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'~ 37 777 Oil\RACrERIZATION ANO USE OF DESMJDlUM SPECIES IN THE CARIBBEAN TROPICS AND SUB1ROPICS

COLEC(ION ~!STOR¡'(A I

, , ,

1-__________ .-.... ! A. M. Thro and K. H. Quesenberry

,

Dep. of Agronorny, D::luisiana state Univ., Baton Rouge, lA 70803, USA (presently Visiting SeL, Cent. Int. Agrie. Trop. (elAT), caH, Colombia);

and Dep. of Agronorny, Univ. of Florida, Ga inesvil le , FL 32611, USA

Forage legume research is active and cooperative in the Garibbean Basin/Central America. Large =llections of rnany Desmodium spEK!ies are available for use. Desmodium. intortum aro Q. uncina.tum have been c:haracterized as unadapted to of our livestock production sites. Native and exotie brawse species (e.g., p. discolor, p. distortum, p. tortuos\lJ11, p. nicaragu~, J). strigillosum, J). velutin\lJ11 and Codariocalvx gyroides (syn. p. gyroides) may be valuable for stock owners with little or no land. Spontaneous species of native pastures (J2. incanuJ!1, n. unHL()I:ulJI, p. pr=umbens D. scomiuru§, D. -ªdscendens, and J). triflonnn) are not presently recommended for cornmercial development. Desmodium parl;¡atlilll is untried. Relative adaptation of the wet tropical lCMland spp. p. heterocaJl2Q!], 1). heterophyllum, and J). ovalifolium varies with site; p. ovalifollum ls more widely adapted. Tbough superior in pasture ~rtment, D. gvaLi1;Qlium provides ¡:xx:¡r animal performance: promising new germplasm is being c:haracterized. AA inl:errlational workshop on Q. ovaUfol1,pm would be timely.

INTROlJJCTION

Tbe follCMing revieltl is the product of research in the CIAT library and the data base of the Red International de EValuation de Pastos Tropicales (RIEPr). HCMever, it has not been possible to circulate this text atrong e>q:Jerienced field researchers of our region. As a result, inaccuracles may have crept in without detection. Moreover, ver¡ recent results may not be included, because research ltIith pasture legumes in the Caribbean Basin is more active at this time than it has ever been.

Fortunately, this Forage Symposium of the 1990 meeting of the Caribbean FC>Od Crops Society brings together many of the very people best qualified te cormnent on a review of Desmodium in the Caribbean¡Central American region. I present this document as a draft, with the sincere request that rny colleagues in atterrlance will ==ect or amplify as 1 proceed, te the ero that our urrlerstanding ef the situation when we leave today will be muc:h better than the ene with whic:h I begin.

'Ihe genus Desmodium is ltIidely distributed in North, Central, and South Arnerica and the Caribbean Islands (Sc:hubert, 1980), and includes many species in all parts of our region (Keoghan, 1980). Sorne species are restricted to natural habitats, but most of those with torage potential are comrnon in unÍlJlproved pastures, road sides, cropped fields, and other disturbed habitats. Several hundred accessions of native Desmodium species have been =llected in the Caribbean and Central America by agencies

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PED. EXTERIOR

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includíng" me university of Florida, CARDljU\'lI, the IDHC (Iazier, 1987), and eIAT.

D?smcx'li~Ll]) spccies that have rúCeivcd attention fram Caribbcan flasin forage researchers can be infonnally dividcd into tour groups. Orle group includes D. intortum (Mill.) Uro. and D. uncinatum (Jacq.) OC., the first De.s!rodium species to be investigated in interdisciplinary resean::h programs with =ntinuity and extensive publication of resulte (Eyles et al., 1985), and merefore the first to receive widespread notice.

BlxMse DesmocHa include the large subshrubs to shrubs D. dis=lor vo::J., D. clistortum Macbr., 1). tortuosum (sw.) oc., D. nicarsguense Oerst., D. strigillosum Sc.hir:rl., D. velutinum oc. and Codariocalyx gyroides (Roxb. ex Link) oc. (syn. D?smodium gyroides oc.). A few species of this second group were being cultivated and studied for browse and cut-and-c.:,rry systems by the early 1900's (calvina, 1952). lIawever, early invcstigators were often isolated and their work was little knawn.

'!he mird group of D?smodium species includes small, grazing tolerant plante native or spontaneous in many "native pastures" oE the caribbean Basin (Sernple, 1962; Meyrat, 1975; Me.rrlenez, 1982a & b; J. Michaud, 1986). 'Ibis group includesJ). incanum (sw.) oc. (syn. D. canum (F.J. Gmell) Shinz & 'Ibell.) , D. uniflorum (n.c.), 11. procumbens (MilI.) Hitchc., D. s=rpiurus (sw.) Desv., D. adscendens (sw.) oc., and1). triflorum (L.) OC.

'Ibe fourth group includes the D. hetercx::arpon (L.)OC.jovalifolium Wall. group, J;l. heterophylllll]l (\'lilld.) OC., and D. j;>¡l@j;:lll]l (L.) Benth. Forage research with these herbaceous species is l'elatively recent (Gro[, 1982).

RIEPr designations will be used for evaluation stages. "A" trials are observations on adaptation oE numerous accessions; "13" trials are clipping trials oE selections frorn A trials; "c" trials evaluate response to grazing af the best B entries, and "D" trials measure perfonnance of animals grazing the best e selections (Toledo, 1982).

