TRABAJOS ORIGINALES

 

The genus Astrocaryum (Arecaceae)

El género Astrocaryum (Arecaceae)

 

Francis Kahn¹

1 IRD, UMR-DIAPC, Casilla 18-1209, Lima – Peru. Email: francis.kahn@ird.fr

Trabajo presentado al Simposio Internacional “Las palmeras en el marco de la investigación para el desarrollo en América del Sur”, del 07 al 09 de Noviembre 2007, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Perú.

 

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Abstract

The palm genus Astrocaryum with 40 species is common in tropical South America extending northwards reaching Central America and Trinidad. Twenty-six species grow in Brazil, 14 in Peru, 11 in Colombia, 9 in Guyana, 9 in Suriname, 8 in Bolivia, 8 in French Guiana, 6 in Venezuela, 4 in Ecuador, 2 in Costa Rica, 2 in Panama, and 1 in Trinidad. The genus includes solitary or caespitose species in the following classes: (i) large palms with tall stem, (ii) palms with large leaves and medium-sized, or short, or subterranean stem, (iii) slender to medium-sized palms, (iv) acaulescent palms with very short leaves. Although most Astrocaryum species are used by humans, only a few may have promising economic potential and are significantly important in the local and regional trade. In this article, I propose a new taxonomic classification based on characters of the fruit, flowers and vegetative parts. The genus is divided in three subgenera: (i) Munbaca with two sections, Munbaca and Mumbacusu, each with 2 species; (ii) Astrocaryum with two sections, Euchambira (new section with 1 species) and Astrocaryum with three subsections — Astrocaryum (9 species), Acaulia (5 species) and Perstaminata (new subsection with 1 species); (iii) Monogynanthus with four sections: Monogynanthus (with 3 species), Ayri (1 species), Guatinajo (new section with 1 species) and Huicungo (new section) that includes three subsections — Huicungo (7 species); Sachacungo (new subsection with 5 species), and Murumuru (3 species). A synoptic review of the genus is presented herein, including descriptions and illustrations as well as data on distribution, habit, ecology and common names for each species. An identification key to all species is also supplied.

Keywords: Astrocaryum, Arecaceae, taxonomy, distribution, ecology, uses, economic potential.

 

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Resumen

El género Astrocaryum con 40 especies es común en las regiones intertropicales de América del Sur y se extiende al norte hasta América central y Trinidad. Son 26 las especies que crecen en Brasil, 14 se encuentran en Perú, 11 en Colombia, 9 en Guyana, 9 en Suriname, 8 en Bolivia, 8 en Guiana francesa, 6 en Venezuela, 4 en Ecuador, 2 en Costa Rica y Panama, 1 en Trinidad. El género produce palmeras solitarias o cespitosas en las siguientes categorias: (i) palmeras grandes de tallo alto, (ii) palmeras de hojas largas y de tallo mediano o corto o subterráneo, (iii) palmeras delgadas a medianas, (iv) palmeras acaulescentes de hojas cortas. La mayoría de las especies son utilizadas por los pobladores de las zonas rurales; sin embargo pocas son las que tienen importancia en los mercados locales y regionales. Se propone aquí una nueva clasificación del género Astrocaryum en tres subgéneros, basada en los caracteres del fruto, de las flores y de las partes vegetativas: (i) Munbaca con dos secciones, Munbaca y Mumbacusu, cada una con 2 especies; (ii) Astrocaryum con 2 secciones, Euchambira (sección nueva con 1 especie) y Astrocaryum con tres subsecciones — Astrocaryum (9 especies), Acaulia (5 especies) y Perstaminata (subsección nueva con 1 especie); (iii) Monogynanthus con cuatro secciones: Monogynanthus (con 3 especies), Ayri (1 especie), Guatinajo (sección nueva con 1 especie), y Huicungo (sección nueva) que incluye tres subsecciones — Huicungo (7 especies), Sachacungo (subsección nueva con 5 especies) y Murumuru (3 especies). Se presenta una tabla sinóptica del género con las referencias de las descripciones e ilustraciones de cada una de las especies, datos sobre su distribución geográfica, forma de vida y ecología, y los nombres vernáculos más comunes. Una clave de identificación de las especies es presentada.

Palabras clave: Astrocaryum, Arecaceae, taxonomía, distribución, ecología, usos, potencial económico.

 

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Introduction

The palm genus Astrocaryum (Arecoideae: Cocoseae: Bactridinae, Uhl and Dransfield, 1987; Dransfield et al., 2005) is commonly found in most tropical ecosystems of South America and in Pacific and Atlantic forests of Central America. It encompasses many life forms, from large palms in the forest canopy to small acaulescent palms hidden in semi-arid shrubby vegetation. Fruits of several species are edible and fiber is extracted from leaves of other species. Moreover all Astrocaryum species are heavily armed with long spines. For all these reasons, these palms cannot escape the eye of the naturalist who traverses South and Central America.

In 1980 it was almost impossible to identify most Astrocaryum species. Existing collections did not provide useful information. Several type specimens were destroyed in the Berlin herbarium during World War II, and Barbosa Rodrigues’ collection had disappeared, probably in a domestic fire (Glassman, 1972). The absence of types was offset, in part, by the excellent descriptions and illustrations published by Martius (1824, 1844), Drude (1881), and Barbosa Rodrigues (1903). Burret (1934) revised the genus and described several new species but did not provide any illustrations. Wessels Boer (1965) treated the indigenous species of Suriname. Granville (1989) described and discussed the distribution of French Guianan species. Several taxa, considered varieties in Henderson (1995) and Henderson et al. (1995), are maintained as valid species in Govaerts and Dransfield (2005).

We have revisited the type localities and collected new material. Once differential characters were identified, we checked their relevance throughout the area of distribution of the species. This fieldwork was the basis for the identification keys. We added to the knowledge of the species and supplemented the description of many of them, re-validating several or justifying synonymy (Kahn, 2001a,b, 2003; Kahn and Granville, 1998; Kahn and Gluchy, 2002). Eight new species have been described. These include two from Colombia, two from Brazil and four from Peru (Galeano-Garcés et al., 1988; Kahn and Millán, 1992; Kahn and Ferreira, 1995). The species of the dry zones of Brazil are still not completely understood and must be further studied.

A new infrageneric classification of Astrocaryum is presented. The genus is divided in three subgenera. Several new sections and subsections are described. A key to species is given.

General data

Number of species and distribution area — The genus Astrocaryum is composed of 40 species distributed in 12 countries (Table 1). It is well represented in Brazil, Peru, Colombia and in the Guianas with 26, 14, 11 and 10 species, respectively. Eight species are endemic to Brazil, 4 to Colombia, and 4 to Peru. Detailed distribution for each species is given under sections and subsections, and recapitulated in Table 2.

