A synopsis of the genus Deguelia (Leguminosae, Papilionoideae, Millettieae) in Brazil

A synopsis of the genus Deguelia (Leguminosae, Papilionoideae,Millettieae) in Brazil

RODRIGO AUGUSTO CAMARGO1

AND ANA MARIA GOULART DE AZEVEDO TOZZI2

1 Programa de Pós-Graduação em Biologia Vegetal, Departamento de Biologia Vegetal,Instituto de Biologia, Universidade Estadual de Campinas, Postal Code 6109, 13083-970,Campinas, São Paulo, Brazil; e-mail: [email protected]

2Departamento de Biologia Vegetal, Instituto de Biologia, Universidade Estadual de Campinas,Postal Code 6109, 13083-970, Campinas, SP, Brazil; e-mail: [email protected]

Abstract. Considering the recent reestablishment ofDeguelia, six new combinations and14 lectotypifications are proposed here. A key is provided for the identification of the 15species of Deguelia that occur in Brazil, along with nomenclatural notes, comments onecology and distribution, and discussions of diagnostic features.

Key Words: Leguminosae, Fabaceae,Deguelia,Derris, Lonchocarpus, lectotypification,Brazil.

Resumo. Considerando o recente restabelecimento de Deguelia Aubl., seis novascombinações são aqui propostas; também estão sendo feitas 14 lectotipificações. Éapresentada uma chave para a identificação das 15 espécies de Deguelia ocorrentes noBrasil, acompanhada de comentários sobre nomenclatura, distribuição e ecologia,caracteres diagnósticos e nomes populares das espécies.

Deguelia Aubl. has a controversial taxonomichistory, and its species have been includedeither in Lonchocarpus ser. Fasciculati Benth.(= Lonchocarpus subg. Phacelanthus Pittier exGeesink) in some published works (Bentham,1839, 1860, 1862; Ducke, 1925; Pittier, 1917,1944; Hermann, 1947; Lemée, 1952) or inDerris ser. Americanae (Bentham, 1860, 1962;Macbride, 1943; Ducke, 1949).The genus Deguelia was described by Aublet

(1775) based on D. scandens Aubl., which wascollected by Aublet in French Guiana andsynonymized with Derris Lour. by Bentham(1860). Under the provisions of the InternationalCode of Botanical Nomenclature, Derris wasconserved against Deguelia (Rickett & Stafleu,1959). Geesink (1984) recognizedDeguelia as agenus distinct from Derris and Lonchocarpus,and Tozzi (1994, 1998) and Sousa (2009) madea new combination and described some newspecies.The species of Deguelia are characterized

mainly by the lianescent habit (less frequentlyarborescent), inflorescences in pseudoracemes

in which the secondary axis is reduced, nodose,and forming a short brachyblast with five ormore flowers, indehiscent pods (less frequentlydehiscent) with 1–3 seeds or up to 11 seeds, andpresence or absence of a vexillary wing.Deguelia is represented by 20 neotropical

species, especially in northern South America(Tozzi, 1998), with only two species reachingCentral America, occurring in Guatemala, Nic-aragua, Costa Rica, and Panama (Sousa, 2009).The Amazon Basin is a center of diversity forDeguelia, mainly between the states of Amazo-nas and Para in northern Brazil, along the RioAmazonas and its tributaries. The southern limitof the genus is in southeastern Brazil, wheresome restricted arborescent species can be found.Most species prefer forested habitats, but onespecies, Deguelia nitidula (Benth.) A. M. G.Azevedo & R. A. Camargo, is widely distributedin the savannas of South America (Tozzi, 1989).Considering the recent reestablishment of

Deguelia, the present paper aims to effect thepertinent combinations and also a taxonomic

Brittonia, DOI 10.1007/s12228-013-9302-4ISSN: 0007-196X (print) ISSN: 1938-436X (electronic)© 2013, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

and nomenclatural synopsis, according to theproposal for the circumscription of the genus.We provide a key for the 15 Brazilian speciesof Deguelia.In all cases where more than one

collection is cited in the protologue ormore than one duplicate of one collectionis deposited in the same herbarium, andthe author has not designated a holotype,we designate the more representative spec-imen as lectotype. We are also designatinglectotypes for all Bentham species, includ-ing the ones for which he cited only onecollection or for which there is only onespecimen deposited at Kew because it isnot explicit in his publications whether hebased his descriptions on only one specimen,and it appears that he saw other specimens ofthese collections too. We have some evidencethat he was visiting other herbaria in Europebefore the original publications, for example onthe first page of his work (Bentham, 1860)where he reports that he was in Vienna workingwith the South American Dalbergieae. Healso mentions on the third page of thissame paper that although he had at hisdisposal a richer collection for editingMartius's Flora Brasiliensis, he did assem-ble a considerable number of notes in theherbaria of Paris, Geneva and others,because he had some doubts due to theinadequacy of some specimens.

Deguelia Aubl., Pl. Guian. 2: 750. 1775.Type: Deguelia scandens Aubl.

Lonchocarpus ser. Fasciculati Benth., J. Linn. Soc., Bot. 4Suppl. 87. 1860. Lectotype, here designated:Lonchocarpus nitidulus Benth. (=Deguelia nitidula(Benth.) A. M. G. Azevedo & R. A. Camargo).

Lonchocarpus subg. Phacelanthus Pittier ex Geesink,Advances Leg. Syst. 1: 257. 1981. Lonchocarpussubg. Phacelanthus Pittier, Contr. U. S. Natl. Herb.20: 45. 1917, nom. illeg. Lectotype, here designated:

Lonchocarpus nitidulus Benth. (=Deguelia nitidula(Benth.) A. M. G. Azevedo & R. A. Camargo).

Derris sect. Derris ser. Americanae Benth., J. Linn. Soc.,Bot. 4 Suppl.: 102. 1860. Deguelia sect. Deguelia ser.Americanae (Benth.) Taub. in Engl. & Prantl., Nat.Pflanzenfam. 3: 345. 1891. Lectotype, here designat-ed: Derris guianensis Benth. (=Deguelia scandensAubl.).

