WAGENINGEN AGRICULTURAL UNIVERSITY PAPERS 90-3 (1990)

Observations on distribution, ecology and cultivation ofth e tuber-bearing legumegenu sPachyrhizus Rich ,e xDC .

Marten S0rensen Department of Botany Royal Veterinary andAgricultural University Rolighedsvej 23 DK-1958 Frederiksberg C (Copenhagen) Denmark

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Wageningen mm Agricultural University

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Sorensen,M .

Observations on distributyion, ecology and cultivation of the tuber-bearing legumegenu sPachyrhizu s Rich,e xDC ./ M.S0rensen .- Wageningen :Agricultura l University. - 111.- (Wageningen Agricultural Universitypapers . ISSN0169-345 X; 90-3(1990) ) Withref . ISBN90-6754-168- 0 SISO632. 4UD C 633.49NUGI83 5 Subject headings:Pachyrhizu s/ tuber-bearin glegume .

ISBN90-6754-168- 0 NUGI 835

© Agricultural UniversityWageningen ,Th eNetherlands ,1990 .

No part of this publication, apart from abstract, bibliographic and brief quo­ tationsembodie di ncritica lreviews ,ma yb ereproduced , re-corded orpublishe d inan yfor mincludin gprint ,photocopy ,microform , elektronico relektromagne - ticrecor dwithou t written permission from thepublishe r Agricultural Universi­ ty,P.O.Bo x9101 ,670 0H BWageningen ,th eNetherlands .

Printedi nth eNetherland sb yDrukkeri j Veenman B.V.,Wageninge n Abstract The distribution, ecology and cultivation of the genus Pachyrhizus is examined on the basis of herbarium taxonomie studies, field observations, growth experi­ ments under greenhouse condition and study of relevant literature. The great potential of the genus as a high-yielding tuber crop for tropical countries isconfirme d and specific areas in need of further study are indicated. Contents

Introduction 1 Materials and methods 1 General remarks on the genus 2 Notes on the species 3 1. Pachyrhizus erosus(L. ) Urban 3 2. Pachyrhizusferrugineus (Piper) Sorensen 6 3. Pachyrhizuspanamensis Clausen 7 4. Pachyrhizus tuberosus (Lam.) Spreng. 7 5. Pachyrhizus ahipa(Wedd. ) Parodi. 9 Biochemical studies of the genus 10 Pests and diseases 14 Growth experiments 16 Conclusions 25 Acknowledgements 26 Lists ofexsiccata e 27 References 36 Introduction

The increase of the general agricultural interest in and knowledge of tuberous legumes during the past decade is very noticeable. This interest has previously been limited to a few research centres in India, Thailand and the Philippines. All these institutions (with one exception in Mexico) are situated outside the natural distribution area of the genus attributed with the highest production of leguminous tubers on a world basis; i.e. the genus Pachyrhizus - the yam beans. A number of countries in the third world has experienced dramatic changes in soil quality due to the application of intensified agricultural practices. This especially applies to areas whereth eus eo fartificia l fertilizer and chemical herbi­ cides and pesticides has increased in order to maintain the productivity level of high-yielding grain crops in monoculture. Thus, the interest in these poten­ tially very valuable crops has recently spread to several developing countries in the tropics. The genus Pachyrhizus has been the subject of acomprehensiv e biosystematic study carried out at the Department of Crop Science; The Institute of Botany, both at the Royal Veterinary & Agricultural University, Copenhagen and the Botanical Laboratory, University of Copenhagen. The objective ofthi s research project has been the examination of the potential of the genus as a high-yielding tuber crop through artificial hybridization and preliminary yield estimates (based on experiments carried out under greenhouse conditions).

Materials and methods

The examination of ecology and distribution of the different species is partly based on the study of herbarium material obtained from the following herbaria: AAU, ARIZ, BAA, BH, BM, BR, BRI, C, COL, CR, F, G, GB, INPA, K, L, LISU, MA, MICH, MO, NY, P, QCA, S, SI, SP, U, UC, US, VEN, and partly onow n observations obtained onfield trip sthroug h theneotropica l distri­ bution area of the genus. Furthermore, the germplasm collection, obtained through field collections and from various botanical and agricultural institu­ tions, forms the basis for the growth experiments (a detailed description of the materials and methods used in these growth experiments is given on page 16). The data on nutritional composition and chemical constituents have been com­ piled from literature and own findings.

WageningenAgric. Univ.Papers 90-3 (1990) Generalremark s onth egenu s

The 5 species(se eSorense n 1988)ar edistribute d from Central America to North and Central South America, see Fig. 1.I n pre-Columbian times the distribution probably extended further south into Argentina. No collections have been seen from Surinam and French Guyana, but the genus was reported to be cultivated in French Guyana by Bertoni (1910)). Therear esom edifficultie s indeterminin g theexten t ofth enatura l distribution of P. erosus and P. tuberosus, as these two species have been cultivated for a very long period, and have been introduced to a great number of areas. It may, however, be deduced that in isolated areas where the species show little or no morphological variation the occurrence may be the result of a single introduc­ tion. E.g. P. erosus in the Yucatan peninsula, where the history of this crop incultivatio n dates back at least onemillennium , and where there are no records ofundoubtedl y wild material. Undoubtedly wild material of P. erosus is available from Guatemala and Costa Rica, where it was collected by the author in 1985. Its wild origin is con­ firmed by the small size of the seeds, legumes and tubers, and in Costa Rica by the absence of records of P. erosus ever having been cultivated (except for a few recent introductions through agricultural institutions). Clausen (1945) claimed tohav esee non ecollectio n from Costa Rica,marke d ascultivated . How­ ever, this specimen has also been examined by the present author, and from theinformatio n given onth elabe li tbecome seviden t that thisparticula r material was introduced from Mexico for cultivation at the agricultural experiment station atTurrialb a (confirmed byDr .J . Leon, CATIE,Turrialba ,pers .comm.) . The altitudinal range of the genus is from coastal plains to Central Mexican and Andean valleys.P. ahipareache s 2.000m a.s.l. ,P.ferrugineus and P. tubero­ susar e restricted to tropical lowlands, P.panamensis occurs in coastal lowlands, and P. erosusrange s from 0-1.750m a.s.l. in Central and coastal Mexico. The habitat ranges from seasonally dry areas with open bush vegetation to dense humid tropical rain forest. The variation in annual mean precipitation rate isfro m 250-500 mm (in the Yucatan peninsula and coastal regions of Ecua­ dor) to over 1.500m m (Central America and the Amazon region). Cultivation practices vary considerably; on larger scale P. erosus is normally cultivated as a monocrop, on smaller scale P. erosusi s often intercropped with Zea mays, thusprovidin g aroo t crop aswel la sa grai n crop.Thi styp eo f cultiva­ tion alsoha sa beneficial effect on soil fertility because of the symbiotic, nitrogen fixating Rhizobium bacteria. Studies carried out at the Botanical Laboratory, University of Copenhagen, proved that plants (P. erosus) grown with the sym­ biotic bacteria to be superior in yield to plants grown without bacteria, but with artificial nitrogen fertilizer (unpubl. manus.). P. tuberosus is grown in shifting cultivation byth e Indians ofth e Amazon region (Brücher 1977an d Duke 1981). Bertoni (1913) reported that the Indians at the Parana river (Paraguay) used an extract of P. tuberosus leaves as an insecticide. P. ahipa is now grown in

2 WageningenAgric. Univ.Papers 90-3 (1990) monoculture in the high lying valleys of Bolivia and Peru, and was formerly also cultivated in the northern provinces of Jujuy and Salta, Argentina (Parodi 1935, Burkart 1952). In agreement with Pinto Cortés (1973), Zinsou (1987a & b and 1988) and Pauli (1988)i tha sbee nobserve d byth epresen t author that maximal tuber devel­ opment is achieved under short day conditions, i.e. an 11-12hou r photoperiod induces flowering aswel la stube r initiation. Tuber initiation isinduce d 4-6wee k after sowing regardless of length of photoperiod. Albeit, daylengths exceeding 12hour swil lencourag e vegetative growth and reduce tuber development. Addi­ tional increase in tuber development may be obtained by the removal of fertile shoots P. erosus as reported by Arguelles (1979), Noda & Kerr (1983) and Zepeda(1985). The considerable number of chemical investigations of the various parts of the plants (with very few exceptions all the material studied has been obtained from onespecie sonly :P. erosus) ca n conveniently beseparate d intotw o subjects: nutritional and insecticidal, i.e. tubers and seeds/pods respectively. The findings ofth e research published prior to thecompletio n ofth epresen t paper are treated in a later paragraph.

Notes on the species

1.Pachyrhizu serosu s(L. ) Urban Morphology: A herbaceous vine, with great variation in the outline of the leaf­ lets, from dentate to palmate. The species is defined by the lack of hairs on the petals, the number of flowers (4-11) per lateral inflorescence axes and the length of the inflorescence, 8-45 cm. Furthermore, morphological characters of the legumes (pods), qualitative as well as quantitative, are used to separate the species. The size, 6-13 cm x 8-17 mm, the reduction of the strigose hairs at maturity and the colour, pale brown to olive-green/brown are characters spe­ cific to the legume of P. erosus. A number of seed characters are also specific, the colour being olive-green, brown or reddish brown, and the shape is flat, square to rounded. Distribution: Known in the wild state from the following areas:Th e Mexican states of Jalisco, Guanajuato, San Louis Potosi, Michoacan, Morelos, Puebla, Guerrero, Oaxaca, Vera Cruz and Chiapas. Central and Western Guatemala, El Salvador, Western Honduras, Western Nicaragua and Northwestern Costa Rica;(figur e 1.). Cultivated inth e Mexican stateso fNayarit , Guanajuato, Yuca­ tan and Quintana Roo, where P. erosus has been introduced for cultivation. Here it is often found as an escape from cultivation. In the states mentioned under wild distribution, different cultivars are often found as an escape, since the plant is widely cultivated in most southern states of Mexico. This situation also applies to El Salvador, where cultivation of this crop is widely practiced.

WageningenAgric. Univ.Papers 90-3 (1990) 3 ^lïiilb

Figure 1. Neotropical distribution ofpachyrhizus species

In Guatemala only limited cultivation is practiced today, the main region of cultivation being the southern states of Santa Rosa, Jutipa and Chiquimula and, as recorded by the present author in 1988, in the occasional field in shifting cultivation in the state of Peten. However, the plant is often found as a relict from earliercultivatio n aswel la snumerou s locationso fwil dmaterial .I n general this situation is probably true for Honduras and Nicaragua as well, where little or no cultivation is now practiced. A number of collections of P. erosus is recorded from Belize (including one by the present author), but the plant has probably beenintroduce d for cultivation from thenorther n parts ofth e Yucatan peninsula, (figure 1.). Iti sknow n tohav ebee ncultivate d byth eAztec si n Central Mexico and by the Mayans in the Yucatan peninsula in pre-Columbian times according to reports by Lundell (1938), and Dibble &Anderso n (1963). Burkill (1935) states that P. erosus has only been known outside the Neotropics after Columbian contacts.

WageningenAgric. Univ.Papers 90-3 (1990) Figure 2. Areas outside the natural distribution area of P. erosus where cultivation has been recorded. Caribbean:Antigua , Cuba, Dominica, Guadeloupe, Hispaniola, Jamaica, Martinique,Puert o Rico, St.Croix , St. Kitt's, St. Thomas, St.Vincen t and Trinidad. SouthAmerica :Brazil , (French Guyana), Paraguay and Venezuela. Africa:Cameroons , Ghana, Senegal, Sierra Leone,Tanzani a and Zaire. IndianOcean :Andamans , Mauritius and Reunion. Asia: Burma, Cambodia, Indonesia, Laos, Malaysia, Northeast India and Sikkim, Singapore, Southeast China, Thailand and VietNam. Pacific Ocean:Formosa , Hawaii, Palau (Carolinas), Philippines, Raiatea (Tahiti Archipelago) and in the Marianas: Tinian and Guam.

Areas outside Mexico and Central America where P. erosus has been intro­ duced for cultivation and where cultivation is still practiced, or where the plant is known to have escaped from earlier cultivation, can be established from her­ barium material and other sources, e.g. in Brazil, (W.E. Kerr, pers. comm.), see figure 2. The history of this crop in cultivation has been recorded from quite an early date as can be seen from the herbarium material, see lists of exsiccatae. The specimens have been listed according to region and year of collection, and it appears that the first area of introduction (and cultivation) was the Far East, i.e. probably old Spanish possessions. The most likely route of introduction would then have been from the Philippines and Indonesia via Ceylon - India to the Mascarenes and then along thewes t coast of the African continent. Furthermore a number of French scientists recorded thecro p in The Far East and Oceania in the beginning of the 19th century, e.g. Gaudichaud-Beaupré (1826) in the Philippines and Oceania, and Perrottet (1821) in the Philippines and Indonesia. Perrottet not only recorded the plant in cultivation in The Far East,wher eh ethough t theplan t indigenous,bu t obtained sufficient seed materi­ al to be able to introduce the crop into the Mascarene islands, Senegambia and French Guyana (Perrottet 1824 and Guillemin et al. 1830). This means that P. erosus returned to the original distribution area after some 300-350 years. Thus, it may be possible that some of the cultivars known from the French Caribbean may have 'travelled' around the world, whereas other may 'only' have crossed over from Central America.

