A New Species of Hymenaea (Leguminosae: Caesalpinioideae) with a Revised Identification Key to the Genus in the Brazilian Atlantic Forest

Abstract

Abstract—
New species of the caesalpinioid legume genus Hymenaea (Detarieae s. str. clade) have not been discovered in the last 40 yr. This study describes and illustrates the striking new species
Hymenaea fariana
from the few known collections
made in the Atlantic Forest biome of the Brazilian state of Espírito Santo. The new species has close morphological affinities to H. altissima, from which it is readily distinguished by the mainly shrubby to small tree habit and lanceolate to oblanceolate petals with a well-differentiated
petal claw. An updated identification key to the Atlantic Forest species of Hymenaea is also provided.

Full text

Systematic Botany (2015), 40(1): pp. 151–156
©Copyright 2015 by the American Society of Plant Taxonomists
DOI 10.1600/036364415X686440
Date of publication February 12, 2015
A New Species of Hymenaea (Leguminosae: Caesalpinioideae) with a Revised Identification
Key to the Genus in the Brazilian Atlantic Forest
Robson Daumas Ribeiro,
1
Domingos Benı
´cio Oliveira Silva Cardoso,
2,3
and Haroldo Cavalcante de Lima
1,4
1
Instituto de Pesquisas Jardim Bota
ˆnico do Rio de Janeiro, Rua Pacheco Lea
˜o, 915, 22460-030, Rio de Janeiro,
RJ, Brazil.
2
Departamento de Bota
ˆnica, Instituto de Biologia, Universidade Federal da Bahia, Rua Bara
˜o de Jeremoabo,
s/n, Campus Universita
´rio de Ondina, 40171-970, Salvador, BA, Brazil.
3
Programa de Po
´s-Graduac¸a
˜o em Bota
ˆnica, Universidade Estadual de Feira de Santana, Av. Transnordestina,
s/n, Novo Horizonte, 44036-900, Feira de Santana, BA, Brazil.
4
Author for correspondence (hlima@jbrj.gov.br)
Communicating Editor: Timothy M. Evans
Abstract—New species of the caesalpinioid legume genus Hymenaea (Detarieae s. str. clade) have not been discovered in the last 40 yr. This
study describes and illustrates the striking new species Hymenaea fariana from the few known collections made in the Atlantic Forest biome
of the Brazilian state of Espı
´rito Santo. The new species has close morphological affinities to H. altissima, from which it is readily distinguished
by the mainly shrubby to small tree habit and lanceolate to oblanceolate petals with a well-differentiated petal claw. An updated identification
key to the Atlantic Forest species of Hymenaea is also provided.
Resumo—Novas espe
´cies de leguminosas do ge
ˆnero Hymenaea (clado Detarieae s.str., Caesalpinioideae) na
˜ote
ˆm sido descobertas nos
u
´ltimos 40 anos. Neste estudo, descrevemos e ilustramos a nova espe
´cie Hymenaea fariana a partir de poucas coletas oriundas apenas do bioma
Mata Atla
ˆntica no estado do Espı
´rito Santo, Brasil. Esta nova espe
´cie apresenta maior similaridade morfolo
´gica com H. altissima,daqualse
diferencia facilmente pelo ha
´bito geralmente arbustivo e pelas pe
´talas lanceoladas a oblanceoladas e distintamente unguiculadas. E
´tambe
´m
apresentada uma chave de identificac¸a
˜o das espe
´cies de Hymenaea ocorrentes na Mata Atla
ˆntica.
Keywords—Caesalpinioideae, Detarieae, Fabaceae, morphology, restinga costal forest, taxonomy.
Palavras-chave—Caesalpinioideae, Detarieae, Fabaceae, morfologia, restinga, taxonomia.
The genus Hymenaea L. (Leguminosae, Caesalpinioideae)
comprises 15 species (Lee & Langenheim 1975; Mackinder
2005; Souza et al. 2014). Hymenaea is one of a number of
legume genera with an Amphi-Atlantic disjunct distribution,
