Phytotaxa 197 (2): 115–124 ISSN 1179-3155 (print edition)

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Article PHYTOTAXA
Copyright © 2015 Magnolia Press ISSN 1179-3163 (online edition)

http://dx.doi.org/10.11646/phytotaxa.197.2.4

A new species of Allagoptera (Arecaceae) from the Cerrado of central Brazil

RENATA CORRÊA MARTINS1, TARCISO S. FILGUEIRAS2, DALVA GRACIANO-RIBEIRO3 & NÁDIA SÍLVIA
SOMAVILLA4
1
Departamento de Botânica, Universidade de Brasília, Brasília, DF, Brazil
2
Instituto de Botânica, São Paulo, SP, Brazil
3
Departamento de Botânica, Universidade Federal de Goiás, GO, Brazil
4
Departamento de Botânica, Universidade Federal de Juiz de Fora, MG, Brazil
1
E-mail: [email protected]

Abstract

A new species of Allagoptera from the Cerrado of central Brazil is described. The new species is distinguished by its solitary
habit, open crown, and very long inflorescence rachis conspicuously curved at maturity. The new species, A. robusta, is
compared morphologically and anatomically with A. campestris and A. leucocalyx.

Key words: anatomy, conservation, Federal District, Goias, Palmae, palms

Introduction

The genus Allagoptera Nees (1821) is distinguished within the subfamily Arecoideae by its spicate inflorescences
bearing congested staminate and pistillate flowers. Allagoptera is highly supported as a monophyletic taxon (Gunn
2004). In addition, it is a sister genus to a clade that includes Attalea Kunth in Humboldt, Bonpland & Kunth
(1815[1816]: 309), Cocos Linnaeus (1735: 1188), Lytocaryum Toledo (1944: 6), and Syagrus Martius (1824: 18–19),
albeit with little support (Dransfield et al. 2008; Meerow et al. 2009). A taxonomic revision of this genus was published
by Moraes (1996), who recognized four species: A. arenaria (Gomes) Kuntze (1891: 726), A. brevicalyx Moraes
(1993: 21), A. campestris (Martius) Kuntze (1891: 726), and A. leucocalyx (Drude) Kuntze (1891: 726). However, in
the Genera Palmarum (Dransfield et al. 2008), Polyandrococos Barbosa Rodrigues (1901: 7) was synonymized under
Allagoptera, adding a fifth species to this genus.
Species in Allagoptera occur on sandy soils in beaches and dunes, open Cerrado areas, grasslands, savannas, and
forest edges, both in Brazil and in Paraguay and Bolivia (Dransfield et al. 2008; Moraes 1996). The estimated number
of species in Brazil ranges from four (Leitman et al. 2012) to five (Lorenzi et al. 2010). Mendonça et al. (2008)
recognized two Cerrado species: A. campestris and A. leucocalyx. In addition to these, a new species was found in the
state of Goiás and Distrito Federal (Martins 2012). This new species (A. robusta) is described in the present paper, and
its morpho-anatomical traits are compared to the traits of two morphologically related species (A. campestris and A.
leucocalyx).