WageningenAgric. Univ.Papers 90-3 (1990) 5 Habitat: In areas with annual dry season; along deciduous forest edges and in scrub vegetation. On soil types ranging from deep clay to sandy loam. Recorded from 0-1.750 m a.s.l. with the majority of records from 500-900 m a.s.l.Th e rainfall range varies from 250-500m m to over 1.500m m mean annual precipitation rate (m.a.p.r.). Flowering season: Flowering specimens from allmonth s except January have been seen,majorit y (90%)wit h flowers inJul y tillOctober , latest inth e southern parts of the distribution area, i.e. at the end of the rainy season. Mature legumes recorded from August- February. In 1985matur e legumeswer ecollecte d during mid-March in Costa Rica. Cultivation: In Mexico and El Salvador P. erosusi seithe r grown in monocul­ ture ori sintercroppe d with maize,cabbag e or onions.Propagate d by seed, sown in April, May, June with harvest beginning in November up to February (infor­ mation on cultivation practices in Mexico and Central America were obtained from growers and local agronomists during thefiel d trip in 1985).I n some areas the removal of the flower buds is practiced in order to improve the tuber size, according to Schroeder (1968), Kay (1973), Duke (1981) and own observations. Recently, the effect of foliar application of different plant hormones, e.g. gib- berelic acid and chlorocholine chloride, on tuber yield has been studied by Zin- sou et al. (1987a & 1987b) and by Dr. Y. Elber, The Volcani Center, Israel (pers.comm.) . Uses: The fresh tubers of P. erosus are used either raw in various dishes or boiled like the roots of Manihot utilissima, cassava. A remarkable feature of boiled or fried tuber slices are the ability of retaining the crunchy quality (Schneider 1986). An excellent flour is obtained from sliced, dried and ground tubers (MacFadyen 1887 and Tadera et al. 1984). If allowed to grow to maxi­ mum size the tubers are used as a fodder for cattle and pigs. The pods can be used asa vegetable when young, but due to the rotenone content they are poiso­ nous when ripe. The rotenone can be extracted from the ripe pods and used as an insecticide. The dried vegetative parts of the plant are used as a forage hay after the harvest of the tubers.

2. Pachyrhizusferrugineu s (Piper) Serensen Morphology: Semi-woody to woody vine; deciduous to semi evergreen (ever­ green in greenhouse cultivation). Like P. erosusthi s species exhibits great varia­ tion in the outline of the leaflets, entire, palmate and lanceolate leaflets occur. The lateral inflorescence axes have the highest number of flowers seen in the genus, 10-37.Th e length of the inflorescence 9-46 cm issimila r that of P. erosus, however, the wing and keel petals are ciliolate with brown hairs. The legumes 8-13 cm long and 12-23 mm wide are distinctly different from the legumes of the other species. The brownish red seeds are flat, square to rounded, larger than those of the other species. Distribution: Known from the states of Vera Cruz, Chiapas and Quintana Roo in Mexico, east through Belize and south through the eastern and central parts of Guatemala, not known from El Salvador. Further south to Honduras,

6 Wageningen Agric. Univ.Papers 90-3 (1990) Nicaragua, Costa Rica, Panama and in Colombia the Department del Chocó. Introduced to theCaribbea n islands ofCuba , Martinique andTrinidad , where itwoul d appear to have escaped from the botanical gardens, (figure 1.). Habitat: In tropical rain forests, both deciduous and evergreen, pine forests and moist thickets. Soil types ranging from deep clay to coarse sand-clay-loam. Recorded from 0-1600 m a.s.l.,distribute d in areas with over 1.500 mm m.a.p.r. Flowering season: The majority of the specimens examined were flowering from November to March, but 6 accessions were flowering in the period from July to October. Mature legumes were found from February to June, with a single specimen in November. Cultivation: This species has never been cultivated. Uses: In Guatemala the ripe seeds were reported to be ground and used as a vermifuge.

3. Pachyrhizuspanamensi s Clausen Morphology: A herbaceous vine recognizable by all parts of the plant being entirely covered by white hairs. The inflorescences are shorter than those of the two previous species, 4-14 cm, and the number of flowers per lateral axis is reduced to 4-7. The wing and keel petals are ciliolate. The legume is 8-13 cm long and 7-12m m wide and the seeds are rounded, olive-green. Distribution: From the Panama Canal in the north, along the northwestern coast of Colombia and along the coastal plain of Ecuador to the province El Oro (Ecuador) in the south, (figure 1.). Habitat: At edges of deciduous forests, in low shrubbery and open grassy slopes. In areas with at least one dry season per year lasting 2-3 months. Recorded from 0-800m a.s.l .wit hth erainfal l rangebein g250-1.50 0m m m.a.p.r. Flowering season: In the north from October to December, with the fruiting season from December to March. A singlespecime n from Colombia had flowers in August. The flowering season is from March to May in Ecuador, with ripe fruits from April to June. Cultivation: P. panamensis is not recorded as being cultivated in Central America or elsewhere. Uses:N o known uses.

4. Pachyrhizustuberosu s(Lam. ) Spreng. Morphology: Also a herbaceous vine, but notably the largest species in the genus, attaining lengths above 10m . The leaflets areentir e (occasionally slightly dentate), but quite uniform. The inflorescence isfro m 7t o 29c m long with 7-33 flowers per lateral axis. The wing and keel petals are usually ciliolate, though glabrous specimens have been recorded. The legumes are the longest found: 13-19 cm, 14-23 mm wide. The seeds are rounded reniform, orange-red, black or black and white mottled. Distribution: Widely cultivated in the Amazonian region of South America and appears to be native to the western part of this region, (figure 1.). Collected in Colombia, Venezuela, British Guyana, Brazil, Bolivia, Per« and Ecuador,

WageningenAgric. Univ.Papers 90-3 (1990) 1 and reported to be cultivated in the eastern provinces of Paraguay (according to Dr. L. Ramella, pers.comm.) . Introduced to some Caribbean islands: Puerto Rico, Jamaica, Hispanola and Trinidad. Seeds of P. tuberosus from Trinidad were,accordin g to notes from Kew (Anonymous 1889a )distribute d to the bota­ nic gardens of Calcutta, Ceylon, Brisbane, Melbourne, Sydney and Adelaide. However, no herbarium specimen has been found of this species from outside the neotropics, and it is doubtful if the seeds distributed mentioned above (Anonymous 1889a )wer e in fact P. tuberosusan d not P. erosus. Habitat: In tropical to subtropical evergreen rain forests, restricted to areas with an annual rainfall of 1.500t o4.10 0m m m.a.p.r. Found at altitudes ranging from 0-1.550 m a.s.l.Occasionall y forming dense tangles. Flowering season: Due to the very heterogeneous origin and the uncertainty of the exact status of the material, i.e. wild or cultivated, the exact time and length of the flowering season cannot be determined. Specimens in full bloom have been registered from all months except February and July, but the bulk ofth emateria l had flowers from October toJune . Mature fruits seenfro m March to December. History: According to Montenegro (1740) P. tuberosus was cultivated by the Guarani Indians in Bolivia, and at the beginning of the century Bertoni (1913) found the plant cultivated in fields along the Parana river in Paraguay. In 1978 P. tuberosuswa s collected near Limoncocha in Ecuador where it was cultivated by the Aucas (according to information on the label of Madison, M.T. et al. 5465). According to Bentham (1859), Pinto (1873) and Warming (1892), P. erosus has been introduced and cultivated in Brazil. According to a recent manual (Anonymous 1979), P. tuberosus may only be a cultivar of P. erosus selected because of its larger roots. All plants of this species grown in greenhouse have produced tubers of the multi tuberous type! In order to elucidate the status of the species, both herbarium material and literature have been consulted. The specimen from Peru, collected by Spruce, R. 4936 in Tarapoto, of which two duplicates exist in Kew, formed the basis for the illustration with the note from Kew(Anonymou s 1889b) .On e sheetha sth eentir etyp eo fleaflet s and the inflor­ escence shown in the illustration, the other sheet has a legume similar to the one in the illustration and a deeply lobed leaf which has not been illustrated. This clearly demonstrates that both leaf types occur within P. tuberosus, maybe even on the same plant! But as the material is supposedly cultivated, and as P. erosusi sknow n to havebee n cultivated inth e area simultaneously, only gene­ tic and ontogenetic studies can reveal whether, considering the 16 specimens, the two species are in fact interfertile or whether the material from Brazil and Peru isactuall y a cultivar of P. erosus,wit h legumes of the P. tuberosustype . Cultivation: P. tuberosusi sknow n to beo r havebee ncultivate d by indigenous people of the Amazon region, occasionally found in 'chacras', i.e. fields, sur­ rounding remote villages located in highland rain forests ('selva alta') of Peru, noticeable in the department of San Martin, or grown in shifting cultivation inth eAmazo n proper. Preferably infertile , light sandy soilswit h good drainage.

8 WageningenAgric. Univ.Papers 90-3 (1990) Propagated by seed at 45-50 kg per hectare. Several bud and flower prunings are believed necessary in order to obtain high tuber quality. The information on cultivation practices are according to Ms. A. Sorensen (pers. comm.), Duke (1981)an d information on the labels of herbarium specimens. Uses: The tubers of P. tuberosus are used in the same way as the tubers of P. erosus, and are recorded to be used for the production of flour in Jamaica (Anonymous 1889a) . Theyoun g podsca n beuse d boiled asa vegetable , howev­ er, somecultivar s do have strigose legumes.

5. Pachyrhizusahip a(Wedd. ) Parodi Morphology: An erect to semi-erect herbaceous plant, 30-40 cm tall, with very short inflorescences, 5-9 cm. The number of lateral axes on the main axis of the inflorescence is greatly reduced, 0-6, with only 2-6 flowers per lateral axis. The wing and keel petals are usually glabrous, but sligthly ciliolated specimens have been seen. The legume is 13-17 cm long and 11-16 mm wide, the seeds arerounde d reniform, black or black and white mottled. Distribution: Widely cultivated in Bolivia and Peru in fertile valleys between 1.500 and 2.500m a.s.l. Herbarium specimens havebee n seen from the provinces of Sorata and Tarija in Bolivia; (figure 1.). Formerly also cultivated in Jujuy (herbarium specimens seen) and Salta in Argentina, but probably from seeds introduced from Bolivia (Bolivian farm labourers working in Argentina recall importing seed material from Bolivia when visiting relatives (pers. obs.)). In Peru tubers are frequently found in funeral bundles from the Paracas Necropolis according to Yacovleff & Muelle (1934), Mangelsdorf (1942), Towle (1952) and Ugent et al. (1986). There are no records of plants undoubtedly wild. Habitat: Asstate d thisspecie si sknow n from cultivation only,i ncoo l tropical/ subtropical valleys,wit h a rainfall ranging from 500t o 1.500 mm m.a.p.r. Flowering season: Usually sown in December, with flowers in February - March and with mature legumes in April. History: Representations of P. ahipa have been found on pottery from the Paracas Necropolis and from the southern coast of Peru (Yacovleff 1933); also found depicted on Nasca embroidery according to O'Neale & Whitaker (1947) and Brücher (1977). Accordingly, substantial evidence exists that P. ahipa was known and cultivated by the Incas in pre-Columbian times, also information in the manuscripts of Oviedo (1535). Cultivation: Grown either asa monocrop or intercropped inmaiz e fields. Uses: The tuber is used as a vegetable, raw or cooked in various dishes. No records on the use of the insecticidal properties of the plant have been seen.

The relationship between the different species iscurrentl y being studied, involv­ ingcross-breedin g and examination of thestabilit y ofvariou s genetic characters, e.g. the relationship between P. ahipaan d the remaining species.Thi s iso f inter­ est as P. ahipapossesse s several desirable characters from an agricultural point of view, e.g. short life cycle and erect to semi-erect habit. Furthermore, as no

WageningenAgric. Univ.Papers 90-3 (1990) 9 wild material of P. ahipa is presently known, new field collections, in order to obtain such material, would certainly constitute a major contribution towards supplying theknowledg e needed to establish the status of this species.