 

 

 

 

 

 

 

 

 

 

Habit and Ecology — Species of Astrocaryum produce solitary or caespitose palms in the following classes (quantitative categories are given in Table 2): (i) large palms with tall stem, (ii) palms with large leaves and medium-sized stem, or short stem, or subterranean stem (considered “acaulescent”), (iii) slender to medium-sized palms, (iv) small, acaulescent palms with short leaves.

Palms of the genus Astrocaryum are found in all parts of the Amazon basin. They commonly form dense stands in riparian or swampy forests, as well as in terra firme forests on well-drained clayey soils, fluvial terraces or on sandy soils; some species are also found in savannas located inside the basin or at its periphery in drier condition. Most of those forest species usually tolerate open areas very well (Kahn and Granville, 1992; Jardim and Stewart, 1994). Several species are limited to the eastern valleys of the Andes under Amazonian influence; they do not grow above 1000 m elevation, except A. faranae collected from 130 m in Amazonian lowlands (Rio Moa, Acre, Brazil) to 1650 m in the eastern cordillera of central Peru. Outside the Amazon basin, Astrocaryum species are found in tropical Pacific or Atlantic rain forests and savannas. The species of subsection Acaulia grow in semi-arid cerrado-vegetation in Brazil. Studies on population dynamics and dispersal have been made in Astrocaryum jauari (Piedade, 1984), A. standleyanum (Smythe, 1989; Hoch and Adler, 1997), A. sciophilum (Sist, 1989a,b; Charles-Dominique et al., 2003), A. paramaca (Forget, 1991), A. aculeatum (Nascimento et al., 1997), A. aculeatissimum (Galetti et al., 2006), and Astrocaryum spp. (as A. murumuru, Cintra, 1997; Cintra and Horna, 1997; Beck and Terborgh, 2002). Leaf production was measured in Astrocaryum paramaca and A. sciophilum in the Kabo and Mapane regions of Suriname (Steege, 1983); Granville (1977) described the trapping system of dead leaves by the funnel-shaped crown of these two palms, and Gasc (1986) studied the herpetofauna that lives in Astrocaryum paramaca. Listabarth (1992) studied the reproductive biology of a species of section Huicungo and several other works dealt with the entomofauna associated with various species (Couturier and Kahn, 1989, 1992; Llosa et al.,1990; Delobel et al.,1995; Couturier et al.,1998).

Uses and Economical potential — All parts of Astrocaryum palms are used:

(i) Fruit and kernel — The fruits of Astrocaryum acaule (Barbosa Rodrigues, 1903) and A. jauari (Wallace, 1853) are used as bait for fishing, and the epicarp of A. aculeatum to smoke-cure rubber (Pinheiro and Balick, 1987). Several species have a fleshy mesocarp that is more or less sweet and edible as in Astrocaryum acaule, A. gratum, A. gynacanthum, A. jauari, A. carnosum, A. murumuru, A. ulei and A. urostachys (Fouqué, 1975; Anderson, 1978; Balick, 1988; Boom, 1988; Kahn and Millán, 1992; Millán, 1998; Balslev et al., 2008). Fruits and seeds of various species are reported to be used for oil production in the Amazon region. Existing analyses of fruit fat content show a relatively homogeneous composition among species, with ca. 20% of fat content in the mesocarp, mostly composed of oleic and palmitic acids, and 20-35% of fat content in the endosperm, with a predominance of lauric acid (Coradin and Lleras, 1983; Pesce, 1985; Pinheiro and Balick, 1987, Lleras and Coradin, 1988; Oboh and Oderinde, 1988). Fruits of A. farinosum are used to prepare meal and starch (Barbosa Rodrigues, 1903). Liquid endosperm of unripe fruit is drunk in most species of subgenus Monogynanthus; that of Astrocaryum aculeatum is used as eye drops. Liquid and solid endosperm of Astrocaryum chambira is drunk or eaten in the same way as coconuts (Mejía, 1988, 1992; Kahn and Millán, 1992; Kahn, 1993). Palikur Amerindians use oil extracted from the seed to cure boils and toothache (Grenand et al., 2004). The Apinayé Amerindians of northeastern Brazil use endocarp of Astrocaryum campestre to make beads and ornaments for necklaces (Balick, 1988). This is a common use of Amazonian species (Lévi-Strauss, 1950). The endocarp of Astrocaryum aculeatum and A. jauari is the raw material for the manufacture of rings, earrings and necklaces (Balslev and Barfod, 1987; Kahn and Moussa, 1999). Shamans use halved endocarps of Astrocaryum aculeatum as bowls to offer the potions to the sick; in Manaus, the halved and well-polished endocarps become game pieces for miniature soccer (jogo do pião) in which each piece is propelled with the help of a comb (Kahn and Moussa, 1999). Larvae collected from fruit (and stem) are eaten in Bolivia (Balslev and Moraes, 1989).

Only a few species are significant in the local or regional trade and may have greater economic potential. Such is the case of Astrocaryum aculeatum and A. vulgare and their edible fruits. Prospections were carried out in Brazil with the purpose of gathering and assessing Astrocaryum vulgare (Lima et al., 1986), and a germoplasm collection is still available in Belém (Lima and Costa, 1991). Fruit centesimal composition is analyzed for this species (Cruz et al., 1984). The mesocarp is rich in provitamin A (Cavalcante, 1974); it provides a fatty, mashed pulp that is used to prepare the very popular French Guianan “bouillon d'awara”, traditionally eaten at Easter time (Fouqué, 1975; Kahn, 1997). Fruits of Astrocarym aculeatum are very much appreciated by the inhabitants of Manaus; they are sold downtown in the local markets as well as on the streets. Juice and ice cream are prepared from the pulp. The fruit is one of the traditional components of the regional breakfast, which has become more and more popular since the 1980’s (Kahn and Moussa, 1999). Schroth et al. (2004) studied extractive use, management and in-situ domestication of this species from a small holding located near Rio Preto da Eva in central Amazonia.