Material & Methods

We used a stereomicroscope at the botany laboratory of the herbarium of Universidade de Brasília, Brazil (UB) for
the morphological analysis. Dissected samples were photographed using a Leica MZ6 digital camera attached to the
stereomicroscope.
For the anatomical study, we sampled individuals of A. robusta (Martins 1131) and A. campestris (Martins 228)
collected in Distrito Federal, Brazil, and of A. leucocalyx (Martins 1093) collected in Cavalcante, Goias, Brazil.
Samples were removed from the middle third of the pinnae at the median portion of the rachis of a mature leaf. Samples
were preserved in 70% ethanol (Jensen 1962) and 25% aqueous glycerin. Cross sections were made free-hand using
a Ranvier microtome and cleared in 30% sodium hypochlorite for five hours and 50% sodium hypochlorite for two
Accepted by William Baker: 13 Nov. 2014; published: 11 Feb. 2015 115
hours (Kraus & Arduin 1997). Double staining was obtained with an ethanol solution of 1:4 safranin and alcian blue
(Luque et al. 1996), both 1%. Slides were dehydrated in an ethyl alcohol series, cleared in butyl acetate, and mounted
with transparent glass varnish (Verniz Vitral Incolor®, Paiva et al. 2006).
The epidermis was studied by removing 1-cm2 samples from the middle third of the pinna. The material was
dissociated using a 50% aqueous solution of nitric acid (Ghouse & Yunus 1972). Samples were rinsed with distilled
water, and the subepidermal residues were removed. Samples were stained with ethanolic safranin and mounted
in transparent glass varnish (Paiva et al. 2006). Photomicrographs were obtained using an Olympus CX31 optical
microscope and an Olympus C-7070 Wide Zoom digital camera. Vascular bundles were classified following Ellis
(1976).
The anatomical analysis was supplemented by observations of micromorphological traits. Samples were dehydrated
in a graded acetone series, and sonicated for 20 minutes after each step of the series to help remove the epicuticular
wax. Samples were critical-point dried, mounted on stubs, coated with gold, and examined with a JEOL JSM-7001F
scanning electron microscope.
Phenological data were obtained from specimens deposited at the herbaria of Cenargen (CEN), Reserva Ecológica
do IBGE (IBGE), Jardim Botânico de Brasília (HEPH), Universidade Federal de Goiás (UFG) and Royal Botanic
Gardens, Kew (K). Uses were observed in the field. The vegetation classification is according to Ribeiro & Walter
(2008).

Taxonomy

Allagoptera robusta R.C.Martins & Filg. sp. nov. (Figs. 1, 2B, 3–5, 6C, 7C)

Species nova affinis Allagoptera campestris (Martius) Kuntze pinnis aggregatis, discoloribus, rigidis, inflorescentia spiciformi, fructibus
congestis, cellulis epidermidis faciei adaxialis 1-seriatis, parenchymate paliformi ad 6 cellulas praedito, fascibus vascularibus
in 3 magnitudine, sed habitu solitario (versus habitu caespitoso), corona foliorum aperta (versus corona foliorum clausa),
inflorescentiae rachis curvata in maturitate, 16–33 cm longa (versus inflorescentiae rachis recta in maturitate, 4.8–15 cm longa),
hypodermidi faciei abaxialis 2–seriata (versus hypodermidi faciei abaxialis 1–seriata), fibris in mesophyllo serie continua in facie
adaxiali et intercepta in facie abaxiali (versus fibris in mesophyllo serie intercepta in ambabus faciebus), fascibus vascularibus
regulariter dispositis (versus fascibus vascularibus irregulariter dispositis) differt.

Type:—BRAZIL. Distrito Federal: Brasília, Park Way, Quadra 21, Conjunto 2, 15.533ºS, 47.557ºW, 1054 m, March 2011 (fl, fr), R.C.
Martins 1131 (holotype UB!)