largely occurring in the Neotropics but with one species in
coastal East Africa, Madagascar, and the Mascarene Islands
(Mackinder 2005; Schrire et al. 2005). Species of Hymenaea can
be small trees, such as the savanna-inhabiting H. stigonocarpa
Mart. ex Hayne, to very large trees in wet forests, such as the
Amazonian H. intermedia Ducke. Hymenaea is easily recog-
nized by the bifoliolate leaves, relatively large, radially sym-
metrical bat-pollinated flowers, and indehiscent pods with a
farinaceous endocarp. Phylogenetic analyses of chloroplast
DNA sequences confirm the monophyly of Hymenaea and
place the genus together with Guibourtia Benn. and Peltogyne
Vogel in a strongly supported Hymenaea clade (Fouge
`re-
Danezan et al. 2007, 2010). This clade is one of several
caesalpinioid radiations of the large resin-producing
Detarieae clade (Fouge
`re-Danezan et al. 2007, 2010; Bruneau
et al. 2008; LPWG 2013).
The diversity of Hymenaea is most geographically concen-
trated in Brazil, where nearly all species occur in almost all
biomes, but mainly in the Atlantic Rain Forest (or Mata
Atla
ˆntica) (Lee and Langenheim 1975; Lima et al. 2010). The
Brazilian Mata Atla
ˆntica originally covered more than
1,000,000 km
2
, but current estimates indicate that deforesta-
tion has severely reduced the biome to less than 8% of its
original vegetation cover (SOS Mata Atla
ˆntica 1998; MMA
2002). Nevertheless, the Mata Atla
ˆntica is still considered
one of the world’s main biodiversity hotspots (e.g. Myers
et al. 2000; Martini et al. 2007), a conclusion supported by
the remarkable number of new species in many different
plant families that have been recently discovered and
described from this biome (e.g. Cardoso and Queiroz 2010;
Fraga and Kollmann 2010; Moraes and van der Werff 2011;
Popovkin et al. 2011; Amaya and Popovkin 2012; Machado
and Vianna-Filho 2012; Mancinelli et al. 2012; Pinto et al. 2012;
Stehmann and Giacomin 2012; Coser et al. 2013; Ferreira et al.
2013; Groppo et al. 2013; Meireles and Lima 2013; Mota et al.
2013; Prado and Hirai 2013; Filardi and Lima 2014; Fraga
et al. 2014; Gil et al. 2014; Goldenberg and Chagas 2014;
Heiden et al. 2014; Riina et al. 2014; Cardoso et al. 2014).
New species of Hymenaea have not been discovered in the
last 40 yr. This study describes and illustrates a striking
new species which came to light as a result of our extensive
fieldwork in the Atlantic Forest of the state of Espı
´rito Santo.
We also present a revised identification key to all known
species of Hymenaea occurring in the Atlantic Forest biome.
Taxonomic Treatment
Hymenaea fariana R. D. Ribeiro, D. B. O. S. Cardoso &
H. C. Lima, sp. nov.—TYPE: BRAZIL. Espı
´rito Santo:
Guarapari, Rodovia do Sol, Setiba, Parque Estadual
Paulo C. Vinha, 205501600S, 402505000 W, 5 April 2007
(fl., fr.), R. D. Ribeiro,H. C. Lima &J. E. Meireles 825
(holotype: RB!; isotypes: CEPEC!, CVRD!, K!, MBM!,
NY!, VIES!).
The new species is similar to H. altissima, but differs in
having a predominantly shrubby or treelet habit and lanceo-
late to oblanceolate and clawed petals. Despite superficial
similarity also to H. rubriflora var. glabra, especially in habit
and fruit characteristics, H. fariana is readily distinguished by
its falcate and non-revolute leaflets, and flowers with white
petals that have a petal claw over 3 mm long.
Shrub to small tree, 1–10 m tall, rarely a large tree to 20 m
tall, branches glabrous, lenticellate, cut branches and trunk
yielding a resiniferous exudate. Leaves bifoliolate, borne on a
151

