A revision of the genus Myroxylon (Leguminosae

KEW BULLETIN (2015) 70:48 DOI 10.1007/S12225-015-9604-7

ISSN: 0075-5974 (print) ISSN: 1874-933X (electronic)

A revision of the genus Myroxylon (Leguminosae: Papilionoideae) Ângela Lúcia Bagnatori Sartori1, Gwilym P. Lewis2, Vidal de Freitas Mansano3 & Ana Maria Goulart de Azevedo Tozzi4

Summary. Myroxylon, an exclusively neotropical legume genus, is taxonomically revised to include two species.

Seven lectotypes, one neotype and three synonyms are designated. M. peruiferum and M. balsamum can be differentiated by pellucid dots and streaks in leaflets, adaxial surface dull or shiny, shapes of calyx lobes and standard, size of flowers and the standard petal blade and seed surface. A key to the two species is presented, together with species descriptions, illustrations and comments on the geographical distribution and ecology of each. Key Words. Amburaneae clade, Fabaceae, Neotropics, Sophoreae, taxonomy.

Introduction Myroxylon L. f. is a neotropical genus of legume (Map 1). Past treatments of the genus have varied greatly in the number of species recognised, from 1 to 12 (Hutchinson 1964; Rudd 1968; Polhill 1981; Pennington et al. 2005) with a correspondingly variable number of synonyms recognised. Affinities between Myroxylon and Myrospermum have been recognised in taxonomic studies (e.g., De Candolle 1825; Hutchinson 1964) and molecular analyses (Doyle 1995; Doyle et al. 1997; Wojciechowski et al. 2004; Cardoso et al. 2012). Anatomical studies carried out by Sartori & Tozzi (2002) reinforced the relationship between Myroxylon and Myrospermum. Recent molecular analyses resolve Myroxylon as a monophyletic genus (Cardoso et al. 2012). Historically, the most comprehensive taxonomic treatment of Myroxylon was published by Harms (1908), who recognised two species and three varieties, although his treatment included no identification key of the taxa, nor information about geographical distribution, and no illustrations of the species. More recently studies by Rudd (1968) and Isely (1981) have been published, but there has been no overview of the genus since 1981.

Even though the genus is well defined, the circumscription of its constituent species has been controversial. The taxonomic revision of Myroxylon presented here re-evaluates the status of the taxa that have been included in the genus, based mainly on morphological characters. A distribution map, species illustrations and descriptions are presented.

Material and methods This study is based on a revision of more than 450 specimens from the following herbaria: B, BA, BHCB, BM, C, CR, CVRD, ESA, FUEL, G, INPA, K, L, MEXU, MO, NY, PEUFR, R, RB, S, SP, SPF, US, UEC, VIC (acronyms according to Holmgren et al. 1990). Morphological, phenological and geographical distribution data were recorded from herbarium labels and by direct field observations. Specimens collected for the study have been deposited in the UEC herbarium. The species descriptions are based on the examination of 40 vegetative and reproductive morphological characters of herbarium specimens; the measurements are always of the subterminal leaflets and of flowers in pre-anthesis. Terms used in the descriptions for leaflet shape and indumentum type follow Radford et al. (1974), those of inflorescence type are based on Lawrence

Accepted for publication 14 October 2015. 1 Centro de Ciências Biológicas e da Saúde, Laboratório de Botânica, Universidade Federal do Mato Grosso do Sul, Campo Grande, MS, Caixa Postal 549, 79070-900, Brazil. e-mail: [email protected] 2 Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK. 3 Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, DIPEQ, Rua Pacheco Leão 915, Jardim Botânico, Rio de Janeiro, RJ, 22460-030, Brazil. 4 Departamento de Biologia, Universidade Estadual de Campinas, Instituto de Biologia, Campinas, Caixa Postal 6109, 13083-970, Brazil.

© The Board of Trustees of the Royal Botanic Gardens, Kew, 2015

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Map 1. Distribution map of Myroxylon.

(1971), and description of seed ornamentation follows the definitions of Stearn (1973). The map was prepared using Versamap version 2.03.

History of the genus Linnaeus described Toluifera balsamum in 1753 (Table 1), based on material collected in Cartagena, Colômbia. Later, Linnaeus filius (1781) described Myroxylon peruiferum based on a specimen collected by Mutis in South America. The circumscription of the genus Myroxylon accepted by Linnaeus filius (l.c.) was based on the 5-toothed campanulate calyx, a pentamerous corolla and a single-seeded legume. Willdenow (1799) expanded the circumscription of Myroxylon to include the synonyms Myrospermum frutescens Jacq. and Myrospermum pedicellatum Lam., based on the same diagnostic characters used by Linnaeus filius (l.c.); the author also recognised Toluifera as a separate monotypic genus. Stokes (1812) described Myroxylon abruptifolium Stokes with abruptly bijugate leaves and Myroxylon trialatum Stokes with a tri-winged stem, but did not explicitly establish the relationship of his two species with any of the previously described species of Myroxylon. Kunth (1823), was one of a few authors who explicitly discussed the morphological flower and fruit differences between Myroxylon and Myrospermum. He published Myroxylon pubescens based on a specimen from Cartagena collected by Humboldt, proposed the new name Myroxylon toluiferum for Toluifera balsamum L., and considered Myrospermum pedicellatum a synonym of Myroxylon peruiferum L. f. De Candolle (1825) placed Myroxylon L. f. and Toluifera L. in synonymy under Myrospermum Jacq., and © The Board of Trustees of the Royal Botanic Gardens, Kew, 2015

KEW BULLETIN (2015) 70:48

recognised two sections in the genus: section Calusia, characterised by persistent stamens, comprised the single species Myrospermum frutescens Jacq., and section Myroxylon, with caducous stamens, which included the species M. peruiferum, M. pubescens Kunth and M. toluifera Kunth. Klotzsch (1843) re-established the genus Myroxylon diagnosed by caducous stamens and sagittate anthers versus the persistent stamens and rounded anthers of Myrospermum. He also described Myroxylon punctatum Klotzsch based on collections from different regions of Peru, but these syntypes lack collector information. Walpers (1843), followed the treatment of De Candolle (1825) and transferred Myroxylon punctatum Klotzsch to Myrospermum sect. Myroxylon (L. f.) DC. Klotzsch (1857) did not accept the treatment of De Candolle (l.c.) nor that of Walpers; he recognised eight species in Myroxylon, proposed a new combination M. pereirae (Royle) Klotzsch and described two new species: M. hanburyanum Klotzsch based on specimens from the Amazonian region, and M. robiniaefolium Klotzsch based on a collection of Warszewicz, from Chongon, Guayaquil provence, Ecuador. Based on the principle of priority, Baillon (1870) adopted the generic name Toluifera, recognising T. peruifera (L. f.) Baill., T. pubescens (Kunth) Baill., T. punctata (Klotzsch) Baill., T. pedicellata (Lam.) Baill., T. balsamum (L.) Baill. and T. pereirae (Royle) Baill. He considered Toluifera to be characterised by its regular corolla, and its seeds and cotyledons kidney-shaped in contrast to Myrospermum with a papilionaceous corolla, the seeds oblong and the cotyledons auriculate. Bentley & Trimen (1880a, b) recognised only two species of Toluifera: T. balsamum and T. pereirae diagnosed by floral characters only. T. balsamum was characterised by its tubular or oblong, sparsely-hispid, 3 – 4-lobed calyx, a standard petal with a linear claw and the stamen filaments not exserted from the calyx. T. pereirae was differentiated by its campanulate, rufous-tomentose, 5-lobed calyx, a standard petal with a linear-triangular claw and the stamen filaments exserted from the calyx. Three years later, Baillon (1883) differentiated Toluifera peruifera, with rough seeds, from T. pereirae, with smooth seeds. The author placed Myroxylon pubescens in synonymy under T. peruifera, citing the fact that neither produce the ‘Balsam of Peru’. He also proposed varietal names in Toluifera balsamum, namely: T. balsamum var. pereirae, T. balsamum var. punctatum and T. balsamum var. balsamum, all three producers of the ‘Balsam of Tolu’. The treatment of Myroxylon by Harms (1908) mostly was based on the studies of Baillon (1883). Harms recognised M. peruiferum and M. balsamum, the latter