Biochemical studies ofth egenu s

1. Nutritional composition of the tubers, table la-b. Although the tubers of the yam beans are known to possess a nutritional com­ position superior to that found in most non-leguminous root crops (Lynd & Pur- cino 1987), the number of analyses of the chemical composition of these tubers does not compare with equivalent studies of e.g.cassav a roots. The first compre­ hensive study of the constituents of tubers and seeds of P. erosus was carried out by T. Peckolt (1880 & 1883) and G. Peckolt (1922). The findings of these studies, confirming the high nutritional value of the tuber, is in agreement with later analyses conducted by Porterfield (1951), Aguilar (1958) and more recently with Duke (1981). Aguilar (1958) presents figures for nutritive value of tubers from 4 different species:P. angulatus,P. palmatilobus, P. tuberosus and P. erosus; however, later studies indicate that these 'species' were in fact different cultivars of P.erosus an d probably material collected from thewild . Ghosh &Guh a (1934) studied the tuber content of vitamin Bl & B2 and reported that P. angulatus( = P. erosus)di dconstitut e a good source ofthes e vitamins. The quality of the protein and starch of yam bean tubers has recently been examined by Tadera et al. (1984). The extremely low values of the protein con­ tent of the tubers reported in that paper were based on a single cultivar only, and, as the tubers tested were immature, the findings are not in agreement with the above mentioned findings of Lynd & Purcino (1987). However, Tadera et al.(1984 )foun d that thestarc h quality ofth etubers ,usin gexperimenta l enzyma­ ticmetabolism , wasindee d of a highly digestible nature. The examination of a possible increase in sugar constituents due to storage and heat treatment has been the objective of the experiments carried out by Kawabata et al.(1986) .The y reported that the sugar content ofya m bean tubers was significantly higher when compared to the sugar content of cassava roots tubers. Furthermore, regardless of the higher moisture content of the yam bean tubers the decrease in moisture content due to prolonged storage was less in Yam Bean tubers than in Cassava roots. The analyses for amino acidconten t arecompare d to the amino acid composi­ tion of the African yam bean. Although the protein content of the African yam bean (approx. 16% on fresh weight basis) is considerably higher than that of yam beans(1-2%) ,th edifference s inamin o acidcompositio n (ing/10 0g protein ) on adr ymatte r basisi sno t ofsignifican t magnitude. Finally, theprotei n content of the tubers must be seen in relation to the tuber yield per hectare (i.e. 70-90 tonnes!).

10 WageningenAgric. Univ.Papers 90-3 (1990) Source 1 2 * 3 4 5e 6 * 7e 8 9 10

Species ero ero# tub? ero ero ero ero? ero ero ero? Moisture 86.9 - 82.25 88.5 82.38 88.65 86.1 78.09 87.49 90.05 Protein 1.4 1.26 1.05 - 1.47 1.03 0.24 2.18 0.83 1.02 Lipids 0.1 0.11 0.30 - 0.09 0.1 0.9 0.80 - 0.14 Carbohydr 11.0 10.67 - - - 9.9 - 14.00 - - Starch - - 8.46 - 9.72 - 6.51 - 2.01 - Red. sug. - - - - 2.17 - - - 3.39 1.45 Nonred.s. - - - - 3.03 - - - 1.02 - Sucrose - - 1.29 0.49 - - - - - 1.43 Glucose - - 0.26 1.30 - - 2.26 - - - Fructose - - - 1.03 ------Pectose - - 1.62 - - - 0.83 - - 2.34 Resin - - 0.13 - - - 0.23 - - 0.17 Fiber 0.6 - 0.66 - 0.64 0.6 - 1.43 - 0.75 Ash 0.6 - - - 0.50 0.3 1.73 0.80 - 0.36 Tot. dig. - 12.04 ------Tot.nodig - - - - - 10.0 - - - 7.68 cal - - - - 16.0 13.3 - - - - pl - - - - - 16.3 - - - - Fel - - - - 1.13 0.53 - - - - Cu1 0.43 ------Ascorb.a. - - - - 14.0 15.67 - - - - Tot. min. - - 1.84 - - - 0.73 - - - Table la. Nutritional values and chemical composition of yam bean tubers.

Source: 1. Duke (1981); 2. Aguilar (1958); 3.Anonymou s (1895); 4. Kawabata et al. (1986); 5. Kay (1973); 6. Montaldo (1972); 7. Peckolt (1880); 8. Porterfield (1951); 9. Tadera et al. (1984); 10. Yacovleff(1933). *: modified from more than one analysis. @: analyses of peeled tubers. # : the species cited in the analyses - P. palmatilobus, P. angulatus, P. tuberosus and P. erosus are all considered as belonging to P. erosus only, as this is the only species used for human consumption in Guatemala. ?: the exact identification of the species remains uncertain as both P. erosus and P. tuberosus wererecorde d ascultivate d in Brazil and Guyana. ': mg/100g

2. Chemical compounds with insecticidal or fungicidal properties in the seeds, pods, stems and leaves,tabl e 2. As is clearly demonstrated in the table (Tab. 2.) practically all studies of the insecticidalcompound s and theirpropertie s havebee nconducte d on one species only, i.e. P. erosus. The only exception known to the author (the list of studies appearingi nth etabl erepresent s themajorit y ofanalyse spublished )i sth eexami -

WageningenAgric. Univ.Papers 90-3 (1990) 11 Source la 5- 6- lib la

Amino acid Afr.Y.-B. Thiamine - 0.05 0.05 - Riboflav. - 0.02 0.04 Niacine - 0.2 0.27 - Asparagine 12.8-55.1 - - 7.40 11.3-13.8 Threonine 2.7-5.0 - - 2.83 4.1-4.5 Serine 3.3-6.3 - - 2.86 5.1-6.4 Glutam. ac. 5.4-11.7 - - 15.36 11.6-12.3 Proline 2.5-4.4 - - 3.51 7.5-7.8 Glycine 1.9-5.0 - - 20.61 4.6-5.2 Alanine 2.5-5.6 - - 2.91 5.3-5.6 Valine 3.2-8.2 - - 2.64 5.2-5.9 Methionine 0.8-1.5 - - 0.81 1.5-1.9 Isoleucine 2.1-4.8 - - 3.54 4.2-4.9 Leucine 2.9-7.9 - - 5.80 7.0-8.4 Tyrosine 1.4-4.7 - - 2.89 3.0-3.4 Phenylalan. 2.3-5.1 - - 3.71 4.2-4.9 Histidine 2.5-3.2 - - 2.51 4.4-5.0 Lysine 3.3-7.8 - - 5.75 7.1-8.3 Arginine 5.0-7.4 - - 4.71 5.2-6.5 Cystine 0.4-1.0 - - + 1.6-2.0 Triptophan. - - - 1.02 - Ammonium - - - 2.22 - Table lb. Amino acid content.

Source: 1.Duk e (1981); 5.Ka y (1973); 6.Montald o (1972); 11. Vidal &Pimente l (1985). a - per 100g tube r meal • - per 100g peele d tuber b - g/16g N , total content Afr.Y.-B. = African yam bean (Sphenostylis stenocarpa) nations published by Hansberry et al. (1947).I n their study the species P. strigo- sus (now considered conspecific with P. erosus), P. ahipa and P. tuberosus were included. Although this extension of the material has increased the knowledge of the properties of the genus in general (the differences between the species found by Hansberry and his associates are remarkably few) it would indeed be valuable and possibly of commercial interest to include the two wild species in new studies of the insecticidal properties. Some findings are shared by all the examinations of the insecticidal com­ pounds: 1. rotenone isno t the sole agent isolated having insecticidal properties, at least two other compounds do possess such properties (erosone and pachyrhizid). 12 WageningenAgric. Univ.Papers 90-3 (1990) 2. onceisolate d the LD50 value of these compounds (including rotenone) com­ pares fairly with other known herbal insecticides. 3. as the majority of the examinations presented here are some 30+ years old, the scientists agree that extraction on a commercial basis cannot be consid­ ered feasible asth eroteno n content rarelyexceed s 1.0%o n adr ymatte r basis. Duke (1981)state stha t extraction ispracticed , but theautho r hasbee n unable to verify this. Ingham (1979) considered the presence of isoflavonoid phytoalexins as vali­ dating atransfe r ofth e genusfro m the subtrib e Diocleinae to the subtribe Glyci- ninae, due to the close chemical relationship with particular Neorautanenia and Glycine. The author of the present paper disagrees with this perception, not only due to the shared presence of canavanine of the members of the Diocleinae asdemonstrate d by Lackey (1977)an d others, but primarily because ofth e mor­ phological, chromosomal, and distributional characters of the genus.

Author species compound(s)isolate d organ property and/orexamine d exam. examined Baker& Lyn n (1953) erosus saponins,fatt yoi l seeds - Cruz (1950) erosus glycerideso foleic ,li - seeds - nolic& saturate dacid s Hansberry &Le e(1943 ) erosus rotenone seeds insecticidal Hansberrye tal . (1947) erosus rotenone,pachyrhizid , seeds insecticidal strigosus erosonean d4 othe r ahipa unnamedcompound s tuberosus Holz& Hon g (1964) erosus undetermined seeds insecticidal Ingham (1979) erosus isoflavonoidphyto ­ stems fungicidal alexins Jakobs (1949) erosus fattyacid s seeds - Krishnamurtie tal .(1970 ) erosus rotenoids seeds - Lamazee tal . (1985) erosus nitrogenouscompound s nodules, ammonia leaves, assimila­ pods tion Lee& Hansberr y (1943) erosus undetermined seeds insecticidal Lepagee tal . (1946) tuberosus undetermined (oils) seeds insecticidal Matthysse& Schward t (1943) erosus undetermined seeds insecticidal Norton (1943) erosus rotenone seeds insecticidal Norton& Hansberr y (1945) erosus rotenone,pachyrhi zid , seeds insecticidal erosonean d4 othe r unnamedcompound s Plank (1944) erosus undetermined seeds, insecticidal pods Shangraw& Lyn n (1955) erosus pachysaponinsA & B , seeds piscicidal pachysapogeninsA & B Sillevoldt (1899) erosus pachyrhizid ,anhydro - seeds piscicide, derivateso fpachyrhi - amphibieide zid Vidal& Pimente l (1985) erosus proteinbodies ,amin o coty­ proteinmo ­ acids ledons bilization Table 2. (chemical compounds with insecticidal or fungicidal properties in the seeds, pods, stems and leaves)

WageningenAgric. Univ.Papers 90-3 (1990) 13 Pests and diseases

Very few pests have been reported to attack this crop: Duke (1981) states that the nematode Heterodera marioni has been isolated from the roots, but whether such an attack does in fact pose a threat to the yield expectancy remains unclear until field tests have been conducted. In agreement with Duke (1981)an d Clau­ sen (1945) the author has found the mealy bug (Ferrisia virgata) to be a severe pest of yam beans grown under greenhouse conditions. No records have been made of this particular pest from field trials, but the author has seen Ferrisia sp. effectively destroying inflorescences of P'.ferrugineus in Belize.Anothe r pest of increasing importance in greenhouse cultivation is the American Thrips: Frankliniella occidentalis which causes the premature abscission of flower buds and thusjeopardiz e anycros sbreedin gexperiments . Bothpest sca nb e controlled through the application of biological control, i.e. the severity of the attack can be minimized. However, only chemical control will completely eliminate the pests, but the damage to fertile shoots, due to the high dosages needed in order to combat thesepests ,ha s been found to be extensive. Fungal attacks causing root rot in P. erosuswa s reported by Yu et al. (1945), who isolated the fungus Pythium aphanidermatum as the disease causing patho­ gen. Both fungal and bacterial leaf spot have been recorded in P. erosus. Mohanty & Behera (1961) isolated the fungus Cercospora canescensfro m P. erosus leaves at the State Agricultural Research Station, Bhubaneswar, India. This fungus was found to cause severe damage - defoliation of a crop of P. erosus at the research station. The bacterial leaf spot caused by the pathogen Pseudomonas syringae pv.phaseolicola was first recorded by Birch et al. (1981) when working with P. erosus at the Hawaii Agricultural Experiment Station, Honolulu. The conclusion of their finding was that P. erosusma y constitute an alternative host for the halo-blight pathogen of beans in tropical regions. Two diseases of P. erosus caused by viral attack have been identified. The sincama mosaic disease was first described by Fajardo & Maranon (1932) and the witches' broom disease of legumes was recorded in P. erosus by Thung & Hadiwidjaja (1957). The frequency of the latter disease appear to be rather low; only one plant wasinfecte d in the trialsdescribe d by the authors. The symptoms of the disease are excessive branching, dwarfed leaves or atrophy of the leaves, and antholysis, i.e. deformation of the flowers. Witches' broom was found not to be transmittable through sap inoculations, but there are some indications that the disease may possibly be transmitted by insects such as Orosius argenta- tus. The sincama mosaic disease is doubtless the most serious disease of the yam bean; it is so far restricted to P. erosus cultivars and escaped material of that species in the Far East. Fajardo & Maranon (1932) stated that the disease may indeed be specific to P. erosus (only this species was examined in their study). However, the present author has found the disease capable of infecting all other