Fig. 1. Hymenaea fariana. A. Flowering branch. B. Bracteoles (adaxial and abaxial views). C. Floral bud. D. Petal when in floral bud. E. Flower. F. Petal
from a mature flower. G. Gynoecium. H. Fruit [All from the holotype, R. D. Ribeiro et al. 825 (RB)].
152 SYSTEMATIC BOTANY [Volume 40

thin, glabrous, 10–20 mm long petiole; pulvinules 2–5 mm
long, contorted; leaflets 2–6
+
1.5–3 cm, usually broader on
vegetative branches, chartaceous, glabrous, falcate, apex acute
to cuspidate, base oblique, the inner side acute, the outer side
rounded, midvein and secondary veins conspicuous on both
surfaces, the reticulate tertiaries conspicuous beneath. Inflo-
rescence a short corymbose panicle, 2–4 cm long, comprised
of several racemose shoots, these 3–18-flowered, axes and
pedicels covered by brownish, puberulent indumentum;
pedicel 3–5 mm long; bracts and bracteoles 2 – 4 mm long,
ovate, glabrescent, caducous. Flowers 1.5–2.2 cm long, radi-
ally symmetrical; floral buds 10–15 mm long, ochraceous;
calyx with a hypanthium ca. 7 mm long, campanulate,
4-lobed, the lobes 10–15
+
5–8 mm, oblong to ovate,
ochraceous-pubescent outside, golden-villous inside; petals
5, 13–18
+
3– 5 mm, lanceolate or oblanceolate, whitish,
glabrous, prominently clawed, the claw 3–4 mm long;
stamens 10, the filaments 25–30 mm long, white to cream,
anthers 3– 5
+
1– 2 mm; intrastaminal nectariferous disc
present; gynoecium 2.5–3.5 cm long, shortly stipitate, the
stipe 2–3 mm long, ovary oblique to oblong, glabrous, ovules
5– 8, style 2–3 cm long, stigma capitate. Fruit 5 –12
+
4–5 cm,
indehiscent, oblong to ovate, laterally compressed, woody,
sometimes with slight depressions between the seeds, sur-
face greyish brown, slightly opaque, verrucose, with numer-
ous conspicuous resiniferous pockets and minute lenticels,
endocarp farinaceous. Seeds 3–6 per fruit, 2 – 3
+
1– 2 cm,
ellipsoid to obovoid. Figures 1 and 2.
Additional Material Examined—BRAZIL. Espı
´rito Santo: Guarapari,
Rodovia do Sol, Setiba, Parque Estadual Paulo C. Vinha, 5 April 2007 (fl),
R. D. Ribeiro et al. 826 (RB); Vila Velha, Barra do Jucu
´, Parque Municipal
de Jacarenema, 30 Aug 2008 (fr), H. C. de Lima et al. 7004 (RB); Vila Velha a
Guarapari, ca. 25 km na Estrada do Sol, 21 May 1990 (fr), A. M. de Carvalho
3162 (CEPEC, RB); Presidente Kennedy, pro
´ximo a
`Praia das Neves,
propriedade da FERROUS, 14 Jan 2010 (fl), L. Maielo-Silva et al. 97 (RB);
Vito
´ria, 20 Mar 1934 (fl), J. G. Kuhlmann 1 (RB, UEC).
Distribution and Habitat—The new species has been
recorded only in a few localities of the municipalities of
Guarapari, Presidente Kennedy, Vila Velha, and Vito
´ria in
Fig. 2. A. The more common shrubby habit of the new species Hymenaea fariana. B –C. Inflorescences. D. Close-up of the flowers [All from the
holotype, R. D. Ribeiro et al. 825 (RB)]. All photographs by Jose
´Eduardo Meireles.
2015] RIBEIRO ET AL.: NEW SPECIES OF HYMENAEA 153

Espı
´rito Santo state, southeastern Brazil (Fig. 3). It grows in
arboreal or scrub white-sand restinga vegetation, a character-
istic plant formation scattered along the Atlantic coast.
Conservation—Hymenaea fariana is only known from a
few collections, in botanically poorly explored areas. Any
effective conservation assessment of this apparently nar-
rowly restricted endemic species requires a detailed survey
of the many other species-diverse areas of restinga vegeta-
tion in southern Bahia and southeastern coastal Brazil.
We temporarily assess H. fariana as data deficient (DD)
(IUCN 2001).
Phenology—Flowering and fruiting from April to August.
Etymology—The specific epithet of the new species
honours Sergio Miana de Faria, a Brazilian scientist at
Embrapa Agrobiologia in Serope
´dica, Rio de Janeiro, who
has greatly contributed to the study of nitrogen fixation in
nodulating legumes.
Taxonomic Comments—Hymenaea fariana is morphologi-
cally similar to the recently reestablished species Hymenaea
altissima Ducke (Souza et al. 2014), both having falcate, gla-
brous leaflets that are relatively small (up to 6 cm long), small
flower buds (less than 2 cm long), and a glabrous ovary.
However, H. fariana usually grows as a shrub or small tree
and has lanceolate to oblanceolate, clawed petals, with the
claw 3–4 mm long, in mature flowers, and has somewhat
laterally compressed, oblong-ovate pods (versus large trees
up to 40 m tall, ovate to obovate petals with claw absent or
very short, less than 2 mm long, and cylindrical pods in
H. altissima). Additionally, H. fariana is ecologically confined
to sandy coastal restinga vegetation in Espı
´rito Santo,
whereas H. altissima is more widespread in wet forests along
the slopes of the Serra do Mar mountain range in the states
of Rio de Janeiro and Sa
˜o Paulo, as well as in the tabuleiro
coastal moist forests that occur on clay soils from northern
Rio de Janeiro to southern Bahia.
Hymenaea fariana also has some morphological affinity with
H. rubriflora var. glabra, especially the glabrous leaflets and
flowers with oblanceolate petals and a glabrous ovary. The
main differences between these two taxa are the falcate
and non-revolute leaflets and flowers with white petals in
H. fariana (vs. oblong, usually revolute leaflets and flowers
with reddish petals in H. rubriflora var. glabra). Our study of
herbarium collections and of many different populations in
the field also revealed that H. rubriflora var. glabra has its
geographical range limited to the Atlantic coastal forest of
the states of Paraı
´ba and Sergipe, in northeastern Brazil.
The genus Hymenaea is represented in the Brazilian Atlantic
Forest biome by nine species (11 taxa). Phylogenetic analyses
of the resin-producing Detarieae that best sampled within
Hymenaea so far (Fouge
`re-Danezan et al. 2007, 2010) did not
resolve a monophyletic group of Atlantic Forest species.
Some taxa occur in the Atlantic Forest, but they have an
Fig. 3. Geographical distribution of known specimens of Hymenaea fariana. The new species has been recorded only in the state of Espı
´rito Santo,
southeastern Brazil.
154 SYSTEMATIC BOTANY [Volume 40

