The Diversity of Machaerium (Leguminosae: Papilionoideae) in the Atlantic

Forest: Three New Species, Nomenclatural Updates, and a Revised Key

Author(s): Fabiana Luiza Ranzato Filardi and Haroldo Cavalcante de Lima
Source: Systematic Botany, 39(1):145-159. 2014.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1600/036364414X678026

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Systematic Botany (2014), 39(1): pp. 145–159
© Copyright 2014 by the American Society of Plant Taxonomists
DOI 10.1600/036364414X678026
Date of publication 02/05/2014

The Diversity of Machaerium (Leguminosae: Papilionoideae) in the Atlantic Forest:

Three New Species, Nomenclatural Updates, and a Revised Key

Fabiana Luiza Ranzato Filardi1,2 and Haroldo Cavalcante de Lima1

1
Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Diretoria de Pesquisa Cientı́fica, Rua Pacheco Leão 915,
CEP: 22460-030, Rio de Janeiro, Rio de Janeiro, Brazil.
2
Author for correspondence (ffilardi@jbrj.gov.br)

Communicating Editor: Austin Mast

Abstract—Machaerium is a predominantly Neotropical genus with around 130 species of trees, shrubs, and lianas occurring from southern
Mexico to northern Argentina. In Brazil there are about 80 species, with the highest diversity occurring in the Atlantic and Amazonian forests.
Our taxonomic revision of Machaerium in the Atlantic Forest included field studies in the northeast, southeast, and south of Brazil as well as
the analysis of collections that belong to 40 herbaria, which has allowed the study of about 10,300 specimens. For the recognition of taxa, the
material studied was compared with the original descriptions of the species and its nomenclatural types. As a result we recognize 41 species
of Machaerium in the Atlantic Forest, including the three new species described here: M. androvillosum, M. aureum, and M. robsonnianum.
We also designate two neotypes, 20 lectotypes, seven epitypes, and 23 new synonyms for 20 species of Machaerium. In addition to the
descriptions and illustrations of the new species and nomenclatural updates, a revised key for all species of the Atlantic Forest with their
geographical distributions is presented.
Keywords—Atlantic Domain, Brazil, conservation, endemism, Fabaceae, typification.

Machaerium Pers. is among the major genera of woody
Papilionoid legumes, with around 130 species (Klitgaard
and Lavin 2005) and a predominantly Neotropical distribu-
tion, from southern Mexico to Argentina, though one species,
M. lunatum (L. f.) Ducke, is amphiatlantic, extending to the
west coast of Africa (Lozano and Klitgaard 2006). The genus
consists of trees, shrubs, and lianas, which are commonly
found in forest formations (Hoehne 1941; Ducke 1949, 1953;
Dwyer 1965; Rudd 1977; Bastos 1987; Lewis 1987; Mendonça
Filho 1996; Sartori and Tozzi 1998; Lozano and Klitgaard
2006; Meléndez González 2009; Ribeiro and Lima 2009;
Filardi 2011).
Brazil is the center of Machaerium diversity, containing
approximately 80 taxa, of which 50 are endemic. Machaerium
occurs in most of the phytogeographic areas of the country
(Filardi 2011), especially in seasonally dry forests (Lima 2000;
Ribeiro and Lima 2009; Filardi 2011). The greatest richness
and endemism are found in the Atlantic and Amazon forests,
with 41 and around 30 species, respectively; while the
Cerrado (ca. 20 species) and Caatinga (10 species) contain
representatives that can also be found in Amazonia as well
as in the Atlantic Forest (Filardi 2011). The tree species of
Machaerium are popularly known as jacarandá (rosewoods)
because of their timber with characteristics that have been
emphasized since the sixteenth century (Hoehne 1941), and
these species were much used for construction, luxury fur-
nishings and by luthiers (Coimbra Filho 1950; Mainieri and
Chimelo 1989).
Several species are capable of establishing symbiotic asso-
ciations with nitrogen-fixing bacteria (Mckey 1994; Sprent
2001; Faria and Lima 2002; Faria et al. 2006), and the charac-
teristics of this symbiotic association provide synapomor-
phies for the clade to which Machaerium belongs (Lavin et al.
2001). Machaerium form a clade with Aeschynomene sect.
Ochopodium Vogel and Dalbergia L. f. in the much larger
“Dalbergioid” clade—a clade for which aeschynomenoid
root nodule forms a synapomorphy (Lavin et al. 2001). The
monophyly of Machaerium was supported by phylogenetic
analyses of chloroplast DNA and of the internal transcribed
spacers region of the nuclear ribosomal DNA (ITS) (Ribeiro
et al. 2007; Filardi 2011). However, recent analyses of the
145
ITS region and of morphological data concluded that
Machaerium is paraphyletic with respect to Aeschynomene sect.
Ochopodium (Lewis et al. 2012). These authors presented a
clade formed by Aeschynomene sect. Ochopodium and by spe-
cies of Machaerium that have leaflets with cladodromous
venation, and a clade formed by species of Machaerium that
have leaflets with brochidodromous venation. The paraphyly
was reported well after the beginning of our study and
because of this reason the four species of Aeschynomene sect.
Ochopodium known for the Atlantic Forest (Rudd 1955) were
not included here. The pattern of secondary venation of leaf-
lets is a very important character for the infrageneric classifi-
cation of Machaerium (Bentham 1860, 1862; Rudd 1987a).
Machaerium sect. Lineata (Benth.) Taub., which represents
about 20% of the genus and includes species that have
cladodromous venation, was the only one of the traditional
sections that was supported as monophyletic in phylogenetic
analyses (Ribeiro et al. 2007; Filardi 2011).
Bentham (1862) published a treatment of Machaerium in
“Flora Brasiliensis” and recognized 43 species in the genus.
Some features of Machaerium were also characteristic of spe-
cies of Drepanocarpus G. Mey (Bentham 1860, 1862), and the
latter was eventually included in synonymy of Machaerium
(Ducke 1922). In a monographic study, Hoehne (1941)
accepted the synonyms proposed by Ducke (1922) and recog-
nized 121 species of Machaerium, among them 78 species
from Brazil. Rudd’s (1987a) treatment of M. sect. Machaerium
recognized eight species and seven varieties among which
seven and four, respectively, occur in Brazil. Just after, Rudd
(1987b) published the treatment for species with wingless
fruit of M. sect. Lineata and recognized five species with three
of them present in the Brazilian Amazon. The revision
presented by Mendonça Filho et al. (2007) of M. sect. Oblonga
(Benth.) Taub. recognized 12 species, and among the nine
species that occur in Brazil six are endemic, as well as
the two species described later for this section (Mendonça
Filho et al. 2011).
The tremendous habitat heterogeneity of the Atlantic For-
est and the many collection gaps pose a great challenge to the
study of species-rich groups like Machaerium. The plasticity of
vegetative characters across the habitats leads to overlapping
146 SYSTEMATIC BOTANY [Volume 39

morphologies and complicates species delimitations. A stereomicroscope. Data on geographic distribution and habitat were
thorough study of herbarium collections coupled with obtained from the labels, literature, and field studies.
extensive fieldwork is essential to understand the variation
in vegetative morphology and produce informed decisions Results and Discussion
on species delimitation. This study is part of our taxo-
nomic revision of Machaerium in the Atlantic Forest biome. A revised identification key for the 41 species of
Here we describe three new species of the genus that are Machaerium currently known in the Atlantic Forest biome is
endemic to the Atlantic Forest, provide the typification presented. The key is augmented with the geographical dis-
and synonymization for 20 other species, and present an tribution of each species. Among these species, 22% are
updated identification key for all 41 species of Machaerium widely distributed and also occur in other countries of South
occurring in this biome. America, while 78% are endemic to Brazil and 56% are
restricted to the Atlantic Forest. General information about
the distribution and habitat for all Brazilian species of
Materials and Methods Machaerium was provided elsewhere (Filardi 2013).
Herbarium materials were analyzed from B, BHCB, BM, CEPEC, Three new species are also described and illustrated here:
CVRD, E, ESA, FI, G, GFJP, GUA, HB, HBR, HST, HUEFS, IAC, ICN, Machaerium androvillosum is known only from herbarium mate-
IPA, JPB, K, M, MAC, MBM, MBML, NY, OUPR, P, PACA, PEUFR, PI, rial collected in seasonal forests in southeastern Brazil;
R, RB, SP, SPF, UEC, UFP, US, VIC, VIES, and W. Images in high resolu-
M. aureum is endemic to southern Bahia state in northeastern
tion sent for the authors by herbaria BR and UB were also analyzed. The
delimitation of the study area followed the concept of Oliveira Filho et al. Brazil and occurs in wet forests with intense anthropic pres-
(2006) that combines seasonally dry forests and extra-Amazonian wet sure; and M. robsonnianum is restricted to southeastern Brazil
forests, including Araucaria forests, mangroves, restingas, and high alti- and has the largest distributional range, occupying areas of
tude grassland. Field studies were carried out between February 2007 and restinga along the coast, wet forests, and seasonally dry forests.
May 2010 in Brazil, especially in the states of Alagoas, Bahia, Espı́rito
Santo, Minas Gerais, Paraı́ba, Paraná, Pernambuco, Rio de Janeiro, and
The nomenclatural update here presented includes two
Rio Grande do Norte. Approximately 10,300 specimens were studied, neotypes, 20 lectotypes, seven epitypes, and 23 new syno-
and the taxonomic treatment included comparisons with the species nyms related to the following 20 species of Machaerium:
descriptions in the protologues as well as analysis of the type material of M. acutifolium Vogel, M. brasiliense Vogel, M. cantarellianum
accepted names and synonyms. The species concept here adopted recog-
Hoehne, M. condensatum Kuhlm. & Hoehne, M. debile (Vell.)
nizes species as metapopulations with independent evolutionary lineages
over time, and one species would be a segment of a particular lineage (De Stellfeld, M. declinatum (Vell.) Stellfeld, M. firmum (Vell.)
Queiroz 2007). From among operational criteria listed by De Queiroz Benth., M. glabrum Vogel, M. incorruptibile (Vell.) Benth.,
(2007) for the delimitation of species we used the diagnostic criterion, M. lanceolatum (Vell.) J. F. Macbr., M. legale (Vell.) Benth.,
which considers qualitative differences measurable by gaps of variation M. leucopterum Vogel, M. nigrum Vogel, M. oblongifolium
between different species. The terminology used for the shape and
indumentum of the structures follows Radford et al. (1974) and Hickey
Vogel, M. paraguariense Hassl., M. pedicellatum Vogel,
and King (2000), and illustrations of the three new species were made M. punctatum (Poir.) Pers., M. reticulatum (Poir.) Pers.,
from dry and hydrated materials using a drawing tube attached to a Zeiss M. stipitatum Vogel, and M. uncinatum (Vell.) Benth.