14 WageningenAgric. Univ.Papers 90-3 (1990) specieswit h theexceptio n oîP.ferrugineus. Furthermore, though cross infection with Phaseolus vulgaris was unsuccessful in the experiments carried out by Fajardo & Maranon (1932), this author finds the postulated specificity of the disease hard to accept when considering the origin of the crop, i.e. neotropical. The symptoms of the disease are according to Fajardo & Maranon (1932) and confirmed by own observations in thegreenhouse : varying degrees of chlor­ osis and mottling of the leaves depending on the time of infection. If infected at an early stage the plants may be stunted/dwarfed and/or spindly with brittle vegetative parts. Infected plant may seem to recover as leaves produced at a later stage may have a healthy looking appearance, but, as will be discussed later, when fertile shoots are produced the presence of the infection will be revealed. Thediseas eha s onlyprove d fatal to thehos t plant ifinfectio n occurred at a very early stage, i.e. when seedlings are infected. The virus is seed borne according to Fajardo & Maranon (1932), a postulate not confirmed by this au­ thor. However, as it is systemic the disease will be spread with tuber material, e.g. for later production of seeds. As the crop is usually produced from seeds (occasionally from tubers in P. tuberosus - a multi tuberous species (Peckolt 1880)) it is of vital importance to use uninfected plants for the production of seed material. Transmission of the disease appear to be restricted to contamination through mechanical wounding and insect attack. The first method of transmission will indeed be the cause of a virulent attack if the practice of staking fertile shoots inorde r toimprov etube rdevelopmen t iscarrie d out. Fajardo &Marano n (1932) were unable to demonstrate any transmission by the mealy bug Ferrisia virgata; own observations have indicated that aphids and possibly spider mites are cap­ able of transmitting the disease. Transmission does not occur through physical contact between healthy and infected plants neither above nor below ground level (Fajardo &Marano n 1932). Both own observations and information according to Fajardo & Maranon (1932) have demonstrated that the tuber yield will be affected by viral infection. Depending on the time of infection and the species infected the development of the tuber will be reduced. Seedling of P. tuberosus, material originating in Brazil,prove d extremely sensitivet o viral attack. These observations were made duringgrowt h experiments under greenhousecondition s overa five month s peri­ od (August-December 1987)a t the Royal Veterinary and Agricultural Universi­ ty, Copenhagen. Tuber development ceased completely, ifth e P. tuberosus seed­ lingswer e infected within a two week period following germination. Examinations of possible changes in the chemical composition of infected tubers,whe n compared with healthy ones,wer einclude d inth e study by Fajardo & Maranon (1932). In spite of the limited material studied, they demonstrated that a reduction of reducing sugars, total sugars, pentosans and dry matter had taken place in infected material, but the starch percentage had apparently increased. Their results warrant verification, but may serve as a basis for com­ parative studies of infected plants belonging to the remaining species. The changes observed in the morphology and fertility of fertile shoots and

WageningenAgric. Univ.Papers 90-3 (1990) 15 organs caused by the mosaic virus are numerous. The abscission rate of floral budsan d offlower s has been found to increasedramatically . Eventh e abscission rate of developing legumes increases, i.e. a rare phenomenon in healthy plants. However, themos t dramatic effect of this disease isth e decrease inpolle n fertili­ ty.Tw o collections of P. tuberosusfro m Brazil (observations based on ten plants of each collection), which prior to the virus infection had pollen fertilities from 88-97%, were practically sterile following infection with the virus, i.e. pollen fertilities of 20% to below 10% were observed. Control measures needed in order tocomba t infection with thesincam a mosa­ ic virus are of course of vital importance when assessing the performance of different cultivars and speciesi nfield trials .Th eimplementatio n of prophylactic control will be an obvious advantage to such tests. All seed material must be germinated and kept under quarantine for a minimum of one month prior to transfer to final growth site. If tuber material is to be included in field trials, they should likewise be kept under quarantine until their disease free condition can be ascertained.

Growth experiments

1. Comparison of tuber growth in species and interspecific hybrids. Seeds were germinated in 11c m plastic pots, 3/4 filled with non-fertilized and non-limed peat and the top 1/4 filled with pure quarts sand, in order to provide optimal drainage. Insufficient drainage proved greatly to reduce or completely inhibit germination in P. ahipa and P. ferrugineus, i.e. the seeds were either attacked byfung i orthe yappeare d to soak tooquickl y and became soggy instead of firm. The pots were placed in a plastic tent within the greenhouse and kept at a temperature not below 25°C during the day and not below 20°C at night. 14 days following germination, seedlings were transferred to 10 cm pots and grown in standard soil (see Appendix l.a). The pot size was increased after two months to 15 cm until the plant could be transferred to the final pot size of 19 cm (3.5 1) after 4 1/2 months. During the entire growth period the plants were watered with a standard nutrient solution (see Appendix l.b). Additional light (growth lamps) was provided for a 12 hour period during autumn and winter. Flowering was induced in plants belonging to P. erosus, P.panamensis (buds only; abscized prior to full development) by reducing the daylength to 8 hours for a three week period. Plants belonging to P.ferrugineus did not respond to this short-day treatment, (manus. in prep.). The tubersneede d for the experimental evaluation ofyiel d sizewer e harvested when plants had completed a full growth period, i.e. once the above ground parts had wilted. Prior to weighing the tubers were washed and cleaned from adhering soil particles. In order to obtain a standardized growth season for the

16 WageningenAgric. Univ.Papers 90-3 (1990) comparison of both species and hybrid tuber material, the tuber weight at har­ vest was divided by the number of days in growth and multiplied by a standard growth season of25 0days .

Tuber morphology. Tubers from all five species and a number of tubers from interspecific hybrids areshow n infigure s 3-6. Themulti-tuberou s nature ofth etw ospecie sP. tubero­ sus and P. panamensis is apparent, also the hybrids involving P. tuberosus (whether serving as 'female' or as pollen donor) are of the multi tuberous type. P. erosus(cultivars ) and P. ahipagenerall y produce only a singletube r per plant. However, P. erosus wild forms are of the multi tuberous type, a fact indicating that 'man made' selection has taken placei nth ecreatio n ofth e present cultivars. It may be argued that wild forms of P. ahipa may have belonged to the multi- tuberous type,confirmatio n ofthi shypothesi smust ,however , awaitth elocaliza ­ tion of wild forms of this species. The difference in the appearance between thetuber s from P. tuberosusan d P. ahipai sremarkabl e and supports the conclu­ sion arrived at by Ugent et al. (1986) that the mummified tubers found in the South American funeral bundles are P. ahipatubers .Althoug h Yacovleff (1933) is his paper on 'La jiquima' refers the pre-Columbian depictions of a tuberous legume to the species P. tuberosus, the correct identity of this plant becomes obvious when compared to the tubers from P. ahipa, i.e. identical. The cause for the prevalent uncertainty in the correct determination of these pre-Colum­ bian materials are due to the extremely limited cultivation and knowledge of P. ahipai n Peru. The last species, P.ferrugineus is either single or multi tuberous (the illust­ ration are of tubers obtained in the natural distribution area of the species), the multi-tuberous type has been observed in material grown from seed (ace. no. FW044/385) in the greenhouse. The tubers are quitewood y and asth e mate­ rial available for potential yield estimates has sofa r been insufficient, additional information on the chemical constituents and growth rate must await the addi­ tion of further material.

Results &discussion. The results are presented in figure 7. The results of this experiment need to be validated through growth experiments, carried out under practical field condi­ tions. The remarkably high yields of the wild species P. panamensis and the wild form of P. erosus, exhibiting yield sizes equal or superior to the cultivated species: P. erosus, P. tuberosus and P. ahipa, seem, however, to indicate that the cultivars presently available have not resulted from a 'man made' selection based on yield performance alone. It appears probable that selection on the basis of plants being mono-tuberous, in the case of P. erosus and P. ahipa, has played a major role. Uniformity partly due to selfpollination and cultivation practices, most cultivars being entirely local, could explain the comparatively low yield performance when compared to wild material. The fact that the four cultivars of P. tuberosus seen have all been of the multi-tuberous type can be

WageningenAgric. Univ.Papers 90-3 (1990) 17 Figure 3. Tubers of P. erosus 1. Wild form from Nentón, Depto. Huehuetenango, Guatemala; 2. Wild form from Finca Pacifica, Prov. Guanacaste, Costa Rica; 3. Cultivar from San Ixtan, Mun. Jalpatalgua, Depto. Jutiapa, Guatemala; 4.Cultiva r from Celaya, Edo. Guanajuato, Mexico.

Figure 4. Tubers of P. ferrugineus 1. From Mountain Pine Ridge, Distr. Cayo, Belize; 2. From Gamboa, Canal Zone, Panama. Tubers of P. panamensis 3. & 4. Madden Dam, Canal Zone, Panama.

WageningenAgric. Univ.Papers 90-3 (1990) Figure 5. Tubers of P. luberosus 1. Cultivar from Pare National Historique Citadelle Sans Souci Ramiers,Dept . Nord-Este, Haïti; 2.Cultiva r from Mun. Uberlandia, Edo. Minas Gérais, Brazil. Tubersof P.ahipa 3 .& 4 .Cultiva r from Prov.Tarija , Bolivia;5 .Cultiva r ofunknow n origin obtained from the Devonian Botanical Garden, University of Alberta, Cananda; 6. Cultivar of unknown origin obtained from Jardin Botanique National de Belgique, Meise, Belgium.

;Hfcf' A

Figure 6. Hybrid tubers from Frgeneration. 1., 2. &3 . P. erosus x P. ahipa (1., 2.& 3. EC032/285 x AC102/485); 4. & 5. P. erosus x P. tuberosus (4. & 5. EC503/882 x TC118/186); 6. P. tuberosus x P. erosus (TC118/186 x EC032/285); 7. P. tuberosus x P. erosus, wild form (TCI 18/186 x EW522/ 386). WageningenAgric. Univ.Papers 90-3 (1990) 19 Il ©

S" 300- ©

© 200- ©

100-

Ho. of seeds par plant

30 i 20 40 50 60 70 80 ®

100- ©

©t 200- ® © © ©

ï" 300J

Figure 7. Vertical axis: tuber weight in grammes at 250 days in growth. Horizontal axis: mean number of seeds per plant in test. 1. P. erosus x P. tuberosus (3).2., 2a. and 2b. P. tuberosus x P. erosus (2. = total (7); 2a. = cross no. 89 (3) and 2b. = cross no. 68 (4)). 3. P. panamensis (18). 4., 4a. and 4b. P. erosus (4. = total (143); 4a. = cultivated material (102) and 4b. = material from wild forms (41)). 5. P. ahipa (5). 6. P. tuberosus (6). 7. P. erosus x P. ahipa (14). The number of tubers tested aregive n in brackets. explained by the status of this crop in cultivation. P. tuberosus is only known to be or have been cultivated by indigenous people of the Amazon region, most commonly grown in shifting cultivation. The morphological variation of the

20 WageningenAgric. Univ.Papers 90-3 (1990) cultivars examined from herbarium specimens indicate that a considerable number of local cultivars exist. Furthermore, thefe w introductions to areas out­ sideth e original distribution area and thepresen t status ofth ecultivatio n taking place inthes e areas,coul d explain the lack ofmor e uniform 'sophisticated' culti­ vars. I.e. only known from Haïti and Puerto Rico,wher e the tubers are gathered and consumed by humans, but the cultural practice is minimal, the plant has been allowed to escape and the tubers are collected from 'wild' populations, according to T.A. Zanoni, Santo Domingo (pers. comm.). This means that P. tuberosus has probably been subjected to very little selection, even in areas of introduction, and cultivars will therefore be almost indistinguishable from known wild forms.

2. Comparison of chemical constituents in plant material (leaves) collected in the natural distribution area and plant material (leaves) from plants grown under greenhouse conditions, tables 3a-e .

Materials & methods. Leaves from Central American material were obtained during a field trip in February - April 1985. The leaves were taken at a median position on the stem and dried. Material from greenhouse plants were taken when the plants had reached maximum growth rate,i.e .4 month s after germination. The leaves were from a position similar to the one in the Central American material. The green­ house material was dried and both materials were ground; 6 part samples of 0.3 g from each plant were separated for the analyses. The part samples were prepared according to themetho d suggested by Allen (1974)usin g 8.8 ml H2S04 and H202. The nitrogen content was determined using the method described by Hinds & Lowe (1980); a spectrophotometric measurement of the intensity ofth eblu ecolou r resulting from theBerthelot s phenol-hypochlorite-ammonium reaction. The rapidity of the reaction and the intensity of the colour were increased by the addition of natrium-nitro-prusside. The phosphor content was determined using the spectrophotometric method described by Allen (1974) by measuring the blue colour resulting from the reduction of the phosphor-molyb- date complex by adding stanno-chloride. The content of calcium, magnesium and natrium was determined through atomic absorption spectrophotometry (Allen 1974). Lathane was added to the diluted part samples, to be tested for calcium and magnesium content, at a concentration of 500 ppm in the diluted samples.

Results &discussion. The analyses, ofwhic h the results are presented inTable . 3.a-e. , were conducted inorde r toexamin epossibl edifference s (ordeficiencies ) inth echemica l constitu­ ents of leaves from plants grown under greenhouse conditions when compared to leaves from plants collected in the original distribution area of the genus. Only the material of P.ferrugineus iso f Central American origin, the remain­ ingmateria l has allbee n obtained from plants grown in thegreenhouse . Howev-

WageningenAgric. Univ.Papers 90-3 (1990) 21 Pachyrhizus erosus Table 3a

Herb, no./ ECO06/ EC116/ EC117/ EC503/ EC503/ EW115/ EW115 EW522/ EW522/ Accession no. 285* 186* -86* 882* 882* 1285* 1285* 386* 386*

Cult./wild cult. cult. cult. cult. cult. wild wild wild wild Locality Oaxaca Jutia. - Gto. Gto. Huehu. Huehu Reduit Reduit Country Mexico Guate. Thail. Mexico Mexico Guate. Guate Mauri. Mauri.