Ecosystem designations (RIEPI', 1982) are used when knawn for a site. 'Ibese include: TRF, tropical rain forest, SESF, semi-evergreen seasonal forest; and WIES, well-drained isohyperthermic savannahs. TWo have been added: ECjD, Eastern caribbean Islandsjdrier; ECjH, Eastern caribbean Islandsjmore humid. 'Ibese are differentiated because " the slight decrease in rainfall north aE 17 N Iat ..• appropriate to unite the islands between PUerto Rico and Guadeloupe into one [rainfall] regime" (Portig, 1976) aro by soil t:ype: many northern islands of the eastern caribbean have calcareous soil parent material and neutral to msic soils (Portig, 1976). Guadeloupe and islands south are mostly of volcanic origin with a variety of soils. EC/D and ECjH designations are irrpromptu. Sails and climate af most caribbean islands are so variable within short distances that agroecological mapping on a regional scale is almost impossible (P. Jones, CIATjAEU, pers. com., 1990).

DESl>'K)!)IDM INI'ORItJr.! ANO DESM:IDIDM UNCINA1UM 'Ihese are species of central (D. intortum) and South (.!? 1!!1cinatum)

American up1ands, first collected in the 1950's (Bryan, 1969). 'Ibey are adapted to high rainfall zones (1,100 mm or more) but unsuited to hot wet

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lowlarrl tropics. '!hey have little drought tolerance. Desrnodiurn intorturn is better adapted to lighter soils arrl 12. intoJj;um to heavier soils. For persistence, both reguire h1gh soil fertility; l:x:>th are to1erant of lTlCXlerate soil acidity. Bryan (1969) aro Tmrie et al. (1983) have provided roviews of these species and the DIO released cultivars, 'Silverlea[' ([) .. -1IDCiTl<"ltUl)l) and 'Greenleaf' (!;J • .intortum).

Hybrídízatíon research fo11owed the idffiltification of the forage va1ue of )2. intortum arrl)2. intortum (Rotar and O1ow, 1971; Stobbs and rmrie, 1976; O1ow and Crowder, 1973). However, roany researchers later concluded that hybridization is efficient for p1ant improvement only after naturally-cx::curring gennplasm has been thorough1y evaluated, if needed genotypes are not fourO (cameron 1983). In conseguence, resources were shlfted from hybridization of Desrnodium to gennplasm research with many legume species (Eyles et al., 1985).

Desrnodium intortum in the caribbean Basin. Positive results with )2. intortum in caribbean and central America have been mostly at high elevations or high latitudes. Rattray (1972), workin;J in Panama, found that )2. intortum was not as well adapted at Gualaca (TRF) as)2. ovalifolium, but made good pasture with 1'. maximum on black soils at 0001 high altitude sitos. In Louisiana arrl Florida, USA, Greenleaf is '" high yielding summer legume (Kretsc:hm2.r, 1970; '!hro et al., in press). It has persisted four years in clipping tria1s on fertile sitos in lDuisiana, dornant during winter (-13 C) ('Ihro arrl Shock, 1987).

Desrnodiuminto¡j::um was considered "promising" in Costa Rica (Orozco, 1939; Pacheco, 1957), but at a time when few other species were recognlzed alternatives. In a palatability trial in PUerto Rico with 11 other leguJlV2S, it was lTlCXlerately palatable (Warmke et al., 1952). In north Andros Island, Bahamas, )2. intortum was used with Macrqptilium atropurpure\1l)l (OC.) Uro. cv. Siratro and Neonotonia wightii (Lackey) in a study of nutrition levels of grazing sheep (Dorsett et al., 1980), but the authors made no carnment on the relative values oi the leguJlV2S. In a one-year trial in Jalisco, Mexico, )2. intorttnn and D. uncinatum yielded less than three N. ~i~ht.U cultivars but llOre than siratro (Sanchez, 1985). In the Isla de la Juventud (WUIS), Olbu, D. intortum and M. ªt!:Oourpure\1l)l were the highest yielding legumes and responded to Ca, P, and K fertlllzation with 200 to 400% yield increases on these sandy SOil5 (lDpez et al., 1978).

At Central Farro, Belize ('l'RF) , .!d. intortum died out in the dry season (Lazier, 1980). It was unadapted in Baya¡ro arrl San Jase de las lajas (Wll1S), CUba (lDpez et al., 1978; Monzote et al., 1982); and in Guadeloupe (xan:1e, 1985). In College station (upland) and Beeville (Gulf coast), TeXas, USA (Ho1t and Allisan, 1979j80), :Q. intortum did not survive the first winter (to -7 C), though it has survived similar winters in lDuisiana.

Qe!i!!11Q"!ium uncinatum in tila caribbean R.¡"Ün. In IDuisiana, USA, Silvadeaf has persisted on eroded soi15 with pH 4.5, and on fertile, neutral soi1s in association with cynodon dacty10n (L.) Pers. CV. Alicia, for eight years ('Ihro, pers. 000.; 'l'hro and Shcx::k, 1987). ln Belize (TRF) (razier 1980), Guadaloupe (ECjH) (Xande, 1985), Antigua (EC/D) (CARDI, 1980), Panama (Rattray, 1972), and Texas (Holt and Allisan, 1979/80), )2. uncinatum has su=umbed to drought, cornpetition, cold stress, and unspecifed causes. It

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was the highest yielding of tour J)esmodil,lll1 species (Q. inc<'\ru.II11 aro two unspecified) during the wet season on a heavy clay site in Antigua (Ee/D) (Keoghan and Devers, 1977), but yielded far less than Teramrrus labüüis (L.f.) Sprerq., !:!. atropurpureum, Q¡mtrosema ~ Benth., and Stylosanthes hamata. (L.) Taub.

Co!mnerCial tL..<;efulness ot )2. illtm:tur)\ aro Q. unqinatum in high latitudes is limited. '!bey are in competition tor tertile, moist, coa1 environrnenta where foad crops and high-value cash crops can be grown. '!beir OO5t use in such naturally favored locations rnay be in soil-building rotations (Lazier, 1980), Le., sugarcane land rotated to pasture in Louisiana. AIso, these late-flowering, short day planta do oot set soed before frost (Kretschmer and Snyder, 1979; Thro, pers. 000.). In the absence of a regional soed production site, illIported soed is prohlbitively expensive.