Scandent shrubs to lianas, less frequentlytrees. Leaves alternate, imparipinnate, 3–17foliolate; leaflets opposite, sometimessubopposite, frequently increasing in size to-wards the distal portion of the rachis; stipulesoften caducous; stipels present or absent. Inflo-rescence pseudoracemose, with secondary axisshortened and nodose forming a brachyblast,frequently long, terminal or axillary, solitary or2–7 clustered at each node; brachyblasts withmore than 5 flowers each; bracts and bracteolesusually caducous. Hypanthium generally poorlydeveloped; calyx campanulate, subtruncate todentate, usually with 3 distinct carinal lobes and2 broader vexillary lobes partially connate;corolla white, yellow or purple to magenta,unguiculate; standard petal orbicular, oblong toobovate, usually emarginate at the apex, some-times with two basal small appendages; wingslightly adnate with the keel petals above theclaw; keel petals often connate at the carinalmargin; stamens pseudomonadelphous, thevexillary stamen free at the tube base butconnate higher up with the reminder; anthersversatile, oblong, bilocular; intrastaminal discoften present; ovary sessile or short-stipitate,(1–)2–15 ovulate; style filiform, curved; stigmacapitate, short. Pod generally compressed,suborbicular, oblong to linear-oblong,membranaceus to coriaceus, rarely woody, themargins nerviform, the vexillary marginsometimes well developed, forming a narrowlongitudinal wing, indehiscent, rarely de-hiscent, the remaining portion of the stylepersistent. Seeds 1–12, oblong-reniform;hilum short.

Key to the Brazilian species of Deguelia

1. Standard glabrous on the outer surface.2. Inflorescence with brachyblasts distant from each other, lacking on the first 6–14 cm. . . . . . . . . . 9. D. amazonica2. Inflorescence with brachyblasts near to each other, appearing close to the base.3. Pod linear-oblong. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. D. scandens3. Pod oblong-ovate to suborbicular. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7. D. negrensis

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1. Standard sericeous to variably pilose on the outer surface.4. Calyx truncate or obsolete dentate, the teeth shorter than ¼ of its length.

5. Lower surface of the leaflets with prominent secondary veins.6. Leaflets 5; vines or scandent shrubs. . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. D. glaucifolia6. Leaflets 9 or more; trees. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. D. costata

5. Lower surface of the leaflets with impressed secondary veins.7. Intrastaminal disc absent; southeastern Brazil. . . . . . . . . . . . . . . . . . . . . .13. D. hatschbachii7. Intrastaminal disc present; northern Brazil, in the Amazon Basin.

8. Trees; bracteole ovate, less than 1.3 mm long; pod compressed, cartaceous, tardily dehiscent.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14. D. spruceana

8. Lianas or scandent shrubs; bracteole suborbicular, more than 1.4 mm long; pod thickened, woody,indehiscent and slightly constricted between the seeds, breaking transversally into 1–2-seededsegments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. D. densiflora

4. Calyx teeth distinct, greater than ¼ of its length.9. Inflorescence congested, shorter or equal to the length of the leaf.

10. Staminal tube strongly recurved, rigid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. D. dasycalyx10. Staminal tube slightly recurved, membranaceous.

11. Lower surface of leaflet adpressed pilose to glabrous with prominent veins. . . . . . 10. D. utilis11. Lower surface of leaflet tomentose to villose with strong prominent veins.. . . . . . 3. D. urucu

9. Inflorescence sparse, longer than the leaf and generally about twice of its length.12. Standard shorter than 1 cm and wings less than 0.9 cm long.

13. Lower surface of leaflet villose, with subuncinate hairs; pod indehiscent. . . . . . . . . . . . . . . . . . 5. D. rariflora13. Lower surface of leaflet tomentose or sometimes with appressed hairs; pod tardily dehiscent. . . .6. D. duckeana

12. Standard and wings more than 1 cm long.14. Standard sericeous on the whole external surface; calyx with lobes acute, similar in size and glabrous onthe internal surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. D. nitidula14. Standard glabrescent, sericeous only at the apex of the external surface; calyx with lobesacuminate, gradually longer at the carinal side, puberulent on the internal surface. . 11. D. angulata

1. Deguelia dasycalyx (Harms) A. M. G.Azevedo & R. A. Camargo, comb. nov.Lonchocarpus dasycalyx Harms, Repert.Spec. Nov. Regni Veg.17: 321. 1921.Derris dasycalyx (Harms) N. F. Mattos,Loefgrenia 98: 4. 1989. Type: Brazil. Riode Janeiro: São Cristóvão, Quinta da BoaVista, Feb–Mar 1887, A. Glaziou 15916(holotype: B [photo & fragment: F];isotypes: C, F, K [image; photos: C, F,UEC], P). (Fig. 1)

Lonchocarpus sarmentosus Glaz., Bull. Soc. Bot. France53: 150. 1906, nom. nud.

Distribution and ecology.—Collected in Ba-hia, on Serra Geral do Caetité, near Brejinhosdas Ametistas, Minas Gerais, at the locality ofMariana. The type from Rio de Janeiro, SãoCristóvão, in Quinta da Boa Vista, was culti-vated. It occurs in Bahia on peaks of thesandstone cliffs with non-disturbed vegetation,in forests to scrub vegetation or inMinas Geraisin forests. Deguelia dasycalyx is mainly asso-ciated with forested habitats, and may be foundin other localities of Serra do Espinhaço, ataltitudes close to 900 m.

Nomenclatural notes.—Although Glaziou(1906) identified the type specimen as L.floribundus, he also suggested a new species,L. sarmentosus, for another herbarium spec-imen (Glaziou 12599b), but this name wasnever validly published.Diagnostic features.—This species is easily

recognized by the aureate velutinous vestitureof its flowers and inflorescences, and by itsshort-petiolate and predominantly oblongleaflets, which are rusty-tomentose on thelower surface.

Selected specimens examined. BRAZIL. Bahia:Serra Geral de Caetité, 1,5 km S. de Brejinhos dasAmetistas, 11 Apr 1980, Harley 21253 (CEPEC). MinasGerais: Mariana, próximo a Ouro Preto, Feb–Mar 1882,Glaziou 12599b (K, P).