Taxonomic Treatment
Key to the Species of Machaerium in the Atlantic Forest
1. Armed plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Branches with only spinescent stipules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Leaflets with cladodromous pattern, the parallel secondary veins branching toward the margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Leaves (3–)5–11(–13)-foliolate, leaflets (1.5–)2–4.5(–7) cm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Leaflet apex retuse; petals pale purple (Brazil: Minas Gerais, Espı́rito Santo,
Rio de Janeiro, and São Paulo) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. reticulatum (Poir.) Pers.
5. Leaflet apex acuminate; petals yellowish-white to purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Leaflets 3–7, adaxial surface glabrous and abaxial surface sparsely sericeous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Lianas to scandent shrubs; petiole and rachis glabrous; petals yellowish-white;
banner petal externally glabrous; stamens monadelphous
(Brazil: Minas Gerais, Espı́rito Santo, Rio de Janeiro, and São Paulo) . . . . . . . . . . . . . . . . . . . . M. declinatum (Vell.) Stellfeld
7. Lianas to small trees; petiole and rachis puberulent; petals purple;
banner petal externally strigose; stamens diadelphous 9 + 1
(Brazil: Bahia, Espı́rito Santo, and Minas Gerais) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. caratinganum Kuhlm. & Hoehne
6. Leaflets 9–13, adaxial and abaxial surfaces aureo-tomentose
(Brazil: Pernambuco, Alagoas, Sergipe, Bahia, Espı́rito Santo,
Minas Gerais, and Rio de Janeiro) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. condensatum Kuhlm. & Hoehne
4. Leaves (9–)13–43-foliolate, leaflets 0.2–2 cm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Trees; spinescent stipules parallel or perpendicular to the branch, usually overlapping;
leaflets narrowly oblong, up to 0.7 cm wide; flowers up to 1.6 cm long
(Panama, Guyanas, Venezuela, Colombia, Brazil, Peru, Bolivia, Paraguay, and Argentina) . . . . . . . . . . . M. hirtum (Vell.) Stellfeld
8. Lianas or escandent to sarmentose shrubs; spinescent stipules recurved, non-overlapping;
leaflets oblong to obovate, up to 2 cm wide; flowers up to 1.2 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Branches glabrous to puberulent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Branches squamous; leaflet base rounded; inflorescence rachis puberulent to glabrous;
flowers up to 12 mm long, calyx purple, corolla pale purple (Bolivia and Brazil) . . . . . . . . . . . . . . . . . . M. amplum Benth.
10. Branches not squamous; leaflet base cuneate; inflorescence rachis tomentose; flowers up
to 9 mm long, calyx green, corolla yellowish (Brazil: Alagoas, Bahia, Espı́rito Santo,
Minas Gerais, Rio de Janeiro, São Paulo, Paraná, and Santa Catarina) . . . . . . . . . . . . . . . . . . . . M. uncinatum (Vell.) Benth.
9. Branches villous to tomentose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
2014] FILARDI AND LIMA: MACHAERIUM IN THE ATLANTIC FOREST 147

11. Leaf rachis and inflorescence rachis tawny villous; leaflets with abaxial
surface villous (Brazil: Bahia and Minas Gerais) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. sericiflorum Vogel
11. Leaf rachis and inflorescence rachis ferruginous tomentose; leaflets with abaxial surface sericeous
only on the primary vein (Brazil: Paraı́ba, Pernambuco, Alagoas, Bahia, Distrito Federal, Goiás,
Mato Grosso do Sul, Minas Gerais, Espı́rito Santo, and Rio de Janeiro) . . . . . . . . . . . . . . . . . . . . . . . . . M. aculeatum Raddi
3. Leaflets with brochidodromous pattern, the secondary veins uniting in simple or compound arches
toward the margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12. Leaflets with adaxial surface glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13. Leaves (7–)9–15-foliolate, leaflets (0.6–)1–2 cm wide; bracteoles with the same length
as the calyx; petals purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14. Leaf rachis glabrous; leaflets with abaxial surface glabrous; pedicel up to 2 mm long;
bracteole strigose (Brazil: Rio de Janeiro) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. obovatum Kuhlm. & Hoehne
14. Leaf rachis sericeous; leaflets with abaxial surface sericeous; pedicel up to 0.5 mm long;
bracteole sericeous (Brazil: Bahia, Espı́rito Santo, and Minas Gerais) . . . . . . . . . . . . . . . . . . . . . . M. ovalifolium Glaz. ex Rudd
13. Leaves 15–50-foliolate, leaflets 0.2–0.7 cm wide; bracteoles shorter than calyx;
petals yellowish to greenish-white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15. Leaflets narrowly oblong and revolute; inflorescence
congested; flowers sessile; bracteole ovate; petals
yellowish, the banner internally sericeous (Brazil:
Bahia, Espı́rito Santo, and Minas Gerais) . . . . . . . . . . . M. jobimianum C. V. Mendonça, Forni-Martins & A. M. G. Azevedo
15. Leaflets oblong and plane; inflorescence lax; flowers pedicellate, the pedicel 1.5–2 mm long;
bracteole orbicular; petals greenish-white, the banner internally glabrous
(Brazil: Espı́rito Santo, Minas Gerais, and Rio de Janeiro) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. gracile Benth.
12. Leaflet adaxial surface with indumentum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16. Bracteoles shorter than calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17. Leaflets with brochidodromous pattern, the secondary veins uniting
in simple arches toward the margin; calyx teeth triangular; samara
with seed chamber reticulous and wing with homogeneous color
(Brazil: Espı́rito Santo, Minas Gerais, and Rio de Janeiro) . . . . . . . . . . . M. ruddianum C. V. Mendonça & A. M. G. Azevedo
17. Leaflets with multiple-brochidodromous pattern, the secondary veins uniting in compound
arches toward the margin; calyx teeth obtuse; samara with seed chamber striate and wing
with heterogeneous color (Bolivia, Brazil, and Paraguay) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. scleroxylon Tul.
16. Bracteoles with the same length as the calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18. Leaflets 0.2–0.7 cm wide; bracteoles and calyx teeth without strigose indumentum on margin . . . . . . . . . . . . . . . . . . . . . . . . . 19
19. Branches glabrous; inflorescence with lax racemes and panicles; spinescent bracts;
petals purple (Brazil: São Paulo, Paraná, and Santa Catarina) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. hatschbachii Rudd
19. Branches puberulent; inflorescence with congested
panicles; foliar bracts; petals pale purple
(Brazil: Rio de Janeiro) . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. macaense C. V. Mendonça, A. M. G. Azevedo & H. C. Lima
18. Leaflets 0.6–2 cm wide; bracteoles and calyx teeth with
strigose indumentum on margin (Brazil and Argentina) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. nyctitans (Vell.) Benth.
2. Branches with spinescent and foliar stipules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20. Leaflets 5–11, with multiple-brochidodromous pattern, the secondary veins uniting in compound arches toward the margin . . . . . . . . . 21
21. Trees; spinescent stipules perpendicular to the branch, reddish, up to 1 cm long,
without strigose indumentum; leaflets 5–7 (Brazil: Pernambuco, Bahia, and Rio de Janeiro) . . . . . . . . . . . . . M. leucopterum Vogel
21. Lianas to scandent shrubs; spinescent stipules uncinate, up to 0.4 cm long,
with strigose indumentum; leaflets 7–11 (Brazil: Minas Gerais and Rio de Janeiro) . . . . . . . . . . . . . . . . . . . . . . . . . M. glabrum Vogel
20. Leaflets 11–25(–29), with brochidodromous pattern, the secondary veins uniting in simple
arches toward the margin (Brazil: Minas Gerais, Espı́rito Santo, Rio de Janeiro, and São Paulo) . . . . . . . . . . . . M. cantarellianum Hoehne
1. Unarmed plants, rarely armed only on terminal branches of individuals with scandent habit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22. Leaflets with weakly brochidodromous pattern, the parallel secondary veins uniting in inconspicuous
arches that end in margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23. Leaflets with adaxial surface sparsely villous to glabrescent; flowers pedicellate, the pedicel
up to 1.5 mm long; stamens diadelphous 9 + 1, filaments villous (Brazil: Minas Gerais) . . . . . . . . . . . . . . . . . . . M. androvillosum sp. nov.
23. Leaflets with adaxial surface glabrous and lustrous; flowers sessile; stamens monadelphous,
filaments glabrous (Brazil: Minas Gerais and Rio de Janeiro) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. nigrum Vogel
22. Leaflets with brochidodromous pattern, the secondary veins uniting in simple or compound arches toward the margin . . . . . . . . . . . . . . . . . 24
24. Leaflet apex obtuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25. Leaflets (7–)9–15, base acute to attenuate, adaxial surface sparsely-sericeous to glabrous;
flowers 5–7 mm long; petals cream to greenish; stamens monadelphous; samara with seed
chamber glabrous and striate (Bolivia, Brazil, Argentina, and Paraguay) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. stipitatum Vogel
25. Leaflets 17–27, base rounded, adaxial surface tomentose, puberulent to glabrescent;
flowers 11–13 mm long; petals purple; stamens diadelphous 9 + 1; samara with seed
chamber villous and verrucose (Brazil: Rio de Janeiro) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. firmum (Vell.) Benth.
24. Leaflet apex acute or acuminate to cuspidate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26. Petals purple to vinaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
27. Trees; stamens diadelphous 9 + 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
28. Leaflets tomentose-villous on both surfaces; samara with seed chamber striate
(Brazil: Bahia, Espı́rito Santo, and Minas Gerais) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. fulvovenosum H. C. Lima
28. Leaflets glabrous adaxially; samara with seed chamber striate or verrucose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
29. Leaflets 5–9(–11), multiple-brochidodromous pattern, the secondary veins uniting
in compound arches toward the margin (Brazil: Bahia, Espı́rito Santo,
Minas Gerais, and Rio de Janeiro) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. pedicellatum Vogel
29. Leaflets 11–25(–29), brochidodromous pattern, the secondary veins uniting in simple
arches toward the margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30. Leaflets 0.2–1(–1.5) cm wide; stipules and bracts caducous; samara with seed chamber
striate or verrucose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
148 SYSTEMATIC BOTANY [Volume 39