Ca ppm 20.2 16.3 26.2 21.6 20.0 26.9 26.0 22.7 25.9

Mg ppm 2.9 4.0 5.1 3.9 3.3 4.3 4.0 3.9 3.9

K ppm 36.3 25.1 24.3 21.8 24.4 23.8 24.8 21.9 21.9

N ppm 41.8 48.0 41.0 43.6 42.5 48.8 47.4 44.1 42.2

P ppm 3.2 4.5 2.3 5.3 4.0 2.2 3.5 4.0 2.8

i=Jï Table 3b,c Pachyrhizus ferrugineus P. panamensis

Herb, no./ 35# 36# 44# 64# FW044/ PW055/ PW055/ Accession no. 385* 385* 385*

Cult./wild wild wild wild wild wild wild wild

Locality Cayo Cayo Izaba. Gamboa Izaba. Canal Canal

Country Belize Belize Guate. Panama Guate. Panama Panama

Ca ppm 27.2 29.5 21.0 48.4 24.8 26.0 26.4

Mg ppm 2.0 3.3 2.5 1.4 3.2 4.0 4.0

K ppm 13.3 14.0 13.9 0.3 25.1 25.1 21.1

N ppm 36.7 36.0 46.6 41.3 45.7 48.0 51.5

P ppm 2.7 3.0 2.5 5.6 2.7 4.5 4.8

Table 3d,e Pachyrhizus ahipa P. tuberosus

Herb. no./ AC102/ AC102/ AC102/ TC118/ TC118/ Accession no. 485* 485* 485* 186* 186*

Cult./wild cult. cult. cult. cult. cult.

Locality Tarija Tarija Tarija N.Esté N.Esté

Country Boliv. Boliv. Boliv. Haiti Haiti

Ca ppm 22.2 27.9 17.9 9.5 16.2

Mg ppm 3.6 4.2 3.6 3.5 3.9

K ppm 25.4 18.3 30.7 26.3 22.8

N ppm 49.5 45.8 52.4 31.6 35.9

P ppm 2.9 2.9 4.4 3.8 4.9

Table 3a. (plant constituents) * - plant material (leaves) from plants grown under greenhouse conditions.

# - plant material (leaves)collecte d in the original distribution area.

Table 3b& c (plant constituents)

Table 3d& e .(plan t constituents) 22 WageningenAgric. Univ.Papers 90-3 (1990) er, a number of the results are of interest when contemplating possible selection of material for breeding new cultivars having optimal adaptability for a specific region. E.g. Lynd & Purcino (1987) demonstrated the importance of available phosphorus in order to maximize yield performance, P.ferrugineus appears to have a somewhat lower content of P in the leaves,whe n compared to the other species; however as will be seen from the soil analyses, presented below (Table 4), the species exhibits a clear preference for soil types suffering from an almost total depletion of available phosphorus. Hence, new cultivars for tuber produc­ tion in areas with soil types low in available P, are likely to benefit from charac­ teristics present in P. ferrugineus. The Caconten t of theleave si sanothe r factor exhibiting an interesting pattern ofvariation . The leaves from wild forms of P. erosusan d of the two wild species have rather high amounts of Ca present in the leaves.

Analyses of soil samples from natural distribution area, table 4.

Materials & methods. The soil samples were all obtained during the field trip to Central and South America in 1985. A minimum of 100 g per sample taken at 4-5 different places within 1 m2o fth esit ewher eth etubers/root s ofth eplan twer elocated , preferably at depths corresponding with the depth of the tuber (in tuberous plants). Themethod s used for thedeterminatio n ofpH ,conductivity , calcium, magne­ sium, potassium, sodium, manganate, phosphor, copper, iron and zinc were according to Hansen (1976): pH - 20 g soil + 100 ml H20. Conductivity - 20 g soil + 100 ml H20, corrected to 20°C umho. Exchangeable Ca, Mg, Mn, K andNa-1 0 gsoi l + 100m lNNH 4Ac(ph 7.0);atomi cabsorptio n spectropho­ tometry (AAS-Model 360). Means of duplicate analyses, results in meq/100 g soil. Analysis of Ca with 5% La203 added. P- 10g soil + 100m l 0.2 N H2S04, spectrophotometric ammonium molybdate method; results in ppm. Cu, Fe and Zn - 20 g soil + 80 ml 0.02 M EDTA, 2 Na; atomic absorption spectrophoto­ metry, duplicate analyses; the results in ppm.

Results and discussion. The quality of the soil of fields with commercially cropping of P. erosus, based on the analyses of the samples presented below, appears quite superior in terms of fertility when compared to the samples originating from localities whith wild forms of P. erosus (Costa Rica) and P. ferrugineus (Belize and Panama). Of special interest are the extremely low values of available phosphor and potassi­ um in soils from P.ferrugineus localities. Other significant features of the soils from wild material, demonstrated when comparing the differences in the analy­ ses, are the increasing acidity, the considerable increase in manganese and iron content and the decrease of the magnesium content. As mentioned previously thisspecie sdemonstrate s ahig htoleranc e towards soilcondition s oflo w fertility. Although, this characteristic will be of obvious advantage when selecting culti­ vars for marginal areas, further collections of material throughout the distribu-

WageningenAgric. Univ.Papers 90-3 (1990) 23 Species Pachy- Pachy- Pachy- Pachy- Pachy- Pachy- rhizus rhizus rhizus rhizus rhizus rhizus ferrugin. ferrugin. ferrugin. ferrugin. ferrugin. ferrugin.

Cult./ wild wild wild wild wild wild wild

Loca­ Cayo Cayo Cayo Gamboa Gamboa Colon lity Distr. Distr. Distr.

Country Belize Belize Belize Panama Panama Panama

PH 6.32 6.38 6.34 7.88 5.18 5.60

Conduc­ 124.1 148.9 148.9 179.6 90.0 175.2 tivity

Ca meq/ 21.36 22.95 24.66 18.37 5.06 22.27 100 g

Mg meq/ 7.67 6.64 8.59 4.98 3.87 14.02 100 g

K meq/ 0.62 0.66 0.78 0.05 0.12 0.12 100 g

Na meq/ 0.52 0.61 0.74 0.52 0.47 0.73 100 g

Mn meq/ 0.36 0.25 0.26 0.13 0.15 0.13 100 g

P ppm 4.00 4.00 2.00 417.58 21.85 36.00

Cu ppm 2.54 2.94 3.72 13.24 4.06 5.75

Fe ppm 273.45 219.54 335.61 175.65 147.69 336.79

Zn ppm 9.08 7.18 12.99 4.33 2.63 2.19

Species Pachy- Pachy- Pachy- Pachy- Phaseolus Calopogo- Calopogo- rhizus rhizus rhizus rhizus hetero- gonium gonium erosus erosus erosus erosus phyllus coeruleum coeruleum

Cult./ wild cult. wild wild wild wild wild wild

Loca­ Morelos Puebla Chiapas Guana- Michoacan Oaxaca Baj a lity caste Verapaz

Country Mexico Mexico Mexico C. R. Mexico Mexico Guatemala

pH 7.02 7.66 7.74 6.50 6.73 6.77 7.60

Conduc­ 538.7 281.8 265.7 147.5 289.1 90.5 140.2 tivity

Ca meq/ 53.25 23.20 46.59 13.25 7.11 3.89 10.24 100 g

Mg meq/ 13.30 10.90 6.42 7.24 9.29 1.98 5.66 100 g

K meq/ 2.23 1.07 0.85 1.31 1.69 0.30 0.18 100 g

Na meq/ 0.48 1.07 0.35 0.44 0.39 0.48 0.25 100 g

Mn meq/ 0.06 0.04 0.03 0.08 0.11 0.14 0.06 100 g

Ppp m 94.86 282.28 139.72 57.94 31.97 253.75 121.88

Cu ppm 2.35 7.64 1.29 4.09 2.77 1.97 3.21

Fe ppm 57.07 89.30 16.35 171.45 169.95 161.08 195.87

Zn ppm 9.29 6.24 2.38 2.19 7.14 19.19 1.98 1 1 Table 4. (soilanalysis )

24 WageningenAgric. Univ.Papers 90-3 (1990) tion area will have to be conducted in order to evaluate the overall tolerance of the species. The fact that wild forms of P. erosus is capable of tolerating soils of low fertility could possibly be of usei n future breeding. The soil samples from Phaseolus heterophyllus and Calopogonium coeruleum localities are included in order to serve as a comparison between other related genera and the wild material of Pachyrhizus. The most apparent difference is the high level of phosphorus of these samples when compared to P. ferrugineus samples;agai n the tolerance of P.ferrugineus is demonstrated.

Conclusions

Thefinding s ofth ebiosystemati c investigations have sofa r clearly demonstrated the considerable variation present within the genus despite the fact that only five species are recognized. Of the three cultivated species P. erosus, P. ahipa and P. tuberosus, the first two must be regarded as the most selected types when seen from an agronomic view, mainly due to the uniform appearance of their tubers and the non-twining habit of P. ahipa. Furthermore, these two species have the longest recorded period in cultivation. The status of P. tuberosus indi­ cate that this species has never been subjected to intensified selection, not to mention actual breeding; the major advantage of this species is its tolerance towards high precipitation rates. Although no interspecific crosses involving the two wild species have been carried out (thetw o specieshav eno t flowered), theinterfertilit y ofth e remaining speciesindicat e that incompatibility on the species levelca n be overcome. The possible extraction of insecticidal compounds present in the mature legumes (seeds)mus t await further studiesinvolvin g the two wild species. An encouraging observation, established through the interspecific breeding involving the three cultivated species (and a wild form of P. erosus), is the yield performance of the interspecific hybrids, both F, and F2. The high seed yields of these hybrids have apparently no negative effect on the yield performance. Furthermore, itha sbee n established that interspecific crossesinvolvin g the erect P. ahipawil lproduc e a number of plants exhibiting theerec t growth habit. In conclusion, the yam beans constitute potentially highly valuable tuber crops for awid e range ofclimati c and ecological conditions, due to the consider­ able variation within the genus. Both cultivated species as well as wild species and thewil dform s ofcultivate d speciesar ehig hyieldin g and exhibit remarkably great tolerances towards differences in soil fertility, altitude and variations in precipitation rate.

WageningenAgric. Univ.Papers 90-3 (1990) 25 Acknowledgements

The author isgratefu l toth e head and staff ofth e Botanical Laboratory, Univer­ sity of Copenhagen for good working facilities, to Dr lb Friis for his generous assistance, valuable comments and for correcting the English. I gratefully ac­ knowledge the kind cooperation of Professor L.J.G. van der Maesen who made the publication of thispape r possible.Th e cooperation of the curators who pro­ vided the material, and the institutions and individuals who have kindly pro­ vided live material, green house facilities, etc. needed to complete this study are likewise gratefully acknowledged. Finance for the investigation was provided by the EEC programme - STD I and STD II, subprogramme Tropical Agriculture, the Danish International Development Agency and the Danish Natural Science Research Council.

26 WageningenAgric. Univ.Papers 90-3 (1990) Listo fexsiccata e 1 Collections ofPachyrhizus speciesi n the natural distribution area. Abbreviations of the taxa - P. ahipa(Wedd. ) Parodi = ahi P. erosus(L. ) Urban = ero P.ferrugineus (Piper) Sorensen = fer P.panamensis Clausen = pan P. tuberosus (Lam.) Spreng. = tub (Hybrids between P. erosusan d P. tuberosuso r presumed cultivars of P. tubero­ sus = cfr. tub). Allneotropica l material isliste dexcep t for material of P. erosuswhic hi s marked 'cultivated' on thelabe lo fth especime nan d material ofP . erosusfro m theCarib ­ bean and Florida, i.e. material introduced to areas in the neotropics outside the original distribution area of this species.