Mm. erythroxylum

Allemão 1857

Mn. peruiferum

Linnaeus f. 1781

Lamarck 1797

Mm. pedicellatum

T=Toluifera, Mm. = Myrospermum, Mn. = Myroxylon.

Mm. pereirae

Royle 1853

T. balsamum

Linnaeus 1753

Table 1. Historical changes in the classification of Myroxylon.

T. pubescens

Mn. pubescens Mn. punctatum

Mn. robiniaefolium Mn. hanburyanum

T. pereirae

T. peruifera T. pedicellata

Mn. peruiferum Mn. pedicellatum

Mn. pereirae

T. balsamum

Baillon 1870

Mn. abruptifolium Mn. trialatum

Stokes 1812

Mn. toluiferum

Klotzsch 1857

T. balsamum Mn. peruiferum Mn. pedicellatum

Willdenow 1799

T. balsamum var. punctatum T. balsamum var. pereirae

Mn. balsamum var. balsamum

Mn. peruiferum Mn. balsamum var. punctatum Mn. balsamum var. pereirae Mn. peruiferum

Mn. balsamum var. balsamum Mn. peruiferum Mn. peruiferum Mn. peruiferum

T. balsamum var. balsamum T. peruiferum

Mn. punctatum

Klotzsch 1843

Harms 1908

Mm. pubescens

Mm. toluiferum Mm. peruiferum

De Candolle 1825

Baillon 1883

Mn. pubescens

Mn. toluiferum Mn. peruiferum

Kunth 1823

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with three varieties: var. balsamum, var. pereirae and var. punctatum. He did not see the type specimen of M. abruptifolium Stokes, but, nevertheless, placed the name in synonymy under M. peruiferum. Harms (l.c.) considered Myroxylon robiniaefolium Klotzsch as a species of uncertain status and proposed it be treated as a synonym of M. peruiferum. In the Index Nominum Genericorum (Farr et al. 1979) Myroxylon L. f. is a conserved name (“nomina conservanda”) against Toluifera L. which is rejected (“nomina rejicienda”). In addition, there exists a homonymic genus name Myroxylon J. R. Forst. (in family Flacourtiaceae); this name is also rejected against Xylosma G. Forst. The orthographic variant “Myroxylum” of Schreber was placed in synonymy under Myroxylon L. f. by Farr et al. (1979). In nomenclatural summary, Myroxylon L. f. is a conserved name with Toluifera L. as a synonym. The genus comprises two species: M. peruiferum and M. balsamum.

Ethnobotany and phytochemistry of balsam The term myron is Greek for sweet oil or perfume (Stearn 1973) and xylon is Greek for wood, the two together combined to refer to the aromatic wood of Myroxylon, from which both the Balsam of Peru and Tolu balsam of commerce are extracted (Pennington et al. 2005). Balsam, mainly extracted from the stems of Myroxylon is included as a component of healing ointments, and as a flavouring in syrups and soaps (Fluckiger & Hanbury 1874). It is also used as a remedy for nasal congestion, stomach pains and rheumatism and in the manufacture of perfumes (Linares & Bye 1987). Historically, different species of Myroxylon had their branches deliberately cut to obtain balsam, which was exported mainly to Europe. Fluckiger & Hanbury (1874) cited the first published reference about Tolu Balsam to be 1574, when Monardes described local Amerindians collecting the balsam. Used by the Spanish, this balsam was considered to be as good a quality as


the famous ‘Balsam of Meca’, used in religious ceremonies. The exudate obtained from Myroxylon balsamum is chemically characterised by a benjoin or vanillin fragrance and it crystallises (Fluckiger & Hanbury 1874). The exudate obtained from M. pereirae (in this study treated as a synonym of M. balsamum) stems has been characterised as a mixture of aromatic esters and alcohol, including styracin, nerolidol, vanillin, benzoic and cinnamic acids (Biemer et al. 1992). Chemical studies of M. peruiferum have reported the presence of essential oils composed principally of nerolidol, together with other components including vanillin and a number of flavonoids (Maranduba et al. 1979).

Taxonomic treatment Myroxylon L. f. (Linnaeus filius 1781: 233). Type: Myroxylon peruiferum L. f. Tree. Branches terete, lenticellate, stipules caducous. Leaves imparipinnate, to 15-foliolate; petiole flattened, rough; rachis square or terete in cross section, sulcate on the adaxial surface; petiolules wrinkled, leaflets alternate, venation brochidodromous, pellucid gland dots and streaks conspicuous in the leaflet blades (on herbarium specimens and in the field these are best seen when a leaflet is held up to the light). Racemes axillary and/or terminal; a bract inserted at the base of the pedicel; minute bracteoles inserted at the base of the calyx, caducous; calyx campanulate, 5-lobed (the lobe apices obtuse) or the lobes obsolete; the sub-papilionaceous flowers spirally arranged along the inflorescence axis, petals white; standard petal with a greenish central blotch; stamens 10 (5 longer), all united at base, filaments and anthers glabrous, anthers dorsifixed, oblong, apiculate, sagittate, dehiscence longitudinal; gynoecium uniovulate, glabrous, stipitate, stigma punctiform. Fruit a 1-seeded samara, wing proximal, veins crowded submedially, seed-chamber distal; testa smooth or rough.

Key to the species of Myroxylon 1.

Pellucid dots and streaks inconspicuous unless leaflet held up to the light, adaxial surface dull; calyx 5-lobed; standard petal depressed-ovate; seed surface smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. M. balsamum 1´ Pellucid dots and streaks conspicuous on leaflet, adaxial surface shiny; calyx lobes obsolete; standard petal broadly ovate; seed surface rough . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. M. peruiferum

1. Myroxylon balsamum (L.) Harms (1908: 94). Basionym: Toluifera balsamum L. (Linnaeus 1753: 384). Type: Venezuela: Puerto Cabello, s.d.,


A. Goering 3/69? (neotype K, selected here, isoneotype NY 02256475). We have not been able to locate any original material of Toluifera balsamum L. (pers. comm.