~E OR CUl'-AND-CARRY SPECIES Because rnuch livestock in sorne countries ls owned by producers with

little or no land who use incidental areas such as roadsides for grazing (Paterson, 1986), high-yielding, N-fixing, cut-and-carry 1egurres may have greater value in sorne situations than legurres tor improved grazed pastures. Sorne interesting species are native, including D. tortuosurn (Lasseigne, 1973; Schubert, 1980) and D. distortum (CARDI, 1980; Schubert, 1980). Others are of Old World origin, e.g., º. ~.9e§, 12. l?1;:rigjJJ,(JSUl]1 and )2. velutinum (Oh<1shi, 1973).

In a study in which D. uncinatum, D. intortum, and Q. ovalifolium had tannin =ntent in the range of 3.5 to 4% and protein =ntent froro 15 to 21%, sorne browse DesmocHa had less tannin and more protein: )2. distorturn (tannin, 1.7 - 2.2%; protein, 21.8 - 26.1%); Q. nica:raguense (tannin, 1.9%; protein, 24.7%); .!d. tortuosum (tannin, 2.0 - 3.3%; protein 16.0 - 29.8%), and -º. gyroides (tannin, 2.8 - 3.2%; protein, 22.0 - 23.6%). On the other harrl, .o. discolor had high tannin (3.4%) and medium protein (21.9%) (Rotar, 1965). Surprisingly,.!d. nicaraguense was less palatable to cattle than D. intortum in a trial in Puerto Ri= (Warmke et al., 1952). In a trial in Colombia, browse species D. velutinurn and D. strigillosum were both moderately palatable in the dry season; in the wet scason, ]J. strigil:Losum was highly palatable and.o. velutinurn was unpalatable (Schultze-Kraft et al., 1989). Work is needed to sort out palatability differences and nutritional value in Desmodium.

Browse or cut: and carry Species in me caribbean Basin. Desmodium nicaraguense. '!be ear1lest recorded cu1 tivator of I:Je-<;JOCxlium in the caribbean Basin was Francis= Giusti in Guatemala (calvina, 1952), who planted Meibomia rensoni Paynter (syn. !1. nicaraguense) for his hornes. Local farmers called it engorda caballo (harse fattener). Razeta (1932) stated that it was used by farmers in Nicaragua and had nore protein and 1ess fiber than alfalfa. Later workers in El Salvador tound that D. nicaraguense had outatanding drought tolerance, with dry season yields superior to Lablab purpureus (L.) SWeet, cajanus cajan (L.) Millsp., and Pueraria phaseolaides (Rom.) Benth., and proposed it as a protein source (Choussey, 1943; Watkins and Chavez Viaud, 1950; watkins and Lewy-van Severen, 1951). It did not persist in trials on an extre.mely p:x:>r soil in Belize (Valencia, 1989).

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De$mOdium dj,sco19r. This species, also relíshed by hor<'....cs, has i::Jc<'Jl kney,;n as mannelada de caballo (horse roarm'l.lade). Calvina (1952) regarded ti highly in Mexico (1908 to 1914) and Cuba (1914 to 1922). Razeta (1932) considered ¡;J. discolor less valuable in Nicaragua tban Q. nicaragu~.

DE!smOdium gistortum. 'Ihis legume looked promising in grazed mixtures with !,. maximum in Jamaica (Jam. Dept. of Ag., 1953). In Antigua CEC/D), ClIRDI evaluated ¡;J. gistortum CIAT 335 in grazed plots between 1977 and 1980 (CAROI, 1980). Keoghan (1980) recormnerrled it for c1ay, calcarecus, and volcanic soils of dryer parts of the carIDbean, when short term pasture with high early productivity is required. To facilitate testing and comrnercial use of ¡;J. distortum, CAROI included it seed production projects in Antigua (EC/O) , Trinidad (ECjH or TRF) (ClIRDI, 1980), and Guyana (CAROI, 1984/85). In Dominica (ECjH) (CAROI 1983/84), Trinidad (ECjH or TRF) (CAROI, 1980; Ahmad and Huggins, 1986; Ahmad, 1986), and French Guyana (Bereau, 1986), ¡;J. distortum established rapidly and gave high early yields. In Dominica (ECjH) , Q. distortum was second in productivity after an unspecified Stylosanthes in cut and carry plots with PenrlisetpJn ~eum (CAROI, 1987).

At Central Farm (TRF), Belize, Valencia (1989) cited "high perfonnance" of D. distortum in introdtiction trials of 40 legumes. HCMever, in TRF sites at Centeno, Trinidad, and Nueva Guinea, Nicaragua, D. gistortum rated only "fair" after 18 and 17 months of eva1uation (CAROI, 1980; RIEPr, unpub. data, 1981-82).

Desmodium tortuosum. 'Ihis species has been outstanding for early production both at Central Farm, Belize (TRF) (lazier 1981a) and in Antigua (EC/D) (CAROI, 1980). In Antigua (EC/O) t ¡;J. tortuosum behaved as an annual and was rec:ommenjed only for short term pastures (CAROI 1980). In the Zona Huenequenera (WCHS), Yucatan, Mexico, ".0. ,(;l!.ll:'tJlIrUlll" (syn. ¡;J. tQrtuOSUlll) was rated "very good" but data is available only from the first 12 weeks of the trial (RIEPr, 1988).

Desmodium dichotomum and Qesmodium virgatunl. Accession CPI 47186 of ]2 •. dichotomum did not persist and was not recormnerrled after testing in Antigua (EC/O) (CAROI 1980). In Trinidad, D. vi¡:gattJl!1. was more productive tban Leucaena leucocephala (lam.) de Wit, T. labialis, ~. pubesgms, Stylosanthes guianensis (Aubl.) SW., 12. hanata, and Centrosema pascuorum Mart. ex Benth. on sandy sails (Ahmad and Huggins, 1986). HCMever, it was less persistent than siratro and Pesmanthus virgatus (L.) Willd.