(ii) Leaf — Leaflets of several species are commonly used for basketry (Astrocaryum jauari, A. aculeatum, A. vulgare). Young leaves of Astrocaryum farinosum are used by the indigenous people for making hats and for basketry, and large leaves are used for thatching (Barbosa Rodrigues, 1903); leaves of Astrocaryum huaimi are used to make hats (Martius, 1844). Astrocaryum standleyanum is used for basketry in Panamá (Velásquez Runh, 2001). Petioles and rachis of Astrocaryum murumuru have been suggested as an alternative source for producing paper but physical-resistance studies are needed to evaluate the potential of this idea (Rocha and Potiguera, 2007). Fiber is extracted from leaflets of Astrocaryum aculeatum, A. chambira, A. jauari, and A. vulgare in the Amazon basin (Archer and Hooker, 1855; Wheeler, 1970; Schultes, 1977; Pinheiro and Balick, 1987). It is used to make bags, hammocks, and fishing nets (Balslev and Barfod, 1987; Kahn, 1997; Vormisto, 2002); fiber from leaves of Astrocaryum arenarium is used to make rope in central Brazil (Barbosa Rodrigues, 1903). Astrocaryum standleyanum (Borgtoft Pedersen, 1994) and A. chambira (Kahn, 1988; Holm Jensen and Balslev, 1995; Gomez et al., 1996) are locally significant resources for fiber extracted from the leaves; by-products (bags, hammocks) are marketed through craft industry network at regional and national level in Colombia, Ecuador, and Peru.

In northeastern Brazil, leaves of Astrocaryum campestre are used by Apinayé Amerindians to prepare a cure for venereal disease (Balick, 1988).

(iii) Palm heart of most species is edible. Some attempts have been made in Brazil to exploit Astrocaryum jauari for palm heart canning in Barcelos region (Kahn, 1997). Borgtoft Pedersen and Balslev (1990) proposed this species as a potential component of agroforestry systems. Extracts from palmito of several species are said to cure hepatitis and fibers, and used as cataplasm for calming back pains (Balslev et al., 2008).

(iv) Stem of Astrocaryum jauari (Mejía, 1988), A. aculeatum (Balslev and Moraes, 1989) and A. triandrum (Galeano-Garcés et al., 1988) is known to be rot-resistant and serves as building material. Bows and arrow points are made from wood of Astrocaryum aculeatum (Balslev and Moraes, 1989); canes are made from wood of Astrocaryum standleyanum (Bailey, 1933).

(v) Root — Palikur Amerindians prepare a decoction of the roots of Astrocaryum vulgare, which is said to have an effect against furunculosis (Grenand et al., 2004). Extracts from roots of Astrocaryum chambira have an effect on hepatitis, malaria and yellow fever (Balslev et al., 2008)

(vi) Sap of A. ciliatum is said to cure snakebite (La Rotta et al., 1989).

Common names — One or several common names are reported for 38 species (Table 2). Two very rare species do not have common names — Astrocaryum pygmaeum, known only from the type specimen, and A. minus, known from the type specimen and two individual palmtrees.

Taxonomy

Astrocaryum G.Mey. (conserved name) — Meyer (1818:265).

Type species: Astrocaryum aculeatum G.Mey.

Avoira Giseke (1792:53). Name rejected in favor of Astrocaryum. Lectotype: A. vulgaris Giseke (=Astrocaryum sp.), see Cook (1940:299).

Toxophoenix H.W. Schott (in Schreibers, 1822:12). Type: T. aculeatissima H.W. Schott (=Astrocaryum aculeatissimum (Schott) Burret).

Astrocaryum mexicanum and A. alatum are excluded. These two species are treated in the genus Hexopetion (Pintaud et al., 2008). They differ from all the other species of Astrocaryum in having (i) multifold lateral segments in adult palms, (ii) staminate flowers covering the whole rachilla, without a sterile part, (iii) rachilla woolly-white between the flowers, (iv) stigmas much shorter than ovary, and (v) perivascular sclerified sheath continuous (discontinuous in Astrocaryum).

Taxonomic history of Astrocaryum — Martius (1844, 1845) divided the genus Astrocaryum in § I Caudescentia and § II Acaulia. Under Caudescentia he separated the species according to the number of pistillate flowers, either one inserted at the base of the rachilla, or several superposed in the proximal part of the rachilla.

From the characters of the fruit, Drude (1881) divided the genus in four sections: Munbaca, Ayri, Tucuma (with two subsections, Caudescentia and Acaulia), and Malybo. This latter section is inconsistent joining together Bactris humilis and two Astrocaryum species (A. acaule and A. caudescens=A. aculeatum) that clearly belong to section Tucuma. Drude (1889, 1897; see Burret, 1934, footnote p.115) considered two subgenera, Munbaca and Tucuma (this including the sections Ayri, Tucuma and Malybo). Barbosa Rodrigues (1888, 1891, 1902, 1903) reduced the number of sections to three: (i) Leiocarpeae with two subsections, Yauary and Chambira; (ii) Astrocarpeae with two subsections, Mumbaca and Mumbacuçu; and (iii) Acanthocarpeae with two subsections, Ayry and Murumuru.

Burret (1934) recognized two subgenera from the number of pistillate flowers at the base of the rachilla: subgenus Pleiogynanthus with two or more pistillate flowers per rachilla, and subgenus Monogynanthus with only one pistillate flower per rachilla. Monogynanthus is divided in two sections, each of them defined from the fruit (epicarp splitting or not): (i) Munbaca with two subsections, Eumunbaca and Mumbacuçu, these defined by characters of the pistillate flower and fruit, and (ii) Ayri with two subsections, Symphyodon and Hexodon, defined by the staminodes either connected and forming a staminodial ring or reduced to six free teeth. Subsection Symphyodon includes two series, Plicata and Eplicata, according to whether the pinna possesses or lacks prominent secondary nerves.

If the number of pistillate flowers per rachilla easily identifies the subgenera, it is also true that several species of section Ayri, which are supposed to have no more than one pistillate flower per rachilla, may be found with several pistillate flowers. These exceptions to the rule are rather frequent when the species grow in open areas, pastures or riverside.

The classification proposed by Barbosa Rodrigues (1903) in defining three groups from the characteristics of the fruit appears to be the most efficient system (Fig. 1–3). This leads us to consider those three groups at the subgeneric rank: subgenus Astrocaryum with two sections, one of them with three subsections; subgenus Munbaca with two sections; subgenus Monogynanthus with four sections, one of them with three subsections. Burret’s division of section Ayri in two subsections, Symphyodon and Hexodon is abandoned; this latter being treated under synonymy of the genus Hexopetion (Pintaud et al., 2008).

 

 

 

 

 

 

Astrocaryum aculeatum G.Mey., the type species of the genus Astrocaryum, was misinterpreted. Drude (1881) and Burret (1934) treated the species as incertae sedis. Wessels Boer (1965) applied the name A. aculeatum G.Mey. to A. tucuma Mart. putting this latter into synonymy. Kahn and Millán (1992) and Henderson (1995) followed this position. Bernal rediscovered the type specimen (E. Rodschied s.n.) at GOET. It corresponds to the species A. gynancanthum Mart. described later. Barbosa Rodrigues (1903) was right in classifying the species described by Meyer in section Astrocarpeae subsection Mumbaca. The correct name for A. gynacanthum Mart. should be A. aculeatum G.Mey. to comply with the rule of priority. However, Bernal (2008) proposes to conserve the name A. aculeatum for A. tucuma — a position I follow here (see nomenclatural notes under subgenus Astrocaryum and subgenus Munbaca if Bernal’s proposal is rejected). Astrocaryum aculeatum (syn. A. tucuma) is the type of subgenus Astrocaryum, section Astrocaryum and subsection Astrocaryum.