Cafferty, Linnean Plant Name Typification Project) or any type material studied by Harms in Berlin. In consequence, we here designate as a neotype a specimen collected in Puerto Cabello, which best fits (as to geographical locality) the species description published in the protologue. Myroxylon balsamum (L.) Harms var. balsamum. (1908: 94). Myroxylon balsamum var. pereirae (Royle) Harms (1908: 95). Myroxylon balsamum var. punctatum (Klotzsch) Harms (1908: 97). synon. nov. Myroxylon hanburyanum Klotzsch (1857: 275). Type: America do Sul, Rio Amazonas, Maranon, P. A. J. de Warszewicz s.n. (lectotype BM53843!, selected here, Herb. J. J. Triana!). The material chosen as a lectotype is one of the originally cited syntypes and has the characteristics described in the protologue of the species. Myrospermum pereirae Royle (1853: 414). Type: Guatemala, s.d., Skinner, s.n. (lectotype K! [Herb. Hookerianum], selected here). The specimen chosen as a lectotype is material collected by Skinner from Guatemala and analysed by Pereira (1851). Myroxylon pereirae (Royle) Klotzsch (1857: 274). Myroxylon punctatum Klotzsch (1843: 12). Type: Peru, Chicoplaya, Pozuzo, Pavon s.n. (lectotype G 00404304!, selected here). Klotzsch (1843), when describing his new species cited a number of localities from Peru: Pozuzo, Muna and Cuchero, in the region of the Maranon river, but provided no additional data about collectors or collection numbers. In 1857, the same author cited Ruiz & Pavon as the collectors of M. punctatum. The material designated here as a lectotype is one of the original syntypes. Myrospermum punctatum (Klotzsch) Walp. (Walpers 1843: 805). Myroxylon toluifera Kunth (1823: 375). Type: Colombia, Turbaco, s.d., Bonpland 1405 (lectotype P 6710270 selected here, photo!). The specimen chosen as a lectotype best fits the original description and is in a good state of preservation. Myrospermum toluifera (Kunth) DC. (De Candolle 1825: 95). Myrospermum balsamiferum Ruiz & Pavon apud Lopez (1956: 717). Type: Ruiz et Pavon, l.c. tab. 373 synon. nov. Materials collected in Peru by Ruiz & Pavon were identified by them as Myrospermum balsamiferum (BM 53811, 53814, 53815, 53826, 53831). Although the name was a nomen nudum at the time it was synonymised by Harms (1908) under M. punctatum. According to Stafleu (1965) and Stafleu & Cowan (1983) the work of Ruiz & Pavon was only partially published in the late eighteenth century; this included an illustration

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of M. balsamiferum, but no species diagnosis. Much later the name was effectively published (Ruiz & Pavon apud Lopez 1956) with a species description. Based on the morphological similarity we propose the name as a synonym of Myroxylon balsamum. Myrospermum of Sonsonate, Pereira (1851: 281). nom. inv. The author doubted the validity of his new species: “For the present, therefore, I shall designate the plant which I have received as the Myrospermum of Sonsonate”. This invalidates the name, according to McNeill et al. (2011), article 36.1 b. Myrospermum sonsonatense Oerst. (Oersted 1855: 27). nom. superfl., illegit. Oersted based his name on the same material as Pereira (1851). Toluifera balsamum Baill. var. balsamum (Baillon 1883: 676). Toluifera balsamum var. pereirae (Royle) Baill. (Baillon 1883: 676). Toluifera pereirae (Royle) Baill. (Baillon 1870: 383). Toluifera balsamum var. punctata Baill. (Baillon 1883: 677). Tree (8 –) 15 – 50 m tall. Leaves 5 –11-foliolate, petiole, rachis and petiolules sparsely ferruginous-tomentose; petiole 1 – 2.7 cm long; rachis 2.7 – 12 (– 17) cm long; petiolules 2 – 6 mm long; leaflets 5.5 – 10.5 2 – 5 cm, oblong to elliptic, apex acuminate or long-acuminate, base attenuate or rounded, terminal leaflets elliptic to rhombic, the lower pairs ovate, margin sinuous, entire, concolorous or discolorous, both surfaces glabrous or sparsely sericeous on the blade and midrib, adaxial surface dull (not shiny), veins prominent on both surfaces, pellucid dots and streaks inconspicuous unless leaflet held up to the light. Racemes terminal and axillary, equalling or longer than the leaves, the axis ferruginoustomentose or greyish tomentose, (4.5 –) 16.5 – 19 cm long; bract lanceolate, apex acute, base truncate, concave or not, ascending to reflexed, c. 1 mm long; pedicel 5 – 15 mm long. Flower buds globose or cylindrical, apex acute or obtuse; bracteole ovate or narrowly oblong, up to 0.5 mm long. Flowers c. 12 – 14 mm long; calyx 5 – 8 11.5 – 15 mm, greyish tomentose to puberulous externally, the lobe apices obtuse; standard petal 10.5 – 14.5 mm long, blade 2.8 – 5.5 4.6 – 8.8 mm, claw 7.2 – 9.3 mm long, depressed-ovate, apex rounded or emarginate, base rounded to subcordate, margin entire, externally glabrous; other petals (6 –) 8.7 – 11.5 (0.36 –)1.1 – 2.6 mm, lanceolate to linear, apex acute, base attenuate, margin rarely sinuous, glabrous; stamens (4.7 –) 7 – 10 mm long, filaments 1.5 – 5 mm long, anthers 2.3 – 5.5 mm long, ovary 10 – 12.6 mm long, stipitate. Fruit a 6 – 12 cm long samara; seed-chamber 2 – 3 cm long, sub-globose; upper margin 1 – 17 mm wide; lower