Acaulescent to short-stemmed palm, solitary, 0.25–1.5 m tall. Stem 0–25 × 15 cm. Leaves 3–6, 100–180 cm long;
sheath 9–30 × 4.5 cm long; petiole 18.5–46 × 0.8–1.4 cm; rachis 68.5–98 cm long; pinnae (16–)48–76 (per side),
inserted in groups of 2–8 in different planes; basal 17–41 × 0.2–1.5 cm, median 15–39.5 × 1–1.6 cm, apical 3.1–31 ×
0.2–1.1 cm. Prophyll 15–26 × 1.5–3 cm. Peduncular bract 23–60 cm long, rostrum 1–5 cm long, 0.2–0.65 cm thick;
inflorescence spicate, 16–70 cm long; peduncle 1.7–25 × 1–2.5 cm; rachis 16–33.5 cm long (9–14.5 cm pistillate
portion + 5–7.5 cm staminate portion); rachillae null. Staminate flowers 0.8–1.5 × 1.5 cm; sepals joined at base,
free at apex, margin entire or slightly ciliate on apex, 0.7–1 × 0.1–0.5 cm; petals free, margin entire, 0.5–2 × 0.35–
0.6 cm; stamens (7–)8–10(–11), 0.4–0.5 cm long, anthers straight, 0.2–0.45 cm long, filaments 0.2 cm long, pistillode
inconspicuous, 1–5 lobed. Pistillate flowers 0.7–1.9 × 0.6–0.9 cm; sepals free, margin entire to irregular, 0.7–0.9 ×
0.45–0.65 cm; petals free, margin entire to slightly ciliate, 0.7–0.9 × 0.65 cm; stigma trifid ca. 0.3 cm long; staminodal
ring inconspicuous, 0.1 cm tall. Fruit obovoid, yellowish-green to orangish-green, 2.5–2.8 × 1.2–1.3 cm; seed 1.
Specimens examined:—BRAZIL. Distrito Federal: DF-100, May 1992. Dias 98 (CEN); Ecopousada Terraviva,
15.404ºS, 48.115ºW, 1117 m, April 2008, Proença & Zanatta 3517 (UB); Estação Ecológica do Jardim Botânico
de Brasília, 15.521ºS, 47.507ºW, 1092 m, April 2011, Martins 1137, 1138 (UB); Fazenda Água Limpa, 15.455ºS,
47.523ºW, 1086 m, February 1999, Martins 225 (UB); May 1989, Scariot 549, 550 (CEN); February 1987, Proença
683 (UB); November 1982, Ratter et al. 4780 (K, UB); October 1982, Ratter et al. 4712 (UB); Park Way (Q.21),
15.533ºS, 47.557ºW, 1054 m, March 1999, Martins 223, 240 (UB); Park Way (Q.18), 15.536ºS, 47.568ºW, 1062 m,
September 2010, Martins 1094 (UB). Goiás: Alto Paraíso, 14.075ºS, 47.303ºW, 1186 m, December 1965, Irwin et al.
9366 (IBGE); Luziânia, 16.150ºS, 47.571ºW, 930 m, September 1978, Brandão 2098 (PAMG); Mineiros, 17.341ºS
52.330ºW, 750 m, May 1992, Noblick 4871 (K, UFG).

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FIGURE 1. Allagoptera robusta. A. Habit; B. Foliar rachis and insertion of pinnae; C. Pinna apex; D. Inflorescence with open staminate
flowers; E. Pistillate flower; F. Sepals of pistillate flower; G. Petals of pistillate flower; H. Gynoecium with trifid stigma; I. Staminate
flower; J. Calyx of staminate flower; K. Corolla of staminate flower; L. Androecium; M. Fruit. (Martins 1131). Illustrator: Renato
Moraes.

ALLAGOPTERA (ARECACEAE) Phytotaxa 197 (2) © 2015 Magnolia Press • 117

FIGURE 2. A. Allagoptera campestris (closed crown); B. Allagoptera robusta (open crown).

Distribution and habitat:—Three populations were located in the state of Goiás (Alto Paraíso, Luziânia and
Mineiros). The species is also found in preserved and remnant Cerrado areas of the Distrito Federal. In shaded
environments, the stem can grow to 1 m tall, with a decumbent habit. In sunny environments, however, the stem
remains almost completely underground.
Etymology:—The specific epithet refers to the robust appearance of this plant in the field.
Vernacular name:—(proposed here) licuri solitário.
Phenology:—Collected with flowers and fruits in the months of February, March, May, June, August, September,
and November. It is important to note that flowers and fruits occur concomitantly.
Uses:—The sweet flesh of the mature fruit and the seeds may be eaten in natura. In the Park Way region of the Distrito

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Federal, the vegetation in public spaces near buildings is frequently removed. However, a few A. robusta individuals
are incidentally preserved in these locations, rendering the plant visually conspicuous and highlighting its ornamental
value. The discolorous leaves confer special aesthetic appeal to this plant, contributing to its potential use in regional
landscaping projects.

FIGURE 3. Habit and inflorescence of A.robusta (Martins 1131).

FIGURE 4. Infructescence of A. robusta (Martins 1131).