31.
Leaflets abaxial surface villous, tomentose, or puberulent; samara
with seed chamber striate and wing apex obtuse (Brazil: Bahia,
Espı́rito Santo, Rio de Janeiro, and São Paulo) . . . . . . . . . . . . . . . . . . . . . . . . M. incorruptibile (Vell.) Benth.
31. Leaflets abaxial surface sericeous; samara with seed chamber verrucose
and wing apex acute (Brazil: Rio de Janeiro) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. legale (Vell.) Benth.
30. Leaflets 1.2–4.5(–5.5) cm wide; stipules and bracts persistent; samara with
seed chamber verrucose (Brazil: Espı́rito Santo, Minas Gerais,
and Rio de Janeiro) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. robsonnianum sp. nov.
27. Lianas, scandent shrubs, rarely small trees; stamens monadelphous (Brazil: Paraı́ba,
Pernambuco, Alagoas, Bahia, and Espı́rito Santo) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. salzmannii Benth.
26. Petals white, cream, yellowish to greenish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
32. Adaxial surface of mature leaflets glabrescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
33. Trees with trunk not ramified at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
34. Bracteole 0.2–0.4(–0.6) mm wide, linear, rarely oblong (Bolivia, Brazil, and Argentina) . . . . . . . . . M. brasiliense Vogel
34. Bracteole 0.5–2 mm wide, ovate or obovate to orbicular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
35. Leaf rachis puberulent to glabrous; inflorescence pendent; flowers 8–12 mm long;
banner petal internally sericeous; ovary uniformly villous to tomentose
(Venezuela, Peru, Bolivia, Brazil, Paraguay, and Argentina) . . . . . . . . . . . . . . . . . . . . . . . . M. acutifolium Vogel
35. Leaf rachis villous to tomentose; inflorescence erect; flowers (6–)7–7.8 mm long;
banner petal internally glabrous; ovary villous only on the margins
(Brazil, Paraguay, and Argentina) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. paraguariense Hassl.
33. Lianas to scandent shrubs, rarely small trees with trunk ramified at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
36. Bracteole 0.2–0.4(–0.6) cm wide, linear, rarely oblong (Bolivia, Brazil, and Argentina) . . . . . . . . . . M. brasiliense Vogel
36. Bracteole 0.8–2 cm wide, ovate, obovate to orbicular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
37. Inflorescence congested, rachis puberulent, tomentose to sericeous; calyx tomentose to sericeous . . . . . . . . . 38
38. Leaflets (3–)5–7; flowers 5–6 mm long; calyx campanulate
(Brazil: Pernambuco, Bahia, Goiás, Distrito Federal, Minas Gerais,
Espı́rito Santo, Rio de Janeiro, São Paulo, and Paraná) . . . . . . . . . . . . . . M. lanceolatum (Vell.) J. F. Macbr.
38. Leaflets (3–)5–25(–29); flowers 8–10 mm long; calyx cylindrical
to campanulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
39. Leaflets (3–)5–9; erect inflorescence; calyx cylindrical, sericeous
(Brazil: Minas Gerais, Rio de Janeiro, São Paulo, Paraná,
and Santa Catarina) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. debile (Vell.) Stellfeld
39. Leaflets (7–)11–25(–29); pendent inflorescence; calyx campanulate,
tomentose (Brazil: Minas Gerais, Espı́rito Santo, Rio de Janeiro,
and São Paulo) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. cantarellianum Hoehne
37. Inflorescence lax, rachis glabrous; calyx glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
40. Lianas with angular branches; leaves 3(–5)-foliolate,
leaflets (2.5–)3–9.5 cm wide; bracteole 0.8–1 mm wide,
tomentose to sericeous; calyx campanulate
(Brazil: Espı́rito Santo, Minas Gerais, and Rio de Janeiro) . . . . . . . . . . . . . M. ternatum Kuhlm. & Hoehne
40. Lianas with cylindrical branches; leaves 7–11-foliolate, leaflets (1.2–)1.5–4 cm wide;
bracteole 1.2–1.5 mm wide, glabrous; calyx cylindrical (Brazil: Minas Gerais,
and Rio de Janeiro) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. glabrum Vogel
32. Adaxial surface of mature leaflets villous to tomentose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
41. Trees with trunk not ramified at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
42. Branches densely villous; leaves usually with numerous leaflets (11–25);
inflorescence congested and pendent; calyx teeth triangular (Bolivia, Brazil,
and Paraguay) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. villosum Vogel
42. Branches tomentose or puberulent to glabrescent; leaves usually with fewer
leaflets (7–13); inflorescence lax and erect; calyx teeth obtuse (Brazil, Paraguay,
and Argentina) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. paraguariense Hassl.
41. Lianas to scandent shrubs, rarely small trees with trunk ramified at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
43. Leaflets sericeous beneath; calyx dark-brown-tomentose; banner petal externally
dark-brown-sericeous; nectary crateriform; ovary predominantly glabrous,
villous-strigose only at base and apex (Brazil: Bahia, Minas Gerais,
and Rio de Janeiro) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. punctatum (Poir.) Pers.
43. Leaflets villous to tomentose beneath; calyx dark-red-tomentose or aureo-sericeous-tomentose;
banner petal externally dark red or aureo-sericeous-tomentose; nectary cupuliform;
ovary glabrous and sparsely villous only on margins, or completely sericeous-villous . . . . . . . . . . . . . . . . . . . . . . .44
44. Branches, leaf and inflorescence rachis, and abaxial surface of leaflets covered
with dark-red indumentum; bracts caducous; bracteole 1.8–2.2(–3) mm long;
calyx campanulate; stamens with filaments glabrous; ovary glabrous and
sparsely villous only on margins (Brazil: Goiás, Bahia, Minas Gerais,
Espı́rito Santo, Rio de Janeiro, and São Paulo) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. oblongifolium Vogel
44. Branches, leaf and inflorescence rachis, and abaxial surface of leaflets covered
with golden indumentum; bracts persistent; bracteole (3–)3.8–5 mm long;
calyx cylindrical; stamens with filaments villous; ovary completely
sericeous-villous (Brazil: Bahia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. aureum sp. nov.

Machaerium androvillosum Filardi & H. C. Lima, sp. nov.—
TYPE: BRAZIL. Minas Gerais. Municı́pio de Viçosa,
Escola Superior de Agronomia e Veterinária (ESAV), 22
Feb 1936, J. G. Kuhlmann s. n. (holotype: BHCB 65912!;
isotypes: UB!, VIC 1326!).
Machaerium androvilosum sp. nov.; M. nigrum Vogel affinis,
sed foliolum indumento supra villosus, floris pedicellata et
androecium diadelphus (9 + 1) et etiam villosus differt.
Trees 5–6 m tall; trunk and bark not seen; branches
2–4 mm diam., sparsely tomentose, glabrescent, conspicuous
2014] FILARDI AND LIMA: MACHAERIUM IN THE ATLANTIC FOREST
lenticels, unarmed. Stipules ca. 11 1 mm, lanceolate,
+
caducous. Leaves 10– 18.5 cm long, imparipinnate; petiole
2–3.4 cm, ca. 1 mm diam., fulvo-villous, glabrescent; rachis
8–15 cm long, 1–2 mm diam., fulvo-villous, glabrescent;
pulvinule 2–3 mm long. Leaflets 11–15, sub-opposite to
opposite, 3.5–9 1.5–3.2 cm, elliptic to oblong-elliptic, apex
+
acute to acuminate, mucronulate, base obtuse to acute, rarely
truncate, adaxial surface sparsely villous to glabrescent,
abaxial surface fulvo-villous. Panicles ca. 20 cm long, axil-
lar or terminal; peduncle 1.5–2 cm long, 2.5–3 mm diam.,
fulvo-tomentose; rachis ca. 18 cm long., fulvo-tomentose;
lateral axis 4.5–9 cm long, fulvo-tomentose; bracts 3–3.8
+
1.8–2 mm, oblong-obovate, fulvo-villous; pedicel ca. 1.5 mm
long, villous; bracteoles 2.8–3 2.1–2.5 mm, ovate, sericeous.
+
Flowers ca. 10 mm long; calyx ca. 5 3.2 mm, brown-
+
yellowish, campanulate, sparsely villous, striate, adaxial teeth
ca. 0.8 1.8–2 mm, apex obtuse, abaxial teeth ca. 1 2 mm,
+
+
apex triangular; petals purple, striate; the banner petal
9–10 ca. 9 mm, glabrous internally, fulvo-sericeous exter-
+
nally, apex retuse, base truncate to attenuate, claw ca. 1
+
1.5 mm; the wing petals 8–9 3–3.5 mm, oblong, gla-
+
brous, apex obtuse, base obtuse to attenuate, claw ca. 3 + Machaerium aureum sp. nov. similis M. oblongifolium Vogel
0.5 mm, sparsely villous; the keel petals ca. 8.2 3.8 mm,
+
oblong, plicate, glabrous, apex obtuse, base obtuse, claw ca.
3 0.1–0.2 mm, sparsely villous; stamens diadelphous 9 + 1,
+
filament 6.5–7.8 mm long, filament sheath ca. 4.2 mm
long, fulvo-villous; anthers ca. 0.4 0.2 mm, oblong; nectary
+
ca. 1 1 mm, cupuliform; ovary ca. 3 1–1.8 mm, fulvo-
+
villous; stipe 3.8–4 mm long, villous; style 1.6–1.8 mm long,
glabrous. Samaras 7–8 cm long; stipe 5–6 mm long, fulvo-
villous; seed chamber 1.5–1.6 1.3–1.4 cm, oblong, brown,
+
villous, longitudinally striate; wing 5.2–6 ca. 2.8 cm, oblong-
+
ovate, sparsely villous, reticulate, apex obtuse. Figure 1.
Habitat and Distribution—Machaerium androvillosum is
only known from a few collections in seasonally semi-
deciduous and deciduous forests at 200–600 m altitude in
the Brazilian state of Minas Gerais.
Phenology—The species flowers between October and
March, and the fruits were collected in March.
Conservation Status—Machaerium androvillosum is assessed
here as endangered (EN), within the criterion B1ab (i, ii, iii,
iv, v) (IUCN 2001). Despite extensive fieldwork and analy-
sis of representative collections, this species is known only
from five herbarium specimens collected between 1930
and 1942. In addition, the two municipalities where the
species was collected, Viçosa and Governador Valadares,
are now extensively deforested, with most of the original
vegetation destroyed (e.g. Fundação SOS Mata Atlântica
1998; MMA 2002).
Etymology—The specific epithet refers to the androecium
with villous filaments, one of the characteristics used to rec-
ognize this new species.
Notes—The type material designated for Machaerium
androvillosum was cited by Hoehne (1938) in his combination
of M. fruticosum (= M. nigrum) based on Nissolia fruticosa sensu
Vellozo (1831). Machaerium androvillosum differs from the
morphologically similar M. nigrum by leaflets villous on both
surfaces (vs. adaxial surface glabrous and lustrous), pedicel-
late flowers (vs. sessile), and stamens diadelphous with villous
filaments (vs. stamens monadelphous with glabrous fila-
ments). Machaerium spicatum Kuhlm. & Hoehne can also be
considered as morphologically similar to M. androvillosum.
However, the analysis of the material originally cited for
149
M. spicatum (RB 5071) by Hoehne (1938) revealed that it
readily differs from M. androvillosum by having leaflets with
base rounded to cordate and shorter length of floral parts,
and especially because of the dense hair covering the imma-
ture fruits. The isotype designated for UB was sent as inter-
change, in 1968, by RB herbarium and its label includes
“RB 45168”. However, despite this number being registered
as “Machaerium fruticosum (Vell.) Hoehne, J. G. Kuhlmann
s. n., 22/II/1936”, unfortunately this material was not found
in RB.
Additional Specimens Examined—BRAZIL. Minas Gerais: Governador
Valadares, 29 Oct 1942, M. Magalhães 2248 (BHCB); Viçosa, Escola Superior
de Agricultura e Veterinária (ESAV), 9 Mar 1930, Y. Mexia 4446 (P, US,
VIC); Viçosa, Escola Superior de Agricultura e Veterinária (ESAV), 22 Mar
1930, Y. Mexia 4501 (G, P, US, VIC); Viçosa, Escola Superior de Agricultura
e Veterinária (ESAV), 22 Mar 1930, Y. Mexia 4504 (G, P, US, VIC).
Machaerium aureum Filardi & H. C. Lima, sp. nov.—TYPE:
BRAZIL. Bahia. Municı́pio de Mucuri, Rodovia BA 698,
Km 35, 18 020 5000 S, 39 370 2900 W, 22 Jul 2008, F. L. R.
Filardi, H. C. Lima, R. D. Ribeiro, A. Silva & A. Calvet 848
(holotype: RB!; isotypes CEPEC!, K!, NY!).
habitu, sed foliis et inflorescentiae indumento conspicuo
colore aureum et etiam floris majoribus abunde differt.
Lianas to scandent shrubs up to 3 m tall; bark flat, gray-
brown to brown, puberulent to glabrescent; branches 3–
4 mm diam., striate, brown, densely aureo-tomentose, incon-
spicuous lenticels, unarmed or armed in the apical branches.
Stipules not seen. Leaves 12.5–23.5 cm long, imparipinnate;
petiole (1–)2–4 cm long, 1–2 mm diam., aureo-villous-
tomentose to glabrescent; rachis (3.5–)5–8.7(–10.3) cm long,
1–1.5 mm diam., aureo-villous-tomentose to glabrescent;
pulvinule 0.5–3 mm long. Leaflets 7–9, opposite, (3.5–)5–
10.5(–12) (1.5–)2.5–5.6 cm, obovate, elliptic-obovate, ellip-
+
tic, oblong-elliptic, apex acuminate, mucronulate, base
rounded, attenuate to oblique, adaxial surface sparsely
aureo-villous to glabrescent and lustrous, abaxial surface
aureo-villous mainly on the midrib. Inflorescence a fascicu-
late raceme or panicles, 3–9(–15) cm long, axillar; peduncle
0–0.8 cm long, ca. 3 mm diam., aureo-villous-tomentose to
glabrescent; rachis 3.3–7(–13.8) cm long, aureo-villous-
tomentose to glabrescent; lateral axis 0–4 cm long, aureo-
villous-tomentose to glabrescent; bracts 5.5–6 2–3.8 mm,
+
ovate to deltoid, aureo-villous-tomentose; pedicel 0–0.5-mm
long, tomentose; bracteoles (3–)3.8–5 1.8–2.8 mm, oblong-
+
ovate, aureo-villous. Flowers (8–)9–12 mm long; calyx 4.2–
5.5 2.8–3.2 mm, brown-green, cylindrical, aureo-villous-
+
tomentose, adaxial teeth 1–1.3 1.5–2 mm, abaxial teeth
+
0.7–1.8 0.7–1.1 mm, both with apex triangular; petals
+
white-green, striate; the banner petal 9–11 6–7 mm,
+
glabrous and with pale purple striations internally, aureo-
sericeous-tomentose externally, apex rounded to retuse, base
attenuate, claw 2.8–3 0.8–1.3 mm; the wing petals 8.2–
+
10.2 2–2.6 mm, oblong-elliptic, aureo-sericeous-tomentose
+
dorsally, apex obtuse, base truncate, claw 3.2–4 0.2–
+
0.3 mm; the keel petals 8–9.5 2–2.5 mm, oblong-elliptic,
+
aureo-sericeous-tomentose dorsally, apex obtuse, base auric-
ulate, claw 3.3–4 0.1–0.3 mm; stamens diadelphous 9 +
+
1, filaments 6–8 mm long, filament sheath 3–5.5 mm long,
sparsely villous; anthers 0.4–0.5 0.3–0.4 mm, oblong; nec-
+
tary 0.3–0.4 0.3–0.4 mm, cupulate, oblique; ovary 3–4
+
0.5–0.8 mm, aureo-sericeous-villous mainly at the base, mar-
gins, and apex; stipe 2–3.3 mm long, aureo-villous mainly on
150 SYSTEMATIC BOTANY [Volume 39