Abbott, R.Q., 404 ero (GH); Aguilar, I., 113er o (F); Aguilar, J.I., 1712 ero (F), s.n./19 Sep 1943 ero (BH), Alduvin, C, 133 fer (MO); Alexander, E.J. & Hernandez X., E., 2030 ero (MICH), 2065 ero (MICH), 2072 ero (NY); Allen, P.H., 2027 pan (F, MO, NY, US); Anthony, H.E. & Tate, g.h.h., 49 pan (US); Andrieux, G., 492 ero (G), s.n./1833 ero (G), s.n./1833 ero (G), Asplund, E., 15461 tub (S), 15854 pan (G, NY), 19837 tub (S); Aureliano Juan & Marcelo, A., 4 fer (XAL); Bang, A.M., 1430 tub (ARIZ, BM, F, G, GH, MICH, NY); Barientos, L., s.n./1943 ero (BH); Bartlett, H.H., 11664fe r (MICH, NY, US), 12407fe r (F,MICH) , 12799fe r (MICH);Bellini , CF501 fer (K); Bernoulli, CG. & Cario, R., 1162 ero (K); Bornemicza, C, 34451 ero (CR); Bourgeau, E?, 2917 ero (C, G, GH, K, S), 3171 ero (G, GH); Breckon, G.J. & Breckon, M.E., 2163 ero (F, NY); Breedlove, D.E., 6465 ero (MICH, NY), 7441 fer (F), 7575 fer (F, MICH, US) 7656 fer (F, MICH), 7698 fer (F), 10267 ero (F, MICH), 11838 ero (F, MICH, US), 20131 ero (MO), 28571 ero (MO, NY); Broadway, W.E., s.n./10 Jan 1916 fer (BH), 5000 fer (BM, NY), 7572 fer (NY), s.n./29 Nov 1932 tub (BM); Bunting, G.S. & Licht, L., 645 fer (F, US), 677 fer (F, NY); Burchell, W.J., 6742 cfr. tub (K); Burkart, A., 882/Herb. Darw. No. ahi (SI), 13314 ahi (K, P); Calderon, G.M., 1309 ero (F, GH, MICH, MO, NY); Calderón, S., 766 ero (GH, NY), 1108er o (US), 2258 ero (US); Callaway, R., 306 ero (US); Calzada, J.I., 3467 (coll. Calzada, J.I. et al.) ero (XAL), 3900 (coll. Calzada, J.I. et al.) ero (XAL), 4071 (coll. Calzada, J.I. et al.) ero (XAL); Carreira, L., 146 eft. tub (INPA), 147 tub (INPA); Castillo C, G., 872 (coll. Castillo C, G. & Tapia, L.) ero (XAL), 2970 (coll. Castillo C, G. & Medina, M.E.) ero (XAL); Chanek, M., 75 fer (MICH); Clarke, O.F., 444 ero (NY); Clausen, R.T., C42-40 ahi (BH, C), C43-25 tub (BH, C), 6048 (coll. Clausen, R.T. et al.) ero (BH), 6049 (coll. Clausen, R.T. et al.) ero (BH), 6098 (coll. Clausen, R.T. & Cervantes G., R.) ero (BH), 6118 (coll. Clausen, R.T. & Cervantes G., R.) ero (BH, MO), 6130 (coll. Clausen, R.T. et al.) ero (BH, UC), 6130A (coll. Clausen, R.T. et al.) ero (BH), 6150 (coll. Clausen, R.T. & Ippisch, F.) ero (BH), 6158 (coll. Clausen, R.T. & Ippisch, F.) ero (BH), 6158 (coll. Clausen, R.T. & Ippisch, F.) ero (BH), 6159 (coll. Clausen, R.T. et al.) ero (BH), 6159A (coll. Clausen, R.T. et al.) ero (BH), 6166 (coll. Clausen, R.T. et al.) ero (BH), 6169 ero (BH), 6171 fer (BH), 6173 fer (BH), 6175 fer (BH, MO), 6182 fer (BH, MO), 6183 fer (BH, MO), 6190 ero (BH), 6193 (coll. Clausen, R.T. & Kovar, A.) ero (BH), 6198A (coll. Clausen, R.T. Kovar, A.) ero (BH), 6201 (coll. Clausen, R.T. & Kovar, A.) ero (BH), 6202 (coll. Clausen, R.T. & Kovar, A.) ero (BH), 6205 (coll. Clausen, R.T. & Kovar, A.) ero (BH), 6207 (coll. Clausen, R.T. & Kovar, A.) ero (BH), 6209 (coll. Clausen, R.T. & Kovar, A.) ero (BH), 6212 (coll. Clausen, R.T. & Kovar, A.) ero (BH), 6214 (coll. Clausen, R.T. & Cervantes G., R.) ero (BH), 6222 (coll. Clausen, R.T. & Cervantes G., R.) ero (BH), 6223 ero (BH); Contreras, E., 5598 fer (F, MICH, US); Cortés, A., 31 tub (COL); Croat, T.B., 469 ero (NO), 22071 fer (C, F, MO); Cuatrecasas, J., 23073 tub (F); D'Arcy, W.G., 9721 fer (MO), 12061 ero (MO, NY); Darwin, S.P., 2363 ero (CR); Delgado

WageningenAgric. Univ.Papers 90-3 (1990) 27 S., A. et al., 695 ero (F), 822 ero (F. MO); Diguet, L., s.n./1959 ero (MICH, P); Dodds, D., 31 ero (MICH); Dodson, C. & McMahon, M., 4344 tub (QCA); Dodson, C.H., 5446 tub (US); Dor­ antes, J. et al., 4130 fer (XAL); Dunn, D. et al., 32 ero (NY); Duss, P.A., 1086 fer (NY); Dwyer, J.D. & Liesner, R., 12028 fer (MO), 12236 fer (MO), 12240 fer (MO), 12245 fer (MO); Eiten, G. & Eiten, L.T., 10637 cfr. tub (MO, US); Fendler, A., 74 fer (MO), 75 fer (F, GH, K); Ferreyra, R. et al., 10865 tub (US); Field, H., s.n./Sep 1946 ero (F); Flores, J.S., 10101 ero (XAL); Forero, E. et al., 1773 fer (COL, MO); Fraser, L., 216 pan (BM); Freyreis, G.W., s.n./s.a. cfr. tub (S); Gadow, H. & Gadow, M., s.n./1902-1908 ero (BM); Galeotti, H., 338S ero (BR), 3167 ero (BR, G(, 3278 ero (BR); Gaumer, G.F., 812 ero (F, GH, MICH), 1977 ero (C, F, GH, MA, S), 23409 (coll. Gaumer, G.F. & sons) ero (F, G, GH, MA), 23426 (coll. Gaumer, G.F. & sons) ero (F, G, GH, MA), 24161 ero (F, G, GH, MA, MO), 24282 ero (F, G, GH, MA), 24437 ero (F); Gentle, P.H., 272 ero (F), 559 ero (F, MICH), 622 fer (F, MICH), 1046 fer (MICH, NY), 1047 fer (F, MICH, NY, S), 1815fe r (F, GH, MICH, NY), 2279 fer (F, GH, K, MICH), 2431 fer (GH, MICH), 3118 fer (MICH), 7854 fer (S), 8063 fer (S), 8087 fer (S, US), 8502 fer (S), 8513 fer (S), 9469 fer (S); Gentry, H.S., 22379 ero (GH); Ghiesbreght, A.B., 595 ero (BM, GH); Gilly Sr., CL., 32 ero (MICH); Gilmartin, A.J., 653 pan (US); Gómez-Pompa, A., 1704 ero (XAL); Greenman, J.M. & Greenman, M.T., 5053 pan (MO); Guerrero, C, B. et al., 448 ero (XAL); Gurgel, 195 tub (SI); Hagen, C. von & Hagen, W. von, 1337 fer (F, NY); Hallberg, B., 990 ero (MICH); Hansen, B. et al., 7945 tub (MO); Harling, G., 3547 tub (S); Harling, G. & Andersson, L., 11589 tub (GB), 14047 tub (GB, MO); Harmon, W.E. & Dwyer, J.D., 3199 ero (MO), 3683 ero (MO), 3852 ero (MO); Harmon, W.E. & Fuentes, J.A., 4723 ero (MO); Harris, J.I., 90 (MICH); Harris, W., 3180 tub (NY);Hazlett , D., 763er o (MO); Heithaus, E.R., 366er o (F, MO, NY); Herb. Dir. Gen. Agric. Guatemala, s.n./10 Apr 1943 ero (BH); Heriberto, Bro., 117 pan (GH, US); Heringer, E.P., 992 tub (SP);Hernande z X., E., 2030er o (F, UC); Hernandex X., E. et al., ES-239 ero (MICH); Heyde, E.T. & Lux, E., 3306 ero (G), 3729 ero (G, GH, US), 3759 ero (G, GH, US); Hinton, G.B., 1205 ero (G, NY), 4424 ero (G, GH), 9367 ero (BM, F, G, GH, MO, NY), 9540 (coll. Hinton, G.B. et al.) ero (BM, F, G, K, MO, NY, US), 11486 (coll. Hinton, G.B. et al.) ero (GH, NY, US), 11776 (coll. Hinton, G.B. et al.) ero (GH, MICH, NY, U, US), 13152 (coll. Hinton, G.B. et al.) ero (F, GH, MICH, NY, S, U, UC, US), 14478 (coll. Hinton, G.B. et al.) ero (F, GH), 14673 (coli. Hinton, G.B. et al.) ero (GH); Holm-Nielsen, L. & Jeppesen, S., 34 pan (AAU); Hunnewell, F.W., 18277 ero (GH); Hunt, D.R., 260 fer (BM, F); Iltis, H.H. & Koeppen, R.C., 1267 ero (UC); Innes, R.R., 166 fer (MO); Irby, H.D. & Rowell Jr., C.M., 3597 ero (MICH); Irwin, H.S. et al. 21114 tub (COL, F, K, MICH, NY, US); Jameson, W., s.n./s.a. pan (K); Janzen, D.H., s.n./20 Nov 1963er o (MICH, UC), s.n./4Ja n 1964 ero (MICH, UC), 10142/20 Sep 1970 ero (MO), 17/14 Nov 1971 ero (MO), T-l 11/Dec 1976 ero (MO), 10635/3 Dec 1977er o (MO); Jéllez, O. & Cabrera, E., 1687 fer (BM, MO); Johnson, CD., 215-68 ero (MO); Johnson, H., fer (F); Jones, M.E., 213 ero (MO); Juan, A. & Marcelo, A., 4 fer (XAL); Kayap, R., 45 tub (F, MO); Kellerman, W.A.,

7260 fer (F, NY); Kenoyer, L.A., s.n./18 Jul 1938 ero (ARIZ), A187/18 Jul 1938 ero (F), C248 ero (GH); Kerr, W., 53497/6 Nov 1975cfr . tub (INPA); Killip, E.P., 12080 pan (C, GH, NY, US); Klug, G., 885 tub (NY), 2225 tub (BM, F, GH, MO, NY, S, US);Kriegel , O., 5140tu b (SP); Lang­ lasse, E., 137 ero (K), 323 (G, GH), 436 ero (G, GH), 437 ero (G, GH); Laughlin, R.M., 1281 ero (US), 2853 fer (US); Lazier, J. & Keoghan, CF408/8 Mar 1976 ero (K); Lazier, J., CF515/30 May 1976 fer (K); Leibert, R.F., 1738 tub (US); Lent, R.W., 2483 fer (F, MO); León, H., 458 fer (MO); Lewis, W.H. et al., 29 pan (MO), 102 fer (GH, MO, US); Liebman, F.M., 5261 ero (C), 5262 ero (C), 5264 ero (C), 5264A ero (C), 5265 ero (C); Long, L.E., 222 fer (F); Luirós, M., 1320A fer (F); Lundell, CL., 122-79 ero (MICH), 12313 (coll. Lundell, CL. & Lundell, A.A.) ero (MICH, NY), 12423 (coll. Lundell, CL. & Lundell, A.A.) ero (MICH, NY, UC, US), 12575 (coll. Lundell, CL. & Lundell, A.A.) ero (MICH); MacDougall, T., s.n./Sep 1969 ero (NY); Macedo, A., 1052cfr . tub (NY, US), 1674 cfr. tub (BM, MO, S, SP); Madison, M.T. et al., 5465 tub (AAU, NY, QCA); Mandon, G., 747 ahi (P); Marshall, S.A. & Neill, D.A., 6479 fer (F, GH); Martinez, M., 15011 ero (BH); Matthews, A., 3258 tub (BM, GH, K), s.n./1840 tub (S); Matuda, E., 1464 ero (MICH), 17035er o (F); Maxon, W.R., 7685er o (US); McVaugh, R., 16059er o (MICH), 18548 ero (MICH), 25546 ero (MICH); Melhus, I.E. & Goodman, G.J., 3656 ero (F); Messer, E., 206 ero (MICH); Mexia, Y., 1295 ero (G, GH, MO, NY, UC, US), 4700 tub (US), 6717 tub (F, UC), 7261 tub (GH, UC, US), 7310 tub (UC); Mille S.J., L., 962 pan (F); Molina R., A., 1149 ero (F,