2. Deguelia glaucifolia A. M. G. Azevedo,Revista Brasil. Bot. 17 (1): 46. 1994. Type:Brazil. Amapá: Rio Araguari, FerreiraGomes, 29 Oct 1957, E. Pereira & W. A.Egler 3371 (holotype: RB; isotype: HBG).

Distribution and ecology.—This speciesprobably occurs only in Amapá. It is

CAMARGO & TOZZI: DEGUELIA (LEGUMINOSAE)2013]

FIG. 1. Deguelia dasycalyx. A. Flowering branch. B. Flower bud. C. Flower. D. Calyx. E. Standard. F. Wings. G.Keel petals. H. Staminal tube. I. Pistil. (A from Glaziou 15916, C; B–I from Glaziou 12599b, K.)

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cultivated at the Museu Paraense; the informa-tion from the herbarium label says that it wasbrought from the Rio Purus, which is likelyincorrect. In order to confirm the distribution ofthis species, more collections are needed.Diagnostic features.—Deguelia glaucifolia

is a distinctive species with elliptic leaflets thatare rounded and slightly emarginate at the apex,and glaucous beneath.Illustration.—Tozzi (1994: 49).

Selected specimens examined. BRAZIL. Pará:Belém, Museu Paraense, cultivated, brought from Rio Purus,20 Jan 1936, R. Siqueira s.n. (RB 35505, UEC).

3. Deguelia urucu (Killip & A. C. Sm.) A. M.G. Azevedo & R. A. Camargo, comb.nov. Lonchocarpus urucu Killip & A. C.Sm., J. Wash. Acad. Sci. 20, 81. 1930.Derris urucu (Killip & A. C. Sm.) J. F.Macbr., FieldMus. Nat. Hist., Bot. Ser. 13:266. 1943. Lonchocarpus nicou var. urucu(Killip & A. C. Sm.) F. J. Herm., J. Wash.Acad. Sci. 37: 112, 1947. Type: Brazil.Pará: Gurupá, 29 Sep 1916, A. Ducke s.n.MG 16561 (holotype: US [photo: UEC];isotypes: BM, NY [image], P). (Fig. 2)

Vernacular names.—Timbó-urucu, timbó,timbó-vermelho, dooui, e-yoo, koo-na, barbasco,timbó-uassú.Distribution and ecology.—From grasslands

to dense forests, from “terra firme” to perma-nently inundated areas, between 100–120 m.Because this species is cultivated, its distribu-tion has the potential to expand, but it seems notto have spread beyond where it is cultivated.This species was cited by Ducke (1949) withpossible occurrence in Amapá and byMacbride(1943) for Peru on the border with Brazil.Diagnostic features.—This species is closely

related to Lonchocarpus rufescens Benth. (stillnot transferred to Deguelia), which occurs inGuyana. Deguelia urucu can be distinguishedbecause it has broader leaflets with brownishhairs underneath and because it has inflores-cences that are smaller than the leaves.

Selected specimens examined. BRAZIL. Amazonas:Vic. of Jamamadi Indian village, Rio Apitua, tributary ofRio Purus, 1 Jul 1971, Prance et al. 13930 (INPA).Pará: Belém, s.d., Pires & Black 831 (IAC).

4. Deguelia nitidula (Benth.) A. M. G.Azevedo & R. A. Camargo, comb.nov. Lonchocarpus nitidulus Benth.,J. Linn. Soc., Bot. 4 Suppl.: 98.1860. Derris nitidula (Benth.) N. F.Mattos, Loefgrenia 98: 3. 1989. Type:Venezuela. “Ad Casiquiarem, in sylvisrecentioribus (Ad flumina Casiquiari,Vasiva et Pacimoru)”, Jan 1854, R.Spruce 3203 (lectotype, here desig-nated: K-117570 [image; photos: C,F]; isolectotypes: BM, P [photo:UEC]). (Fig. 3)

Lonchocarpus floribundus Benth., Ann. Nat. Hist. Ser. 1. 3:432. 1839. Derris floribunda (Benth.) Ducke, Bol. Técn.Inst. Agron. N. 18: 197. 1949, non Blume ex Miq., necNaves ex Villar. Type: Guiana, 1834, M. R. Schomburgk238 (lectotype, here designated: K [image; photos: C, F];isolectotypes: BM, L).

Vernacular names.—Timbó, timbó-rana,timborana, timbó-venenoso.Distribution and ecology.—Widely distrib-

uted in northern South America. In Brazil itoccurs in the states of Amazonas, Goiás,Maranhão, Mato Grosso, Pará, Piauí, Roraima,Minas Gerais, and São Paulo. Ducke (1925)recorded this species for Amapá. It also occursin Colombia, Guyana, French Guiana andVenezuela, and Macbride (1943) recorded itfor Peru. It is the only species in the genus thatgrows in “cerrado”, “campo cerrado” andsavannas. In the remaining vegetation types italways occurs in open areas, such as on theborder of secondary forest or on river banks,“capoeira” or “carrascal”. This species growsbetween 80–700 m, and it prefers sandy soils.Nomenclatural notes—Although the epithet

“floribundus” is older, having priority over“nitidulus”, it cannot be used because it isalready in use, existing for a Java vine asDeguelia floribunda (Miq.) Taub. (Taubert,1891). There are two different collections thatare considered types of L. nitidulus at K, andonly one of them was cited by Bentham in theprotologue (R. Spruce 3203).Diagnostic features.—Deguelia nitidula is

a polymorphic and widely distributed species;it can be identified by its chryso-sericeousindument on the flower, fruit, and lowersurface of the leaflet (where hairs are gener-ally short and adpressed).


FIG. 2. Deguelia urucu. A. Flowering branch and one leaf. B. Flower bud. C. Bracteole. D. Calyx. E. Standard. F.Wings. G. Keel petals. H. Staminal tube. I. Pistil. (From Pires & Black 831, IAC.)