Fig. 1. Machaerium androvillosum. A. Inflorescence. B. Detail of unarmed branch. C. Leaflets, base of abaxial surface. D. Bracteole. E. Calyx. F. Banner
petal, adaxial surface. G. Wing petal. H. Keel petals. I. Stamens. J. Gynoecium. K. Fruit. (A–J from Kuhlmann s. n. (BHCB 65912); K from Mexia 4501).
2014] FILARDI AND LIMA: MACHAERIUM IN THE ATLANTIC FOREST
upper half; style 1.5–2 mm long, glabrous. Samaras (imma-
ture) 5.2–6.8 cm long; stipe 5–8 mm long, glabrous; seed
chamber 2–2.4 1–1.1 cm, oblong to slightly falcate,
+
glabrous, uniformly proeminent, reticulate; wing 4.3–8
+
1.2–1.4 cm, oblong, yellow-brown, glabrous, reticulate, apex
obtuse, short-apiculate. Figure 2.
Habitat and Distribution—Machaerium aureum is endemic
to southern Bahia and occurs in wet forests on formations of the
Tertiary and Quaternary, including the vegetation types locally
known as “mussununga” (e.g. Simonelli 2007) and restinga.
The species grows from sea level to 180 m in elevation.
Phenology—The species flowers between June and
August, and the fruits were collected in June and July.
Conservation Status—Machaerium aureum occurs in areas
of the state of Bahia that have been heavily impacted by the
replacement of natural vegetation by plantations of Eucalyp-
tus L’Her. for the paper industry. Thus, the species is
assessed here as vulnerable (VU), within the criterion B1ab
(ii, iii) (IUCN 2001).
Etymology—The epithet refers to the golden-colored hairs
present on the branches, leaves, inflorescence axes, bracts,
bracteoles, petals, and gynoecium.
Notes—Both Machaerium aureum and M. oblongifolium
Vogel have a scandent habit, but the former is distinctive
because of the golden color of the hairs present on its
branches, leaves, inflorescences, and floral parts (vs. dark-
red color of hairs in M. oblongifolium). Other distinctive char-
acters of the new species are leaflets in greater number (7–9
vs. 3–5 (–7)), flowers up to 12 mm long (vs. up to 9 mm long),
and calyx cylindrical with triangular teeth (vs. campanulate
with obtuse teeth).
Additional Specimens Examined—BRAZIL. Bahia: Alcobaça, 11 Aug
1995, G. Hatschbach 62966 (HUEFS, MBM); Caravelas, 29 Jul 2009, N. C.
Fraga et al. 2784 (RB); Medeiros Neto, 4 Feb 2009, D. Cardoso et al. 2405
(HUEFS, RB); Porto Seguro, 5 Jul 1990, D. Folli 1195 (CVRD, K); Porto
Seguro, 21 Jun 2004, V.C. Souza 29976 (ESA); Porto Seguro, 2 Jul 2006,
M. M. M. Lopes et al. 929 (CEPEC, HUEFS, NY, RB); Santa Cruz de Cabrália,
16 Jul 1981, H. S. Brito & P. C. Vinha 15 (CEPEC, K, RB); Santa Cruz de
Cabrália, 25 Aug 1988, L. A. Mattos Silva et al. 2526 (CEPEC, K, RB, SPF).
Machaerium robsonnianum Filardi & H. C. Lima, sp. nov.—
TYPE: BRAZIL. Rio de Janeiro. Municı́pio de Macaé,
Praia de Imbetiba, Ilha Santana (Força Aérea Brasileira),
distante aproximadamente 8 km da orla, na subida para
o Farol [approximately 8 km from the shore, on the
ascent to the Lighthouse], 13 Feb 2007, F. L. R. Filardi,
H. C. Lima, R. D. Ribeiro & V. Maioli 706 (holotye: RB!;
isotypes: BHCB!, MBML!, K!, NY!).
Machaerium robsonnianum sp. nov.; M. fulvovenosum H. C.
Lima affinis, sed habitu, foliolum supra glabrum, bracteolis
majoribus et fructus circa semina verrucosus abunde differt.
Trees (2–)3–7(–10) m tall, trunk straight or angled, usually
divided from the base, (5–)10–25 cm diam., unarmed, bark
squamous, brown to brown-gray, inconspicuous lenticels;
branches 4–8 mm diam., striate, brown-gray, brown-tomentose,
conspicuous lenticels, unarmed. Stipules 8–18 2–5 mm,
+
spathulate, oblong, oblong-lanceolate to lanceolate, persistent.
Leaves 12–27 cm long, imparipinnate; petiole 2.3–3.5 cm long,
1–2 mm diam., brown-tomentose; rachis 7.2–14.4 cm long, 1–
1.5 mm diam., red-brown-tomentose; pulvinule 2–4 mm long.
Leaflets 11–17, sub-opposite to alternate, 2.2–8.8 1.2–4 cm,
+
obovate, oblong-elliptic or elliptic, apex obtuse, acute to acu-
minate, mucronulate, base rounded, cuneate to oblique, adax-
ial surface glabrescent to glabrous, abaxial surface villous,
151
sericeous to glabrescent, secondary veins prominent beneath.
Inflorescence paniculate or a fasciculate raceme, 5.5–18 cm
long, axillar, supra-axillar or terminal; peduncle 0–2.1 cm
long, 1–4 mm diam., puberulent to brown-tomentose; rachis
4–17 cm long, brown to red-brown-tomentose; lateral axis (3–)
4.5–9 cm long, brown to brown-tomentose; bracts 3.5–10 1–
+
3.2 mm, spathulate, lanceolate, oblong-lanceolate, villous to
tomentose, rarely caducous; bracteoles 2–2.2 2–2.5 mm,
+
large-obovate to orbicular, tomentose. Flowers sessile, 8–
10 mm long; calyx 3.5–4.2 2.8–3.2 mm, red-brown, campan-
+
ulate, sericeous-tomentose, adaxial teeth ca. 1 1.5–2 mm,
+
apex obtuse, abaxial teeth 0.8–1 1–1.4 mm, apex triangular;
+
petals rose-purple to white, striate; the banner petal 7.2–8.2
+
8–9.2 mm, glabrous internally, sericeous-tomentose exter-
nally, with a central white macula, apex tomentose, emargin-
ate, base attenuate to truncate, claw 1.5–1.8 1–1.6 mm; the
+
wing petals 9–10 3.5–3.8 mm, oblong, apex obtuse, sparsely
+
tomentose, base slightly auriculate to truncate, claw 3.8–4
+
0.3–0.5 mm; the keel petals 8.5–9 3–3.2 mm, elliptic, gla-
+
brous, apex obtuse, base auriculate, claw 3.5–4 0.2–0.3 mm;
+
stamens diadelphous 9 + 1, filaments 6.2–8.2 mm long, fila-
ment sheath 3.2–4.8 mm long, sparsely-villous to glabrescent;
anthers ca. 0.3 0.3 mm, oblong; nectary 0.9–1.3 1.1–
+
1.6 mm, cupulate, oblique; ovary 2–2.3 1.1–1.5 mm,
+
vellutinous; stipe 3–3.5 mm long, vellutinous to sparsely vil-
lous; style 1.3–1.8 mm long, glabrous. Samaras 6.8–8.5 cm
long; stipe 7–10 mm long, brown-tomentose, puberulent to
glabrescent; seed chamber 2–3 1.3–1.8 cm, oblong-falcate,
+
brown-tomentose, sericeous to puberulent, verrucose; wing
4.6–6.3 1.5–2.8 cm, oblong-falcate, fulvo-tomentose, seri-
+
ceous to puberulent, reticulate, apex obtuse to acute. Figure 3.
Habitat and Distribution—Machaerium robsonnianum is
restricted to southern Brazil, in the states of Espı́rito Santo,
Minas Gerais, and Rio de Janeiro, where it predominantly
occurs in seasonal semideciduous forests and restinga vegeta-
tion, mainly along the coast from sea level to 670 m elevation.
Phenology—Flowers have been collected mainly between
October and December, and fruits have been collected through-
out the year, but especially between March and August.
Conservation Status—Machaerium robsonnianum is cate-
gorized here as least concern (LC) because it does not
match any of the IUCN categories of threat (IUCN 2001).
Although the species occur in areas that suffer with
intense real estate speculation, M. robisonnianum is abun-
dant throughout its distribution and also is present in pro-
tected reserve areas.
Etymology—The epithet is a posthumous tribute to the
botanist Robson Daumas Ribeiro (JBRJ) for his sincere friend-
ship, teachings, encouragement and valuable collections.
Notes—Machaerium robsonnianum is a tree with a tortuous
trunk that is generally ramified at the base reaching 3–7 m in
height, while the most morphologically similar species,
M. fulvovenosum H. C. Lima, has a straight trunk not rami-
fied at the base and usually exceeds 10 m in height. Other
diagnostic characters for this new species are the persis-
tent stipules measuring 8–18 mm long (vs. early caducous
stipules in M. fulvovenosum), the leaflets with glabrous
adaxial surface (vs. tomentose adaxial surface) and generally
larger (up to 4 cm wide vs. up to 2.5 cm wide), the sessile
flowers (vs. pedicellate flowers), the stamens with villous
filaments and anthers up to 0.3 mm long (vs. puberulent
filaments and anthers up to 0.7 mm long), and the verrucose
seed chamber (vs. striate).
152 SYSTEMATIC BOTANY [Volume 39