Codarioca1vx gyroides. 'I'his Desmodium ally is included by tradition. In a trial in Colombia, it had lCMer tannin content and higher palatability than ¡;J. ovalifolium (CIATjTPP, 1981).

Codariocalvx gyroides in the caribbean Basin. 1. WLHS. In lDs Santos, Panama, !;. gyroige§ CIAT 3001 had dry natter yields egual to or higher than all species tested in the first wet season of a "B" trial (RIEPI, 1982). In Isla de la Juventud, Cuba, !;. gy:roides CIAT 3001 performed well through the second dry season but did poorly the second wet season of a B trial; it was COllSidered "adapted" (RIEPr, 1985). HCMever, in an A trial at Moblissa, Guyana, !;. gy:roides was judged unadapted (RIEPr, unpub. data, 1982). At El Cllepo, Panama, very poor stands of !;. gyroides CIAT 3001 increased in yie1d fraro very lCM to lCM over one cycle (RIEPr, 1982).

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11. 1RF. At Central Fann, Belize, j:;. GYroÜ!9S Cl" 29 (CIAT 3001) was fourd to be palatable, prcrluctive, and persistent ([azier, 1981 a and b; Neal, 1981; Ahmad, 1986). These trials included an 18-Jronth experiment with intensive rapid grazirr;J every six weeks (Lazier, 1981a). Valencia (1989) fourd that~. gyraides did well in en-farro tria1s on crackirr;J b1ack clay soils in Belize, but that ~. ~, j". 1eucccephala, and F.. pha5e9loides were Jrore a::>rnpetitive with volunteer vegetation. In costa Rica (San carlos, 'I'l.l=ialba, Guapiles), ~. gyraides CIAT 3001 was rated "good" to "exce11ent" in type A tria1s reportirr;J observations over periods that ranged froro 3 to 28 Jronths (RIEPr unpub. data, 1981-87).

On the other hand, í:;. gyraides CIAT 3001 was very low yieldirr;J at calabacito, Panama (first wet season data only) (RIEPr, 1982). In Nicaragua, í:;. gyraides CIAT 3001 was rated good after 17 Jronths of an A tria1 at Nueva Guinea but produced low yields in a type B trial at the saIne

site (RIEPr, 1982); and at El Recreo was one of the few accessions 1eft by the second wet season, thought its yie1ds were far inferior to those of .Q. ovalifolium CIAT 350 and were dec1inirr;J (RIEPT, 1982). In Honduras (la eeiba) I ~. gyraides CIAT 3001 was rated poor after 15 Jronths observation (it had been rated excellent at this site in the first wet season) (RIEPr, unpub. data, 1982).

III. SES!". In a B tria1 in San Isidro, Costa Rica, r;;;. gy¡;::piqe..5 CIAT 3001 was low yie1dirr;J in its first cycle but by its second dry season had very high yie1ds (RIEPT, 1982). However lit was 1CNer y ie1dirr;J than .Q. ova1ifolium CIAT 350.

Pesmodium sp. CIAT 3019. This accession was rated "good" after 14 Jrontha in A tria1s in Guapiles ('IRF), costa Rica (RIEPr, unpub. data, 1982-1984) but "fair" te "poor" in Nueva Guinea (1RF), Nicaragua (lliEPI', unpub. data, 1981-82) •

Desrnodium strigi110sum and Desrnodium velutinum. CIAT accessions 13153, 13155, and 13158 of .Q. strigillosum and CIAT 13218 .Q. velutinum were tested in Guapiles (1RF) and San Isidro (SESF), costa Rica. At both sites, .Q. strigi110sum yielded less than!;l. ovalifolium but more than D. heterophy11um (CIATjTPP, 1989). Yield of .Q. velutinum at Guapiles was higher than a11 but 000 of more than 30 accessions of !}. ovalifolium.

SFúNTANEXXJS r.mJMES OF NKrIVE PAS'IURES American species in this group include Desrnodium incanum, D.

procurnbens, .Q. scorpiurus and.Q. adscenclens (Schubert, 1980). De..'?l\'lOdiurn triflorum is pantropic (Verdoourt, 1979). Al1 are small plants that survive grazirr;J by becoming prostrate, an attribute that makes their contribution to pasturas difficult to measure (Lazier, 1981a). In sorne cases they comprise froro 10 to 20% (CAROI, 1980; J. Michaud, 1986) or even one-third (Dirven, 1954) of the plant population and may make a valuable contribution to the biotic system. Valencia (1989) noted that animal average daily gain increased in certain heavily-grazed padd=ks at Central Farro (TRF), Belize, after Q. adscenclens and Q. heterophyllum volunteered in those padd=ks.

Desmoé!ium incanum in the caribbean Basin. 1. 1RF. Lazier (1981a, b) fourd D. incanum per5istent at Central Farro, Belize, but selected~. gyraid~ for

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further research because of its equal persistence and greater prcduction and animal use. In A trials in PUerto cabezas (RIEPr, unpub. data, 1983-85) and Nueva Guinea (RIEPl', unpub. data, 1981-82), Nicaragua, Q. }."(l<;;anUlll accession CIAT 13032 was ratcd "fair" avcr 12 to 17 monUls o[ CVilltliltiol1. In a cdnqlc early rating of an A trial in El Bongo, Panama, D. incanum "261" was rated "fair" and CIAT 13032 "p::x:lr" (RIEPl', unpub. data, 1983). In Calabacito, Panama, "252" and CIAT 13032 were both rated "fair" to "p::x:lr" in an A tria1; on1y the first five months of data fram this tria1 are availab1e (RIEPl', unpub. data, 1984). After 15 months of evaluation in an A trial at la eeíba, Honduras, CIAT 13032 was rated "fair" (RIEPr, unpub. data, 1982-84).

