Chapter 1
The Botany of Anadenanthera
Taxonomic Background
The genus Anadenanthera Spegazzini is in the section Mimosoideae of the family Fabaceae, order Fabales. Its first scientific description appeared in 1737 in the Hortus Cliffortianus, from a specimen growing in the Clifford Garden in Holland. Safford (1916) and Reis Altschul (1964) are of the opinion that the seed from which the type specimen was grown came from the West Indies or northern South America. The earliest valid name is Mimosa peregrina, from Linnaeus's Species Plantarum (1753; Safford 1916). Linnaeus did not specify why he applied the epithet peregrina. His description was based on the more complete entry in the Hortus Cliffortianus (Reis Altschul 1964). The 1806 Willdenow edition of Species Plantarum identified as Inga niopo the material collected by Humboldt and Bonpland along the Orinoco River. In 1814, Humboldt corrected Willdenow and identified the plant as an Acacia and not an Inga, labeling it Acacia niopo (Humboldt and Bonpland 1971). In her taxonomic study of the genus, Siri von Reis Altschul (1964) considers this specimen to be equivalent to Anadenanthera peregrina var. peregrina (see Appendix).
The genus Anadenanthera as established by J. P. M. Brenan (1955) and Reis Altschul (1964, 1967, 1972) was originally conceived by Bentham (1840, 1841-1842, 1874-1875) as section Niopa of the genus Piptadenia. Bentham placed 25 species under Piptadenia, all of them American. He divided the genus into three sections, Eupiptadenia and Pityrocarpa with spicate inflorescences, and Niopa with globose inflorescences (Reis Altschul 1964; Brenan 1955). Bentham originally distinguished five species in section Niopa: Piptadenia peregrina, P. macrocarpa, P. falcata, P. colubrina, and P. microphylla. Bentham (1874-1875) later proposed only four species, placing P. microphylla in synonymy with P. macrocarpa. Brenan (1955) indicated that Niopa is a natural group, whereas the species included in the other two sections were very mixed and should not be classified as proposed by Bentham.
In 1922, Carlos Spegazzini founded the genus Anadenanthera to accommodate P. peregrina and P. falcata. He based the genus on the flowers lacking anther glands, and on the elliptical and elongated anthers that differ from those found in the species with anther glands (Spegazzini 1922). According to Reis Altschul (1964) and Brenan (1955), this is insufficient to justify generic separation. In 1927, Britton and Rose proposed section Niopa as a genus (Reis Altschul 1964). However, Anadenanthera antedates Niopa as a generic name.
In 1955, Brenan revised the genus Piptadenia according to criteria that included the mode of dehiscence of the pod and the structure of the seed. Brenan proposed eight genera with species formerly included in Piptadenia. The genus Anadenanthera as proposed by Brenan (1955) consists of the four species formerly in section Niopa of Piptadenia. He distinguished Anadenanthera by the following characteristics: globose inflorescence; dehiscence of the pod along one suture; suborbicular, narrowly or not winged seeds lacking endosperms; and strictly American distribution (see also Reis Altschul 1964). Differences in wood anatomy coincide with the eight genera proposed by Brenan and support the distinctiveness of Anadenanthera as a genus (Reis Altschul 1964; Brazier 1958; see also Tortorelli 1948).
Siri von Reis Altschul (1964), in her taxonomic revision of the genus, considered Anadenanthera to consist of two species, A. peregrina (L.) Speg. and A. colubrina (Vell.) Brenan, each species having two varieties. The two varieties of Anadenanthera peregrina are A. peregrina (L.) Speg. var. peregrina Reis Altschul and A. peregrina (L.) Speg. var. falcata (Benth.) Reis Altschul. The varieties of A. colubrina are A. colubrina (Vell.) Brenan var. colubrina Reis Altschul and A. colubrina (Vell.) Brenan var. Cebil (Griseb.) Reis Altschul.
Reis Altschul distinguished these two species by means of a few consistent morphological characters and their correlation with particular geographical locations. The taxonomy was based partly on the presence or absence of a gland on each anther, and on the position of the involucre surrounding the peduncle. The two species can be identified most easily on the basis of the pod texture, since the flowers of the different genera that compose the Mimosoideae are so similar. A. peregrina has dull, scurfy to verrucose pods and it is the more northerly of the two species. A. colubrina has nitid, smooth to reticulated pods, and it is limited to the southern hemisphere (Reis Altschul 1964). A. peregrina var. peregrina (Plates 1, 2) and A. colubrina var. Cebil (Plates 3-5) are the two species employed as the source of psychoactive preparations, most notably in the form of snuff powders.
The genus Anadenanthera has a wide distribution on the South American continent and in the Greater Antilles. The two species have a preference for savanna habitats, although they seem to be adaptable to a variety of altitudes. Siri von Reis Altschul (1964: 40) stated:
Neither species of Anadencinthera is represented, to my knowledge, in western Amazonas nor in the Andes of northern Peru, Ecuador and Colombia. This circumstance is probably due to an inability of the representatives of A. peregrina var. peregrina to tolerate the true rain-forest conditions through which they would have to pass in a westward dispersal. There is equally little reason to believe that representatives of A. colubrina var. Cebil would descend from the Andes into lowland rain-forest.
In a study of plant domestication among the Yanomamö, Napoleon Chagnon reported the presence of A. peregrina var. peregrina in the upper Orinoco region. Chagnon emphatically stated that in this region Anadenanthera is a jungle plant ("hisioma es una planta selvática en esta región"). All Yanomamo groups know this tree and refer to it using the same term, hisioma, suggesting a wide distribution of Anadenanthera in the upper Orinoco region (Chagnon et al. 1970: 188-189).
Species Descriptions
Anadenanthera peregrina (L.) Speg. (Reis Altschul 1964:45-46):
Shrub to tall tree, 3-27 m high. Trunk 20-40 cm in diameter at breast height, usually leaning, twisted, sometimes divided at the base into several shafts; more frequently the contorted, irregular branches spread out above the middle of a solitary trunk into an umbrella-like crown. Bark gray to nearly black with many small lenticels; unarmed or lower trunk producing conical thorns or wedge-shaped projections, sometimes intensely so when young, becoming tubercular-verrucose, corky, rugose and, in drier climates, very thick. Young twigs and foliage puberulent, occasionally glaucescent; mature foliage glabrous or nearly so. Leaves, including petioles, 12-30 cm long, the main rachis more or less channeled ventrally. Petioles somewhat darkened at their bases, 5-15 mm above which each bears a flattish, oval or oblong gland .5-5 mm long; 1-4 similar, smaller glands borne one between or just below each of the ultimate pinna pairs. Pinna pairs 10-30 or more, each pinna 2-5 cm or more long, opposite or subopposite leaflets usually imbricate 25-80 pairs, 2-8 mm long, .5-1.5 mm wide, linear, oblong or lanceolate, straight to falcate, at the base oblique or truncate, at the apex acute to acuminate to apiculate; venation obscure except for a single, nearly straight, slightly excentric mid-vein; membranaceous to coriaceous and nitid, sometimes differing in color and texture dorsiventrally. Heads 10-18 mm in diameter, including stamens, greenish white to creamy yellow, in fascicles of 1-5, puberulous to glabrous in the bud; the heads axillary to the leaves and subterminal, rarely becoming arranged in racemose patterns in the branch apices. Peduncles 1.75-4 cm long, puberulous, filiform or thicker, each bearing about three-quarters of the way up the axis a puberulous bi-dentate, campanulate involucre which becomes detached and slides down to encircle loosely the base of the peduncle. Cal...