Fig. 2. Machaerium aureum. A. Inflorescence. B. Detail of armed branch. C. Leaflets, apex of adaxial surface. D. Leaflets, apex of abaxial surface.
E. Bracteole, adaxial surface. F. Bracteole, abaxial surface. G. Calyx. H. Banner petal, adaxial surface. I. Wing petal. J. Keel petals. K. Stamens.
L. Gynoecium. M. Fruit. (A and E–L from Filardi 848; B–D from Cardoso 2405; M from Souza 29976).
2014] FILARDI AND LIMA: MACHAERIUM IN THE ATLANTIC FOREST 153

Fig. 3. Machaerium robsonnianum. A. Inflorescence and immature fruits. B. Detail of the unarmed branch with persistent stipules. C. Leaflets, abaxial
surface. D. Bracteole. E. Calyx. F. Banner petal, adaxial surface. G. Wing petal. H. Keel petals. I. Stamens. J. Gynoecium. K. Fruit. (A–K from Filardi 706).
154 SYSTEMATIC BOTANY
Additional Specimens Examined—BRAZIL. Espı́rito Santo: Governador
Lindemberg, 26 Apr 2007, V. Demuner et al. 3886 (MBML, RB); Presidente
Kennedy, 6 Feb 1988, J. M. L. Gomes 460 (RB, VIES); Presidente Kennedy,
10 Apr 1993, J. M. L. Gomes et al. 1848 (RB, VIES); Santa Teresa, 20 Jun
2000, V. Demuner et al. 1157 (MBML, RB); Serra, 5 Apr 2007, F. L. R. Filardi
et al. 763 (RB); Serra, 5 Apr 2007, F. L. R. Filardi et al. 764 (RB); Vitória,
26 Jun 1991, E. Pereira 86 (RB, VIES); Vitória, 26 Jun 1991, E. Pereira 87 (RB,
VIES). Minas Gerais: Dionı́sio, 13 Feb 1986, W. G. Campos s. n. (BHCB
16865); Ipaba, 9 Oct 2004, G. S. França et al. 650 (BHCB); Leme do Prado,
16 Mar 2001, E. Tameirão Neto 3287 (BHCB); Marliéria, 24 Oct 1982,
P. Heringer 18612 (IBGE, RB); Marliéria, Nov 1993, L. V. Costa et al. s. n.
(BHCB 30731); Muriaé, 17 Apr 1999, A. Salino 4610 (BHCB); Rio Casca,
28 Feb 1998, A. Salino 4074 (BHCB); São Pedro dos Ferros, 9 Dec 2000,
F. M. B. Pereira 25/56 (RB, RFA). Rio de Janeiro: Abre Campo, 20 Dec
2000, E. Pereira 22159 (RB, RFA); Armação dos Búzios, 19 Oct 1993, H. C.
Lima et al. 4803 (RB); Armação dos Búzios, 6 Dec 1998, D. Fernandes et al.
192 (RB); Armação dos Búzios, 20 May 1999, A. Lobão et al. 423 (RB);
Armação dos Búzios, 22 Aug 2001, C. Farney 4380 (RB); Armação dos
Búzios, 29 Aug 2003, H. G. Dantas et al. 13 (RB); Armação dos Búzios,
31 Aug 2003, H. C. Lima et al. 6117 (RB); Armação dos Búzios, 17 May
2005, R. D. Ribeiro et al. 468 (RB); Armação dos Búzios, 17 May 2005,
R. D. Ribeiro et al. 470 (RB); Armação dos Búzios, 18 May 2005, R. D.
Ribeiro et al. 483 (RB); Armação dos Búzios, 22 Feb 2006, R. D. Ribeiro
et al. 601 (RB); Armação dos Búzios, 15 Feb 2007, F. L. R. Filardi et al.
718 (RB); Armação dos Búzios, 15 Feb 2007, F. L. R. Filardi et al. 722
(RB); Armação dos Búzios, 5 Jul 2008, R. D. Ribeiro et al. 993 (RB);
Armação dos Búzios, 26 Apr 2010, R. D. Ribeiro et al. 1461 (CEPEC, K,
NY, RB); Armação dos Búzios, 23 May 2010, R. D. Ribeiro et al. 1539
(K, NY, RB); Cabo Frio, 19 Oct 1993, H. C. Lima 4803 (RB); Cabo Frio,
26 May 1995, S. M. Faria et al. 930 (RB); Cabo Frio, 27 May 1995, S. M.
Faria et al. 940 (RB); Carmo, 28 Jul 2010, M. Barros & M. P. Morim 84
(RB); Macaé, 7 Mar 2004, R. D. Ribeiro et al. 96 (RB); Macaé, 26 Mar
2005, R. D. Ribeiro & V. Maioli 419 (RB); Macaé, 1 Mar 2006, R. D.
Ribeiro & V. Maioli 624 (RB); Rio das Ostras, 28 Feb 1999, J. M. A.
Braga 56 (RB); Rio das Ostras, 18 Mar 2000, H. C. Lima et al. 6050 (RB);
Rio das Ostras, 18 Mar 2003, H. C. Lima et al. 6069 (RB); Rio das
Ostras, 18 Mar 2003, H. C. Lima et al. 6073 (RB); Rio das Ostras,
18 Mar 2003, H. C. Lima et al. 6075 (RB); Rio das Ostras, 18 Mar 2003,
H. C. Lima et al. 6076 (RB); Rio das Ostras, 6 Dec 2008, R. D. Ribeiro &
V. Maioli 1050 (RB).
MACHAERIUM ACUTIFOLIUM Vogel, Linnaea 11: 187. 1837
[March].—TYPE: BRAZIL. “Brasil merid.,” F. Sellow s. n.
(holotype: B, destroyed; lectotype here designated:
K 000530222!; photo at NY 2645!; isolectotype: E!).
Machaerium muticum Benth., Comm. Legum. Gen. 36. 1837
[June]. Machaerium acutifolium var. muticum (Benth.)
Benth., Fl. bras. 15 (1b): 245. 1862.—TYPE: BRAZIL.
Minas Gerais: São João Baptista, 1837, J. E. Pohl 560
(holotype: W!; isotypes: BR, K!, M!, NY!, US!). syn. nov.
Machaerium juglandifolium Rusby, Bull. New York Bot. Gard.
6(22): 513–514. 1910.—TYPE: BOLIVIA. Tumpasa, 19 Jan
1902, R. S. Williams 529 (holotype: NY!; isotypes: BM!,
US!). syn. nov.
Machaerium pseudacutifolium Pittier, Bol. Soc. Ven. Cien. Nat.
7: 148. 1941. Machaerium acutifolium var. pseudacutifolium
(Pittier) Rudd, Phytologia 25 (6): 399. 1973.—TYPE:
VENEZUELA. Guárico, Mesa de El Sombrero, in
savanna, 10 Sep 1927, H. Pittier 12484 (holotype: VEN;
isotypes: A, G!, M!, MAD, NY!, US!). syn. nov.
Notes—Two varieties proposed for Machaerium acutifolium
by Bentham (1862) and Rudd (1973) were included in its
synonymy because of the wide morphological variation
observed in the shape and dimensions of the leaflets; how-
ever, some doubt about M. acutifolium var. enneandrum
(Hoehne) Rudd still remain. Initially, Hoehne (1938) consid-
ered as diagnostic characters for M. enneandrum the leaves
paripinnate and nine monadelphous stamens, and pointed
[Volume 39
out variations of its characters, considering M. acutifolium
var. muticum as very closely related taxon (Hoehne 1941).
On the other hand, Ducke (1949) included M. enneandrum
under M. hoehneanum Ducke, the most common liana species
of the genus in remnant forests in the vicinity of the city of
Manaus in Amazonas State. Unfortunately, the holotype of
M. enneandrum (R 2580) was not located, and this question
remains unresolved.
MACHAERIUM BRASILIENSE Vogel, Linnaea 11: 185. 1837
[March].—TYPE: BRAZIL. Rio de Janeiro: “Brasil prov.
Rio de Janeiro, Corcovado,” 1834, Luschnath s. n. (Martius
Herbarium Florae Brasiliensis 285) (lectotype here desig-
nated: M!; isolectotypes: BR, G!, P!, W!).
Machaerium vestitum Vogel, Linnaea 11: 190. 1837 [March].—
TYPE: BRAZIL. “Bras. merid.,” F. Sellow s. n. (holotype: B,
destroyed; photo at F 2293!, G!; lectotype here desig-
nated: K 000530231!; photo at NY 2645!). syn. nov.
Machaerium triste Vogel, Linnaea 11: 416. 1837 [March].—
TYPE: BRAZIL. Rio de Janeiro: “pr. Rio Jan.,” Luschnath
s. n. (holotype: B, destroyed; isotype: probably LE),
ex descr.
Machaerium ciliatum Benth., Comm. Legum. Gen. 35. 1837
[June].—TYPE: BRAZIL. Rio de Janeiro: “Brasiliae,
Corcovado,” C. F. P. Martius s. n. (holotype: M!).
Machaerium densicomum Mart. ex Benth., Comm. Legum. Gen.
35. 1837 [June]. Machaerium brasiliense var. densicomum
(Mart. ex Benth.) Hoehne, Revista Mus. Paul. Univ. São
Paulo 10: 697. 1918.—TYPE: BRAZIL. Maranhão: “In
sylvis ad flumen Itapecures,” C. F. P. Martius 3–44c.
(holotype: M!). syn. nov.
Machaerium erianthum Benth., Comm. Legum. Gen. 35. 1837
[June]. Machaerium brasiliense var. erianthum (Benth.)
Rudd, Phytologia 25(6): 399. 1973.—TYPE: BRAZIL.
“Brasilia,” J. E. Pohl s. n. (holotype: B, destroyed; photo
at F 2281!, US!; lectotype: K 0000530246!, designated
by Rudd 1973; photo at NY 2644!; isolectotype: W!).
syn. nov.
Machaerium luschnathianum Presl, Abh. K. Bohm. Ges. Wiss.,
5(3): 61. 1843.—TYPE: BRAZIL. Rio de Janeiro: “habitat
ad Rio de Janeiro Brasiliae,” Luschnath s. n. (holotype:
probably PR; isotype: probably LE), ex descr.
Machaerium acutifolium var. clausseni Benth., Fl. bras. 15 (1b):
245. 1862.—TYPE: BRAZIL. Minas Gerais. “Brasilia,
Minas Gerais,” 1840, P. Claussen s. n. (lectotype here
designated: K 000530223!; photo at NY 2637!;
isolectotype: BR 5196838). syn. nov.
Notes—The circumscription of Machaerium brasiliense
presented here conforms to the wide morphological variation
observed for the vegetative characters, including habit
(lianas, scandent shrubs and trees), and the number, dimen-
sions, and indumentum of the leaflets, that varies according
to the place along the branch and with the maturation of