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margin 1 – 5 mm wide, apiculate; stipe 6 – 13 mm long; bracts reflexed, persistent, or bracts and calyces caducous; seed testa smooth. Fig. 1. DISTRIBUTION. Myroxylon balsamum occurs in Central America, and northern and western South America (Map 1). SELECTED SPECIMENS EXAMINED. BELIZE: Cayo, July 2006, T. Sarkinen 236 (K). BOLÍVIA: Beni, Gral José Ballivian, June 1989, D. N. Smith, M. Buddensiek & V. Garcia 13578 (K, LPB, MO); Vale do Rio Bopi, Oct. 1921, H. Rusby 668 (K, NY). BRAZIL: Acre, próximo ao Rio Macauhan, tributário do Rio Yaco, Aug. 1933, B. A. Krukoff 5484 (B, BM, G, K, NY, US); Marechal Taumaturgo, April 1993, D. C. Daly et al. 7715 (MEXU, NY); Porto Alegre, rio Acarí, March 1948, R. L. Fróes 23112 (NY); Rio Branco, BR 317 KM 68, Sept. 1990, R. S. Saraiva et al. 1069 (NY); Sena Madureira, Rio Macauhan, March 1994, L. Lima et al. 535 (NY); Seringal Iracema, March 1933, A. Ducke 23786 (G, K, S, US); Xapuri, Sept. 1989, K. A. Kainer 104 (NY); Pará, Belém, Jardim Botânico Museu Goeldi, Oct. 1942, W. A. Archer 7714 (NY). COLÔMBIA: Boyaca, Tuaguagui, vale do Rio Negro, July 1917, H. N. Whitford & J. Pinzon 9 (US); Bolívar, entre Sincelejo e Colosó, Dec. 1962, R. R. Castaneda 9250 (NY); Magdalena, Oct. 1959, R. Romero-Castañeda 7671 (NY); Vale do Cauca, 1846, Gourlat s.n. (G 8203108). COSTA RICA: Cantón Central, San Jose, Jardim do Museu Nacional, Dec. 1995, A. Cascante 922 & R. Chacón (CR), Puntarenas, Osa Peninsula, Feb. 1995, R. T. Pennington, N. Zamora & R. Aguilar 588 (K). CUBA: Havana, Lomas de Canoa, Sept. 1923, E. L. Ekman 17553 (G, NY, S). EL SALVADOR: La Libertad, San Andrés, Feb. 1959, P. H. Allen 7216 (US); San Francisco, Finca Altamira, sul de San Salvador, April 1958, P. H. Allen & R. Armour 6856 (NY); San Salvador, Nov. 1921, S. Calderón 275 (NY, US); Sonsonate, Chilata, Dec. 1921, P. C. Standley 19310 (NY); Las Tabelas, próximo ao rio Acachapa, May 1942, J. M. Tucker 1356 (G, K, NY, US). ECUADOR: Eala, 1936, J. Leemans 236 (BM, C); El Oro, June 1943, E. L. Little Jr. 6620 (NY, US); Napo, Aguarico reserva étnica Huaorani, Km 79 – 82, March 1994, A. Dike & R. Enomega 1165 (MEXU, NY); Coca, Oct. 1987, D. Neill et al. 7872 (NY); Coca-Canon Monos, 4 km ao norte de Coca, Sept. 1986, W. Palacios & D. Neill 1292A (K, MEXU, NY,S, US); Pastaza, Pastaza Canton, VillanoPandanuque, junto ao rio Villano, July 1992, W. Palacios 10313 (NY); Francisco de Orellana, Rio Tiputini, Aug. 2003, G. Villa 1921 (K, QCA); El Oro, June 1943, E. L. Little Jr. 6620 (K, US). GUATEMALA: El Petén, carretera Tikal km 48, June 1965, Aguilar 64 (NY, US); Esquintla, s.d., S. Haynes s.n. (K [Herb. Hookerianum]); Guatemala, Jardim Botânico, May 1942, J. A. Steyermark 46533 (US). GUYANA: s.l., noroeste das montanhas de Kanuku, April 1938, A. C.



Smith 3533 (B, G, NY, S); Wabuwa Kanuru, Sept. 1948, depto. florestal 5713 (NY). HONDURAS: Comayagua, near La Paz, Jan. 1983, C. E. Hughes 283 (K); El Cayo, Valentin, July 1936, C. L. Lundell 6338 (NY); El Paraíso, Depto. El Paraíso, localidade escuela de Rio Arriba, c. 2 km da cidade, June 1996, J. L. Linares 3438 (K, MEXU); Tegucipalga, May 1983, C. Nelson 8610 (K); MÉXICO: Campeche, El Tormento, campo experimental, Dec. 1965, E. Hernández et al. ES291 (MEXU); Chapingo, March 1968. T. D. Pennington & J. Sarukán 9567 (K); Cordova, s.d., H. Finck 1865 (BM, K); Chiapas, Cacaluta, May 1982, E. Matuda 16529 (K, NY); Cosolapa, entre Cosolapa e El Palmar, May 1966, J. Chavelas ES-1693 (US); El Faro, distrito Cuicatlán, Nov. 1919, C. Conzalli 3820 (MEXU); El salto de Eyipantla, Nov. 1973, J. I. Calzada 1058 (BM, MEXU, NY); Escarcega, entre Escarcega e Silvituc, Jan. 1966, A. G. Pompa 1327 (US); Huastepec, Sept. 1946, Helbinger s.n. (MEXU 444444); Jalisco, Amatitlán, 1946, G. Aguirre 553 (MEXU); Los Tuxtlas, June 1975, M. Souza 4668 (MEXU); Morelos, Cuernavaca, Aug. 1961, H. W. Schwabe 1987 (B); Minatitlán, Sept. 1971, T. Wendt et al. 4028 (MO, NY); Yucatan, s.d., F. Gaumer s.n. (BM53808); Motozintla de Mendonza, March 1981, Ramamoorthy et al. s.n. (MEXU 286700); Oaxaca, Acatlán, Distr. Tuxtepec, rincon del Tigre, May 1986, M. Souza et al. 13036a (MEXU); Ocosingo, porção N de la omega, monumento Natural Yaxchilán, Dec. 1997, A. R. Gutiérrez 860 (MEXU); Piedras Rajada, Dec. 1950, Enriquez 6740 (MEXU); Rosário, a 21 km NE Chilillos, July 1983, E. Martinez S. et al. 4051 (MEXU); Sta. Maria, June 1990, J. I. Calzada et al. 15694 (MEXU); Tabasco, Tezonapa, Sierra Cruz Tetela a 3 km de Vista Hermosa, Aug. 1986, R. Robes G. 984 (MEXU); Vera Cruz, San Andrés, Jan. 1978, J. I. Calzada 4240 (NY). PANAMÁ: Balboa, zona canal, Jan. 1924, P. C. Standley 30861 (US); Buena Vista, July 1959, W. L. Stern et al. 856 (US); Chepo, Oct. 1911, H. Pittier 4765 (C, NY); Darien, ao redor de Pinogana, June 1914, H. Pittier 6995 (NY); San Blas, vila de Cangandi, Feb. 1985, G. de Neves 4932 (MEXU). PERU: Amazonas, Pongo de Manseriche, 1924, G. Tessmann 4480 (G); Coronel Portillo, Calleria, April 2003, J. G. Graham 2588 (K); Arequipa, Maucallacta, Rio Paranapura, Jan. 1935, G. Klug 3947 (K); Cuzco, La Convencion, Palma Real to Quillabamba, April 1995, T. D. Pennington, J. Leon & M. Cavero 15059 (K); Huánuco, Leoncio Prado, RupaRupa, July 1968, L. T. R. del Aguilla 36 (NY); Leoncio Prado, Rupa Rupa, April 1968, L. T. Rodriguez 25 (F, K); Pachieta, Panao, July 1962, R. L. Magín 20 (K); Puerto Inca, April 1989, Kroll 223 (G, K, NY); Yuyapichis, April 1989, Kroll 222 (K, MO, NY); Pasco, Oxapampa, March 1984, D. N. Smith, S. Knapp, J. Mallet & M. López-M. 6405 (K); Monte Virgem, rio Santiago, Sept. 1979, V. Huashikat 326 (MO); Maynas, Caseario


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Fig. 1. Myroxylon balsamum. A branch with inflorescence; B flower; C calyx; D standard; E petals; F androecium; G gynoecium; H fruits. (A from L. de Lima et. al. 620; B – G from A. Ducke 23787; H from W. Palacios & N. Neill 1292). DRAWN BY LAVINIA M. JOLY.