Description of the genus — Solitary or caespitose, acaulescent or stemmed, spiny, pleonanthic, monoecious palms. Stem short to very large, slender to robust, 3–40 cm in diameter, erect, covered with leaf bases, or becoming bare and conspicuously tinged with leaf scars, unarmed or armed with usually black flat spines in rows or groups, pointing in several directions. Leaf from less than 1 m to 8 m long, pinnate, reduplicate; sheath, petiole and rachis covered with a dense indument, armed with large and small spines; petiole very short to long, adaxially channeled near the base, distally ±flattened or angled, abaxially rounded; rachis usually much longer than the petiole, adaxially ±angled, abaxially rounded; leaflets numerous, regularly arranged in one plane or grouped and oriented in several directions, single-fold or secondarily plicate, linear acute, adaxial face dark green and shiny, abaxial face usually with abundant white indument, rarely pilose, margins usually armed with short spines or bristles. Inflorescence branching to one order, interfoliar, erect, arching or pendulous, protogynous; peduncle usually elongated, circular to oval in cross section, densely covered in indument, often heavily armed with spines; prophyll short, bicarinate, fibrous, unarmed or armed with small spines or bristles, hidden in the leaf bases; peduncular bract much exceeding the prophyll, spindle-shaped, often rostrate, splitting longitudinally along the abaxial face, persistent or eroding, usually densely tomentose, heavily armed with spines or unarmed; rachis shorter than the peduncle bearing spirally arranged rachillae, each sustented by a narrow triangular bract; rachillae ca. 10 to numerous, proximal part glabrous, setose or spiny, bearing 1–5 triads, distal part catkinlike, bearing densely packed staminate flowers partially sunken into pits. Staminate flower symmetrical, trimerous, from barely to widely open at anthesis; sepals 3 short, ±triangular, sometimes basally connate; petals 3, much exceeding the sepals, valvate, boat-shaped, straight or reflexed, basally connate; stamens 3–12, anthers small, ±oval-linear, versatile, latrorse; pistillode present, minute, trifid, or absent. Pistillate flower much larger than the staminate ones; calyx subglobose, urn-, vase-, cask- or cup-shaped, or tubular, truncate, tridenticulate or shallowly to deeply tridentate, with limb straight or curved inwards, or pleated horizontally, unarmed or armed with small spines, glabrous, glabrate or tomentose, limb margin sometimes bristly or spiny; corolla shorter or longer than the calyx, or subequal, globose, cup-, urn-, pitcher-, or vase-shaped, or tubular, regular or contracted, truncate, tridenticulate or shallowly to deeply tridentate, with limb straight or curved inwards or outwards, or pleated horizontally, sometimes carinate vertically, tomentose, bearing bristles or spines, these flattened, flexuose, yellowish to brownish, black, limb margin sometimes bristly or spiny; staminodes connate into a ring, this adnate in the corolla or free and membranous, entire, sometimes deeply laciniate, or reduced to 6 small teeth, or absent; pistil tomentose, setose, sometimes armed with whitish to brownish spinules, tricarpellate, triovulate, styles connate, straight or recurved, tomentose, rarely spiny, usually short, sometimes long and trifid, stigmas large, fleshy, papillate, sometimes neatly tongue-shaped. Fruit 1(–2) seeded with apical stigmatic remains, subglobose, obovoid, ellipsoid, top-shaped, pear-shaped, short to long beaked; epicarp brown, yellowish or orange-red, not splitting when ripe, or splitting and spreading to expose the endocarp, tomentose, setulose or glabrous, unarmed or armed with flattened, brown or black, up to 1,5 cm long spines; mesocarp 2­–10 mm thick, fleshy or dry and starchy, ±fibrous; endocarp thick, stony, with flattened black fibers forming a star-like pattern around the germinating pores, these located in the distal part; endosperm homogeneous. Eophyll bifid, usually bristly.

Subgenera, Sections and Subsections

1. Subgenus Astrocaryum

Subgenus Pleiogynanthus Burret (1934:119). Section I Leiocarpae Barb. Rodr. — Barbosa Rodrigues (1888:47, 1891:102, 1902:80, 1903:62).

Type: Astrocaryum aculeatum G.Mey.

Nomenclatural note: If Bernal’s proposal to conserve Astrocaryum aculeatum with conserved type is rejected, the name Pleiogynanthus Burret must be applied to this subgenus. Astrocaryum tucuma Mart. is then a valid name and the type of subgenus Pleiogynanthus Burret, section Pleiogynanthus and subsection Pleiogynanthus.

Epicarp smooth (rarely with indument, scale-like spines, or bristles when unripe), not splitting when ripe; pistillate flowers 2–5 per rachilla.

1.1. Section Astrocaryum

Section Tucuma Drude (1881:367). Section Malybo Drude (1881:368, excluded A. humile).

The fruit characterizes this section. The epicarp is yellowish, orange or red and the perianth is cup-shaped, more or less widened, to wheel-shaped. The staminodial ring is adnate, high or low in the corolla. The petiole in young plant is armed with black to reddish spines. The pinnae are more or less neatly arranged in groups and oriented in several directions, except for Astrocaryum malybo (subsection Perstaminata), whose pinnae are arranged in one plane.

1.1.1. Subsection Astrocaryum

Subsection Caudescentia Drude (1881:368). Subsection Chambira Barb. Rodr. — Barbosa Rodrigues (1902:80, 1903:62). Subsection Yauary Barb.Rodr., pro parte — Barbosa Rodrigues (1902:80, 1903:62).

This subsection brings together solitary or caespitose, medium-sized to large, or subacaulescent, or acaulescent palms, all of them with large leaves (4–8 m long), and two slender species with leaves 2–3,5 m long and a stem up to 3-4 m in height and up to 15 cm in diameter. The pinnae are usually grouped and oriented in several directions. The stem is heavily armed with long spines at the internodes. The subsection includes nine species. Five are found in the Amazon basin (Astrocaryum acaule, A. giganteum, A. jauari, A. vulgare and A. aculeatum, the type of the genus); the latter three reach the Guianas; Astrocaryum aculeatum is also reported from Trinidad (Wessels Boer, 1965). Two species are found in the northwestern part of South America and in Central America (Astrocaryum standleyanum on the Pacific slope and A. confertum on the Atlantic slope). And the two slender species grow in the southern drier periphery of the Amazon basin (Astrocaryum echinatum in Brazil and A. huaimi in Bolivia, Brazil and Peru).