leaflets. Sessile and pedicellate flowers and linear to oblong
bracteoles were also observed in the same specimen,
depending on their position along the inflorescence rachis.
However, future studies with collections from the Cerrado
biome might reveal the necessity of infraspecific treatment
for M. brasiliense.
2014] FILARDI AND LIMA: MACHAERIUM IN THE ATLANTIC FOREST
Machaerium luschnatianum and M. triste were previously
cited under M. brasiliense (e.g. Bentham 1862 and Mendonça
Filho 1996, respectively). We also believe they are homotypic
synonyms of M. brasiliense, although we could not see their
types. Only one specimen collected by Luschnath in Rio de
Janeiro was cited in the protologue of Machaerium brasiliense
and M. triste by Vogel (1837), as well as of M. luschnatianum by
Presl (1843). Whereas Vogel (1837) used only fruits in the
description of M. brasiliense and flowers in the description of
M. triste, Presl (1843) described characters from both flowers
and fruits for M. luschnatianum. According to Urban (1906),
Luschnath was in Rio de Janeiro between 1831 and 1834,
and it might be possible that the same specimen was
accessed in different phenological stages. Thus, probably
because of the wide morphological variation in vegetative
characters, Vogel (1837) recognized two different taxa. Fur-
thermore, Presl (1843) described flowers, fruits, and wide
variation of the leaves of M. luschnatianum, based on infor-
mation from distinct materials that had been collected from
the same individual plant.
MACHAERIUM CANTARELLIANUM Hoehne, Arq. Bot. Estado São
Paulo 2(1): 30, Table 29. 1938.—TYPE: BRAZIL. São
Paulo. Municı́pio de São Paulo, Parque e Jardim
Botânico de São Paulo, 20 Jan 1932, F. C. Hoehne s. n.
(lectotype here designated: SP 28731!; isolectotypes: F,
G!, HB!, K!, MBM!, NY!, P!, SPF!, US!, W!).
Notes—Two of five specimens cited by Hoehne (1938) as
Machaerium cantarellianum are specimens of M. stipitatum
( J. G. Kuhlmann s. n. VIC 2225 and J. G. Kuhlmann s. n.
VIC 2233). Specimens of M. cantarellianum were also erro-
neously described by Bentham (1860, 1862), Hoehne (1941)
and Ribeiro and Lima (2007) as M. legale (see notes on
this species).
MACHAERIUM CONDENSATUM Kuhlm. & Hoehne, Arq. Bot.
Estado São Paulo 1: 31, Table 31. 1938.—TYPE: BRAZIL.
Minas Gerais. Municı́pio de Caratinga, Bom Jesus, 30 Jul
1928, J. G. Kuhlmann 56 (holotype: RB!; isotype: SP!).
Machaerium viridipetalum Ducke, Mem. Inst. Oswaldo Cruz
51: 444. 1953.—TYPE: BRAZIL. Pernambuco. Municı́pio
de Recife, Estrada da Aldeia, 28 Aug 1950, D. A. Lima
50–625 (lectotype here designated: IPA!; isolectotypes:
PEUFR!, SP!, SPF!, US!). syn. nov.
MACHAERIUM DEBILE (Vell.) Stellfeld, Tribuna Farm. 12: 131.
1944. Nissolia debilis Vell., Fl. Flumin. 279. 1829 {1825};
Icon. 7: Table 81. 1831 {1827}.—TYPE: BRAZIL. Rio
de Janeiro, “habitat silvis maritimis ad ripas fluvii
Taguahy”. (lectotype: Vellozo, Icon. 81. 1831 {1827}, des-
ignated by Stellfeld 1944). EPITYPE (here designated):
BRAZIL. Rio de Janeiro. Municı́pio do Rio de Janeiro,
Matas da Gávea, 24 Oct 1925, A. Ducke s. n. (RB 2404!;
isoepitypes: K!, P!).
Machaerium violaceum Vogel, Linnaea 11: 186. 1837.—TYPE:
BRAZIL. “Brasilia,” F. Sellow s. n. (holotype: B!; photo
at G!; isotype: K 000530221!; photo at NY 2641!).
syn. nov.
Machaerium eriostemon Benth., J. Linn. Soc. Bot. (4 suppl.): 68.
1860.—TYPE: BRAZIL. Rio de Janeiro. “Habitat in
fruticetis inter Mandioca et Joze Dias,” L. Riedel s. n. (holo-
type: LE; photo at K!, US!). syn. nov.
155
Machaerium dimorphandrum Hoehne, Arq. Bot. Estado São
Paulo 1: 32, Table 33. 1938.—TYPE: BRAZIL. São Paulo.
Municı́pio de São Paulo, Jardim Botânico do Parque do
Estado de S. Paulo, 2 Apr 1936, O. Handro & F. C. Hoehne
s. n. (holotype: SP 35668!; isotypes: K!, US!). syn. nov.
Notes—Rudd (1956) listed Nissolia debilis as the basionym
of Machaerium debile. The scandent habit, morphological var-
iation of leaflets, laxly paniculate inflorescences, and flowers
with tubular calyx and white corolla all agree with the syno-
nyms here presented. The designation of the epitype is
important to reestablish the original concept of N. debile.
MACHAERIUM DECLINATUM (Vell.) Stellfeld, Tribuna Farm. 12:
131. 1944. Nissolia declinata Vell., Fl. Flumin. 296. 1829
{1825}; Icon. 7: Table 77. 1831 {1827}.—TYPE: BRAZIL.
Rio de Janeiro. “habitat silvis maritimis”. (lectotype:
Vellozo, Icon. 77. 1831 {1827}, designated by Stellfeld
1944). EPITYPE (here designated): BRAZIL. Rio de
Janeiro. Silva Jardim, Reserva Biológica de Poço das
Antas, trilha do Morro do Calcário, 9 Jan 1993, H. C. Lima
4586 (RB!; isoepitypes: ESA!, NY!)
Machaerium discolor Vogel, Linnaea 11: 204. 1837 [March].—
TYPE: BRAZIL. “in itinere inter Campos et Victoria,” F.
Sellow s. n. (holotype: B!). syn. nov.
Machaerium lineatum Benth., Comm. Legum. Gen. 34. 1837
[June].—TYPE: BRAZIL. “Brasilia,” M. W. Schott s. n.
(holotype: K 000530182!; photo at NY 2624!; isotypes:
NY 00016232!, W!).
Notes—The designation of the epitype with floral char-
acters was considered appropriate because the lectotype of
the basionym is an illustration with branches, leaflets, and
fruit only.
MACHAERIUM FIRMUM (Vell.) Benth., Comm. Legum. Gen. 37.
1837. Nissolia firma Vell., Fl. flumin. 297. 1829 {1825};
Icon. 7: Table 83. 1831 {1827}.—TYPE: BRAZIL. Rio de
Janeiro. “Habitat silvis maritimis Pharmacopolitanis”.
(lectotype: Vellozo Icon. 7: Table 83. 1831 {1827}, desig-
nated by Bentham 1837). EPITYPE (here designated):
BRAZIL. Rio de Janeiro, Martius Herbarium Florae
Brasiliensis 1109 (M! sheet #167/52; isoepitypes: G!, K!,
M! sheet #167/53, P!).
Notes—The designation of the epitype with floral charac-
ters was considered appropriate because the lectotype of the
basionym is an illustration with branches, leaflets, and fruit
only. Complementing the concept of Machaerium firmum
based on more complete type material is important because
this species is difficult to be distinguished from M.
incorruptibile and similar species.
MACHAERIUM GLABRUM Vogel, Linnaea 11: 187. 1837.—TYPE:
BRAZIL. “Bras. Merid.,” F. Sellow 1020 (holotype: B!; photo
at F 2283!; isotype: K 000530245!; photo at NY 2640!).
Machaerium papilisetosum Hoehne, Arq. Bot. Estado São Paulo
1: 34; Table 38. 1938.—TYPE: BRAZIL. Rio de Janeiro,
Parahyba do Sul, 22 Nov 1936, M. Kuhlmann s. n. (lecto-
type here designated: SP 37012!; isolectotypes: SPF!,
US!). syn. nov.
MACHAERIUM INCORRUPTIBILE (Vell.) Benth., Comm. Legum.
Gen. 37. 1837. Nissolia incorruptibilis Vell., Fl. flumin.
156 SYSTEMATIC BOTANY
297. 1829 {1825}; Icon. 7: tab 82. 1831 {1827}.—TYPE:
BRAZIL. Rio de Janeiro. “Habitat silvis mediterraneis
transalpinis”. (lectotype: Vellozo Icon. 7: Table 82. 1831
{1827}, designated by Bentham 1837). EPITYPE (here
designated): BRAZIL. Rio de Janeiro. Municı́pio do Rio
de Janeiro, matas do Horto Florestal, 9 Sep 1947, J. G.
Kuhlmann s. n. (RB 61383!; isoepitype: NY 603936!).
Machaerium mucronatum Vogel, Linnaea 11: 191. 1837.—
TYPE: BRAZIL. “Brasilia,” F. Sellow s. n. (holotype: B,
destroyed; lectotype here designated: K 000530249!;
isolectotypes: BM!, G!).
Machaerium fluminense Rudd, Phytologia 25 (6): 400. 1973.—
TYPE: BRAZIL. Rio de Janeiro. Municı́pio do Rio de
Janeiro, Morro da Viração, 28 Dec 1871, A. F. M. Glaziou
5807 (holotype: P!; isotypes: G!, K!). syn. nov.
Notes—Rudd (1973) indicated Machaerium incorruptibile as
the most similar to M. fluminense, distinguishing it by the
larger leaflets. However, this variation can be observed among
different specimens, probably as a result of different habitats,
as well as in the same specimen, depending on the branch
position sampled. The designation of the epitype with flowers
was considered appropriate because the lectotype of the