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Alianza, Rio Manití, Aug. 1980, A. Gentry et al. 29339 (MEXU, NY); Madre de Dios, Tambopata, Las Piedras, Reserva Amazonica, April 2005, R. Vásquez, R. Rojas & L. Valenzuela 30896 (K, MO); San Martin, Alto do rio Huallaga, Dec. 1929, L. Williams 6230 (NY); Distr. Chazuta, May 2005, J. S. Biset & J. C. de la Cruz 368 (K); Tumbes, Jan. 1968, D. R. Simpson & J. Schunke 518 (G); Ucayali, Pucallpa,Tingo-Maria, Sept. 1990, T. D. Pennington, A. Daza & W. Upiachiua 16981 (K); Purús, ditr. Purús, Rio Curanja, Colombiana, Oct. 1997, J. Graham & J. Schunke Vigo 289 (K). VENEZUELA: Carabobo, Guaramales, s.d., H. Pittier 9094 (G, NY); Caura, La Prisión, Medio Caura, March 1939, W. Llewelyn 11661 (US); Falcón, Depto. Buchivacoa, Caserio Los Chucos, Feb. 1982, T. Ruíz et al. 3699 (NY); Miranda, Cerros del Bachiller, March 1978, J. A. Steyermark & G. Davidse 117026 (MEXU); Yaracuy, Selva de Yumare, colônia I.A.N., Feb. 1959, A. L. Bernardi 7008 (G, NY); Zulia, Depto. Bolívar, via entre carretera Lara-Zulia e El Zamuro, entre km 5 – 8, Nov. 1979, G. S. Bunting 8297 (US). HABITAT. The species is fairly common in tropical forest at 200 – 690 m altitude. In Peru and Brazil the species is mostly associated with rivers, and sometimes grows on lateritic soil. CONSERVATION STATUS. Myroxylon balsamum is more restricted in its geographical distribution than M. peruiferum. It is known from a number of localities in Central America and northern South America. At present it is assessed as being of least concern (LC), although the species is under threat from logging, especially in the Amazon Forest. In remnants of mesophyllous forest it is likely that logging will become a threat to the species in the near future and the conservation assessment would then have to be changed to Vulnerable. PHENOLOGY. Flowering and fruiting throughout the year. VERNACULAR NAMES. Brazil: Bálsamo (A. Ducke 23787), Cumarú (D. C. Vasconcelos 11036), Costa Rica: Bálsamo (W. H. Hatheway 1660), Cuba: Guatemala (E. L. Ekman 17553); Colômbia: Bálsamo (R. Romero 7008); Tolu or Bálsamo de Tolu (H. N. Whitford & J. Pinzon 9); El Salvador: Bálsamo (G. P. Lewis & C. E. Hughes 1736), Bálsamo-negro (P. C. Standley 19310); Bálsamo do Peru (Natural History Museum: BM 53809), Crusnagua (S. Haynes 25); Ecuador: Bálsamo (A. Dike & R. Romero 1165), Pido-quera (J. A. Duke 8391), Guatemala: Nabá (Aguilar 64); Honduras: Bálsamo (C. E. Hughes 283); Panamá: Barba (G. de Neves 4932); Peru: Yarina-cocha (G. Tessmann 5470), Estoraque (Kroll 223); Quino-quino (BM53831); México: Bálsamo (Reko 115011), Palo de bálsamo (E. Matuda 16529), Nabá (J. Marroquin 278), Bálsamo-ó-nabá (J. Chavelas & A. Gomez ES 762), Mattaluca (H. Finek 1865); Nicaraguá: Bálsamo (A. Grisalva & D. Soza 3724); Venezuela: Bálsamo (A. L. Bernardi 7008). © The Board of Trustees of the Royal Botanic Gardens, Kew, 2015


NOTES. Myroxylon balsamum is distinguished by its

leaflets with a dull adaxial surface, the leaflet blade with inconspicuous pellucid dots and streaks, the calyx lobes obtuse, longer flowers 10 mm (without pedicel) and seeds smooth. Lanceolate, reflexed bracts and linear petals are frequent characteristics of material from Peru and Brazil. Collections of the species from Venezuela and Panama sometimes have bracts which are not reflexed, but other vegetative characters are as cited above. Historically, infra-specific taxa of Myroxylon balsamum have been accepted in a number of treatments (e.g., Harms 1908; Rudd 1968; Isely 1981). Rudd (1968) recognised infraspecific taxa based mostly on differing chemistry of the balsam, as well as the length and width of the fruits. We do not consider these traits to be consistent and thus do not recognise infraspecific taxa in our treatment. 2. Myroxylon peruiferum L. f. (Linnaeus filius 1781: 233). Type: America meridionalis, s.d., Mutis s.n. (hototype P00679272, photo!). Myrospermum peruiferum (L. f.) DC. (De Candolle 1825: 95). Toluifera peruifera (L. f.) Baill. (Baillon 1870: 383). Myroxylon pubescens Kunth (1823: 374). Type: America equatorial, Cartagena, s.d., M. A. Bonpland 1890 (lectotype P00151682, selected here, photo!). Myrospermum pubescens (Kunth) DC. (De Candolle 1825: 95). Toluifera pubescens (Kunth) Baill. (Baillon 1870: 383). Myrospermum pedicellatum Lam. (Lamarck 1797: 191). Type: Peru, J. Jussieu s.n. (lectotype G00404303!, selected here). The lectotype designated has the morphological characteristics that were described in the species protologue. We agree with the synonymisation proposed by Kunth (1823) and accepted by Harms (1908). Myroxylon pedicellatum (Lam.) Willd. (Willdenow 1799: 96). Toluifera pedicellata (Lam.) Baill. (Baillon 1870: 383). Myroxylon robiniaefolium Klotzsch (1857: 276). Type: Ecuador, A. J. de Warszewicz s.n. (lectotype BM 000053832!, selected here, isolectotype: K500797!, photo UEC!). synon. nov. Myrospermum erythroxylum Allemão (1857: 51). Type: Brazil, Rio de Janeiro, Allemão s.n. (lectotype BM000053824!, selected here). Tree, 6 – 30 m tall. Leaves 5 – 15-foliolate, densely to sparsely ferrugineous-tomentose, petiole 0.5 – 3.4 cm long, rachis 3.2 – 14 cm long; petiolules (1 –) 3 (– 6) mm long; leaflets 2.9 – 4.5 (– 10.5) (1 –) 1.7 – 2 (– 3.3) cm, oblong, elliptic, broadly-elliptic to ovate, apex obtuse, retuse to acuminate, base oblique, rounded or attenuate to subcordate, terminal leaflets