1.1.2. Subsection Acaulia Drude (1881:368).

Type: Astrocaryum campestre Mart.

Subsection Yauary Barb.Rodr., pro parte — Barbosa Rodrigues (1902:80, 1903:62).

This subsection includes five species (Astrocaryum arenarium, A. campestre, A. kewense, A. pygmaeum, and A. weddellii) that grow in the dry regions of Brazil at the southern periphery of the Amazon basin. A. campestre is also found in Bolivia. These five species are small acaulescent palms with very short leaves (usually less than 1,5 m long). Drude (1881) provided a key for identifying Astrocaryum campestre, A. pygmaeum and A. weddellii. Barbosa Rodrigues (1903) listed the differential characters of Astrocaryum weddellii and A. arenarium, respectively. Burret (1934), however, considered Astrocaryum arenarium as a possible synonym of A. weddellii. This group of small acaulescent palms is still poorly known and needs additional study.

1.1.3. Subsection Perstaminata F. Kahn, subsect. nov.

— Pinnis in eadem directione abeuntibus, regulariter dispositis, parte distali rami floriferi staminatis floribus omnino obtecti.

Type: Astrocaryum malybo H.Karst.

This subsection is monotypic. Astrocaryum malybo differs from all the other species of section Astrocaryum in having the pinnae regularly disposed in one plane and in having the distal pistillate flower contiguous with the staminate portion of the rachilla. Astrocaryum malybo is a Colombian species, found in Magdalena, Cauca and Sinu valleys; it reaches the Isthmus of Panama.

The name Perstaminata refers to the distal part of the rachilla, which is entirely covered with staminate flowers.

1.2. Section Euchambira F. Kahn, sect. nov.

— Epicarpio in fructu juveni albido-furfuraceo, nigro-setuloso; demum glabrato, inermi. Perianthio fructifero obconico. Annulo staminodiali altissimo, corollam altitudinae fere aequans.

Type: Astrocaryum chambira Burret.

This section is monotypic. Astrocaryum chambira differs from the other species of subgenus Astrocaryum in having the biggest fruit, with epicarp glabrate when unripe, usually smooth, sometimes sparsely spinulose, greenish to yellowish when ripe, and a distinctive massive obconical perianth, and in having yellowish spines on the petioles of juvenile plants. DNA analysis (AFLP) clearly separates Astrocaryum chambira from the other Amazonian species in subgenus Astrocaryum (Kahn and Second, 1999).

2. Subgenus Munbaca Drude (1889:83, 1897:57)

Type: Astrocaryum gynacanthum Mart.

Section Munbaca Drude (1881:366); Section Astrocarpeae Barb. Rodr. — Barbosa Rodrigues (1888:47, 1891:102, 1902:80, 1903:62).

Nomenclatural note: If Bernal (2008)’s proposal to conserve Astrocaryum aculeatum with conserved type is rejected, the name of subgenus Munbaca Drude and section Munbaca is Astrocaryum. Astrocaryum aculeatum G. Mey. (type specimen E. Rodschied s.n., syn. Astrocaryum gynacanthum) is then the type of genus Astrocaryum, subgenus Astrocaryum and section Astrocaryum. And the name Munbaca Drude will be applied to section Mumbacusu (Barb. Rodr.) F. Kahn.

Epicarp splits and opens at maturity like a flower, showing the yellowish, orange or red mesocarp with the endocarp at the center. This phenomenon never occurs in the other two subgenera. The fruit is prolonged into an elongated rostrum, up to 2 cm long. The pistillate flowers are armed with dense black spines and they are crowded on the rachis that seems to be itself black as a result.

2.1. Section Munbaca

§ Mumbaca Barb. Rodr. (1902:80; 1903:62). § 1. Leiocarpae Drude (1897:57, not Leiocarpeae Barb. Rodr. 1888). Subsection Eumunbaca Burret (1934:139).

This section is distinguished through the following characters: pistillate flower sessile, calyx cup-shaped. It includes two species: Astrocaryum gynacanthum, the type of subgenus Munbaca and section Munbaca, and Astrocaryum minus. The first species, a slender caespitose palm, occurs in all parts of the Amazon basin but less frequently in the northwestern region. The second species, a slender single stemmed palm, was collected only twice: the first time in Brazil, Jutahy river valley, in the western part of the basin (Trail, 1877); the second time, more recently, in French Guiana, on Grand Matoury hill near Cayenne (Kahn and Granville, 1998). The fruit of Astrocaryum minus is still unknown. Morphological characters (Kahn and Granville, 1998) as well as DNA analyses (Kahn and Second, 1999; Pintaud, in prep.) clearly show that this species is closely related to A. gynacanthum.

2.2. Section Mumbacusu (Barb. Rodr.) F. Kahn, stat. nov.

Type: Astrocaryum paramaca Mart.

Subsection Mumbacuçu Barb. Rodr. — Barbosa Rodrigues (1902:80, 1903:62), Burret (1934:141). § 2. Acanthocarpae Drude (1897:57, not Acanthocarpeae Barb. Rodr. 1888).

The section is distinguished through the following characters: pistillate flower pedicelled, calyx vase-shaped, deeply tridentate. It includes two species: Astrocaryum paramaca, an acaulescent palm with large leaves, and A. rodriguesii, a tall, single stemmed palm. Both species grow in terra firme forests in the Guianas. Astrocaryum rodriguesii is also found in the central region of the Amazon basin where it is a medium-sized palm.

3. Subgenus Monogynanthus Burret (1934:139, excluding Sect. Munbaca Drude)

Type: Astrocaryum sciophilum (Miq.) Pulle

Section Ayri Drude (1881:366, pro parte). Section Acanthocarpeae Barb. Rodr. — Barbosa Rodrigues (1888:47, 1891:102, 1902:81, 1903:62). Subsection Symphyodon Burret (1934:143).

Epicarp covered in indument, setose to spiny; usually with only one pistillate flower inserted at the base of the rachilla (2–3 flowers per rachilla can be occasionally found in palms growing in open areas).

3.1. Section Monogynanthus

Series a Plicata Burret (1934:143).