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FIG. 3. Deguelia nitidula. A. Flowering branch. B. Flower bud. C. Calyx. D. Standard. E. Wings. F. Keel petals.G. Staminal tube closed, side view. H. Staminal tube longitudinally opened, front view. I. Pistil. (A from Andrade3309, UEC; B–I from Spruce 3203, P.)


Selected specimens examined. BRAZIL. Amazonas:Iranduba, Cacau Pereira, lado esquerdo, 1 km após o Porto,14 Ago 2003, Souza & Assis 248 (INPA).Goiás: Filadélfia,Rio Tocantins, 8 km N. da cidade, 4 Ago 1964, Prance &Silva 58555 (K, NY).Mato Grosso: Patronal, Vila Bela daSantíssima Trindade, BR-364 Cuiabá–Porto Velho, 15 kmda divisa comRondônia, 9 Jul 1984,Cid et al. 4381 (INPA).Pará: Porto Trombetas, mineração Rio do Norte, 1990,Soares & Knowles 841 (INPA). Roraima: Boa Vista, BR-174, 5 km à esquerda do km 20 km, direção Boa Vista–Mucajai, 23 Mar 1995, Miranda 488 (INPA). São Paulo:São Simão, em direção a Luiz Antônio, 12 May 1981,Leitão F. et al. 12531 (UEC).

5. Deguelia rariflora (Mart. ex Benth.) G. P.Lewis & Acev.-Rodr., Smithsonian Contr. Bot.98: 422. 2012. Lonchocarpus rariflorus Mart.exBenth., J. Linn. Soc., Bot. 4 Suppl.: 98. 1860.Derris rariflora (Benth.) J. F. Macbr., FieldMus. Nat. Hist., Bot. Ser. 13: 264. 1943. Type:Brazil. “Prope Barra, Prov. Rio Negro”, Aug1851, R. Spruce 1726 (lectotype, here desig-nated: K [image; photos: C, F]; isolectotypes:BM, M, NY [image], P). (Fig. 4)

Lonchocarpus cearensis Glaz., Bull. Soc. Bot. France 53:150. 1906, nom. nud.

Vernacular names .—Timbó-bravo,cipozinho, cipó-cururú, timbó-cururú e cipó-envira, timbó-rana, cururu, timbó-amarillo.Distribution and ecology.—Distributed in

northern Brazil (Amazonas state), Guyana andPeru (Bentham, 1862; Macbride, 1943; Ducke,1949; Tozzi, 1989), along the Rio Negro andRio Solimões basins, in open areas or “capoei-ra”, occasionally in “terra firme” forests, onsandy soils, at an altitudinal range of 25–200m.Although the cultivated specimen from Rio deJaneiro, Glaziou 9719 (C, K, P), has beenreferred as collected from Ceará state, there aresome questions about the correctness of thisinformation, and the specimen may have beenmislabeled.Diagnostic features.—Deguelia rariflora is

distinguished from the other species by itssmall flowers and pedicels, and by thevestiture of the corolla, subsericeous on thetop of the standard, becoming glabroustowards the base, and fruits. The majorvegetative character used for its identifica-tion is the rusty-villous lower surface ofthe leaflets. It is closely related to D.duckeana, which has similar leaf shape

and long sparse inflorescences but differsmainly by its leaf vestiture, smaller numberof leaflets (3 or 5), and indehiscent pod.

Selected specimens examined. BRAZIL. Amazonas:Rio Urubu, Manaus–Caracarai km 15, 4 Aug 1975, Prance& Forero 23580 (INPA, MO).

6. Deguelia duckeana A. M. G. Azevedo,Revista Brasil. Bot. 17 (1): 48. 1994.Type: Brazil. Amazonas: Rodovia Ma-naus–Itacoatiara, km 19, 15 Nov 1963,E. Oliveira 3033 (holotype: IAN;isotype: UB).

Vernacular names.—Cipó-cururu, timbó.Distribution and ecology.—Along the Rio

Amazonas basin and its tributaries (RioMadeira, Rio Urubu, and Rio Tapajós) inAmazonas state, reaching Pará state, inforests or dense “capoeiras”, often in non-inundated forests.Diagnostic features.—Deguelia duckeana

is very similar to D. rariflora, but it differs insize, number and vestiture of leaflets, and bythe presence of latex dots on the flower andthe tardily dehiscent pods.Illustration.—Tozzi (1994: 51).

Selected specimens examined. BRAZIL. Amazonas:Itacoatiara, AM-010, km 215, Faz. Aruanã, 7 Aug 2007,Souza B17 (INPA). Pará: Porto Trombetas, mineraçãoRio do Norte, Mina Saracá, 1999, Barbosa et al. 304(INPA).

7. Deguelia negrensis (Benth.) Taub., Bot.Centralbl. 47: 387. 1891. Derris negrensisBenth., J. Linn. Soc., Bot. 4 Suppl.:107. 1860. Lonchocarpus killipiiDucke, Trop. Woods 69: 5. 1942.Type: Brazil. Amazonas: “Secus RioNegro, inter Barcellos et San Isabel”,Dec 1851, R. Spruce 1926 (lectotype,here designated: K [photos: C, F];isolectotypes: B [photo: F], BM, NY[image], P). (Fig. 5)

Derris longifolia Benth., J. Linn. Soc., Bot. 4 Suppl. 107.1860. Deguelia longifolia (Benth.) Taub., Bot. Centralbl.47 (13): 387. 1891. Lonchocarpus longifolius (Benth.)Ducke, Trop.Woods 69: 5. 1942. Type: Brazil. Amazonas:Gapo do Rio Negro, Mar 1851, R. Spruce 1169 (lectotype,here designated: K-535168 [photo: UEC]; isolectotypes:K-535169 [photos: C, UEC], P).

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FIG. 4. Deguelia rariflora. A. Flowering branch. B. Flower bud. C. Calyx. D. Standard. E. Wings. F. Keel petals.G. Staminal tube. H. Pistil. I. Pods. (A–H from Glaziou 9719, P; I from Killip & Smith 30074, US.)


FIG. 5. Deguelia negrensis. A. Flowering branch. B. Flower bud. C. Flower. D. Calyx. E. Standard. F. Wings. G.Keel petals. H. Staminal tube closed, side view. I. Staminal tube longitudinally opened, front view. J. Pistil. (A fromPrance et al. 8029, US; B–J from Spruce 1926, P.)