11. SESF. In a B tria1 in la Esperanza, Honduras, D. incanum CIAT 13032 was one of few legumes to persist tw"O years and one of very few to produce any forage in the dry season (RIEPI', 1988). Severe powdery mi1dew was noted on D. incanum at la Esperanza. In B trials in the D::lm.inican Republic (Pedro Brand, Haras Nacional), CIAT 13032 was re1atively very 1CM yielding even in the dry season, over two cyc1es (RIEPl', 1985).

III. Wl:ltS. In Bauta, Ulba, D. incanum was defoliated, cause unidentified, and disappeared froro p10ts in a B tria1 (RIEPl', 1988).

IV. EC/D. Keoghan and Devers (1977) reported very lCM yie1ds for D. incanurn on a heavy clay site in Antigua (EC/D), both wet and dry season, re1ative to yie1ds of 20 legume species. HOI-leVer, D. incanum persisted when p1anted on a similar Antigua site (heavy c1ay, high pH soil) in mixtures with~. 1abia1is, the highest-yielding legume in this environment, and after five years made up half of the mixture (Cl\RDl, 1980).

In Louisiana, USA, where D. incanum is neither native or naturalized, CIAT 13032 was not winterhardy (Thro, 1984), but Tifhardy-1, a selection froro Georgia, USA (MiUer and WeUs, 1989) has persisted and spread on a very acid, ercded site with 1CM soil water holding-capacity. .!!;)"11uodiUl1l incanum is a highly persistent legume in Florida, but is not used due to lCM palatability and lCM yields.

Desmodium scorpiurus, D. adscendens, and D. triflorum were ubiquitous and well-nodulated in Belizean (TRF) pastures (lazier, 1981a), and were likely making a nitrogen contribution. HCMever, D.ª~ens. did not compete well with Cynodon nlemfuensis Vanderyst, Bothriochloa pertusa, or f'. maxÍInUm at Central Farm (TRF) in Belize (Valencia, 1989). D. scorpiurus, !l. adscendens, and D. triflorum have been considered as "promising" in COsta Rica (Orezco, 1939; Semple, 1962).

~i\Jlll uniflorum accessíon "26062" had 1CM prcductivity in Antigua (EC/D) (CAROl, 1980). Desmodium procumbens behaved as a transítory annual in Antigua (CAROI, 1980), and was not considered useful.

DESMJDIUM HETEROCARFON/OVALIFOLIUM, l). HEl'EROrnYUDM, ANO DESMJDIUM l3lI.RBl\IUM 'llle first three species are Asían legumes considerad by Ohashí (1973)

to be one species wíth several botanical varieties. HCMever, forage researchers have found consistent morphological differences between the species (SChultze-Kraft and Benavides, 1988; Quesenberry et al., 1989). AH three tolerate shade and heavy soils (Schultze-Kraft, pers. com., 1990) and were :reconmended by Keoghan (1980) for long-term pastures in wet regions of

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the caribbean. Desmcdium barbatum is a pantropical species (Schubert, 1980; Ohashi, 1973). Although it is a distinct species, sorne acces5ions oE ¡;¡. barbatum are similar to ¡;¡. heteroca:tJX)n and !d. ové!lJfolj..lllJl in grcMth habito

DesmcdiUl!l heterocarpon. ¡;¡. heterocat:pOn CV. Florida, a selection from USD/\ PI 217910, i5 released in southern Florida, USA (Kretscl:nner et at., 1979), and has been tested by RIEPI' as CIAT 365. It has better drought tolerance than ld. heterpPlwllum, better seed production, and gacd grazing tolerance in spite of an upright growth habit in the absence of grazing, but is susceptible to root knot nematode. Additional collections of .R. heteroca:tJX)n and Q. ovalifoliUl1) haye been tested in Florida (Kretschmer et al., 1980; Quesenberry and Dunn, 1987; Kretschmer et al., 19CiO) ard accessions haye been identified with greater resistance. Root knot nematode-resistant accessions of ¡;¡. ovaliíolilml have been crossed with Q. peterocarpon (Quese.nberry et al., 1989) and advance::l generation hybrids froro these crosses appear to combine ne.matode resistance of D. ovalifoliUl!l with characteristics of p. heteroca~ such as earlier seed production.

Desmcdium heterocarpon in thª-caribbear¡. L TRF. In Nueva Guinea, Nicaragua, .R. heterocalJ2Qn CIAT 365 was rated "gacd" for 17 lrOnths (RIEPr, unpub. data, 1982). In San carlos, Guapiles, and TUrrialba, Costa Rica, CIAT 365 (at Guapiles, CIAT 3787, as well) was rated "gacd" ovar observation intervals of 3 to 14 rnonths (RIEPI', unpub. data, 1982-84). In Belize, l). hete~ was rec:ommen:led for poorly drained sails (Ahmad, 1986). At Central Fann, Belize, l). het§~lJ29D CF 1037 was one of 12 of 28 legume entries tbat persi5ted for three years (Valencia, 1989). In recent work in costa Rica, p. heteroca~ was outstanding at Guapiles but not at san Isidro (WIl:lS) (CIAT/TPP, 1989).

n. SESF. In San Ram:>n, costa Rica, ¡;¡. hete~rpon CIAT 3787 was rated "poor" after 14 rnonths of observation (RIEPI', unpub. data, 1985-87).