This section is characterized by having remarkable tongue-shaped stigmas. The basal (sterile) part of the rachillae is pilose. Leaf segments are plicate longitudinally (i.e. secondary nerves are prominent); the spines on the petiole are regularly arranged in horizontal or oblique parallel rows. The inflorescence and infructescence are erect. The section includes three gregarious species that form dense stands in the understory of terra firme forests. Astrocaryum sociale, an acaulescent species with large leaves, is common in the central region of the Amazon basin (Brazil, Amazonas and west of Pará). Astrocaryum farinosum, a subacaulescent palm, is found in northern Brazil (Roraima, north of Amazonas and northwest of Pará), in Guyana and in the central region of Suriname. Astrocaryum sciophilum, the type of subgenus Monogynanthus, produces an unarmed stem with sheaths of dead leaves persistent in the upper part under the crown; this palm occurs in Suriname, Guyana, French Guiana and in the neighbouring regions of Brazil, in Pará and Amapá.

3.2. Section Ayri Drude (1881:366, pro parte; reduced to A. ayri Mart.= A. aculeatissimum (Schott) Burret).

This section is monotypic. Burret (1934) treated Astrocaryum aculeatissimum in the series Eplicata (subsection Symphyodon). The middle pinnae are plicate, however. Astrocaryum aculeatissimum differs from the other species of subgenus Monogynanthus in the following characters: sheaths of dead leaves not persistent under the crown, stem heavily armed with dense spines falling with the age, infructescence pendulous, petals of the staminate flower reflexed, pistil with a long style. These differences are important enough to justify its treatment in a section apart. Astrocaryum aculeatissimum grows in the Brazilian Atlantic forest.

3.3. Section Guatinajo F. Kahn, sect. nov.

— Pinnis in nervis secundariis haud plicatis; parte proximali sterili rami floriferi setosa; floribus masculis tristaminibus; staminodiis nullis.

Type: Astrocaryum triandrum G. Galeano, R. Bernal and F. Kahn

This monotypic section is characterized by the number of stamens being reduced to three, and the absence of staminodes. The basal part of the rachillae is covered with yellowish bristles. Leaf segments are not plicate longitudinally. Astrocaryum triandrum is known from the middle Magdalena river valley in Colombia.

“Guatinajo” is a vernacular name for this palm species (Ranghel, 1941).

3.4. Section Huicungo F. Kahn, sect. nov.

— Pinnis in nervis secundariis haud plicatis; parte proximali sterili rami floriferi glabra.

Type: Astrocaryum huicungo Dammer ex Burret

Series b Eplicata Burret (1934:146, excluding A. aculeatissimum).

This section differs from the three others in the basal (sterile) part of the rachilla that is not pilose. Leaf segments are not plicate longitudinally. The subsections are distinguished on the basis of characters of the pistillate flower.

“Huicungo” is the vernacular name given to these species in Peru.

3.4.1. Subsection Huicungo

The calyx of the pistillate flower is armed with flat, flexuose, yellowish, brownish or black spinules or spines. It is shorter or longer than the corolla, or subequal, cask-, cup-, or vase-shaped, or tubular, with limb straight or pleated horizontally, truncate, tridenticulate or tridentate. Subsection Huicungo includes seven Amazonian palms. Three species are endemic to Peru (Astrocaryum carnosum in the upper Huallaga river valley, Astrocaryum huicungo, the type of section Huicungo, in the lower Huallaga river valley and Astrocaryum scopatum in the upper Marañon and lower Cenepa, Morona, Nieva and Santiago river valleys). One species is endemic to Colombia (Astrocaryum ciliatum known from the middle Caquetá river valley and from the Amazonas river valley, near Leticia). Two species grow in Brazil and Peru (Astrocaryum faranae is found from the upper Moa river in Acre, the westernmost region of Brazil, to the upper Huallaga river valley in Peru, and Astrocaryum javarense, found in the lower Jauari river valley in Brazil, extends to the lower Ucayali river valley in the neighbouring Peruvian region). And one species (Astrocaryum ferrugineum), frequent in the central part of the Amazon basin in Brazil (Manaus, Balbina, Borba), extends westwards to Colombia (Leticia).

3.4.2. Subsection Sachacungo F. Kahn, subsect. nov.

— Calyce floris pistillati glabro vel glabrato, urceolato, elongato-urceolato, corolla longiore.

Type: Astrocaryum macrocalyx Burret.

The calyx of the pistillate flower is glabrous or glabrate (with bristles in Astrocaryum gratum and A. urostachys, and spinules in A. perangustatum), longer than the corolla, urn- or vase-shaped, very contracted at the orifice, the limb straight or turned inwards, pleated horizontally (in Astrocaryum macrocalyx, A. perangustatum and A. urostachys), shallowly tridentate or tridenticulate. Subsection Sachacungo includes five species. Astrocaryum gratum is found from Santa Cruz de la Sierra in Bolivia to Madre de Dios in Peru. Astrocaryum macrocalyx is found from Iquitos, Peru, northwards to Colombia. Astrocaryum urostachys is found throughout Amazonian Ecuador; it reaches the neighbouring regions of Colombia, and extends in Peru to the Tigre river valley. Astrocaryum perangustatum, endemic to Peru, grows in the Pozuzo, Palcazu, Pichis and Perene river valleys. Astrocaryum cuatrecasanum is only known from Colombia (Caquetá Department).

“Sachacungo” is a contraction of sacha (that means “-like” in quechua) and huicungo.

3.4.3. Subsection Murumuru Barb. Rodr. — Barbosa Rodrigues (1902:80, 1903:63).

Type: Astrocaryum murumuru Mart.

The calyx of the pistillate flower is glabrous, cup-shaped, shallowly tridentate, shorter than corolla, rarely hardly subequal, never covering it. Subsection Murumuru includes three species: Astrocaryum chonta, found from Santa Cruz de la Sierra in Bolivia to the lower Ucayali river valley in Peru; Astrocaryum ulei, found from north of Bolivia to the southern bank of Rio Solimões in Brazil (Acre, Amazonas, Rondônia), and Astrocaryum murumuru, found in the Guianas and in the northeastern and central regions of the Amazon basin in Brazil (Amapa, Amazonas, Pará, Roraima, Rondônia, it reaches Acre westwards in the southern region).

Key to species

(References with illustrations, as well as data on distribution, habit and ecology are given for each species in Table 2).