basionym is an illustration without floral characters. Improve-
ment of the accuracy of the circumscription of Machaerium
incorruptibile is important, because this species is easily con-
fused with M. firmum, M. fulvovenosum, and M. legale.
MACHAERIUM LANCEOLATUM (Vell.) J. F. Macbr., Field Mus.
Nat. Hist., Bot. Ser. 13 (3–1): 281. 1943. Nissolia lanceolata
Vell., Fl. flumin. 299. 1829 {1825}; Icon. 7: Table 87. 1831
{1827}.—TYPE: BRAZIL. Rio de Janeiro. “Habitat silvis
maritimis Regii Praedii S. Crucis” (lectotype: Vellozo
Icon. 7: Table 87. 1831 {1827}, designated by Macbride
1943). EPITYPE (here designated): BRAZIL. Rio de
Janeiro. Municı́pio do Rio de Janeiro, matas do Horto
Florestal, 28 Jul 1968, D. Sucre et al. 3333 (RB!;
isoepitypes: B!, F, NY!).
Machaerium leiocarpum Vogel, Linnaea 11: 203. 1837
[March].—TYPE: BRAZIL. Rio de Janeiro. “Prope Rio de
Janeiro,” 1836, B. Luschnath s. n. (lectotype here desig-
nated: P 153–170/71!). syn. nov.
Machaerium secundiflorum Mart. ex Benth., Comm. Leg. Gen.
36. 1837 [June].—TYPE: BRAZIL. Rio de Janeiro. “Prope
Rio de Janeiro,” J. E. Pohl s. n. (lectotype here designated:
NY 00016261!; isolectotype: W!).
Machaerium secundiflorum var. laxiflorum Benth., Comm. Leg.
Gen. 36. 1837 [June]. Machaerium secundiflorum var. majus
Benth., Fl. bras. 15 (1b): 247. 1862.—TYPE: BRAZIL. Rio
de Janeiro. “Prope Sebastianopolis in sylvis,” 1839,
Martius Herbarium Florae Brasiliensis 160 (lectotype here
designated: M!; isolectotypes: BM!, BR, E!, FI!, G!, K!, P!,
W!; photo ex K 2642 at NY!, US!). syn. nov.
Machaerium secundiflorum var. minus Benth., Fl. bras. 15 (1b):
247. 1862.—TYPE: BRAZIL. Rio de Janeiro. Municı́pio do
Rio de Janeiro, “Rio de Janeiro, monte Corcovado,” 1839,
A. Guillemin 251 (holotype: K!; isotypes: FI!, G!; photo ex
K 2643 at NY!, US!). syn. nov.
Notes—The combination of Machaerium lanceolatum had
been published for the “Flora of Peru,” but Macbride (1943)
[Volume 39
cited its occurrence in Brazil and expressed doubt about its
presence in Peru. The specimen deposited at P with label
information identical to the protologue and indicated as the
type material of M. secundiflorum var. minus was not included
in the typification presented here because the material actu-
ally represents a specimen of M. paraguariense. The designa-
tion of the epitype was considered appropriate because the
lectotype of the basionym is an illustration. The varietal epi-
thets of M. secundiflorum var. laxiflora, M. secundiflorum var.
major, and M. secundiflorum var. minor were corrected to con-
form to the neuter gender of the genus, according to the
Articles 32.7 and 60.11 of ICBN (McNeill et al. 2006).
MACHAERIUM LEGALE (Vell.) Benth., Comm. Legum. Gen. 37.
1837. Nissolia legalis Vell., Fl. flumin. 297. 1829 {1825}; Icon.
7: Table 84. 1831 {1827}.—TYPE: BRAZIL. Rio de Janeiro.
“Habitat silvis maritimis ad ripas fluvii Taguahy” (lecto-
type: Vellozo Icon. 7: Table 84. 1831 {1827}, designated by
Bentham 1837). EPITYPE (here designated): BRAZIL. Rio
de Janeiro. Municı́pio de Petrópolis, Rodovia Washington
Luis (BR 040) próximo ao Mirante do Cristo, 22 Nov 2008,
H. C. Lima 7014 (RB!; isoepitypes: K!, NY!).
Notes—Bentham (1862), Hoehne (1941), and Ribeiro and
Lima (2007) presented the treatment for Machaerium legale
based on specimens of M. cantarellianum, and because of this,
the designation of the epitype is important to determine the
true circumscription of M. legale. Filardi (2011) provided the
first description of M. legale after “Florae Fluminensis”
(Vellozo 1831). The high commercial value of the wood of
M. legale, commonly called “jacarandá” (a vernacular name
well-known for the highly threatened Dalbergia nigra
Allemão ex Benth.), is certainly the cause of its exploitation,
which is reflected in the scarcity of herbarium materials for
the species.
MACHAERIUM LEUCOPTERUM Vogel, Linnaea 11: 189. 1837.—
TYPE: BRAZIL. “Bras. merid.,” F. Sellow 454 (holotype:
B, destroyed; photo at F 2285!, G!, US!; lectotype here
designated: K!; isolectotypes: BM!, BR; photo ex K 2639
at NY!, US!).
Machaerium pungens Allemão, Trab. Soc. Vell. 56. 1852. nom.
nud.
MACHAERIUM NIGRUM Vogel, Linnaea 11: 188. 1837 [March].—
TYPE: BRAZIL. Rio de Janeiro, 1833, C. Gaudichaud 869
(lectotype: G!, here designated; isolectotypes: P!, US!).
Machaerium velutinum Benth., Comm. Leg. Gen. 36. 1837
[June].—TYPE: BRAZIL. Rio de Janeiro. “Ad Pertinja, et
inter Porto d’Estrela et Mandioca,” 1817–1818, J. C.
Mikan s. n. (lectotype: W!, designated by Rudd 1987;
isolectotypes: F, K!).
Machaerium fruticosum (Vell.) Hoehne, Arq. Bot. Estado
São Paulo 1: 33. Table 35. 1938. Nissolia fruticosa Vell.,
Fl. Flumin. 298. 1829 {1825}; Icon. 7: Table 86. 1831.
{1827}. (nom. illeg.).—TYPE: BRAZIL. Rio de Janeiro.
“Frequentissme habitat fruticetis maritimis” (lectotype:
Vellozo, Icon. 7: Table 86. 1831. {1827}, designated by
Lima 1995). syn. nov.
Notes—The illustration of branches, leaflets, and fruits,
as well as the habit varying from lianas to small trees, and
the geographical distribution presented by Vellozo (1831) for
2014] FILARDI AND LIMA: MACHAERIUM IN THE ATLANTIC FOREST
Nissolia fruticosa match well with the overall morphology
and distributional range of Machaerium nigrum. However,
N. fruticosa sensu Vellozo (1831) is an illegitimate name,
since it is a homonym of N. fruticosa Jacq. (1760). The com-
bination M. fruticosum (Vell.) Hoehne was validly published
(Hoehne 1938), but according to the Article 58.1 of ICBN
(McNeill et al. 2006), its priority is not retroactive to the
publication of the illegitimate basionym N. fruticosa Vell.
Thus, M. nigrum should be the accepted name.
MACHAERIUM OBLONGIFOLIUM Vogel, Linnaea 11: 184. 1837.—
TYPE: BRAZIL. “Bras. merid.,” F. Sellow s. n. (holotype:
B, destroyed; lectotype here designated: K 000530165!
isolectotypes: E!, W!; photo ex K 2646 at NY!, US!).
Machaerium heterophyllum Presl., Abh. K. Bohm. Ges. Wiss.,
5 (3): 61. 1843.—TYPE: BRAZIL. Rio de Janeiro. “habitat
in Brasilia ad Rio de Janeiro,” Lhotsky s. n. (holotype:
probably PR), ex descr.
MACHAERIUM PARAGUARIENSE Hassl., Bull. Herb. Boissier,
ser. 2, 7: 358. 1907.—TYPE: PARAGUAY. “in regione
fluminis Yhú,” Nov 1905, E. Hassler 9617 (lectotype
here designated: G!; isolectotypes: K!, MO, P!, W!;
photo ex G at F 28179!, US!).
Machaerium paraguariense var. cuspidatum Hassl. ex Tamayo,
Darwiniana 7: 126. 1945.—TYPE: PARAGUAY. “Prope
San Bernardino,” Jan 1913, E. Hassler 12408a (holotype:
G!; isotypes: E!, K!, NY!, US!). syn. nov.
Notes—The variation of the apex of the leaflets is con-
tinuous throughout the species distribution, justifying the
synonymy of the variety cuspidatum. The type materials
referred above have glabrous leaflets, however individ-
uals with leaflets varying from villous to tomentose at
abaxial surface also occurs along the distribution of
Machaerium paraguariense.
MACHAERIUM PEDICELLATUM Vogel, Linnaea 11: 202. 1837.—
TYPE: BRAZIL. Rio de Janeiro. “Rio Jan. prop.
Corcovado, c. fl. mense Novembr.,” Nov 1832, A. L. P.
Silva Manso & J. Lhotzky 52 (holotype: B, destroyed;
lectotype here designated: G!; photo ex B at F
2286!, NY!).
Machaerium scleroxylum Allem., Trab. Soc. Vell. 56. 1852.
nom. nud.
Machaerium allemani Benth., J. Linn. Soc. Bot. (4 suppl.): 63.
1860.—TYPE: BRAZIL. Rio de Janeiro. “Habitat in prov.
Rio de Janeiro in sylvis. Jacarandá tan,” F. Freire Allemão
s. n. (holotype: BM!). syn. nov.
MACHAERIUM PUNCTATUM (Poir.) Pers., Syn. Pl. 2 (2): 276. 1807.
Nissolia punctata Poir., Lam. III, Icon. 600: Table 1. 1794.
Encyclopédie Méthodique Botanique 4: 492. 1797.
Nissolia stipitata DC., Prodr. 2: 258. 1825. (nom. illeg.).—
TYPE: BRAZIL. Rio de Janeiro. “Brasilia circa Rio de
Janeiro, caule sarmentoso, legumine longo stipitato, ala
