Morphological analyses suggest a new taxonomic circumscription for Hymenaea courbaril L. (Leguminosae, Caesalpinioideae)
Author
Souza, Isys Mascarenhas
Universidade Estadual de Feira de Santana, Programa de Pos-graduacao em Botanica, Herbario, Km 03 - BR 116, Campus. 44031 - 460, Feira de Santana, Bahia, Brasil
Author
Funch, Ligia Silveira
Universidade Estadual de Feira de Santana, Programa de Pos-graduacao em Botanica, Herbario, Km 03 - BR 116, Campus. 44031 - 460, Feira de Santana, Bahia, Brasil
Author
Queiroz, Luciano Paganucci de
Universidade Estadual de Feira de Santana, Programa de Pos-graduacao em Botanica, Herbario, Km 03 - BR 116, Campus. 44031 - 460, Feira de Santana, Bahia, Brasil
luciano.paganucci@gmail.com
text
PhytoKeys
2014
2014-06-18
38
101
118
http://dx.doi.org/10.3897/phytokeys.38.7408
journal article
http://dx.doi.org/10.3897/phytokeys.38.7408
1314-2003-38-101
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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:
Mexico
, 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:
Mexico
, 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,
Para
,
Belem
(Ilha do
Marajo
?),
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:
Mexico
, 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,
Paraiba
, 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)
x
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
Parana
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
"Waescht
in Brasilien in
Waeldern
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
Sao
Paulo (Santana farm) with an attached label indicating that this plant was distributed over the "Prov. Rio de
J°
., 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 (
"Waechst
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
Para
Botanical Garden (now Museu Paraense
Emilio
Goeldi,
Belem
, state of
Para
, 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.