1a. Epicarp splitting and spreading to expose the endocarp when ripe; pistillate flower one at the base of the rachilla — 2

1b. Epicarp not splitting when ripe; pistillate flowers one or several at the base of the rachilla — 5

2a. Pistillate flower inserted on the rachilla at some distance (2—5 mm) from the rachis; calyx vase-shaped (resembling a long inverted trunkate cone), deeply tridentate, up to 18 mm long — 3

2b. Pistillate flower sessile, inserted on the rachis or very close to it; calyx cup-shaped, minutely 3-denticulate, up to 9 mm long — 4

3a. Acaulescent palms with large leaves; inflorescence erect; fruit with a crown of black bristles in the distal third — Astrocarym paramaca

3b. Medium-sized to large palms with stem up to 20 m long, more than 12 cm in diameter, internodes spiny, spines falling in old palms; inflorescence pendulous; fruit not bristly — Astrocaryum rodriguesii

4a. Caespitose, with stems less than 10 cm in diameter, internodes up to 5 cm long, spiny; leaves with less than 50 pinnae per side; inflorescence pendulous; pistillate flower 8—11 mm long, calyx and corolla armed with flat, flexuose spines long enough to hide the floral parts; petals of the staminate flower strongly reflexed — Astrocaryum gynacanthum

4b. Solitary, with stem 10—15 cm in diameter, internodes up to 10 cm long, spiny; leaves with more than 50 pinnae per side; inflorescence arching horizontally; pistillate flower 13—20 mm long, the spines not hiding the floral parts completely; petals of the staminate flower slightly reflexed — Astrocaryum minus

5a. Epicarp smooth (rarely with indument, scale-like spines or glabrate when fruit unripe); pistillate flowers 2—5 per rachilla — 6

5b. Epicarp covered in indument, setose or spiny; pistillate flower usually one per rachilla (some rachillae, usually those located at the base of the rachis, are sometimes found with 2—3 flowers in the most vigorous plants) — 21

6a. Epicarp greenish to yellowish; corolla obconical in fruit; staminodial ring almost as high as the corolla; stamens 6; limb of the pistillate corolla turned inwards girdling the style; fruit 5—7 cm long, 4,5—5 cm broad; petiole armed with yellowish spines in young plant — Astrocaryum chambira

6b. Epicarp yellowish, orange or red; corolla cup-shaped to flat in fruit; staminodial ring high or low in the corolla; stamens 6—12; limb of the pistillate corolla not turned inwards, not girdling the style; fruit usually up to 5 cm long (6,5 cm in A. aculeatum), up to 4,5 cm broad; petiole armed with black to reddish spines in young plant — 7

7a. Length of the leaves more than 2,5 m; tall or short-stemmed, subacaulescent or acaulescent palms — 8

7b. Length of the leaves up to 2 m; small, acaulescent palms — 17

8a. Pinnae arranged in one plane; distal pistillate flower contiguous with the staminate part of the rachilla — Astrocaryum malybo

8b. Pinnae pointing in several directions; distal pistillate flower not contiguous with the staminate part of the rachilla — 9

9a. Infructescence pendulous, the peduncle recurved — Astrocaryum standleyanum

9b. Infructescence erect — 10

10a. Epicarp with small scale-like spines — Astrocaryum confertum

10b. Epicarp smooth, without scale-like spines — 11

11a. Length of the leaves up to 3 m; slender to medium-sized palms, stem up to 15 cm in diameter — 12

11b. Length of the leaves more than 4 m; palms with stem more than 15 cm in diameter, or subacaulescent, or acaulescent palms — 13

12a. Pinnae markedly curved downwards; peduncular bract curved downwards, dark rusty tomentose; pistillate flowers up to 3 at the base of the rachilla; fruit oval, 4,2 cm long, 3,5 cm broad; solitary palms — Astrocaryum echinatum

12b. Pinnae straight, or slightly curved; peduncular bract spindle-like, not dark rusty-tomentose; pistillate flowers up to 5 at the base of the rachilla; fruit obovato-subglobose, 3,5—3,9 cm long; caespitose palms — Astrocaryum huaimi

13a. Acaulescent or subacaulescent (to short-stemmed) palms; petiole rusty-red tomentose; rachis up to 30 cm long, bearing up to 70 rachillae up to 20 cm long — 14

13b. Palms with large stems; petiole not rusty-red tomentose; rachis usually longer than 40 cm, bearing more than 90 rachillae usually longer than 25 cm — 15

14a. Acaulescent palms; leaves up to 5 m long; infructescences erect; fruit obvoid slightly asymmetrical — Astrocaryum acaule

14b. Subacaulescent or with a short stem up to 4 m in height, 22 cm in diameter; leaves up to 7 m long; infructescence erect to arching horizontally; fruit obovoid, symmetrical — Astrocaryum giganteum

15a. Stamens 9—12; staminodial ring 2/3—4/5 as long as the corolla; caespitose, riparian palms — Astrocaryum jauari

15b. Stamens 6; staminodial ring up to 1/2 as long as the corolla; solitary or caespitose, not riparian palms — 16

16a. Solitary palms; pistillate flower with calyx vase-shaped, corolla clearly shorter than the calyx; ripe fruit greenish to brownish, 4,5—6,5 cm long, 3,5—4,5 cm broad — Astrocaryum aculeatum

16b. Caespitose palms; pistillate flower with calyx urn-shaped, corolla as long as the calyx; ripe fruit orange to reddish, up to 4 cm long and 3 cm broad — Astrocaryum vulgare

17a. Peduncular bract woolly, rusty-red or dark flesh-coloured; rachis length more than 10 cm — 18

17b. Peduncular bract not woolly; rachis length up to 10 cm — 19

18a. Peduncular bract obtuse, dark flesh-coloured; leaves crisp-like; pinnae long acuminate; leaf rachis rusty-red tomentose, setose and spiny; peduncle of the inflorescence with rusty-red prickly bristles; pistillate flowers 3—4 per rachilla; fruit obovoid, 3,5—4 cm long, 2,5—3 cm broad — Astrocaryum weddellii

18b. Peduncular bract acute, mucronate, curved downwards, rusty-red; leaves not crisp-like; pinnae obliquely acuminate; leaf rachis whitish tomentose, spiny, not setose; peduncle of the inflorescence glabrous to slightly tomentose with sparse spines; pistillate flowers 1—3 per rachilla; fruit globose, 3,5 cm long, 3,3 cm broad — Astrocaryum arenarium

19a. Apical pinnae thread-like; middle pinnae long acuminate; peduncular bract with sparse or dense spines — 20

19b. Apical pinnae not thread-like; middle pinnae obliquely long acuminate; peduncular bract brown-tomentose with dense spines — Astrocaryum kewense

20a. Length of the leaves up to 1 m; peduncular bract covered with velvet, not spiny; rachillae ca. 10, each with 1—2 pistillate flowers —Astrocaryum pygmaeum

20b. Length of the leaves up to 2 m; peduncular bract tomentose with sparse to dense spines; rachillae more than 20, each with 2—4 pistillate flowers — Astrocaryum campestre

21a. Basal part of the rachilla pilose — 22

21b. Basal part of the rachilla glabrous — 26

22a. Stamens 3; staminodes absent; middle pinnae not plicate; fruit armed with up to 15 mm long spines; solitary palms — Astrocaryum triandrum

22b. Stamens 6; staminodes forming a ring; middle pinnae plicate; fruit armed with short or long spines; solitary or caespitose palms — 23