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Vernacular names.—Timbó-rana, timbó-de-jacaré, barbasco-de-lagartillo, bejuco-guacuraya,uña-murcielago.Distribution and ecology.—In the states of

Pará, Rondônia, and Amazonas, mainly inthe basins of the Rio Negro and RioSolimões, reaching Venezuela, Colombia,Bolivia, and Peru; also cited for Piauí state(Bentham, 1862). Deguelia negrensis occursalong river banks, “igarapé”, “igapó”,“capoeira” or forests, in seasonally orpermanently inundated areas, and in non-inundated forests, in clay or sandy soils, up to450 m in altitude.Nomenclatural notes.—Derris negrensis

Benth. was based on two specimens collectedby Spruce (1906 and 1926) and the second isdesignated here as the lectotype, because it hasflowers at anthesis while the first collection hasflowers buds.Diagnostic features.—The suborbicular

fruit is the main character that distin-guishes this species from the closelyrelated species, Deguelia scandens. Inaddition, the inflorescence vestiture isbrownish tomentose becoming nigrescent,and the flowers are generally larger and thebracteoles usually smaller than in D.scandens. Sterile specimens of D. scandensand D. negrensis are difficult to distin-guish.

Selected specimens examined. BRAZIL. Amazonas:Manaus, próximo ao Rio Arara, 26 Apr 1973, Loureiro etal. s.n. (INPA 37770). Pará: Rio Trombetas shore,Porteira cemetery, 3 Jun 1974, Campbell et al. P22498(INPA, NY).

8. Deguelia scandens Aubl., Pl. Guian. 2:750, tab. 300, excl. fruit illustration.1775. Derris scandens (Aubl.) Pittier,Contr. U. S. Natl. Herb. 20: 41. 1917.non Benth., 1860. Lonchocarpusscandens (Aubl.) Ducke, Trop. Woods69: 5. 1942. Type: French Guyana, s. d.,J. B. Aublet s. n. (holotype: BM [photos:BM, MO, NY]). (Fig. 6)

Lonchocarpus pterocarpus DC, Prodr. 2: 260. 1825.Derris pterocarpus (DC) Killip, J. Wash. Acad. Sci.26: 360. 1936. Type: French Guiana, 1821, G.Perrottet 10 (holotype: G [photos: F, G]).

Derris guianensis Benth., J. Linn. Soc., Bot. 4 Suppl. 106.1860. Type: Guyana. High banks of Quitaro river, 1838,

M. R. Schomburgk 564 (lectotype, here designated: K-535165 [image; photos: C, F]; isolectotypes: BM, K-535166 [image], L, P).

Vernacular names.—Timbó, timbó-rana,timbó-assu, timbó-de-jacaré, assa-há pagaraundeguelé.Distribution and ecology.—Colombia,

Guyana, Venezuela, and Brazil (Amazonas,Maranhão, Pará and Roraima states). Don(1832) and Lemée (1952) cited this speciesfor French Guiana, Miquel (1844) forSurinam, and Macbride (1943) for Peru. Itgrows in “terra firme” forest, generally onriver banks and “igapó”, from 150–700 m.Diagnostic features.—Deguelia scandens is

closely related to D. negrensis based on itssimilar leaves and flowers, and to D.amazonica, based on the similar fruitshape. It can be distinguished from thelatter by the number of leaflets, inflores-cence size, flower color and size, corollashape, and fruit vestiture, which is sparselypubescent. Its flowers, like those of D.negrensis, are atypical in the genus, withnarrow, long-unguiculate petals, membranaceous,reddish brown in dried specimens, and some-times with yellow veins and margins. Degueliascandens is easily differentiated from D.negrensis by its linear-oblong pod with anacute to short acuminate apex.

Selected specimens examined. BRAZIL. Amazonas:Manaus, Estação Experimental de Silvicultura ZF2, 1997,Lemos 156 (INPA). Pará: Porto Trombetas, Mineração Riodo Norte, 1991, Evandro & Knowles 555 (INPA).

9. Deguelia amazonica Killip, J. Wash.Acad. Sci. 24: 48. 1934. Lonchocarpusnegrensis Benth., J. Linn. Soc., Bot. 4Suppl.: 98. 1860. Derris amazonicaKillip, J. Wash. Acad. Sci. 24: 48.1934. Type: Brazil. Amazonas: “RioNegro, ad oram septentrionalem flum.Amazonum, ad ostium Rio Negro”,Aug 1851, R. Spruce 1671 (lectotype,here designated: K-535171 [image];isolectotypes: B-n.v. [photo: F], BM,G, K-535170 [image; photos: F,UEC], K-535172 [image], LE, NY[image], P). (Fig. 7)

Vernacular names.—Urubu-timbó, timbó-da-caatinga, timbó-bravo, cipó-timbó, timbó,gwe gwe tete.


FIG. 6. Deguelia scandens. A. Flowering branch (apical leaflet missing). B. Flower bud. C. Calyx. D. Standard. E.Wings.F.Keel petals.G. Staminal tube.H. Pistil. (A fromDucke second collection 167, IAN; B–H from Schomburgk 564, K.)

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FIG. 7. Deguelia amazonica. A. Flowering branch. B. Flower bud. C. Flower. D. Calyx. E. Standard. F. Wings. G.Keel petals. H. Staminal tube. I. Pistil. (Drawn from Irwin et al. 55486, SP.)


Distribution and ecology.—In the states ofAmapá, Amazonas, Mato Grosso, Pará andRondônia, as well as in Colombia and Suriname.It has been cited for Peru and Guyana(Macbride, 1943), and in Venezuela (Bentham,1860) and probably also occurs in FrenchGuyana and Bolivia. This species grows inundisturbed forests, river banks and “igarapé”,generally preferring “terra firme”with clay soils,in non-inundated areas, although it can occur insandy soils and in seasonally inundated lowland.Nomenclatural notes.—Bentham (1860)

created the oldest specific epithet, and observa-tion of the vexilar margin in its fruit supportedits transfer to Derris, by Killip (1934), whocreated a new name for the species becauseTaubert (1891) had already proposed Deguelianegrensis for a different species.Diagnostic features.—Deguelia amazonica

is easily recognized by the shape and vestitureof leaflets, which are often oblong to oblong-ellipitic and ferruginous-tomentellous under-neath, by the suborbicular and glabrous stan-dard, and also by the indehiscent pods, whichare reddish tomentose with vexilar margin.

