Hymenaea courbaril L., Sp. Pl. 1192. 1753. Lectotype. "Ceratia diphyllos Antegoana, Ricini majoris, fructu osseo, siliqua grandi incluso" in Plukenet, Phytographia, t. 82, f. 3, 1691 (designated by Lee and Langenheim in Univ. Calif. Publ. Bot. 69: 81. 1975).

Souza, Isys Mascarenhas, Funch, Ligia Silveira & Queiroz, Luciano Paganucci de, 2014, Morphological analyses suggest a new taxonomic circumscription for Hymenaea courbaril L. (Leguminosae, Caesalpinioideae), PhytoKeys 38, pp. 101-118 : 108-110

Hymenaea courbaril L., Sp. Pl. 1192. 1753. Lectotype. "Ceratia diphyllos Antegoana, Ricini majoris, fructu osseo, siliqua grandi incluso" in Plukenet, Phytographia, t. 82, f. 3, 1691 (designated by Lee and Langenheim in Univ. Calif. Publ. Bot. 69: 81. 1975).

Hymenaea resinifera Salisb., Prodr. 327. 1796, nom. nud.

? Hymenaea animifera Stokes, Bot. Mat. Med. 2: 449. 1812, fide Lee and Langenheim in Univ. Calif. Publ. Bot. 69: 81. 1975.

Hymenaea candolleana Kunth [spelled ' candolliana’], Nov. Gen. and Sp. 6: 323, pl 556. 1824. Type: México, Acapulco, Bonpland 3875 (holotype P!).

Hymenaea confertifolia Hayne, Getreue Darstell. Gew., 11, pl. 9. 1830. Lectotype (designated here): Sellow s.n., s.d. (lectotype W 48857!).

Hymenaea stilbocarpa Hayne, Getreue Darstell. Gew., 11, pl. 11. 1830. Hymenaea courbaril var. stilbocarpa (Hayne) Lee and Lang., J. Arnold Arbor. 55: 449. 1974, syn. nov. Lectotype (designated here): "S. Pauli, sylv. ad Faz. de S. Anna", Martius (lectotype M! [barcode n° M-0215314], photo HUEFS!, isolectotype M! [barcode n° M-0215313], photo HUEFS!).

Hymenaea retusa Willd. ex Hayne, Darst. Beschreib. Arzneigew. 11: pl. 12. 1830. Type: México, Acapulco, Humboldt (holotype B!), nom. illegit.

Hymenaea splendida Vogel, Linnaea 11: 409. 1837. Type: Sellow 1025 (holotype B†).

Hymenaea courbaril var. obtusifolia Ducke, Arch. Jard. Bot. Rio de Janeiro, 4: 47. 1925. Type: Brazil, Pará, Belém (Ilha do Marajó?), Ducke RB 16906 (holotype RB!, isotypes A!, MG!, P!, U!, US!).

Hymenaea courbaril var. subsessilis Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 265. 1925, syn. nov. Neotype (designated by Lee and Langenheim in Univ. Calif. Publ. Bot. 69: 89. 1975): Brasil, Amazonas, Flores, Ducke MG 11167 (neotype MG!).

Hymenaea multiflora Kleinhoonte, Recueil Trav. Bot. Neerl. 22: 405. 1925. Type: Suriname, Boschreserve, Kaboeri, Coll. by the Forestry Bureau Herb. 4943 (holotype U!).

Inga megacarpa M.E. Jones, Contr. West Bot. 15: 140. 1929. Type: México, Nayarit, M.E. Jones 23012 (holotype RSA!, isotypes F!, GH!, NY!).

Hymenaea courbaril var. villosa Y.T. Lee and Andrade-Lima, J. Arnold Arbor. 55: 446. 1974, syn. nov. Type: Brasil, Paraíba, Paquevira de Natuba, Lee and Andrade-Lima 83 (holotype UC!).

Description.