narrowly elliptic, obovate, or rhombic, base ovate, margin sinuous, entire, rarely crenate, concolorous, adaxial surface glabrous or sparsely tomentose over the midrib, abaxial surface sparsely-tomentose over the blade, tomentose over midrib, adaxial surface shiny, veins prominent on the abaxial surface, pellucid dots and streaks conspicuous over the entire blade. Racemes axillary or terminal, longer than the leaves, the axis ferruginous-tomentose, 11 – 20 cm long; bract deltate, apex acute, base truncate, concave, ascending, c. 1 mm long; pedicel 9 – 16 mm long, greyish or ferruginous-tomentose. Flower buds globose, apex obtuse; bracteole ovate, under 0.5 mm long. Flowers 7 – 9 mm long; calyx 5 – 7 9.6 – 11.4 mm, greyish tomentose externally, the lobe apices obsolete; standard petal blade (3.3 –) 4.6 (6.5 –) 7.5 – 9.7 mm, claw (4 –) c. 6.6 mm long, broadly-ovate, apex rounded or emarginate, base attenuate to rounded, margin entire, externally glabrous (rarely with trichomes at apex); other petals (5.9 –) 7.6 – 10.3 0.7 – 1.5 mm, lanceolate, apex acute, base attenuate, margin sinuous, sometimes externally sericeous at apex and on margin; stamens c. 9 mm long, filaments c. 4 mm long, anthers c. 5.2 mm long; ovary c. 6.5 mm long, short-stipitate. Fruit a 5.7 – 9.8 cm long samara; seed-chamber 1 – 2 cm long, elliptic; upper margin 1 – 4.5 (rarely up to 7) mm wide; lower margin 1 – 1.8 cm wide; stipe 3 – 9 mm long; calyx sometimes persistent; seed testa rough. Fig. 2. DISTRIBUTION. Myroxylon peruiferum is disjunctly distributed in the Americas, from Mexico to northern Argentina and southern Brazil (Map 1). SELECTED SPECIMENS EXAMINED. ARGENTINA: Jujuy, Capital Zapla, Nov. 1974, A. Burkart 30480 (MEXU); Ledesma, Serra de la Calilegus, Oct. 1927, S. Venturi 5402 (B); Quinta per Laguna, 1901, R. E. Fries 420 (S); Santa Bárbara, Sept. 1971, P. R. Legname & A. C. Cuezze 8581 (C); Salta, Anta, cerro Maldonado, Sierra González, June 1934, Ragonese 13238 (BA); Rio Piedras, Nov. 1913, D. Rodriguez 1110 (BA, US); Orán, Hartagal, Oct. 1940, Schreiter 11434 (S); Livessua, Sierra de Calilegna, Oct. 1927, S. Venturi 5402 (BM, US). BOLÍVIA: Chuquisaca, Calvo, Nov. 1998, J. R. I. Wood 14201 (K); Oropeza, Nov. 2004, J. R. I. Wood 21070 (K); Tomina, Dec. 1999, J. R. I. Wood, D. J. Goyder & M. Serrana 15344 (K); Cochabamba, Cochabamba, Oct. 2003, J. R. I. Wood 19710 (K); Campero Pasorapa, Oct. 1992, C. Antezana 415 (MO); Santa Cruz, Caballero, Nov. 1999, M. Nee 50678 (K); Florida, Nov. 1999, M. Nee 50448 (K); Samaipata, Quebrada de Aohira, Oct. 1928, J. Steinbach 8219 (BA, BM, K, NY, US); Vallegrande, Dec. 1990, I. G. Vargas 849 (NY); Zarija, El Desemboque, Nov. 1974, Z. L. López 5002 (US); Tarija, OĆonnor, Nov. 2003, J. R. I. Wood, H. Huaylla & D. J. Goyder 20093 (HSB, K). BRAZIL: Bahia, Itaju da Colônia, rodovia Palmira/Itaju, Oct. 1967,

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R. S. Pinheiros 280 (K, US); Ceará, Cariri, 25 March 1936, P. Luetzelburg, Comissão de serviços complementares da inspectoria de secas (EAC 37533); s.l., s.d., Fr. Allemão & M. de Cisneiros 445 (R); Distrito Federal, Brasília, Catetinho, Sept. 1980, E. P. Heringer et al. 5436 (NY, US); Brasília, Aug. 1980, E. P. Heringer et al. 5288 (K); Espírito Santo, Linhares, RVRD, June 1995, D. A. Folli 2626 (CVRD); Goiás, Luziânia, June 1972, E. P. Heringer 12159 (K, MO, RB, US); Mato Grosso, Sararé 10 km E do Campo Base, Aug. 1978, J. M. Pires & M. R. Santos 16615 (INPA, NY, R, UEC, US); Minas Gerais, Belo Horizonte estação experimental, Sept. 1937, M. Barreto 7749 (R); Ituiutaba, June 1946, A. Maciedo 787 (BM, K, MO, NY, S, US); Paraíba, Natuba, Nov. 1971, D. A. Lima et al. 1042 (PEUFR), s.l., Aug. 1832, L. Riedel s.n. (MO); Paraná, Ibiporã, fazenda Doralice, Nov. 1991, F. C. Silva 1448 (FUEL, UEC); Santo Antonio Caiua, Rio Paranapanema, June 1966, G. Hatschbach 14495 (C, K, L, US); Sertanópolis, fazenda Ferraz, Oct. 1989, M. Favarão et al. s.n. (FUEL 8089); Pernambuco, Piedade, estrada para Garanhuns, Nov. 1950, D. A. Lima 50-691 (RB); Rio de Janeiro, Itaperuna, Oct. 1984, A. S. Moreira s.n. (RB 256219); Rio de Janeiro, May 2006, R. D. Ribeiro, V. Maioli & R. Mendonça (RB 427618, K); Rio de Janeiro, Feb. 1882, A. Glaziou 12566 (K); São Paulo, Campinas, Faz. São Vicente, Sept. 1989, L. C. Bernacci 25879 (UEC); Campinas, s.d., J. C. de Mello 1864 (K); Cássia dos Coqueiros, Nov. 1994, L. S. Kinoshita & T. G. Guaratini 94-323 (UEC); Pederneiras, estação experimental, Oct. 1980, J. B. Baitello & O. T. Aguiar 5941(F); Penápolis, Faz. São Sebastião, July 1980, J. R. Pirani 780 (UEC); Pindorama, estação experimental, Aug. 1937, O. T. Mendes 2119 (RB); Piracicaba, Sept. 1931, J. G. Kuhlmann 2406 (RB); Pitangueiras, Sept. 1951, B. Costa (B 102648); São João da Boa Vista, 1869, A. F. Regnell 1758 (C, K, R, S); São Paulo, Jardim Botânico, Sept. 1985, O. Handro 2329 (SP, UEC); Tarumã, fazenda Berrante, March 1994, G. Durigan 31691 (UEC); Teodoro Sampaio, Parque Estadual Morro do Diabo, June 1987, A. L. K. M. Albernaz 11668 (UEC). COLÔMBIA: without exact locality, 1760 – 1808, J. C. Mutis 3768 (US); without exact locality, April 1905, R. Blake White s.n. (K), Medellin, 1879, R. A. Coliché s.n. (K); Medellin, s.d., R. B. White 11 (K); Amazonas, Rio Vaupés, desde rio a 5 km da desembocadura de Cafio Arara, Nov. 1975, A. Roa 233 (INPA). COSTA RICA: San Jose, Cantón de Mora, May 1997, A. Cascante 1324 (CR, K). ECUADOR: Las Cochas, a 10 km N de Zapotillo, Aug. 1975, A. Samaniego & F. Vivar 36 (US); Loja, Yamarine, June 1997, B. B. Klitgaard, B. Stahl, P. Lozano, T. Delgado, F. Elizalde, H. Alejandro & F. Tinitana 204 (K, LOJA); Loja, Aug. 1996, G. P. Lewis 2515 (K); in Andibus Ecuadorensibus, Oct. 1857, R. Spruce 5075 (BM, C, G, K, NY). EL SALVADOR: Ahuachapán, El © The Board of Trustees of the Royal Botanic Gardens, Kew, 2015