Selected specimens examined. BRAZIL. Amapá:Rio Araguari, foz do Anicahy, 8 Oct 1969, Pires et al. 51554(K, UB). Amazonas: Manacapuru, Mata do Pagé, RioSolimões, 8 Oct 1935, Krukoff 7705 (BM, K, P). MatoGrosso: Parque Estadual do Cristalino, a leste do RioCristalino, nas proximidades da margem do Rio Teles Pires,18 Jul 2006, Sasaki et al. 97 (INPA). Pará: Gurupá, 14 Aug1918, Ducke 17201 (BM, P, R). Rondônia: Abunã to RioMadeira, km 6, 15 Jul 1968, Prance et al. 6086 (INPA, NY).

10. Deguelia utilis (A. C. Sm.) A. M. G.Azevedo, Revista Brasil. Biol. 58: 512.1998. Lonchocarpus utilis A. C. Sm. inKrukoff & Smith, Amer. J. Bot. 24: 580.1937. Lonchocarpus nicou var. utilis (A.C. Sm.) F. J. Herm., J. Wash. Acad. Sci.37: 112. 1947. Derris utilis (A. C. Sm.)Ducke, Bol. Técn. Inst. Agron. N. 18:197. 1949. Derris nicou J. F. Macbr.,Field Mus. Nat. Hist., Bot. Ser. 13: 263.1943. Type: Peru. Loreto: Iquitos, 3–11Aug 1929, E. P. Killip & A. C. Smith27278 (holotype: US-1356611; isotypes:K [image], NY [image], US). Epitype:Colombia. Intendencia del Amazonas,Dept. Atlántico en el límite con Dept.Bolívar: Los Pendales, HaciendaRiodulce, 21–26 Jan 1946, A. Dugand

& R. Jaramillo 4156 (epitype, designatedby Tozzi, 1998: COL; isoepitype:MEXU-n.v. [image]).

Lonchocarpus nicou var. languidus F. J. Herm., J.Wash. Acad.Sci. 37: 111. 1947. Type: Colombia. Intendencia del Meta:Monte deMachadero, Ocoa, 4 km SEVillavicencio, 28 Jan1944, F. J. Hermann 11123 (holotype: US-2059191;isotypes: C-n.v., COL, US-2249632).

Deguelia alata M. Sousa, Rev. Mex. Biodiv. 80: 304.2009. Type: Colombia. Intendencia del Amazonas, Dept.Atlántico en el límite con Dept. Bolívar: Los Pendales,Hacienda Riodulce, 21–26 Jan 1946, A. Dugand & R.Jaramillo 4156 (holotype: MEXU-n.v. [image]; isotype:COL).

Vernacular names.—Timbó-grande, timbó-legítimo, timbó-macaquinho, timbó-urubu,timbó-verdadeiro, timbó, timbó-branco,barbasco, eobo wa’i, seamba, timu-ambi,timu, cube, cube-de-almedón, coapi, pacai.Distribution and ecology.—Along the Am-

azon Basin, extending from Pará and Amazonasto Acre states, reaching neighboring countries,such as Guyana, Colombia, Peru, and Ecuador.Ducke (1949) cites it for Amapá state. This plantis known from the pre-Columbian cultures andis only rarely found in subspontaneous state or inundisturbed forests (Ducke, 1949). Most speci-mens are sterile and collected from old indige-nous plantations or from cultivation areas called“barbascal”, also from plants growing in densesecondary forests that are not inundated orinundated (Tozzi, 1998). This species usuallyforms shrubs in “capoeiras”, forest gaps orshaded areas, near river banks, in clay soils ornot, neutral to strongly acid, from 20–500(−1300) m in altitude.Nomenclatural notes.—Deguelia utilis is

closely related to D. angulata. The formerwas only known from sterile material untilTozzi (1998) designated as epitype the col-lection A. Dugand & R. Jamarillo 4156(COL), which is fertile. However, Sousa(2009) recognized the same collection as typeof a new species, D. alata M. Sousa, whichhe said was close to D. amazonica. Althoughwe did not have access to the fruitingcollection cited by Sousa (2009), D. alata isput under synonymy here, because we do notagree with his statement based on A. Dugand& R. Jamarillo 4156; the vegetative charac-ters among different sterile collections of D.utilis are highly variable but still close theepitype. In addition, D. amazonica is a very

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distinct species in the genus and presents veryfew morphological affinities to D. utilis.Hermann (1947) designated a specimen atUS as holotype of L. nicou var. languiduswith isotypes in NA and C. We found twospecimens at US: US-2059191 which weassume is the holotype that Hermann (1947)referred to, and the isotype US-2249632,which has a stamp saying that it wastransferred to US in 1953.Diagnostic features.—Deguelia utilis is a

variable species, but some characters are con-stant, such as the venation pattern, and the textureand straight elliptic shape of leaflets, which haveacute bases and long caudate-acuminate apiceswith ferruginous hairs underneath.Illustration.—Tozzi (1998: 513).

Selected specimens examined. BRAZIL. Acre:Região do Rio Jurupari, bacia do Rio Juruá, 1936, Krukoff7718 (K). Amazonas: Lago Marrahã, Seringal SãoClemente, Rio Purus, 10 May 1975, Prance et al. 23414(INPA, NY).Pará:Tapaná, bacia do RioAmazonas, 25Oct1929, Pires et al. 51554 (K, RB, US).