In an adaptation trial in an infertile, acid-sail site in Iouisiana, USA, Q. heterg:;;armn CV. Florida was rated "excellent" during the first summer but wa5 killed the first winter ('Ihro, 1984). In spite of excellent seed set, only isalated seedling5 were observed the following year.

lRsmodium heterophyllum. '!he low grcMth habit of this 1egume gives it tolerance te grazing and compatibility with such grasses as BrachJaria decu:ntJens Staph (Imrie et al., 1983) but linrits seed recovery; fortunately, it can be vegetatively established from sto10ns. It is tolerant oE acid sai15 and grows wel1 on Eerti1e a1luvial sai1s; it is not drought tolerant. 'Ibere is one commercial cultivar, Johnstone, released in Australia in 1973 (Imrie et al., 1983), tested by RIEPI' as CIAT 349.

Pesmodium hete~.llUl]l in the caribbean Basin. 1. TRF. Rattray (1972) cited the ease of vegetative reproctuction of this species in Gualaca, Panama. At central Farro, Belize, D. hete!!>phyllUl!l was recommended for poor1y drained soils (Ahmad, 1986). It survived as a reseeding wet season arnlual at least from 1977 to 1982 (Valencia, 1989). In la CeÍba, Honduras, I). heterophyllum CIAT 349 was ratad "gacd" in the first 15 rnonths oE an A trial (RIEPI', 1985), and CrAT 3782 was mentioned as outstanding for persistence, disease resistance, productiyity and ease of establishment in a B tria1 at tbat site (RIEPT, 1988). In PUerto cabezas, Nicaragua, CIAT 349

8

was r.ated "excellent" during one year of observation (RIEPI', unpub. data, 1984-85). Type B trials including CIAT 349 and 3782 were established at Guapiles, Costa Rica (RIEPI', 1988), where both accessions were well nodulated at 12 weeks, and A triale including CIAT 349 at 'I\.lrrialba (RIEPI', unpub. data, 1983), but later resulta of these triale were not available.

However, the accession CIAT 349 was not outatandíng among entries in an A trial in Sona, Paroma (CIATfTPP, 1987). In Nicaragua, 1;>. heterophyllum CIAT 349 was ratee! "excellent" in the fírst year of an A trial in PUerto cabezas, but in El Recreo (B trial) and Nueva Guinea (A trial), it was ratee! "peor" to "fair" during 17 months of observation, had low yields, and disappeared from clipped plota by the secand wet season (RIEPT 1982 ¡ RIEPT, unpub. data, 1982). In Costa Rica, I). heterophyllum was ratee! "fair" to "poor" in A trials in san carlos (RIEPT, unpub. data, 1983), though CIAT 349 ratee! "good" during 14 months of observation in Guapiles, Costa Rica (RIEPT unpub. data, 1982-1984). In Trinidad (location unspec.), Johnstone was one of the least proctuctive of the legumes testee! (CAROI 1978-79, CAROI, 1980). In El Centeno, Trinidad, first year yields were low (RIEPT, 1982).

11. SESF. At San Rarnon, Costa Rica, 1;>. hetexophyllum was rated "[air" to "poor" (RIEPT 1985-87). In B trials at san Isidro, Costa Rica, yields of D. heterophyllum CIAT 349 were consistently low for two years (RIEPT, 1982). In later work, D. heterophyllum accessions did poorly at both Guapiles (TRF) and at san Isidro campared to D. het~, D. QIIalifolium, !1. velutinum, and D. strigillosum (CIAT/TPP, 1989). CIAT 349 persisted for two years in infertile, eroded, acid soil in lDuisiana, USA), after whích it died out ('Ihro, 1984). However, during two years, D. heterophYllum CIAT 349 was one of the rnost productive legumes in the cool season in a B trial in Tlapocoyan, Veracruz, Mexico (RIEPT, 19(8).

~.:lurn ovalifoliurn. This species has excellent adaptc.tion to high alumínum-content, highly acid soils of the tropi= (Imrie et al., 1983). lt is adapted to wet regions with rainfall of 2000 mm or rnore (Grof, 1982) and with short dry seasons of no more than two or three months (Schultze-Kraft, CIATjTPP, pers. com., 1990). A COV'er crop cultivar from a Singapore-based seed company has been testee! as CIAT 350 (Schultze-Kraft and Giacometti, 1979). It wae found tohave excellent persistence under grazing even with aggressive grasses such as Brachiaria species for which no oti1er companion legumes were available (Grof, 1982). SUbsequent studies have revealed shortcominge of thie otherwise highly pranising legume.

It ie susceptible to a etem nematode, Fterotylel}Chus cecidcg¡mue, found on native D?srnodium hosta, and to a false rust disease, causal organism synchytrium desrnodii, a exotic pathogen introduced here by an unkna.m ÍlnpOrter of Asian seed (CIATjTPP, 1985). When a resistant accession, CIAT 13089, wae grown on two soils in mixture with !}. humidicola (Rendle) Schwe., its vigorous growth habit, freedam from disease, and low palatability allowed it to domínate the grass, yet its presence did not increase animal production, and in sorne cases even depressed it (CIATjTPP, 1989) relative to the grasa control.

A collection of 84 acoessions of ~. ovalifolium contained substantial variation for pest resistanoe, palatability, tannín content, flowering date, growth habit, forage yield, and protein contento Tannin content and

9

palatability were negative1y oorrelated in this oollection (Schultze-Kraft and Benavides, 1988).