23a. Caespitose, stemmed palms; stem heavily armed at the internodes, spines falling with the age; sheaths of the dead leaves not persistent on the stem; spines on the petiole not arranged in parallel rows; infructescence pendulous; petals of the staminate flower reflexed — Astrocaryum aculeatissimum

23b. Solitary, acaulescent, subacaulescent or stemmed palms; stem not spiny; sheaths of the dead leaves persistent on the stem or only in the upper part under the crown; spines on the petiole regularly arranged in horizontal or oblique parallel rows; infructescence erect; petals of the staminate flower not reflexed — 24

24a. Acaulescent palms; spines on lateral and abaxial sides of the petiole arranged in more or less horizontal parallel rows, those on the abaxial side are longer; inflorescence less than 0,6 m long; pistillate flower 8—13 mm long, calyx shallowly tridentate; epicarp with short spines, less than 5 mm long — Astrocaryum sociale

24b. Subacaulescent or stemmed palms; spines on both lateral sides of the petiole clearly arranged in oblique parallel rows; inflorescence more than 0,6 m long; pistillate flower 12—20 mm long; calyx deeply tridentate; epicarp with short or long spines — 25

25a. Stem up to 12 m high, sheaths of the dead leaves persistent only in the upper part; inflorescence about 1 m long; epicarp with 5—12 mm long spines; staminate flower without an apparent bract — Astrocaryum sciophilum

25b Subacaulescent palms, or with a short stem entirely covered with sheaths of the dead leaves; inflorescence up to 2 m long; epicarp with short spines, up to 5 mm long; staminate flower with an acute 2—3 mm long bract — Astrocaryum farinosum

26a. Calyx of the pistillate flower armed with flat, flexuose, yellowish, brownish or black spinules or spines — 27

26b. Calyx of the pistillate flower glabrous or glabrate — 34

27a. Calyx of the pistillate flower with the limb pleated horizontally — 28

27b. Calyx of the pistillate flower with the limb not pleated horizontally — 29

28a. Caespitose palms; calyx of the pistillate flower with spines persistent; fruit top-shaped to obovate, not remarkably long tapering basally; mesocarp very fleshy when ripe, up to 8 mm thick — Astrocaryum carnosum

28b. Solitary palms; calyx of the pistillate flower not spiny or with spines falling in fruit; fruit top-shaped with a remarkably long tapering, often slightly curved base; mesocarp not very fleshy when ripe, up to 3 mm thick — Astrocaryum perangustatum

29a. Abaxial side of the leaf pilose, with brown to rusty-red hairs — Astrocaryum ferrugineum

29b. Abaxial side of the leaf not pilose — 30

30a. Middle pinnae with 1—2 parallel ribs on each side of the midrib near the margins; corolla of the pistillate flower armed with black spinules forming a dense fringe at the margin, persisting in fruit; acaulescent, solitary palms — Astrocaryum ciliatum

30b. Middle pinnae without parallel prominent ribs; corolla of the pistillate flower without a fringe of black spinules at the margin; subacaulescent to short-stemmed, solitary or caespitose palms — 31

31a. Solitary palms — Astrocaryum javarense

31b. Caespitose palms — 32

32a. Pistillate flowers not very crowded on the rachis, subglobose, calyx cask-shaped — Astrocaryum scopatum

32b. Pistillate flowers crowded on the rachis, much longer than broad, calyx cup-shaped or tubular — 33

33a. Epicarp setose; calyx of the pistillate flower hardly shorter than the corolla or equal — Astrocaryum huicungo

33b. Epicarp armed with spines, up to 15 mm long; calyx of the pistillate flower neatly shorter than the corolla, rarely subequal — Astrocaryum faranae

34a. Calyx of the pistillate flower covering the corolla entirely, very contracted at the orifice, urn- to vase-shaped, glabrous or glabrate — 35

34b. Calyx of the pistillate flower not covering the corolla entirely (calyx shorter than the corolla or hardly subequal), cup-shaped to tubular, glabrous — 39

35a. Calyx of the pistillate flower with the limb pleated horizontally — 36

35b. Calyx of the pistillate flower with the limb straight — Astrocaryum gratum

36a. Fruit top-shaped with a long tapering, slightly curved base; leaf segment satiny beneath — Astrocaryum perangustatum

36b. Fruit top-shaped to sub-globose, not remarkably long tapering basally; leaf segment whitish beneath — 37

37a. Epicarp armed with spines up to 15 mm long — Astrocaryum cuatrecasanum

37b. Epicarp setose or armed with spines up to 7 mm long — 38

38a. Solitary palms; calyx of the pistillate flower glabrous, usually shorter than 14 mm; fruit with 3—4 mm long bristles — Astrocaryum macrocalyx

38b. Caespitose palms; calyx of the pistillate flower glabrate (some bristles are always present), usually longer than 15 mm; fruit with 5—6 mm long bristles or armed with spines — Astrocaryum urostachys

39a. Caespitose palms; infructescence often pendulous; fruit up to 9 cm long, 4,5 cm broad; mesocarp very fleshy when ripe, 6—10 mm thick; corolla of the pistillate flower vase-shaped, or tubular often shallowly contracted at the middle or at the distal third (like a 8), large enough to encompass the proximal part of the stigmas — Astrocaryum murumuru

39b. Solitary palms; infructescence usually erect or arching; fruit up to 6,5 cm long, 3,5 cm broad; mesocarp more or less fleshy when ripe, less than 4 mm thick; corolla of the pistillate flower cask- or pitcher-shaped, or regularly tubular, not or hardly covering the base of the stigmas — 40

40a. Leaf rachis up to 4 m long; palms subacaulescent or with a stem up to 4 m long; calyx of the pistillate flower bone-colored, usually 1/4—1/3 as long as the corolla; corolla cask-shaped; fruit obovate to top-shaped, often asymmetrical, 4—5 cm long, with a pedicel up to 3 cm long — Astrocaryum ulei

40b. Leaf rachis up to 7 m long; palms with a stem up to 15 m long; calyx of the pistillate flower tawny to brown, usually 1/3—4/5 as long as the corolla; corolla pitcher-shaped to tubular; fruit obovate, 4—6,5 cm long, sessile or with a 0,5—2 cm long pedicel — Astrocaryum chonta

Acknowledgements

I am especially indebted to Scott Zona who helped me to solve problems of nomenclature and revised the English version, and to Henrik Balslev, Rodrigo Bernal, Jean-Jacques de Granville and Jean-Christophe Pintaud for their useful comments and suggestions on the manuscript. I thank Betty Millán, Irés Paula de Andrade Miranda, Afonso Rabelo and Harri Lorenzi for providing material of several species.

 

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