11. Deguelia angulata (Ducke) A. M. G.Azevedo & R. A. Camargo, comb.nov. Lonchocarpus angulatus Ducke,Arch. Jard. Bot. Rio de Janeiro 4: 89.1925. Derris angulata (Ducke)Ducke, Bol. Técn. Inst. Agron. N. 2:29. 1944. Type: Brazil. Pará: BelaVista, Rio Tapajós 23 Jul 1923, A.Ducke s.n. (holotype: RB-5057 [pho-to: UEC]). (Fig. 8)

Lonchocarpus sylvestrisA. C. Sm., in Krukoff & Smith,Amer. J. Bot. 24: 576. 1937. Derris sylvestris (A.C. Sm.) J. F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13:266. 1943. Type: Peru. Loreto: AlongRioMaranon, nearmouth of Rio Tigre, 19 Aug 1929, E. P. Killip & A. C.Smith 27528 (lectotype, here designated: US-1356619[image]; isolectoypes: US-1356620 [image]; NY[image]).

Vernacular names.—Timbó-rana, timbó-bravo, timbó-do-mato, timbó-branco, sacha-barbasco, barbasco-del-monte.Distribution and ecology.—Distributed

through the Brazilian Amazon (Acre, Amazo-nas, Pará, and Rondônia states), also in thePeruvian, Bolivian and Colombian Amazon.Ducke (1949) cited this species for northern

Mato Grosso state. The species grows inriverine forests, on flooded river banks. Itspreferential habitat seems to be associated with“várzeas”, but it is rarely found in “terra firme”forests, from 20–400 m.Nomenclatural notes.—The synonymy of

L. sylvestris and D. angulata was establishedby Tozzi (1989), based on a lack of consistentdifferences between the two. For the lecto-type of L. sylvestris, we selected the bestquality specimen, which bears flowers, fromamong the duplicates.Diagnostic features.—Flowering material

can be recognized by the inflorescences, whichdevelop on leafless branches, as well as thecalyx with the lower central tooth relativelylonger and the corolla having a yellowish colorin dried material. Sterile material can berecognized by the presence of short adpressedtrichomes on the under surface of the leaflets.

Selected specimens examined. BRAZIL. Acre: RioBranco P. Z., entrada do prédio que dá acesso ao Parque,lado direito, 10 Ago 1992, Claros 170 (INPA). Rondônia:Abunã, E. bank of Rio Madeira, 12 Jul 1963, Prance et al.5903 (INPA, NY).

12. Deguelia costata (Benth.) A. M. G.Azevedo & R. A. Camargo, comb. nov.Lonchocarpus costatus Benth., J. Linn.Soc., Bot. 4 Suppl.: 100. 1860. Derriscostata (Benth.) N. F. Mattos, Loefgrenia98: 3. 1989. Dalbergia costata Mart. exBenth., Fl. Bras. 15: 287. 1862, pro syn.Type: Brazil. Minas Gerais: Cachoeirado Campo, 1839, P. Claussen 181(lectotype, here designated: K [image];isolectotypes: G, P). (Fig. 9)

Vernacular names.—Embira-de-carrapato,pau-de-carrapato, pau-carrapato.Distribution and ecology.— Minas Gerais,

Espírito Santo and Rio de Janeiro states,growing on forested plateaus, in forest rem-nants, and in gallery forests.Nomenclatural notes.—We have chosen the

lectotype Claussen 181 deposited at K, amongthe syntypes cited by Bentham (1862), becausethis material contains vegetative and reproduc-tive characters that best fit the original descrip-tion (Bentham, 1860).Diagnostic features.—Although this species

has some variation in the vestiture of the standard,


FIG. 8. Deguelia angulata. A. Vegetative branch. B. Flowering branch. C. Flower bud. D. Flower. E. Calyx. F.Standard. G. Wings. H. Keel petals. I. Staminal tube closed, side view. J. Staminal tube longitudinally opened, frontview. K. Pistil and disc. (A–B from Prance et al. 5903, COL; C–K from Krukoff 4871, K.)

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FIG. 9. Deguelia costata. A. Flowering branch. B. Flower bud. C. Calyx. D. Standard. E. Wings. F. Keel petals.G. Staminal tube. H. Pistil and disc. (A from Glaziou 12599, C; B–H from Warming 783, C.)


pedicel, and lower surface of the leaflets, thesecharacters show continuous variation. Thisspecies can be recognized because the lowersurface of the leaflets has prominent secondaryveins and dense vestiture, and also the standardis often oblong-obovate with golden hairs.

Selected specimens examined. BRAZIL. MinasGerais: Paraopeba, Faz. do Funil de Guilherme, 31 Mar1960,Heringer 7477 (RB, SP, UB).Espírito Santo:Castelo,Caxixe Quente, 18 May 1999, Hatschbach et al. 69149(CEPEC, MBM, SPF).

13. Deguelia hatschbachii A.M. G. Azevedo,Revista Brasil. Bot. 17: 45. 1994. Type:Brazil. Minas Gerais: Manhumirin, 9Feb 1973,G.Hatschbach& Z. Ahumada31393 (holotype: MBM [photo: UEC];isotypes: C, MEXU [image]).

Distribution and ecology.— Rio de Janeiro,Espírito Santo, and Minas Gerais states, as aheliophyte, in semi-deciduous forests. It isfrequent around the Rio Manhuaçu, in south-eastern Minas Gerais. It also grows in denseombrophilous forest.Diagnostic features.—The major diagnostic

characters are the size and shape of the floralparts and the generally elliptic leaflets, slightlydiscolorous and sericeous on both surfaces.This species is closely related to D. spruceanaand D. costata, and is distinguished from theseby the longer yellowish pods, inflorescenceaspect and leaflets.Illustration.—Tozzi (1994: 47).

Selected specimens examined. BRAZIL. MinasGerais: Descoberto, Reserva Biológica da Represa doGama, 21 Feb 2001, Castro et al. 130 (HUEFS, UEC).Rio de Janeiro: Rio Claro, 17 Mar 1978, Lima 272 (RB,UEC). Espírito Santo: Pancas, 14 Dec 1977, Marx s.n.(RB 180119).