Comments:—Allagoptera robusta can be distinguished by the solitary habit, open crown, and inflorescence
rachis conspicuously curved at maturity. The typical robust appearance of this species can be observed in the group of
structures near the ground: leaf sheaths, prophylls, peduncular bracts, and peduncles.
The species was found in four localities in the Distrito Federal (DF) besides the municipalities in the state of Goiás
(Alto Paraíso, Luziânia, and Mineiros), in Cerrado sensu stricto and campo sujo and as a remnant in deforested areas.

ALLAGOPTERA (ARECACEAE) Phytotaxa 197 (2) © 2015 Magnolia Press • 119

Allagoptera robusta is similar to A. campestris in having rigid pinnae in different planes, but differs in its solitary
habit (vs. caespitose in A. campestris), open crown (vs. closed crown) (Fig. 2), ramenta absent (vs. present), tomentose
peduncle (vs. glabrous peduncle), inflorescence rachis 16–33.5 cm long (vs. 4.8–15 cm) and petals of pistillate flowers
7–9 × 6.5 mm (vs. 4–6 × 3 mm). In Distrito Federal and northeastern Goiás, A. robusta and A. campestris occur
sympatrically in Cerrado sensu stricto and campo sujo. Their underground stems represent a remarkable adaptation
against occasional fires in the Cerrado.

FIGURE 5. Inflorescence and infructescence of A. robusta (A and B: Scariot 550; C and D: Martins 1131).

In the past, the new species has been confused with A. leucocalyx, both in herbarium specimens and in the
literature (Moraes 1996, Martins & Filgueiras 2006). Allagoptera robusta differs from A. leucocalyx by its solitary
habit (vs. cespitose in A. leucocalyx), ramenta absent (vs. present) and its occurence in gallery forests (vs. cerrado s.s).
A comparison of related species is presented in Table 1.
The anatomical description of A. robusta with A. campestris and A. leucocalyx revealed that the three species
were similar in many aspects. The three of them have a thick cuticle, uniseriate epidermis with common triangular
or rectangular epidermal cells on the adaxial surface and square cells on the abaxial surface, stomata with internal
and external stomatal ridges, abundant on the abaxial surface and rare on the adaxial one (Fig. 6B). In addition, the
hypodermis occurs on both sides of the foliar pinnae, formed primarily by parenchyma cells with cellulosic walls, and
a few with lignified walls (Fig. 6), and the mesophyll is homogeneous and formed by palisade parenchyma (Fig. 6A).

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The vascular bundles present up to three orders of branches, with sclerenchyma extensions in the sheath of the only
on the abaxial surface of third-order bundles, and on both surfaces in second- and first-order bundles (Fig. 6); in this
region there are a few stegmata cells containing druse-like silica bodies (Fig. 6B). In all three species, the main vein is
prominent on the adaxial surface, and the vascular system is surrounded by fibers (Fig. 7). The epidermal cells of the
abaxial surface are papillose (Fig. 7), and the vascular bundles are collateral, with phloem separated by parenchyma or
sclerenchyma. Hypodermal cells elongated in the longitudinal axis occur at the junction of the blade with the main vein
of the abaxial surface, forming an expansion tissue with two layers (Fig. 7A,B,C), as described by Tomlinson (1990).
Despite the above descriptions show many similarities among the species, the anatomical traits employed for the
differentiation of this species are exposed in the Table 2. The analysis of the epidermis under SEM revealed the
presence of trichomes on the adaxial surface of A. campestris and A. leucocalyx, while no trichomes were observed in
A. robusta.

TABLE 1. Morphological differences among A. campestris, A. leucocalyx and A. robusta.

A. campestris A. leucocalyx A. robusta
Habit Caespitose Caespitose Solitary
Ramentas over the main vein on the Absent Present Absent
abaxial surface
Inflorescence rachis 4,8–15 cm 3,5–19 cm 16–33,5 cm
Peduncle 10–54,5 x 0,3–1,8 cm 46–100 x 0,4–0,9 cm 1,7–25 x 1–2,5 cm
Habitat Cerrado s.s. Gallery forest Cerrado s.s.