Desmodíum ovalifolium in the caribl:lC'~,n l3<1sin. 1. TRP. In HOlJduras (J il Ceiba) ~ 1>. ~IfoiiÜni eIAT 350 ani3784were rated "excellent" in an 15-month A trial (IUEPr, unpub. data, 1983). In Panama (Sona), 1>. ovalifolium CIAT 350 had outstandirq rosease resistance in the wet season and average yields increasirq over time (RIEPr, 1985). In Costa Rica (San carlos, Guapiles, and Turrialba) 1>. ovalífoUum has advanced fram A trials in which it rated "gaod" after 8 months of observation at 'rurrialba arrl 14 months at Guapiles, but only "fair" after two dry seasons and one wet season in san carlos (IUEPr, 1985) to e trials at Turrialba (IUEPr, 1988). CIAT 1>. ovalifoUum accessions 350, 3673 and 3784, were tested in tilese B trials; CIAT 350 was superior to elAT 3673 for pest resistance aro had better seed production in san carlos. In tila e trials at Turrialba, D. ovalifolium with ,{:. nlemfuensis hae! lower pasture and legumo yiele! aro qua lit Y than associations with Arachis pintoi Greg. & Krap. ane! E. phaseoloides, but only D. ovalifolium increased in tile pasture as more animals were added.

Desmodium ovaUfolium was rated highly in a B trial at Fray Bartolomeo de las casas, Guatemala (eIAT 3793) (IUEPr, 1988); another B trial in Seybo el Valle, r:x:mú.nican Republic (ClAT 350 and 3784) (IUEPr 1985); an A trial in Puerto Cabezas, Nicaragua (CIAT 3784) (IUEPr, 1982) and B trials in El Recreo and Nueva Guinea, Nicaragua (CIAT 350) (IUEPr, 1982). In the B trial at El Recreo, eIAT 350 was rated "best" and was one of few species 1eft by the secorrl rainy season. In Belize, D. ova,lifolium kept a gaod stand in a c-type grazirq trial on a vertíc planosol froro 1977 to 1989 (lIhll1<"1d, 1986) and accession CF 200 (CIAT 350) was rated "best overall performance" on acid, infertile lowland soils in native pastures (Valencia, 1989). However, Desmodium ovalifolium CrAT 350 and 3784 both did equally poorly at Hojancha, Costa Rica (IUEPr, 1985), especially in the dry season.

n. WOOS. In the Dominican Republic (Fi2dro Brand and Raras Nacional), D. ovalifolium CIAT 3784 yielded near the mean of all B trial entries, less than Zomia latifolia Sm. in Rees. aro 1'. phaseoloides entries (IUEPr, 1985) but was rated highly for its dry season value. In Moblissa, Guyana, D. ovaHfolium eIAT 350 is oonsidered adapted (IUEPr, 1985; CUrnerbatch, 1986) . In Surinam, D. ovalífolium did not resporrl when pH of a highly acid soil was increased (simons, 1984). Bere.au (1986) rcportcd (roro Frcnch Guyana that D. ovaHfolium CIAT 350 was slow to establish, but had excellent persistence and disease resistance. CllRDI (1980) included D. ovaHfolium in its seed production project.

lI1. SESF. In a 20 month A trial at San Ramon, Costa Rica, ]:l. ovalifo1ium CIAT 3781 was rated "good" but eIAT 3376 and CIAT 350 were rated "fair" and eIAT 3782 ,3784, and 3787 disappeared before the ene! of the trial (IUEPr, unpub. data, 1985-87). In a B trial at Bauta, CUba, it was defoliated and disappeared fram the plots (accession unspecified) (IUEPr, 1988). At Penonome, Panama, there was no visible pest damage to acoount for the languid perfonnance of CIAT 3784 (IUEPr, 1988). In San Isidro, Costa Rica, CIAT 350 was rated "adapted" but low to average in yield, surpassed by Q. gyroides, Aeschvnomene histrix Poir, aro fi. gyíanensís (IUEPr, unpub. data, 1983). However, when a large collection of !J. ovalifolium (> 30 accessions) was tested at two oontrastirq locations in Costa Rica, Guapiles

10

(TRfl and San Isidro, yield of sU: accessions comrron to both sites was not different between the two sites (CIATjTPP, 1989), and in a Trinidad site with pronounced wet and dry seasons, D. ovaJifolium 1Ie.¡ 350" was superior to 80f 12 other legumes (Davis, 1986).

In Mexico, in B trials in Destierro and loma Bonita (vlIl:fS), Jalapa/El Gusto (TRf), and Jerico (SESF), ranging in length froro one dry season to two years, CIAT 350 and two low tannin, pest resistant se1ections (CIAT 3788 and 3784) were amorq the best entries for yield and persistence (RIEPr, 1982, 1985, 1988). In Jalapa, CIAT 3784 was higher yielding than CIAT 350; in the other locations there was 1itt1e difference.

In a D trial at San JOS€! de las lajas (wrHS), ama, j}. ovalifolium CIAT 350 provided pcorer animal performance than 12. guianensis CIAT 184 in a !;;. dactylon mixture, and less available forage (RIEPr, 1985).

Desmcx:lium ovallfollum CIAT 350 was rated "good" during tile first summer of an adaptation trial in lDuisiana, USA, but did not survive the winter (Thro, 1984).

Desmcx:lium barbatum. This species is also adapted to acid soils (Irorie et al., 1983; Schultz-Kraft and Giacornetti, 1979). Semple (1964) observed it in native pastures in Costa Rica fraro sea level to above 1000 m. It volunteered in Hyparrheniª rufa (C. G. Nees) Staph pastures at Zamorano, Honduras; in sorne places these pastures were 40% D. barbattl!lY'60% grass (Semple, 1964). De..<m1Odium barbatum was equally palatable with the grass when pasture height was about 30 ano Semple reported that D. barbat\JJ11 needed at least a 6-month wet season; SdlUltze-Kraft aOO GÜlcometti (1'l79) describe the species as adapted to high rainfall areas. Semple (1964) aOO Schultze-KJ:aft and Giacometti (1979) cornment on the bread range of morpho1ogical diversity in j}. barbatum for growth habit, foliage density, and productivity.

