Anadenanthera Speg., Physis (Buenos Aires) 6: 313. 1923.

Bruneau, Anne, de Queiroz, Luciano Paganucci, Ringelberg, Jens J., Borges, Leonardo M., Bortoluzzi, Roseli Lopes da Costa, Brown, Gillian K., Cardoso, Domingos B. O. S., Clark, Ruth P., Conceicao, Adilva de Souza, Cota, Matheus Martins Teixeira, Demeulenaere, Else, de Stefano, Rodrigo Duno, Ebinger, John E., Ferm, Julia, Fonseca-Cortes, Andres, Gagnon, Edeline, Grether, Rosaura, Guerra, Ethiene, Haston, Elspeth, Herendeen, Patrick S., Hernandez, Hector M., Hopkins, Helen C. F., Huamantupa-Chuquimaco, Isau, Hughes, Colin E., Ickert-Bond, Stefanie M., Iganci, Joao, Koenen, Erik J. M., Lewis, Gwilym P., de Lima, Haroldo Cavalcante, de Lima, Alexandre Gibau, Luckow, Melissa, Marazzi, Brigitte, Maslin, Bruce R., Morales, Matias, Morim, Marli Pires, Murphy, Daniel J., O'Donnell, Shawn A., Oliveira, Filipe Gomes, Oliveira, Ana Carla da Silva, Rando, Juliana Gastaldello, Ribeiro, Petala Gomes, Ribeiro, Carolina Lima, Santos, Felipe da Silva, Seigler, David S., da Silva, Guilherme Sousa, Simon, Marcelo F., Soares, Marcos Vinicius Batista & Terra, Vanessa, 2024, Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification, PhytoKeys 240, pp. 1-552 : 1

Anadenanthera Speg., Physis (Buenos Aires) 6: 313. 1923.

Figs 164 , 165

Piptadenia sect. Niopa Benth., J. Bot. (Hooker) 4: 340. 1841. Type: Piptadenia peregrina (L.) Benth. [≡ Mimosa peregrina L. (≡ Anadenanthera peregrina (L.) Speg.)]

Niopa (Benth.) Britton & Rose, Addisonia 12: 37, t. 403. 1927. Type: Niopa peregrina (L.) Britton & Rose [≡ Mimosa peregrina L. (≡ Anadenanthera peregrina (L.) Speg.)]

Lectotype

(designated by Altschul 1964). Anadenanthera peregrina (L.) Speg. [≡ Mimosa peregrina L.]

Description.

Unarmed trees or shrubs, 3-30 m high; trunk ± smooth or with mammillate projections; bark often suberose, sometimes thick. Stipules small, bristly, caducous; bracts enclosing new shoots broad, commonly persistent. Leaves feathery; extrafloral nectaries on petiole above base, sometimes on rachis between ultimate pairs of pinnae, small, round; pinnae 7-35 pairs, opposite or almost so; leaflets 20-80 pairs per pinna, opposite or almost so, ± narrowly oblong, lanceolate, cultrate to slightly falcate, sometimes imbricate, main-vein central and straight. Compound inflorescences of pedunculate capitula, in fascicles of up to 7 peduncles inserted in series in successive leaf axils or forming terminal paniculate groups by suppression of leaves; peduncles each with 2 membranous bracts united to form an annular involucre; capitula spherical, 1-2 cm diameter including stamens, each with 35-60 flowers, greenish white to creamy yellow or rarely orange, fragrant. Flowers hermaphroditic or sometimes staminate and hermaphroditic [at least in A. colubrina (Vell.) Brenan], small, sessile; calyx campanulate, loosely gamosepalous, to 3 mm long; corolla tubular-campanulate, to 4 mm long, lobes loosely cohering to the level of the calyx mouth; stamens 10, far-exserted, free distally, adnate to corolla proximally, anther gland present (at least in bud, A. colubrina ) or absent ( A. peregrina ); pollen in polyads of 8, 12 or commonly 16 grains, porate, exine granular, columellae absent ( Altschul 1964; Guinet 1969, 1981b; Caccavari 2002; Soares EL et al. 2022); nectary disc absent; ovary sessile to subsessile. Fruits shortly stipitate, narrowly oblong, ± flattened, sometimes slightly falcate, dry-coriaceous, sometimes falsely septate internally, pulp and gum absent, dehiscent along 1 suture, sutures slightly to ± prominently thickened and sometimes ± contracted between the seeds, surface of valves glabrous, reticulately veined, shiny or scurfy to verrucose. Seeds 8-16 in 1 series, circular, thin-discoid with a thin, sharp rim, dark brown to black, shiny, pleurogram present (Fig. 164 ).

Chromosome number.

2 n = 26 (24) ( Santos et al. 2012).

Included species and geographic distribution.

2-4 species, possibly more ( Altschul 1964; Cacharani et al. 2020; Mangaravite et al. 2023). Endemic to South America (Fig. 165 ), from northern Argentina and Paraguay northwards into Bolivia and Peru, southern and eastern Brazil, southern Guyana and Venezuela, with a few occurrences in Amazonia; scattered in much of the northern Andes; although present in the West Indies (Hispaniola, Puerto Rico, Lesser Antilles, Trinidad-Tobago), it was probably introduced there in the pre-Columbian era ( Altschul 1964).

Ecology.