ecological preference for habitats in Cerrado (savanna) and
Caatinga (seasonally dry woodland) vegetation. We present
below an identification key for all taxa of Hymenaea recorded
from the Atlantic Forest.
Key to Species of HYMENAEA from the Atlantic Forest Biome
1. Leaflets densely pubescent beneath. . . ............................................................................................ 2
2. Flowers with petals distinctly unguiculate (clawed) and spathulate. ................................................... ............. 3
3. Large trees, to 20 m tall, branches with peeling surface; leaflets 5–9 cm long,
golden brown tomentose beneath; ovary evenly tomentose. . . ......................................................... H. aurea
3. Small trees or shrubs, to 5m tall, branches with smooth to rough surface;
leaflets 2– 4 cm long, pubescent beneath; ovary bearing a tuff of hirsute trichomes at the base. ........................... H. eriogyne
2. Flowers with petals not spathulate and without a differentiated claw. . . . ......... .................................................. 4
4. Flowers with reddish, oblanceolate petals. . ............................................................ H. rubriflora var. rubriflora
4. Flowers with whitish, ovate to obovate petals. . ............................................................................. 5
5. Leaflets usually more than 10 cm long; flowers large, ca. 3.5 cm long, the ovary glabrous
or rarely with a few hairs along the margins, or at the base. ...................................... H. stigonocarpa var. pubescens
5. Leaflets less than 10 cm long; flowers less than 2 cm long,
the ovary commonly bearing a tuff of hairs at the base. . . . ...................................................... H. martiana
1. Leaflets glabrous or glabrescent (then indumentum present only along the midvein). . . .................................................. 6
6. Leaflets falcate, the apex usually acuminate. . . ................................................... ............................... 7
7. Mature flowers with lanceolate or oblanceolate and clawed petals, the claw 3 –4 mm long. ................................. H. fariana
7. Mature flowers with ovate to obovate petals, sessile or rarely shortly clawed,
the claw less than 2 mm long. ........................................................................................... 8
8. Leaflets 3–5 cm long, reticulate venation conspicuous beneath; flowers ca. 15 mm long;
pod surface strongly verrucose. ............................................................................. H. altissima
8. Leaflets 4– 10 cm long, reticulate venation inconspicuous beneath; flowers ca. 18 mm long;
pod with inconspicuous resiniferous pockets. . . ............................................................... H. courbaril
6. Leaflets oblong to obovate, the apex usually obtuse. ............................................................................. 9
9. Leaflets usually revolute; flowers with reddish petals. ...................................................... H. rubriflora var. glabra
9. Leaflets never revolute; flowers with white to cream petals. .................................................................. 10
10. Inflorescences largely paniculate, ca. 18 cm long; flowers ca. 15 mm long,
petals spathulate and clawed; pods ovate, opaque, up to 6 cm long, with 1 –3 seeds. . ..... .............. H. oblongifolia var. latifolia
10. Inflorescences shortly paniculate, ca. 8 cm long; flowers ca. 26 mm long, petals obovate,
claw absent; pods oblong, shiny, up to 15 cm long, with 6 –8 seeds. . . ............................ H. stigonocarpa var. stigonocarpa
Acknowledgments. ProjetoBiodiversidadedoBiomaMataAtla
ˆntica/
PROBIO II/MCTI/JBRJ supported all fieldwork that resulted in the dis-
covery of the new species. We thank Rafael B. Pinto and Jorge Fontella
Pereira for helpful comments on earlier drafts of the manuscript; the
curators of the cited herbaria are thanked for kindly permitting access to
their collections; Aline S. Oliveira prepared the line drawing, and Jose
´
Eduardo Meireles helped with fieldwork and kindly provided photo-
graphs of the new species. We also greatly appreciate the careful reading
and suggestions provided by two anonymous reviewers. This study is
wholeheartedly dedicated to the memory of the first author, who tragi-
cally died before he could conclude his master’s dissertation.
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