28 WageningenAgric. Univ.Papers 90-3 (1990) GH), 1809 fer (F), 2952 fer (F, GH), 3196 fer (F), 4129 fer (F, GH), 10487 fer (F, NY), 13591 fer (F, NY), 14628 ero (F, NY), 18197 (coll. Molina R., A. et al.) fer (F, MO, NY), 22450 ero (G, MO, NY), 22935 ero (NY), 23291 fer (F), 24620 (coll. Molina R., A. & Molina, A.R.) fer (F, NY), 26464 (coll. Molina R., A. & Molina, A.R.) ero (MICH, U), 31077 (coll. Molina R., A. & Molina, A.R.) ero (F, MO); Montogomery, G., s.n./30 Jun 1976er o (MO); Moore Jr., H.E., 3er o(BH ,GH) ; Morales R., J., 1173 ero (F);Mosquin ,T .e t al.,670 7er o (GH);Muller ,F. , s.n./1855 ero (NY); Nagel, O., 8076 ero (GH); Nee, M., 7774 pan (MO); Neill, D., 929 ero (US); Nelson, C. & Clewell, A., 0333 fer (MO, NY); Nelson, C. & Vargas, E., 2699 fer (MO); Nelson, E.W., 2882er o (GH), 3002er o (GH), 3248 fer (GH); Nunez, J.S.C., 216er o (MO);Oersted , A.S., 33/1851 ero (K); Opler, P.A., 1909er o (F, MO, UC); Orcutt, CR., 4182 ero (BM, F), 4427er o (F); Orozco, J.M., 282 ero (F); Ortiz, R.T., 53 fer (F, UC), 1532 fer (F, MO, NY), 2215 fer (BM, F, MICH); Palmer, E., 318 ero (BM, GH, K, MO, NY); Parodi, L.R., 11901 (coll. Ragonese, A.) ahi (BAA), 12145 (coll. Ragonese, A.) ahi (BAA), 12146 ahi (BAA), 12217 ahi (GH, K); Pavón y Jiménez, J.A., s.n./s.a./Herb. Pavón ero (G), s.n./1827 (see Note 1)er o (G); Peck, M.E., 963 fe (GH); Pelly, R.S., 22 fer (F); Penland, C.W. & Summers, R.H., 126 tub (GH); Pfeifer, H.W., 1628 ero (US); Philcox, D. & Freeman, B., 4658 cfr. tub (K, NY); Pilz, G.E. & Pilz, M., 1317 ero (MO); Pittier, H., 280 fer (NY), 2918 fer (GH, NY), 3152 fer (CR), 12169 fer (US), 14405 tub (F, VEN); Pringle, CG., 6501 ero (BM, F, G, GH, S);Proctor , G.R., 27386 (coll. Proctor, G.R. et al.) fer (F), 35861 fer (MO); Purpus, CA., 1891 ero (F, GH, NY, UC, US), 1892 ero (F, GH, US), 8079 ero (ARIZ, UC), 9260 ero (F, GH, MO, NY), 10163 ero (NY, US); Record, S?J?, s.n./1926 fer (US); Rieko, B.P., 3623 ero (US); Robles H., L., 360 ero (XAL); Rocha, G.L. da, s.n./Mar 1944 cfr. tub (SP); Rodriguez, J.V., 818 ero (F), 1019 ero (F), 2993 fer (F), 3386 ero (F, UC); Roe, K. & Roe, E., 2262 ero (MICH); Rose, J.N., 5347 (coll. Rose, J.N. & Painter, J.H.) ero (US), 6578 (coll. Rose, J.N. & Painter, J.H.) ero (C, F, GH, US), 6904 (coll. Rose, J.N. & Painter, J.H.) ero (GH, US), 8540 (coll. Rose, J.N. et al.) ero (GH), 9275 (coll. Rose, J.N. et al.) ero (GH), 9346 (coll. Rose, J.N. et al.) ero (GH), 11072 (coll. Rose, J.N. & Rose, J.S.) ero (US), 22581 (coll. Rose, J.N. & Rose, J.S. (tub (NY); Rzedowski, J., 22675 ero (MICH); Salazar, F., s.n./4 Oct 1914 ero (US); Santos, J.V., 2619 ero (MICH); Schipp, W.A., 718/21 Mar 1930 fer (F, G, GH, MICH, NY, S), S-280/4Ja n 1932fe r (F, G, GH, MICH, NY, S);Schomburgk , M.R., s.n./1840-1844 tub (K); Schu­ bert, B.G. & Gomez Pompa, A., 1695 fer (GH); Seeman, B.C., 211 pan (BM, K); Seibert, R.J., 1738 tub (MO); Seier, C. & Seier, E., 2686 ero (GH), 3095 ero (GH); Sessé, M. et al., 3722 ero (F); Shemluck, M., 312 tub (F); Sintenis, P., 5689 tub (BM, GH, K); Skutch, A.F., 2427 fer (GH, GH, K, MICH, NY, US, S), 5311 fer (MICH); Smith, CL., 1310 ero (F, MICH); Smith, H.H., 2503 pan (F, GH, NY); Smith, J.D., 2313 ero (GH, US); Smith, L.C., 99 ero (GH); Sohns, E.R., 1593fe r (MICH); Solano, S.C. & Vara, M.A., 179er o (F, MO, NY); Solomon, J.C, 617 ero (CR), 2422 ero (MO); Sousa, M., 3125 ero (GH, NY), 4215 (coll. Sousa, M. et al.) ero (GH), 4777 (coll. Sousa, M. & Ramos, C.) ero (GH, MO), 96755 (coll. Sousa, M. et al.) ero (MO); Sparre, B., 13805 tub (S), 14791 tub (S), 17245 tub (S), 18475 tub (S); Spruce, R., 4936 cfr. tub (K), 6359 pan (BM, K); Standley, P.C., 25200/26 Nov-9 Dec 1923 (see Note 2) pan (S), 25271/27 Nov-10 Dec 1923 pan (S), 27281/17 Dec 1923 fer (S, US), 28798/28 Dec 1923 fer (US), 30213/9 Jan 1924 fer (US), 72385/26-27 Apr 1939fe r (F), 83885/9 Oct 1940er o (F), 74500/15 Oct 1940er o (F), 75049/24 Oct-5 Nov 1940 ero (F), 76676/7-18 Nov 1940 ero (F), 77451/16 Nov 1940 ero (F), 77939/20-27 Nov 1940er o (F),78849/3 0No v 1940er o(F) ,79434/ 6De c 1940er o(F) , 1172/11De c 1946(coll . Standley, P.C. et al.) fer (F), 6454/25 Mar-5 May 1947(coll . Standley, P.C. & Chacon, J.) fer (F), 7743/22-27 Apr 1947 fer (F), 9462/4-13 Jun 1947 ero (F), 9672/19 May-9 Jul 1947 ero (F), 10950/19 Jun-9 Jul 1947 ero (F), 11592/3-17 Aug 1947 ero (F), 12569/22 Sep 1948 ero (F), 12749/Oct-Nov 1948 ero(F) , 12895/6Oc t 1948er o (F), 13123/18Oc t 1948er o (F), 13922/Oct-Nov 1948er o (F), 16976/21 Feb 1949 fer (F), 17545/5-16 Mar 1949 fer (F), 18226/19 Mar 1949 fer (F), 21331/16 Jul 1949 ero (F), 22258/6Au g 1949er o (F,NY) ; Stevens,W.D. , 1293er o (MICH, MO);Steyermark , J.A., 29481 ero (F), 29625 ero (F), 38311 fer (F), 38450 fer (F), 38553 fer (F), 41833 fer (F), 42969 ero (F), 44047 fer (F),4411 3fe r (BH, F), 45290 fer (F),4970 5 fer (F), 52701 tub (F);Tessmann , G., s.n./s.a. cfr. tub (NY), 3269 cfr. tub (G), 3447 cfr. tub (G, NY), 5439 cfr. tub (NY, S), 5456 cfr. tub (G, NY); Thieme, C, 5205 fer (G); Ton, A.S., 1600 fer (US), 3128 ero (F, NY), 3157 ero (F, MICH), 3308 fer (NY); Tonduz, A., 4147 fer (CR), 4764 fer (G), 12889 fer (BM); Tuerckheim, H. von, II 1671 fer (US), 7617 fer (GH, NY); Tyson, E.L. & Blum, K., 2534 pan (MO); Ucân, E. et al.,

WageningenAgric. Univ.Papers 90-3 (1990) 29 2839 ero (XAL); Valdivia Q., P.E., 2349 ero (XAL); Vazquez, B., 1297 ero (BM, MO); Vazquez, M. et al., 1669fe r (XAL); Ventura A., F., 2880 ero (F, MICH, NY); Ventur, P., 143fe r (F);Vrieze , J.M., 4344 tub (QCA); Wall, E., 429 ero (S); Weberbauer, A., 6416 tub (F, GH); Webster, G.L. et al., 17M°926 ero (F); Wedel, H. von, 2981 fer (GH, MO); West, J., 3540 ero (GH, MO, UC); Williams, L., 416 tub (F); Williams, L.O. & Molina R., A., 10813 ero (F, GH), 13240 ero (F, GH), 13300 ero (F, GH, MO), 16955 ero (F); Williams, L.O. & Williams, R.P., 18332 ero (F, GH); Yuncker, T.G. et al., 5608 fer (MICH); Zelaya, L., 233 ero (MO); Zola B., M.G., 793 ero (XAL); Œrsted, A.S., 5263er o (C, F).

Note 1.Accordin g to Stafleu and Cowan (1983) specimens from the herbarium of Pavón yJimene z marked: NE werecollection s made by Sessé& Moçifi o in Mexico.

Note 2. The collections made by P.C. Standley have two series of numbers. In order to avoid any confusion between the two series the date of collection as stated on the specimen label is given with the number.

LIST OFEXSICCATA E 2

Collections of P. erosusfro m outsideth e natural distribution area ofthi sspecies .

1. NORTH AMERICA

1901-1950 Buswell,W.M. , s.n./7Oc t 1943Florid a BH-s.n./ll Oct 1943 Flor­ ida BH, F, MO, UC - Clausen, R.T. & Buswell 6227/26 Oct 1943 Florida BH - 6236/27 Oct 1943 Florida BH - Lynch, S.J., (coll. Calkins, O.W.) 1/10 Nov 1943 Florida BH - (coll. Krome, W.M.) 2/10 Nov 1943 Florida BH - Piper, C.V., s.n./Mar 1923Florid a US-Watson , M., s.n./21 Sep 1943Californi a BH.

1951- Plowman, T., 13277/19 Dec 1983Florida C.

2. CARIBBEAN

-1850 Agron., s.n./Apr 1820 Antilles G - Benzon, P.E., s.n./1817-1848 St. Croix C- Forsström, J.E., s.n./1803-1815 St. Martini S- Hornbeck, H.B., s.n./ 1825-1844 St. Thomas C - Krebs, HJ., s.n./1843-1870 St. Thomas NY - Mas- son, F., s.n./[1779-1781] West Indies BM - Ravn, P., s.n./1829 St. Croix C - s.n./1828-1839St .Croi x C- Rugel, F.,629/184 9Cub a BM- Ventenant, (Herb.) E.P., s.n./s.a. [1757-1808] St. Domingue G - s.n./s.a. [1757-1808] Puerto Rico C, G - [West, H.,] Ex Herb. Richard, L.C.M., s.n./s.a. [1758-1811] St. John, Tortola, Martinique P.

1851-1900 Belanger, C.P., 567/Jul 1853 Martinique G, P - 715/1860 Martini­ que P - Duss, P.A., 1085/1883 Martinique NY - 3465/1894 Guadeloupe F, NY - 3686/1895 Guadeloupe NY - Fendler, A., 313/1877-1880 Trinidad BM, K,

30 WageningenAgric. Univ.Papers 90-3 (1990) NY - Goll, G.P., 155/10 Nov 1899 Puerto Rico NY, US - Hahn, L., 194/Nov 1867 Martinique BM, G - 1219/Nov 1869 Martinique BM, G, GH - 1316/Nov 1870 Martinique G - Sintenis, P., 5581/4 Dec 1886 Puerto Rico BM, C, G, GH, K, S, US - Smith, H.H. & Smith, G.W., 598a/ Jul s.a. (Reed. 1890) St. Vincents GH - 1916/Jun (Reed. 5 Oct 1890) St. Vincents BM - Wright, C, 2338/1860-1864 Cuba BM, G, GH, NY, S.

1901-1950 Allard, H.A., 13565/22 Nov 1945 Dominican Republic US - 14467/7Ja n 1946Dominica n Republic GH, S,U S- 16100/14Oc t 1946 Domini­ can Republic US - 16227/19 Oct 1947 Dominican Republic F, MO, NY, S - Baker, CF., s.n./l Oct 1907 Cuba UC - Britton, N.L. & Cowell, J.F., 588/8 Sep-5 Oct 1901 St. Kitt's NY - Cobin, M., 16177(Herb. B.A. Krukoff no. 3R4943)/Recd. Mar 1943Puert o Rico U-Ekman, E.L., 2304/28 Oct 1924 Haiti S - 5098/13 Oct 1925 Haiti S, US - H15780/27 Jul 1930 Dominican Republic S Gonzales, A., 145/15 Sep 1941 Cuba BM, GH, MICH, NY - 270/9. Oct 1941 Cuba BM, F, GH, MICH, NY, S- Kosel, G.W., (coll. Fairchild, D. & Dorsett, P.H.) P.I.97754/8 Nov 1943 (Reed. 29 Feb 1932) Antigua BH - Leon, Bro.J.S.S. ,677/1 5Oc t 1908Cub a NY-s.n./l 5Oc t 1908Cub a NY- 14238/Oct 1929Cub a GH-Leonard, E.C., 7173/18No v 1925Haït i NY, US-Nash, G.V., 677/3 Aug 1903 Haïti F, NY - Nash, G.V. & Taylor, N., 1789/14 Aug 1905 Haïti NY - Stehle, M. & Stehle, H., 4807/5 Sep 1939 Martinique US - Stehlé, 5973/8 Dec 1941Martiniqu e P- Stevenson, J.A ,2272/1 2Oc t 1914Puert o Rico NY, US-Wilson, 3520/18 Oct 1905Cub a F, NY, UC.