Large trees, to 30 m tall. Petiole 1.1-1.7 (-2) cm long; petiolule 4-5 mm long; leaflets 6-9 (-10.2) × 2.8-4 (-5.1) cm, elliptic, rarely ovate, straight or slightly falcate due to central vein with an angle c. 10°, apex acute or obtuse, rarely acuminate, base acute along the inner margin and rounded along the outer margin, distance from the inner margin to central vein 7-10 mm in the basal region. Flower buds 2.1-2.5 cm long; flower 2.2-3.1 cm long; hypanthium 7-15 mm long; petals 1.1-2.0 cm long. Fruit cylindrical, of uniform width, apex mostly rounded and apiculate.

Hymenaea courbaril is defined here more narrowly than the circumscription adopted by Lee and Langenheim (1975), as we are proposing the exclusion of the varieties altissima and longifolia and their recognition as distinct species. In this narrower sense, Hymenaea courbaril is characterized by leaflets with slightly convex inner margins and midrib slightly arched, resulting in an elliptic or ovate outline, not or only slightly falcate, apex mostly acute or obtuse ( Figure 3 ). It presents flowers larger than the other species of the Hymenaea courbaril complex, and cylindrical fruits mostly with more than five seeds ( Table 4 ). It has a wide geographical range, mostly in different nuclei of Seasonally Dry Forests in Mexico, Central America, Caribbean, northern South America, Bolivia, and Brazil (from the Amazon region to Paraná State in the south).

Nomenclatural problems with several names associated with Hymenaea courbaril were not amended by Lee and Langenheim (1975) when they undertook their taxonomic revision of the genus. No formal type for Hymenaea stilbocarpa was cited by Lee and Langenheim (1975), who stated that no specimen was cited in the original description of Hayne (1830). However, Hayne (1830) based Hymenaea stilbocarpa on material collected by Martius, citing “Wäscht in Brasilien in Wäldern der Provinzen S. Paulo, Minas Geraes und Bahia (Martius)". We found a specimen in the M herbarium collected by Martius in the Brazilian state of São Paulo (Santana farm) with an attached label indicating that this plant was distributed over the "Prov. Rio de Jº., S. Paulo, Minas Geraes, Bahia", which probably served as the original material for Hayne’s description of Hymenaea stilbocarpa . Thus, we are lectotypifying this species with Martius’ specimen held in M under the barcode number M-0215314.

Hymenaea confertifolia Hayne was based on material collected by Sellow and Olfers in Brazil ( “Wächst in Brasilien (Olfers u. Sellow)"; Hayne 1830: table 9). Lee and Langenheim (1975: 88) stated that this name was based on Sellow 1025 and that the “holotype” in the B herbarium was destroyed. However, as Hayne (1830) did not refer to one particular specimen, all materials collected by Sellow or Olfers that can be linked with Hymenaea confertifolia should be considered syntypes. No such specimens can be found in the B herbarium, and were probably destroyed. A duplicate from B collected by Sellow was found in W annotated as Hymenaea confertifolia , and is designated here as the lectotype of this name.

Hymenaea retusa Willd. ex Hayne was published as a homotypic synonym of Hymenaea candolleana (Hayne 1830). It is thus illegitimate under the Article 52 of the International Code of Botanical Nomenclature ( McNeill et al. 2011).

Hymenaea courbaril var. obtusifolia Ducke was published based on a tree cultivated in the Pará Botanical Garden (now Museu Paraense Emílio Goeldi, Belém, state of Pará, Brazil). Lee and Langenheim (1975) misinterpreted this as evidence that the holotype was the specimen in the herbarium of this museum (MG). However, Ducke (1925: 47) explicitly stated that "specimina florifera and fructus in herb. Jard. Bot. Rio n. 16.906", and thus the RB material should be considered as the holotype of this name.

Ducke (1925: 265) did not cite any specimen when describing Hymenaea courbaril var. subsessilis Ducke. Lee and Langenheim (1975) did not explicitly designate a type for this variety, but chose a "representative specimen" collected "in the same general area believed to be the type locality". We are accepting it as an inferential typification, and the status of this material should be a neotype since no other specimen was refereed in the protologue.

Figures

Figure 3. Leaf morphology of the species recognized in the Hymenaea courbaril complex: A Hymenaea longifolia B Hymenaea courbaril C Hymenaea altissima. Scale bar = 1 cm.