Desmcx:lium barbatum in the caribbean Basin. lDcally collected accessions of D. barbatum were planted in trials at Sona (TRf) and Los Santas (wrHS), Panama (CIATjTPP, 1987); no data are yet available. In Belize (TRf), Valencia (1989) observed good ear1y performance froro j}. barbatum CF 1024, but it did not persisto In Louisiana, CIAT 363 was rated "excellent" in the first summer of an adaptation trial but did not survive the winter (Thro, 1984) •

DISCUSSION "In areas that cannot be cropped, grazing animals will always have a

place, turning plants that humans cannot digest into edible protein" (Keo::¡han and Devers, 1977). It appears that D. int9rtum and D. uncinatum are not useful in sites of the type referred to by Keoghan and Devers.

Desmodium shnlbs and subshrubs are capab1e of providing high dry matter yields for browse or cut and carry situations. In the western caribbean, 5;;. gyroides looks best; and j}. distortum in the eastern caribbean. For short-term plantings with high ear1y yields, D. tqJ;tuosum has been recornmended. For rnost of these species, there is insufficient data to determine the 1imits of their adaptation, but!;;. gyroides is widely adapted and sorne species (e.g., j}. tortuosum) are widespread.

11

In research to date, the small, spontaneous species widcly distributed in native pastures have lacked sufficient advantages to justífy commercíal developrnent. D9sm0dium heterocarpon and p. heterophyllum are adapted to grazed pastures in 'l'RF sites, and 1). heterocarpon in Florida; there is no data for these specíes in other ecosystems. In wet, acid-soil regíons of t:he true tropics, p. ovalifolium has been superior to the other two species.

D9sm0dium has been tested in llOre places in our regíon than anyt:llinJ except CocaCola and has been just as widely successful. However, animal performance in D trials with high-tarmin CIAT 350 and CIAT 13089 has been disappoínting. 1Dw tarmin accessions have been identified.

D9sm0dium barbatum carne to the attention of researchers at about the san\€! time as p. ovalifolium (Schult.:e-Kraft and Giacomettí, 1979) and was eclipsed. An enlarged collection of p. barbatum TlCM exista and might reward examination (Schultze-Kraft, CIATfTPP, pers. com., 1990).

CDNCIDSIONS

Momentum. At this time, there is great momentum in D9sm0dium research, and legume research in general, in our region. fue effects of CATIE, CAROI, and RIEPr, as each has come on the scene, are strikingly evídent in continuity and quality of research and availabilíty of information. fuese coordínators have amplified the value of each local program's contribution and greatly increased the probabílity that we will, together, successfully COIlplete the fírst stages in the development of new D9sm0dium species. 'Ihis first stage is a huge task: thorough evaluation of a representative sarrple of natural genetic varíatíon in a repreSC'J1tative range of cnvíronmonts, from ad;,ptation tríals of many accessions through animal perfo:t:1Tlc":mce trials with a few of the best.

We are in the thick of the job ríght l1O'Ñ. 'Ihe RIEPT data, for example, shows conclusions that are building, not yet formed. Continuity, communication, and cooparation are essential to carrying on froro this high-energy poínt.

What next? Orle priority at this stage should be what to do about p. ovalifolium, the highly p.romising legume with the serioos drawbacks for animal performance. Study of natural variation in this species has received much effort and may be close to COIlpletion, when plant breeding should begin if needed. Breeding programs could be location-specific or aimed at broad adaptation, as desirect. It may be time for a workshop on this species, to consolidate information and together plan the next steps.

A second priority is to meet the needs of special purpose systems and of environmenta where 1). ovalifolium is not superior, such as the eastern arrilibean. For example, unless other genera provide better legumes for these niches, work should be continued with S;;. gyroides and.!;>. distortum. An enlarged collection of S;;. gyroides is l1CM available for research (Schultz-Kraft, CIATjTPP, pars., com., 1990). Early resulta with D. strigillosum and D. velutinum should be pursued.

~/.

Adoption. Closely integrating our research with on-fann biological, economic, and social conditions, though ext.remely costly in time as well as

12

2 7 JUN. 1990

funds, rnay be improve our chances of developing cultivars that are really in denrurl. Working wim our customers, we will be in a better position to develop legumes to fit common types of rnanagement and mismanagement.

Seed availability is also key. Rltential seed production envirornnents for promising species should be scouted throughout me region. Technology adoption prograrns for forage legumes should attempt to integrate seed production and marketing research wim development of market denrurl, so that producers are not frustrated by extension tales of unavailable, unaffordable wonder legumes.

* Iet us keep cooperating * It is time for an international strategy session on !l. ovalifolium * stay clase to our customers

ACI<NI::M.E[X;mENT5 'Ihis review is dedicated to National Agricul tural Research Programs,

and their pasture scientists in Antigua, Antilles Francaises, Bahamas, Belize, Costa Rica, Cuba, Dominica, El Salvador, Guatemala, Guyana, Guyane Francaise, Honduras, Jamaica, Mexico, Nicaragua, Panalra, PUerto Rico, Republica Dominicana, Surinam, Trinidad and Tobago, and US Virgin Islanda.

'Ibe first aumor thanks me staffs of me CIAT Lilirary and Agroecological studies unit and RIEPI' =rdinator L. H. Franco for supero professional assistance; CIAT/TPP scientists R. Schultze-Kraft and carlos Lascano for sharing meir perspectives; and R. Sarria-Marin for =vering in my absence "100% plus".

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Bereau, M. 1986. Comportement de quelques legumineuses in Guyane Francaise. Pasturas Trop. Bol. 8(2) :19-21. Inst. Nat. de la Rech. Agron., Groupe Region. de Guyane, Kourou Cedex, Guyane Francaise.

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13

CARD1. 1980, 1983/84, 1984/85, 1985/86. Annual Report: Research ara Devalopnent SUmmary. Univ. West Irrlies, Sto Augu&tine, Trinidad.

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14

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15

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