Tropical and subtropical seasonally dry forests, along rivers and at forest margins, in woodland, thickets and wooded grassland (cerrado, savanna) and caatinga, often planted near villages, sometimes weedy and found along roadsides and on wasteland; growing on a range of substrates, sometimes dominant; to 2000 (-2700) m. Trees are partly deciduous.

Etymology.

an - (Gr., lacking), aden - (Gr., gland), anthera (L., anther), indicating a lack of anther glands, although A. colubrina has them in bud.

Human uses.

In pre-Columbian times, ground seeds of both species were used by Amerindians as a source of hallucinogenic snuff ( A. peregrina - cohoba, niopo, yopo; A. colubrina var. cebil - Anadenanthera cebil , curupay, vilca), probably for 3000+ years. The northern species, A. peregrina , is still used for magical, medicinal, religious and stimulative purposes ( Altschul 1972; Torres and Repke 2006). Anadenanthera colubrina (angico) is used for timber, paper, and leather-tanning.

Notes.

The long-established treatment by Altschul (1964) recognised only two species ( Anadenanthera peregrina , A. colubrina ) with overlapping distributions, each containing two varieties with a third subsequently added to A. colubrina in Argentina ( Cacharani et al. 2020). A recent genetic study by Mangaravite et al. (2023) elevated Altschul’s varieties to specific level, reinstating Brenan’s (1955) species concepts by recognising A. falcata (Benth.) Speg. and A. macrocarpa (Benth.) Brenan to give four species in total. However, this study, based on material from southern and eastern Brazil, does not cover the entire distribution area of the genus in South America and the Caribbean and may still not reflect the full genetic diversity within the genus.

Bentham (1841b) placed the species he recognised in Piptadenia sect. Niopa Benth. under five names, all based on South American material. One of them, P. peregrina (L.) Benth., was subsequently treated as a distinct but related genus Anadenanthera by Spegazzini (1923), who excluded several un-named Australian taxa that Taubert (1894) had suggested also belonged to sect. Piptadenia Niopa , as well as another that is now a synonym of Schleinitzia novoguineensis (Warb.) Verdc. Britton and Rose (1927) created the genus Niopa to contain N. peregrina , but Brenan (1955) resurrected the name Anadenanthera as it had priority and this was then used by Altschul (1964) in her monograph. This work gave detailed accounts of the taxa and their morphology. The species of Anadenanthera had been atypical within Piptadenia because of their small, globose, rather than spicate, inflorescences ( Bentham 1875) and pods that dehisce along only one suture, rather than more or less along both ( Lewis and Elias 1981). Lewis and Elias (1981) assigned Anadenanthera to the Piptadenia group of tribe Mimoseae , as did Luckow et al. (2003) and Luckow (2005).

In addition to its capitula and dehiscent pods, distinctive characters of Anadenanthera include the thin, rimmed or very narrowly winged seeds that are circular in outline and lack endosperm. The species recognised by Altschul (1964) differ from one another in the texture of the pods, the presence/absence of anther glands and the position of the involucre on the peduncle.

The use of Anadenanthera as an hallucinogen is dependent on the presence in the seeds, fruits and bark of many populations of a range of indole alkaloids derived from tryptamine and related to serotonin; among these, bufotenin is often especially abundant and psychoactive ( Torres and Repke 2006). Some of these compounds are present in other Mimoseae but at much lower concentrations.

Plants of Anadenanthera colubrina are partially self-incompatible and its main pollinators are diurnal eusocial bees; eusocial honey wasps are nectar thieves or sometimes pollinators, and other insects are largely thieves of nectar and/or pollen (Kiill and da Silva 2016; Borges et al. 2017). Unusually, nectar is produced by the corolla lobes in this species ( Borges et al. 2017).

Trees are possibly mast fruiting and seed dispersal is by gravity and probably wind and leaf-cutter ants ( Fredericksen et al. 2000). Insect seed-predators include Curculionidae ( Justiniano and Fredericksen 1998) and bark exudates are consumed by marmosets ( Callitrichidae ) (Fransisco et al. 2017). The nodulated roots form a large tuber in Anadenanthera peregrina ( Gross et al. 2002).

Taxonomic references.

Altschul (1964); Cacharani et al. (2020); Mangaravite et al. (2023); Martinez et al. (2013).

Figures

Figure 164. Morphological features of Anadenanthera (A-I) and Parkia sect. Platyparkia (J-L) A A. colubrina (Vell.) Brenan capitulum cut in half, at anthesis B capitula at anthesis and in bud C shrub D Anadenanthera sp. foliage and petiolar nectary E A. colubrina mature pods F mammillate projections on trunk G dehisced pods H A. peregrina var. falcata (Benth.) Altschul mammillate trunks I A. peregrina (L.) Speg. var. peregrina trees with smooth trunks J Parkia pendula (Willd.) Benth. ex Walp. (Hopkins & Hopkins 273), Neotropics, tree crown K capitulum at anthesis, nectar exuding from apical flowers L pods with enlarged adaxial suture secreting gum into which the seeds have fallen. Photo credits A-I RT Queiroz https: // rubens-plantasdobrasil. blogspot. com / J-L MJG Hopkins and HCF Hopkins.

Figure 165. Distribution of Anadenanthera based on quality-controlled digitised herbarium records. Occurrences in the West Indies may be due to pre-Colombian naturalisation. See Suppl. material 1 for the source of occurrence data.