1951- Augusto, Bro. B., 1068/2 Nov 1963 Hispaniola NY - Liogier, A.H., 20224/20 Sep 1973 Dominican Republic - 30354/29 Jan 1980 Puerto Rico NY - Powell, D.A., 862/16 Oct 1960 Jamaica NY - Proctor, G.R., 8189/15 Nov 1953 Jamaica US - 22824/7 Oct 1962 Jamaica MICH, NY, U - 25579/29 Sep 1964Jamaic a MICH, U.

3. SOUTH AMERICA

1901-1950 Pittier, H,F., 15203/1942 Venezuela G, VEN.

4. AFRICA

-1850 Leprieur, F.R., s.n./Sep 1828 Senegambia G - Perrotet, G.S., 221/1824-1829 Senegal BM.

1851-1900-

1901-1950 Brouns, 14/1933 Belgian Congo BR - Corbisier-Baland, A., 1045/20 Apr 1931 Belgian Congo BR - 1997/5 Aug 1933 Belgian Congo BR - 1999/1933 Belgian Congo K - Deighton, F.C., 2700/2 Jun 1933 Ghana K - Kosel, G.W., (coll.Maitland , T.D.) P.I.88365/8No v 1943(Reed . 17Ju n 1930)

WageningenAgric. Univ.Papers 90-3 (1990) 31 Cameroons BH - Sierra Leone K - Irvine, F.R., 4791/1924-1939 Laurent, E. & Laurent, M.D.J., s.n./1903-1904 Belgian Congo BR - s.n./31 Dec 1903 Bel­ gian Congo BR, G - Laurent, M.D.J., 1207/1 Oct 1909 Belgian Congo - Peter, A., K53A/5 May 1914 German East Africa (Tanzania) BR, K - Vanderyst, H., s.n./22 Feb 1907 Belgian Congo BR - Zeemant, 324/1936 Belgian Congo BR.

1951- -

5. INDIA andINDIA N OCEAN

-1850 Hooker, J.D., s.n./(1847-1852) India K - Kamphövener, B.C., 633/1845-1847 India C - Roxburgh, W., s.n. (Hort. Calcutt. 5526C)/s.a. [1751-1815] India - 1851-1900 Blackburn, J., s.n./17 Jul 1863 Mauritius or Madagascar NY - Bouton, L., s.n./(Recd. 1864) Mauritius K - Clarke, C.B., 13780/23No v 1870Indi a BM-42410A/2 Dec 1885Indi a K-Herb. Hort. Bot. Calcuttensis s.n./1865Indi a G-s.n./Nov 1896Indi a G- Parry, N.E., 1073/Nov 1879Indi a K - Prain, D., (coll. Mani) 4/Dec 1900Andaman s G.

1901-1950 Parkinson, CE., 4306/28 Nov 1934 India (Bangladesh) S- Prain, D., 44/20Ja n 1901Andaman s G.

1951- -

6. THEFA R EAST

-1850 Burmann, J., s.n./1707-1779 Indonesia G - s.n./1707-1779 Indonesia (Java) G - s.n./1759 Indonesia (Java) G - Cuming, H., 650/1841 Philippines BM, G, K, MO - Didrichsen, D.F., 3657/1845-1847 Malaysia C - Horsfield, T., s.n./1802-1818 Indonesia K - s.n./(1802-1818 ) Indonesia BM - Römer, von, s.n./1841 Indonesia G - Thunberg, C.P., 13/1777 Indonesia S - Voigt, J.O., 4/s.a. [1798-1843] Malaysia C - s.n. (Herb. no. 24)/s.a. [1798-1843] Malaysia C - s.n. (Herb. no. 25)/s.a. [1798-1843]Indi a C - Wallich, N., 5526A,B,C,D/22 Nov 1814Indi a BM- Zollinger, H., 2767/s.a. [1818-1859]Indonesi a BM, G.

1851-1900 Balansa, B., 1181/Aug 1885 Viet-Nam GH, P - 1182/1885-1889 Viet-Nam K,P - 2279/Oct 1887Viet-Na m K, P, US- 4734/1890-1891 Viet-Nam G, K, P - Bodinier, E., 17s8/9 Aug 1897 China P - Curtis, C, 1211/Sep 1887 China K-Delavay, P.J.M., 2983/3Oc t 1887Chin aP - 3529/28 Aug 1888 China P- Dupuy, C, 73/Recd. 1899Lao sP - Forbes, H.O., 4110/1881-1882Indonesi a - Godefroy-Lebeauf, A., 720/7 Sep 1878 Viet-Nam P - Henry, A., 1093/Recd. Apr 1895 Taiwan K, MO, S 11005/Pres. 1898 China K - 12539/1896-1899 China NY, US - Junghuhn, F.W., 215/1854 Malaysia K - Loher, A., 2300/Sep 1891 Philippines K- 2301/Oct 1891 Philippines K - 2302/Nov 1891 Philippines K - 2303/16 Nov 1891 Philippines K, US - Pierre, L., 863/Oct 1871 Viet-Nam

32 WageningenAgric. Univ.Papers 90-3 (1990) F, GH, P-Ridley, H.N., s.n./1892 Singapore BM-Thorel, s.n./1862-1866Viet - NamG.

1901-1950 Ahern, G.P., For. Bur. no. 3324/Sep 1905Philippine s K, NY-Arr- henius, O.W., s.n./Apr 1921 Indonesia S- Bakhuizen van der Brink Jr., R.C., 2462/28 Mar 1923 Indonesia U - Barnes, P.T., 11135/Oct 1903 Philippines K, NY - Bloembergen, S., 3539/21 Mar 1939 Indonesia K - Borden, T.E., 24037/Sep-Dec 1904Philippine s F, K, NY - Bradley, C.B., s.n./1908? Thailand UC- Bünnemeyer, H.A.B.W :11340/ 4Ma y 1921Indonesi a K, U- Ching, R.C., 22159/16 Nov 1939 China GH - Chiu, S.F. & Lin, S., H77/22 Oct 1940 China BH - H86/23 Oct 1940Chin a BH - Chun, N.K. &so , C.L., 43602/19 Aug 1932 China MICH, MO, NY, S - Chun, W.Y., 3076/17 Aug 1929 China F, NY - Chung, H.H., 1057/1922 China UC - 3001/17 Aug 1924 China GH, UC - Cle­ mens, M.S., 6997/Sep 1925 Philippines UC - Dahlström, E., 487/10 Sep 1950 China S- Ducloux, F., 6465/22 Aug 1904Chin a P- Dizon, L., 11/23Au g 1950 Philippines US - Evrard, F., 2447/2 Nov 1925Viet-Na m P - Feureu, & Heide, F.F.R., s.n./1921 Indonesia S- s.n./1922 Indonesia S- Fox, R.B., 125/28 Jul 1948 Philippines MICH - Franck, C.W., 155/Oct 1929 Thailand C - Gates, F.C., 7078/29 Sep 1913Philippine s F - Groff, G.W., Herb. no. 5238/8 Oct 1919 China GH - Henry, A., s.n./Purch. 1901 Taiwan NY - s.n./Purch. 1901 China NY - s.n./s.a. Taiwan NY - Hosseus, C.C., 30/30 Sep 1904 Thailand BM, K -Iboet, 189/7Ap r 1925Indonesi a U- Kerr, A.F.G., 2145/30 Sep 1911Thailan d BM - 3836/23 Oct 1919 Thailand BM, C, K - Kjellberg, G., 1202/3 Apr 1929 Indonesia S - Lambert, W.H. & Brunson, L.W., 91/Sep 1945 Philippines US - Lau, S.K., 4214/1-29 Aug 1934 China BM, G, GH - Lei, C.I., 771/21 Jun 1933 China BH, F, MICH, NY, UC - 908/3 Aug 1933 China BH, K, MICH, NY, UC Levine, CO., 1263/30 Jul 1917 China BM, GH, MO - Liang, H.Y., 63168/29 Sep 1933Chin a MICH, NY - MacDaniels, L.H., 52/13Ap r 1943Phi ­ lippines BH,G H- Maire, E.E., Ghe28 SérieB/Oc t 1910Chin a NY, UC- Maire, R.P., 89/Oct 1906 China UC - McClure, F.A., 1773/29 Oct 1925 China UC - 1749/30 Oct 1925Chin a UC - 20603/27 Jul 1937Chin a BH, K, UC - McGre­ gor, R.C., s.n./Nov 1924 Philippines UC - Merrill, E.D., 3098/Oct 1903Philip ­ pines BM, K, NY, US - 570/Sep 1913 Philippines BM, F, GH, K, MO, NY - Nedi, 459/20 Oct 1937 Indonesia (Moluccas Isl.) GH - Ngeou, P., 1478/22 Aug 1909 China UC - Ping, T.K., 11399/11 Oct 1923 China NY - Poilane, E., 9183/15 Dec 1927 Viet-Nam AAU - 14299/20 Oct 1927 Cambodia AAU - 27979/11 Oct 1928 Laos AAU, P - Ramos, M. & Edano, G., s.n.(Herb. no. 49639)/10No v 1927Philippine s BR, UC- Silva,C , s.n./25Se p 1927Philippine s BH - Smith, H.C., 16287/21 Oct 1936 Burma K - Steward, A.N. &Cheo , C.C., 1121/10 Oct 1933 China GH, NY - Tak, T.W., 131/22 Jul 1927 China UC - 720/15 Jun 1928 China UC - Tak, T.W. & Chow, W.K., 2347/2 Oct 1926 UC - Toroes, R.S., 2152/28 Mar-10 May 1932 Indonesia MICH, NY - 2605/28 May-26 Jun 1932 Indonesia MICH, NY, UC - Tsang, W.-T., 131/21 Jul 1927 China GH, NY - 720/15 Jun 1928 China GH - 26733/25-30 Aug 1936 China C, K - Tsang, W.-T. & Chow, W.K., 2347/2 Nov 1926 China GH - Tsiang,

WageningenAgric. Univ.Papers 90-3 (1990) 33 Y., 7098/13 Sep 1930 China GH, NY - 8546/15 Jul 1930 China GH, NY - Vidal,J.E. , 655B/17Oc t 1948Lao sP - Walsh, CG., 171/2 0Ma r 1929Indonesi a BM - Wang, C, 34833/23 Oct 1933Chin a NY - 34940/31 Oct 1933Chin a NY -Wilson, E.H., 3807/1910-1911 China K.

1951- Charoenphol, Ch. et al., 4462/2 Nov 1970 Thailand AAU - Conklin, H.C., 641/2 Oct 1953Philippine s GH - Conklin, H.C. &Buwaya , 1-846/19 Dec 1962Philippine s GH - Kalkman, C, BW3799/11 Aug 1957Dutc h New Guinea GH, K- Maxwell,J.F. , 73-491/14 Oct 1973AA U - Pancho, J.V., 1125/Jul 1953 Philippines BH - Pottier, 532B/27 Dec 1969 Laos P - Tixier, M.P., s.n./16 Oct 1955Lao sP .

7. OCEANIA

-1850 Gaudichaud-Beaupré, C, s.n./1819-1820 The Marianas G.

1851-1900-

1901-1950 Degener, O. & Wiebke, H., 2282/25 Aug 1927 Hawaii NY - Degener, O., 17985/Sep 1927 Hawaii NY, UC - Fosberg, F.R., 24869/8-11 Jun 1946Th e Marianas (Tinian) NY, US - 25385/4 Jul 1946Th e Marianas (Guam) US - 47419/22 Aug 1965 The Carolines T (Palau) GH, NY, US - Meebold, A.K., 12,128(Degener's)/Oc t 1935Hawai i NY-Moore, J.W.,681/2 1 Mar 1927 Society Islands (Raiatea) MO, U.

1951- McKee, H.S.,42606/2 8 May 1985Nouvell e Caledonie AAU-42671/13 Jul 1985Nouvell e Caledonie AAU.

34 WageningenAgric. Univ.Papers 90-3 (1990) Appendix 1.

Appendix La. (composition of standard soil) Appendix l.b. (composition of standard nu­ trient solutions) pH: 5.8; Electric conductivity (EC): l-2mS/cm

Ingredient type/amount Nutrient eq

Basal compound peat and clay granules Nitrogen (N) 10.0000

Granules fine to medium Phosphorus (P) 1.2082

Turn over rate 45-65 Potash (K) 3.2692

PH 5.5-6.5 Calcium (Ca) 1.5623

Conductivity 4.0-6.0 Magnesium (Mg) 0.8961

Nitrogen 40-80 g/kg Sulphur (S) 0.5072

Phosphorus 60-80 g/kg Sodium (N) 0.4430

Potash 25-45 g/kg Chlorine (Cl) 0.5961

Magnesium 15-30 g/kg Iron (Fe) 0.00581

Calcium equival. 175 Manganese (Mn) 0.00788

Added per m3 amount Boron (B) 0.01090

Lime 5.0 kg Zinc (Zn) 0.00323

Clay granules 40.0 kg Copper (Cu) 0.00064

Fe 15.84 g Molybdenum (Mo) 0.00042

Mn 3.96g

Cu 3.30 g

Zn 3.30g

Mo 1.22 g

B 1.98 g

WageningenAgric. Univ.Papers 90-3 (1990) 35 References

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38 WageningenAgric. Univ.Papers 90S (1990)