subpunctata,” Jul 1767, P. Commerson s. n. (holotype: P!;
isotypes: FI!, G!, Herbier Lamarck P00296584!).
Machaerium verrucosum Vogel, Linnaea 11: 183. 1837
[March].—TYPE: BRAZIL: “legit inter Campos et
Victoria,” F. Sellow s. n. (holotype: B, destroyed; lectotype
here designated: K 000530230!). syn. nov.
157
Machaerium puberulum Mart. ex Benth., Comm. Legum. Gen.
35. 1837 [June].—TYPE: BRAZIL. “In sylvis prope
Bahia,” C. F. P. Martius s. n. (holotype: M!). syn. nov.
Machaerium villosulum Mart., Flora 20 (2): Beibl. 118. 1837
[August]. Machaerium oblongifolium var. villosulum Benth.,
Fl. bras. 15 (1): 250. 1862.—TYPE: BRAZIL. Rio de Janeiro.
“Prope praedium Donna rosa, in serra da Broca, prov. Rio
de Janeiro, fl. Jul. Dryas,” Jun 1837, Martius Herbarium
Florae Brasiliensis 159 (holotype: BR; isotypes: G!, K!, M!,
P!, W!; photo ex K 2647 at NY!). syn. nov.
Machaerium oblongifolium var. subcordatum Benth., Fl. bras.
15 (1): 250. 1862.—TYPE: BRAZIL. Bahia: “lecto in silvis
prov. Bahiensis ad Soteropolia,” C. F. P. Martius s. n.
(lectotype here designated: M!). syn. nov.
Notes—Nissolia punctata was published by Poiret (1797), and
Persson (1807) presented it together with two other species of
Nissolia to establish the genus Machaerium; both authors cited a
specimen collected by Philibert Commerson in Madagascar.
However, Brazil and the city of Rio de Janeiro are clear on the
label of the holotype (P), as is the year 1767. According to
Urban (1906), Commerson was in Rio de Janeiro in that year,
and only afterward, from 1770–1773, he went to Madagascar.
The label of Herbier Lamarck isotype is a reproduction of the
prologue, and because of this it has Madagascar. However, a
note questioning this location was added to the label by
Lamarck (Dr. Aymonin pers. comm., May 2010). De Candolle
(1825) relegated Machaerium to a section of Nissolia, and he did
not agree with the epithet of N. punctata in reference to the fruit,
proposing N. stipitata, an illegitimate name and erroneously
treated as a basionym of M. stipitatum (Vogel 1837). The varietal
epithet of M. secundiflorum var. subcordata was corrected to
conform to the neuter gender of the genus, according to the
Articles 32.7 and 60.11 of ICBN (McNeill et al. 2006).
MACHAERIUM RETICULATUM (Poir.) Pers., Syn. Pl. 2 (2): 276. 1807.
Nissolia reticulata Poir., Lam. III, Icon. 600: Table 2. 1794,
Encyclopédie Méthodique Botanique 4: 492. 1797.—
TYPE: BRAZIL. Rio de Janeiro. “caule sarmentose,
foliolis obtusis, leguminibus brevissime stipitatis,” 1767,
P. Commerson s. n. (holotype: P!; isotype: Herbier
Lamarck P00296586!).
Nissolia aculeata Vell., Flumin. 278. 1829 {1825}; Icon. 7:
Table 79. 1831 {1827}.—TYPE: BRAZIL. Rio de Janeiro.
“Habitat silvis Pharmacopolitanis maritimis. Floret Dec.”
(lectotype here designated: Vellozo Icon. 7: Table 79.
1831 {1827}).
Machaerium vellozianum Benth., Comm. Legum. Gen. 34.
1837.—TYPE: BRAZIL. “Brasilia,” M. W. Schott s. n.
(holotype: W 32099!; isotypes: K 000530183!).
Notes—The label of Machaerium reticulatum in P shows
Brazil as the location of Commerson’s collection, whereas
the isotype in Herbier Lamarck indicates Madagascar, but
with no reference of collector0 s name. Bentham (1862) indi-
cated M. vellozianum (synonym) as a common species in Rio
de Janeiro. Thus, we assumed that Commerson made his
collection in Rio de Janeiro, the only Brazilian city that he
visited (Urban 1906).
MACHAERIUM STIPITATUM Vogel, Linnaea 11: 189. 1837.—TYPE:
BRAZIL. Rio de Janeiro. “Bras. merid. Inter Rio de
158 SYSTEMATIC BOTANY
Janeiro et Campos,” 1815, F. Sellow 140 (holotype: B,
destroyed; photo at F 2290!, US!).—NEOTYPE (here des-
ignated): BRAZIL. Rio de Janeiro. Rio de Janeiro, Santa
Teresa, entre Silvestre e Almirante Alexandrino, 3 Aug
1922, A. Ducke & J. G. Kuhlmann s. n. (RB 2256!).
Machaerium minutiflorum Tul., Arch. Mus. Par. 4: 94. 1844.—
TYPE: BRAZIL. Minas Gerais. “Brasilia, Prov. Minas
Gerais, Cachoeiras do Campo,” 1840, P. Claussen 887
(holotype: P!; isotypes: K!, NY!, US!).
Notes—Vogel (1837) proposed Machaerium stipitatum
based on the description of Nissolia stipitata DC. (Prodr. 2:
258. 1825), but stressed differences between the illustration of
N. punctata (Lam. III, Icon. 600: Table 1. 1794) and the speci-
men examined by him (Sellow 140). As discussed above, N.
stipitata is a nomenclatural synonym of M. punctatum, and
probably Vogel did not have the opportunity to see its type
specimen. Therefore, we recognize M. minutiflorum as the
only taxonomic synonym, and the single specimen cited by
Vogel (1837) for the purpose of typification of M. stipitatum.
This type material (B) was destroyed during WWII and like
none isotype was found, and only exist photos was neces-
sary the designation of a neotype. The selected material has
floral characteristics that are important for the recognition
of the species that are added with fruit characteristics in
holotype’s photos.
MACHAERIUM UNCINATUM (Vell.) Benth., Comm. Legum. Gen.
34. 1837. Nissolia uncinata Vell., Fl. Flumin. 295. 1829
{1825}; Icon. 7: Table 76. 1831 {1827}.—TYPE: BRASIL.
Rio de Janeiro. “Habitat fruticetis inundatis Regii Praedii
S. Crucis. Floret sept.”. (lectotype: Vellozo Icon. 7:
Table 76. 1831 {1827}, here designated). EPITYPE (here
designated): BRAZIL. Bahia. Porto Seguro, Reserva
Florestal de Porto Seguro, Companhia Vale do Rio Doce,
11 March 1988, D. A. Folli 692 (CVRD!; isotype: RB!).
Machaerium splendens Vogel, Linnaea 11: 192. 1837.—
TYPE: BRAZIL. “Brasilia,” F. Sellow 5781 (holotype:
B, destroyed; photo at F 2289!, G!).—NEOTYPE (here
designated): BRAZIL. Rio de Janeiro, 1839, A. Guillemin
275 (K!).
Notes—Bentham (1837) has published Machaerium
uncinatum based on Nissolia acuminata and cited the illustra-
tion number 76 published by Vellozo (1831). This illustration
actually belongs to N. uncinata, and thereafter, Bentham
(1860) cited the correct spelling for the basionym of M.
uncinatum. The original illustration of N. uncinata showing
the branches, leaflets, and samaras (Vellozo 1831) was desig-
nated as lectotype, and was also designated an epitype. This
selected material has floral characteristics that are important
for the recognition of the species that are added with fruit
characteristics presented in the illustration. The holotype of
M. splendens deposited in B was destroyed during WWII, and
because of the fact that no isotype was found and only exists
in photos, it was necessary to designate a neotype.
Acknowledgments. This work is part of the first author’s Ph. D.
thesis developed at the Escola Nacional de Botânica Tropical (ENBT) /
Instituto de Pesquisas Jardim Botânico do Rio de Janeiro (JBRJ). The
authors express their gratitude to curators of the herbaria that were vis-
ited; to Maria Alice Rezende for the black-and-white line drawings, to
Erika von Sohsten Medeiros (JBRJ) for high-resolution figures, to Pablo
Pietro Viany (CNCFlora / JBRJ) for his assistance with assessing the
[Volume 39
conservation status of the new species, and to Janet Reid (JWR Associ-
ates) for English revision. We also acknowledge José Eduardo Meireles
(Duke University), Dr. Rafaela Campostrini Forzza (JBRJ), Dr. Jefferson
Prado (Instituto de Botânica de São Paulo), and Dr. Tarciso Filgueiras
(Instituto de Botânica de São Paulo) for valuable discussion of this study,
and especially to Dr. Marli Pires Morim (JBRJ), Dr. Cláudia Franca Barros
(JBRJ), and Dr. Sérgio Ricardo Sodré Cardoso (JBRJ) for suggesting
improvements in the first author’s Ph. D. thesis. The first author extends
her thanks to the Brazilian Research Council (CNPq) for the Doctoral
grant, and also to IAPT for financial resources provided.
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