14. Deguelia spruceana (Benth.) A. M. G.Azevedo & R. A. Camargo, comb. nov.Lonchocarpus spruceanus Spruce exBenth., J. Linn. Soc., Bot. 4 Suppl.:99. 1860. Derris spruceana (Benth.)Ducke, Bol. Técn. Inst. Agron. N. 18:199. 1949. Type: Brazil. Pará: “Invicinibus Santarém”, Apr–Aug 1850,R. Spruce s.n. (lectotype, here desig-nated: K [image]; isolectotypes: BM,C [photo: UEC], F-875625, LE, M,NY-15742 [image], P). (Fig. 10)

Vernacular names.—Favinha, sucupirinha-do-campo, timbó-pau, turiuva, embira-de-sapo, aquiqui, facheiro.Distribution and ecology.—Amazonas, Pa-

rá and Maranhão states, in non-inundated“terra firme” forests, “capoeiras” and grass-lands. A heliophyte species, frequentlyfound close to water streams in sandysoils.Diagnostic features.—A well defined spe-

cies, with an arborescent habit up to 20 mhigh, strongly discolorous leaflets that areshiny and subsericeous on the uppersurface, and glaucous and sericeous ortomentose on the lower surface. In addi-tion, the pedicel is slender and long, thestandard is canescent on the upper surface,and the pods are tardily dehiscent with upto ten seeds and brownish with nerviform,undulate margins.

Selected specimens examined. BRAZIL. Amazonas:Manaus, BR-17 km 7, 18 Sep 1962, Rodrigues & Chagas4645 (INPA, US). Pará: Alto Curuauna, Centro deTreinamento da FAO, 6 Oct 1962,Duarte 7259 (INPA, RB).

15. Deguelia densiflora (Benth.) A. M. G.Azevedo ex M. Souza, RevistaMex. Biodiversidad 80: 306. 2009.Lonchocarpus densiflorus Benth., Ann.Nat. Hist., ser. 1, 3: 437. 1839. Type:Guyana. banks of the upper Rio Essequi-bo and Rio Rupununi, 1837, M. R.Schomburgk 52 (lectotype, here desig-nated: K [image; photos: C, F]; isotypes:BM [photo: UEC], L, P). (Fig. 11)

Lonchocarpus glabrescens Benth., Hooker´s J. Bot. KewGard. Misc. 2: 333. 1850. Derris glabrescens (Benth.)J. F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13: 261.1943. Type: Brazil. Pará: “In vicinibus Pará”, Aug1849, R. Spruce 248 (lectotype, here designated: K-117575 [image; photos: C, F]; isolectotypes: BM[photo: UEC], K [photo: UEC], NY [image]).

Lonchocarpus boliviensis Pittier, Contr. U. S. Natl. Herb.20: 93. 1917. Type: Bolivia. Beni: junction of the riversBeni and Madre de Dios, Aug 1886, H. Rusby 2362(holotype: US-32680 [image]; isotypes: BM, F, GH-n.v.,K [image], MI, MO [image], NY [image], US-204454[image], US-477659 [image], US-1337541 [image]).

Lonchocarpus ulei Harms ex Ule, Bot. Jahrb. Syst. 40:142. 1907, nom. nud.

Vernacular names .—Timbó-der r i s ,barbasco, bejuco, bastard-hiarry.

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FIG. 10. Deguelia spruceana. A. Flowering branch. B. Flower. C. Calyx. D. Standard, side view. E. Standard,front view. F. Wings. G. Keel petals. H. Staminal tube. I. Pistil and disc. (A from Ducke s.n., MG-15275, BM; B–Ifrom Spruce s.n., K.)


FIG. 11. Deguelia densiflora. A. Flowering branch. B. Flower. C. Calyx. D. Standard. E. Wings. F. Keel petalsclosed, side view. G. Keel petals opened, front view. H. Staminal tube. I. Pistil and disc. J. Pod. (A–I from Spruces.n., BM; J from Fróes 23985, IAN.)

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Distribution and ecology.— Amapá, Amazo-nas, Pará and Roraima states in Brazil, and alsoin Bolivia, Colombia, Guyana, Peru and Vene-zuela, extending to Panama and Costa Rica.This species seems to be associated to “várzeas”and flooded or inundated areas, such as riverbanks or in locations around lakes. It is frequentin tall and dense gallery forests or in more orless dense forests such as “igapós”, occasionallyin “capoeiras”, forage grasslands or savannas. Itoccurs from sea level up to 300 m.Nomenclatural notes.—Ule 5541 was cited

as L. ulei for Flora do Amazonas (Ule, 1907),but this name was not validly published. LaterHarms (1921) synonymized this name under L.glabrescens. The analysis of many specimensshows a gradation of morphological characters,and the inflorescence position is very variable,which is why L. glabrescens and L. boliviensisare placed in synonymy here.Diagnostic features.—Deguelia densiflora

is recognized by its pods, which are the onlyones in the genus that are woody, slightlyconstricted between the seeds, and at maturitybreaking transversally into segments with one ortwo seeds each; in addition, its flowers have arigid staminal tube that is strongly recurved.

Selected specimens examined. BRAZIL. Amapá:Baciado Rio Oiapoque, Rio Urucauá, 17 Oct 1960, Irwin 48777 (K,UB).Amazonas: Lago da Cobra, Rio Purus, N bank oppositeBoca do Acre, 19 Sep 1966, Prance et al. 2418 (INPA, K).Roraima: Banks of Rio Mucajaí, between Pratinha and RioApiaú, 26 Jan 1967, Prance et al. 1967 (INPA).

Acknowledgments

The authors thank all herbaria curators of thefollowing institutions for allowing the study ofspecimens via loans or during personal visits:BHCB, BM, C, CEPEC, COL, EAC, F, G,HUEFS, IAC, IAN, INPA, K, L, LE, LP,MBM, MO, NA, NY, P, RB, S, SP, UB, UECand US. We are also thank Esmeralda ZanchetaBorghi for the preparation of the illustrationsand two reviewers of this manuscript for theirevaluation. This paper is part of PhD. thesis inthe Programa de Pós-Graduação em BiologiaVegetal, Universidade Estadual de Campinas,and supported by CNPq and CAPES.

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Documents

A synopsis of the genus Deguelia (Leguminosae, Papilionoideae, Millettieae) in Brazil