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Fig. 2. Myroxylon peruiferum. A branch with inflorescence; B flower bud; C calyx; D standard; E petals; F androecium; G gynoecium; H fruits. (A from S. Honda & M. O. Pedraz 936; B – G from J. R. Pirani 7-80; H from P. Silveira s.n. INPA 15557). DRAWN BY LAVINIA M. JOLY.



Corozo, Oct. 2000, J. M. Rosales 1651 (BM, K); Finca La Esperanza, Nov. 1999, A. K. Monro et al. 3077 (BM); San Benito, April 1993, E. Sandoval 1194 (K); HONDURAS: Comayagua, Pito Solo, Lake Yojoa, Aug. 1932, J. B. Edwards 422 (K, US). MÉXICO: Chiapas, Motozintla de Mendonza, March 1981, M. Souza et al. 11823 (MO); Michoacan, Coalcoman, Villa Vitoria, Sept. 1938, G. B. Hinton et al. 12296 (K); Morelos, Cuernavaca, March 1928, C. D. Mello 1 (NY); Oaxaca, Cerro Espino ao N Pochutla, April 1976, M. Souza et al. 5582 (US); Pochutla, Jan. 1978, E. Makrinius s.n. (MEXU 525161); Yucatan, s.d., G. F. Gaumer 776 (C). NICARAGUÁ: Matagalpa, Santa Maria Ostuma, March 1984, A. Grisalva & D. Soza 3724 (MO). PERU: s.l., 1847, s.c. (Herb. Pavon, K 000111320); Andaloy, Cajamarca, June 1981, A. Sagástegui 9731 (MEXU, MO); Chicoplaya, 1895, s.c. (K000111321); Tumbes, Feb. 1968, D. R. Simpson 518 & J. Schunke (G); s.l., s.d., D. Hanbury, numbers 3, 4 and 7 (K); without exact locality, March 1865, D. Hanbury (Herbarium Hookerianum, K). HABITAT. Myroxylon peruiferum occurs mostly in mesophyllous forest and dry habitats at 540 – 2000 m elevation. In Bolivia, it is known from dry forest on stony, sandy soils and also in remnant forest in the lower valleys. CONSERVATION STATUS. Myroxylon peruiferum is a widely distributed species but is not abundant within its area of occurrence. The impacts of human activities such as deforestation, mainly in areas of Atlantic Forest and mesophyllous forest, have resulted in only small remnants of these vegetation types persisting and thus an associated decline in the populations of M. peruiferum. The species is temporarily assessed to be Near Threatened (NT). PHENOLOGY. Flowering from June to September, fruiting from August to November. VERNACULAR NAMES. Argentina: Quina-quina (Rodriguez 1110), Quina colorada (Cozzo 75982); Brazil: Bálsamo (A. Maciel 787), Cabreúva (A. L. K. M. Alvernaz 11668), Cabreúva-vermelha (O. Vecchi 191), Caburayba (Bacelli R69645), Óleo-vermelho (J. G. Kulmann 16564); Bolívia: Quina-quina (T. Centurión & R. Quevedo 1209), Ecuador: Chaquino (A. Samaniego & F. Vivar 36); Peru: Quinillo (A. Sagastegui 15087); México: Bálsamo (G. B. Hinton et al. 12296), Cedro-quino (R. Conzatti & Makrinius 3079), Quina (Schreiter 11434), Quinacolorada (S. Venturi 5402). NOTES. Myroxylon peruiferum is easily distinguished from M. balsamum by the combination of its leaflet with a shiny adaxial surface, the leaflet blade with conspicuous pellucid dots and streaks, broadly-ovate standard petal (4.6 mm wide), the standard claw 6.6 mm long, the calyx lobes obsolete, smaller flowers 10 mm long (without pedicel) and the rough seed testa. On any one specimen of M. peruiferum one can encounter

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different leaflet shapes. Smaller flowers are seen in specimens from Peru and Bolivia. Glandular trichomes, when present, can be observed on the leaf rachis, the abaxial surface of the leaflets, the inflorescence axis, flower-buds and calyces.

Excluded taxa The two species described by Stokes (1812), are not accepted as species of Myroxylon. The tri-winged branches described for Myroxylon trialatum indicate that it belongs to a genus other than Myroxylon. The name Myroxylon abruptifolium alludes to the presence of abruptly bijugate leaves. The species description is based on a single character which does not distinguish it from other species accepted in the genus. Additionally, the Liverpool Herbaria (LIV), where Stokes worked, apparently do not house the types of his two species (pers. comm. D. Hughes). Acknowledgements The authors thank the directors and curators of Herbaria that loaned specimens or allowed access to collections. We also thank Renee Fortunato, André M. de Carvalho (in memoriam), and Renato de Jesus for assistance during botanical collecting expeditions in Mexico (Renee) and Brazil. Lavinia M. Joly is thanked for the two illustrations. The work was supported by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the doctoral and post doctoral fellowship to ALBS. References Allemão, F. F. (1857). Leguminosae. Myrospermum erythroxylum. Revista Brazil. 1: 51. Baillon, H. (1870). Toluifera L. Histoire des plantes. 2: 369 – 370. Librairie Hachette, Paris. ____ (1883). Toluifera balsamum L. Traité de botanique médicale, phanérogamique. 5: 676 – 677. Librairie Hachette et Cie, Paris. Bentley, R. & Trimen, H. (1880a). Toluifera pereirae. Medicinal Plants 42: 83. J. & A. Churchill, London. ____ & ____ (1880b). Toluifera balsamum. Medicinal Plants 42: 84. J. & A. Churchill, London. Biemer, T. A., Asral, N. & Albanese, J. A. (1992). Simultaneous, stability-indicating capillary gas chromatographic assay for benzocaine and the two principal benzyl esters of Balsamum Peru formulated in a topical ointment. J. Chromatogr. 623: 395 – 398. Cardoso, D., Queiroz, L. P., Pennington, R. T., Lima, H. C., Fonty, E., Wojciechowski, M. F. & Lavin, M. (2012). Revisiting the phylogeny of papilionoid legumes: New insights from comprehensively sampled earlybranching lineages. Amer. J. Bot. 99: 1991 – 2013. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2015