TABLE 2. Anatomical traits of the leaves of three species of Allagoptera (A. campestris, A. leucocalyx and A. robusta).
A. campestris A. leucocalyx A. robusta
Cross section
Trichomes Present Present Absent
(adaxial surface) (adaxial surface)
Adaxial hypodermis Biseriate, the first one continuous Uniseriate continuous and Uniseriate continuous and
and the second one interrupted biseriate in the region of biseriate in the region of
by fibers and sclerenchymatous vascular bundles vascular bundles
extensions.
Abaxial hypodermis Discontinuous near stomata, Interrupted near stomata Discontinuous near stomata
bundles, and extensions and biseriate in bundles
Mesophyll (parenchyma) Unifacial with (5–)6–7 rows of Unifacial with 3–5 rows of Unifacial with 5–6 rows of
cells; lacuna absent cells; lacuna present cells; lacuna absent
Fibers In both surfaces, in the adaxial Only in adaxial surface, isolated In both surfaces, in the adaxial
one or more discontinuous rows; or in small groups one or more continuous
in the abaxial in chains rows, interrupted around the
extensions; in the abaxial,
isolated or in small groups
Number of vascular Up to 8 second-order Up to 2 second-order and Up to 4 second-order and 4
bundles between two Up to 4 third-order 8–9 third-order third-order
consecutive first-order
bundles
Distribution of second- and Second-order bundle flanked by Second-order bundle flanked by Second-order bundle flanked
third-order bundles 0–3 third-order bundles 3 third-order bundles predominantly by third-order
bundles
Shape of the central vein Truncate and ovoid Apiculate and elliptical- Slightly “pointed”
and vascular system irregular and ovate
Paradermal section
Types of stomata Tetracytic and anomocytic Tetracytic Tetracytic

ALLAGOPTERA (ARECACEAE) Phytotaxa 197 (2) © 2015 Magnolia Press • 121

FIGURE 6. Cross-section of pinna (blade region). A. Allagoptera campestris (Martins 228); B. Allagoptera leucocalyx (Martins 1093);
C. Allagoptera robusta (Martins 1131). Scale bars: A. 100μm, B. 50 μm, C. 100 μm. Legends: fi: fibras, ade: adaxial face epidermis, abe:
abaxial face epidermis, sb: silica bodies, st: stomata, ext: extension of sheath vascular, hp: hypodermis, sr: stomatal ridges, vb1 (1st order
vascular bundle), vb2 (2nd order vascular bundle), vb 3 (3rd order vascular bundle).

Some cells of the hypodermis of the central vein of Allagoptera robusta and A. campestris have thickened and
lignified walls, forming a uniseriate or biseriate layer, which may also be triseriate in A. campestris (Fig. 7A). On the
abaxial surface of A. leucocalyx, there is a ramenta with up to six rows of cells with a wall that is thickened at the base,
converging toward a row of cells with thin walls (Fig. 7B).

Identification key for three species of Allagoptera studied.

1. Pinnae flexuous, with ramentas above midrib on the abaxial surface; in gallery forest .................................................. A. leucocalyx
1’. Pinnae rigid, without ramentas on the midrib on the abaxial surface; in cerrado sensu stricto .........................................................2
2. Adult plant caespitose, closed canopy; congested pinnae; rachis of the inflorescence <15 cm.....................................A. campestris
2’. Adult plant solitary, open canopy; lax pinnae; rachis of the inflorescence >16 cm .............................................. A. robusta sp. nov.

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FIGURE 7. Cross-section of pinna (region of the main vein). A. Allagoptera campestris (Martins 228); B. Allagoptera leucocalyx
(Martins 1093); C. Allagoptera robusta (Martins 1131). Scale bars: A. 100μm, B. 200 μm, C. 100 μm. Legends: et: expansion tissue, pc:
papillose cells, ra: ramenta.

ALLAGOPTERA (ARECACEAE) Phytotaxa 197 (2) © 2015 Magnolia Press • 123

Acknowledgments:—To Renato Moraes for the illustration in Figure 1. To Daiane Veras for technical support at the
Plant Anatomy Laboratory of Universidade de Brasília. We thank two anonymous reviewers for constructive comments
on the manuscript. The Electronic Microscopy laboratory of the University of Brasilia and PhD Silvia Fernandes
helped with the handling of equipment and preparation of the samples.

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