACKEY (1977, 1981) stated that the genus was in need of revision, since Bentham's work was the most recent treatment of the genus in itsentirety . In 1868 GOMES (MERRILL, 1935) validated a herbarium name ofLoureiro , Zeydora agrestis Lour, ex Gomes, which is actu ally P. montana (Lour.) Merr., in the present revision referred to a variety of P. lobata (Willd.)Ohwi . 3. RELATIONSHIPS OF PUERARIA IN THE TRIBE PHASEOLEAE AUGUSTIN PYRAME DE CANDOLLE placed his Pueraria in the tribu des Lotees, serie Clitoriees, near Glycine (1825). Pueraria tuberosa, earlier sent to him as Hedysarum tuberosum Roxb. ex Willd., did not actually have the articulated pod (§4)o f the Hedysarieae as the casual observer might seeit . BENTHAM (1867) found that incomplete fertilization frequently caused constrictions in the pods, which caused WIGHT and ARNOTT (1834) to revise their earlier opinion of place ment in Clitorieae. They decided that P. tuberosa might have to be reduced to Desmodium, in Hedysarieae. BENTHAM (1865) referred Pueraria to Phaseoleae Benth.,subtrib e Dioclieae,followe d byTAUBER T(1894 )an d HARMS(wh o spelled 'Diocleinae'). LACKEY (1977) revised the Phaseoleae and added Pueraria to the alliance of the Glycininae Benth., now consisting of 16 genera (LACKEY, 1981).Th e Glycine group(Lackey , 1977a)contain sEminia Taub.,Pseudeminia Verde, Pseudovigna Verde, Pueraria DC, Nogra Merr., Sinodolichos Verde, Glycine Willd., and Teramnus P. Br. The Shuteria group contains Diphyllarium Gagnep., Mastersia Benth., Teyleria Backer, Shuteria W. & A., Dumasia DC, Cologania Kunth. and Amphicarpa Nutt. Neonotonia Lackey (= Glycine wightii (W.& A.)Verde), established in 1977, belongs also in the Shuteria group (LACKEY, 1977c). Once described as a species of Glycine, it is obvious that Teyleria Backer seems closer to Glycine than Pueraria. All genera should be studied to establish their place in one of the genepool s of Glycine, in particular those of the Glycine group. Somespecie sof Pueraria ar edifficul t toplace .Anomalou si nth egenus , they fit nowhere elsesatisfactorily . Specieslik e P. stricta Kurz, P. brachycarpa Kurz and P. bellaPrai n arepossibl y referable to Neonotonia, according to LACKEY (1977c). " Agric. Univ. WageningenPapers 85-1 (1985) 4. MORPHOLOGY OF PUERARIA As can be gleaned from the descriptions, Pueraria spp. are strong climbers, rarely shrubs, creeping in or over low vegetation or climbing high in tall trees. Pueraria lobataha s the ability to smother forests, as in the southeast of the UnitedStates ,whil ei tapparentl yi sno ts oaggressiv ei nit snativ ehabitat , China, Indo China, Japan, and NE India. No other speciesi s aggressive, but they are abundant at times. The trifoliolate leaves, flowers and flat to cylindrical long pods are typical ofmos t Phaseoleae. The stipulesma y beattache d at their base or below the middle, the basal lobe may be single, twin-lobed or fringed. The inflorescence provides important diagnostic characters, but often it is crowded and difficult to analyse in the herbarium. There are either 3, rarely 2, or 4 to 7flower spe rmor eo rles sthickene dnod e(brachyblast) .Flowers ar emostl ysub - sessile,o rhav eshor tpedicels ,tende d byver ycaducou s bractsan d have2 smal l persistent bracteolesnea r thecalyx .Th euppe r calyxlobe sar epartl y or entirely connate. Thevexillum base isclawed , auricled or not, but always inflexed near the base,wit h or without callosities. Puerariapulcherrima and P. mirifica have very small flowers, P. lobata about the largest. Flower colour varies from pure white to all shades ofblu e and purple, sometimeswit h a green or yellow patch on thevexillum . Theflower s are occasionally described asfragran t and beauti ful. Indeed, P. lobata was and still is sold in the USA for gardening purposes. Pods are often flat, hairy or glabrous,pal ebrow n to black, septate or aseptate. Theseeds, u pt o2 0pe rpod ,ar eflattened-oblong , seemsmoot hbu tar eminutel y pittedwhe nviewe dwit ha magnificatio n of40 Xo rmore .