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de Candolle, A. P. (1825). Leguminosae. In: A. P. Candolle (ed.), Prodromus systematis naturalis regni vegetabilis, Vol. 2, pp. 93 – 423. Treuttel & Würtz, Paris. Doyle, J. J. (1995). DNA data and legume phylogeny: a progress report. In: M. D. Crisp & J. J. Doyle (eds), Advances in Legume Systematics, part 7, pp. 11 – 30. Royal Botanic Gardens, Kew. ____, Doyle, J. L., Ballenger, J. A., Dickinson, E. E., Kajita, T. & Ohashi, H. (1997). A phylogeny of the chloroplast gene rbcL in the Leguminosae: taxonomic correlations and insights into the evolution of nodulation. Amer. J. Bot. 84(4): 541 – 554. Farr, E. R., Leussink, J. A. & Stafleu, F. A. (1979). Index nominum genericorum (Plantarum). Regnum Veg. 100. Scheltema & Holkema, Utrecht, Bohn. Fluckiger, F. A. & Hanbury, D. (1874). Balsamum tolunatum. Pharmacographia 15: 177 – 184. Harms, H. (1908). VI Zur nomenclatur des Perubalsambaumes. Notizbl. Königl. Bot. Gart. Berlin 5: 85 – 97. Holmgren, P. K., Holmgren, N. H. & Barnett, L. C. (1990). Index Herbariorum, Vol. I: The herbaria of the world. New York Botanical Garden, New York. Hutchinson, J. (1964). The genera of flowering plants, Vol. 1. Clarendon Press, Oxford. Isely, D. (1981). Leguminosae of the United States III, subfamily Papilionoideae; Tribes Sophoreae, Podalyrieae, Loteae. Mem. New York Bot. Gard. 25(3): 1 – 264. Klotzsch, J. F. (1843). Myroxylon punctatum Klotzsch. In: F. G. Hayne, Getreue Darstellung und Beschreibung der in Arzneykunde Gebrauchlichen Gewachse 14: 12. Auf kosten des verfassers, Berlin. ____ (1857). Eine synoptische Aufzählung der Arten von Myroxylon Mutis und Myrospermum Jacq. Bonplandia 5: 272 – 277. Kunth, C. S. (1823). Voyage de Humboldt et Bonpland. In: F. H. A. von Humboldt, A. J. A. Bonpland & K. S. Kunth (eds), Nova Genera et Species Plantarum, Vol. 6: 371 – 375, 570 – 571. Facsimile ed. 1963, J. Cramer, Weinheim. Lamarck, J. B. A. P. de (1797). Encyclopédie Méthodique. Chez H. Agasse, Paris. Lawrence, G. H. M. (1971). Taxonomy of vascular plants. Macmillan, New York. Linares, E. & Bye Jr., R. A. (1987). A study of four medicinal plant complexes of Mexico and adjacent United States. J. Ethno-pharmacol. 19: 153 – 183. Linnaeus, C. (1753). Toluifera balsamum. Species Plantarum ed. I, 1: 383 – 385. Impensis l. Salvii, Holmiae. Linnaeus filius, C. (1781). Myroxylon peruiferum. Supplementum Plantarum 34: 232 – 234. Impensis Orphanotrophei, Brunsvigae. Maranduba, A., Oliveira, A. B., Oliveira, G. G. de, Reis, J. E. de P. & Gottlieb, O. R. (1979). Iso© The Board of Trustees of the Royal Botanic Gardens, Kew, 2015


flavonoids from Myroxylon peruiferum. Phytochemistry 18: 815 – 817. McNeill, J. C., Barrie, F. R., Buck, W. R., Demoulin, V., Greuter, W., Hawksworth, D. L., Herendeen, P. S., Knapp, S., Marhold, K., Prado, J., Prud'homme van Reine, W. F., Smith, G. F. & Wiersema, J. H. (2011). International Code of Nomenclature for algae, fungi, and plants (Melbourne Code). Regnum Veg. 154. Koeltz Scientific Books. Oersted, W. (1855). Myrospermum sonsonatense. Vidensk Meddel Naturhist. Foren. Kjöbenhavn: 27 – 32. Pennington, R. T., Stirton, C. H. & Schrire, B. D. (2005). Tribe Sophoreae. In: G. Lewis, B. Schrire, B. Mackinder & M. Lock (eds), Legumes of the world, pp. 227 – 249. Royal Botanic Gardens, Kew. Pereira, J. (1851). On the Myrospermum of Sonsonate, from which balsamum of Peru, white balsamum and balsamito are obtained. Pharm. J. Trans. 10: 27 – 29. Polhill, R. M. (1981). Sophoreae. In: R. M. Polhill & P. H. Raven (eds), Advances in legume systematics part 1, pp. 213 – 230. Royal Botanic Gardens, Kew. Radford, A. E., Dickinson, W. C., Massey, J. R. & Bell, C. R. (1974). Vascular plant systematics. Harper & Row, New York. Royle, J. F. (1853). Myrospermum pereirae. A manual of materia medica and therapeutics 2: 414. John Churchill, London. Rudd, V. E. (1968). Leguminosae of MexicoFaboideae. I. Sophoreae and Podalyrieae. Rhodora 70: 493 – 532. Ruiz, H. & Pavon, J. (1956). Flora Peruviana et Chilensis. Anales Inst. Bot. Cavanilles 14: 717 – 784. Sartori, A. L. B. & Tozzi, A. M. G. (2002). Comparative leaflet anatomy in Myrocarpus Allemão, Myroxylon L.f. and Myrospermum Jacq. (LeguminosaePapilionoideae-Sophoreae) species. Bot. J. Linn. Soc. 140: 249 – 259. Stafleu, F. A. (1965). Florae Peruvianae et Chilensis, Vol. 43. Facsimile, J. Cramer, Lehre. ____ & Cowan, R. S. (1983). Taxonomic Literature, Vol. 4. W. Junk, Utrecht. Stearn, W. T. (1973). Botanical Latin. Redwood Press Limited, Trowbridge. Stokes, J. (1812). Myroxylon abruptifolium. A botanical materia medica 2, p. 471. J. Johnson, London. Walpers, W. G. (1843). Myrospermum. Repertorium botanices systematicae 1: 805. Sumtibus Friderici Hofmeister, Lipsiae. Willdenow, K. (1799). Species plantarum, vol. 2. Impensis G. C. Nauk, Berolini. Wojciechowski, M. F., Lavin, M. & Sanderson, M. J. (2004). A phylogeny of legumes (Leguminosae) based on analysis of the plastid matK gene resolves many well-supported subclades within the family. Amer. J. Bot. 91: 1846 – 1862.