Pityrocarpa (Benth.) Britton & Rose, N. Amer. Fl. 23(3): 190. 1928.

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

Pityrocarpa (Benth.) Britton & Rose, N. Amer. Fl. 23(3): 190. 1928.

Figs 172 , 173 , 180

Piptadenia sect. Pityrocarpa Benth., J. Bot. (Hooker) 4: 339. 1841. Type: Piptadenia moniliformis Benth. [≡ Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson]

Monoschisma Brenan, Kew Bull. 10(2): 179. 1955, nom. inval., non Monoschisma Duby, Mém. Soc. Phys. Genève 19: 294. 1868 ( Musci, Meteoriaceae ). Type: Monoschisma leptostachyum (Benth.) Brenan [≡ Piptadenia leptostachya Benth. (≡ Pityrocarpa leptostachya (Benth.) L.P. Queiroz & P.G. Ribeiro)]

Pseudopiptadenia Rauschert, Taxon 31(3): 559. 1982. Type: Pseudopiptadenia leptostachya (Benth.) Rauschert [≡ Piptadenia leptostachya Benth. (≡ Pityrocarpa leptostachya (Benth.) L.P. Queiroz & P.G. Ribeiro)]

Lectotype

(designated by Britton and Rose 1928). Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson [≡ Piptadenia moniliformis Benth.]

Description.

Trees or shrubs; indumentum composed of simple trichomes; brachyblasts absent; branches and leaves unarmed, not odoriferous. Stipules present, sometimes caducous. Leaves bipinnate; extrafloral nectaries between or just below the first pair of pinnae; pinnae 1-4 (10) pairs, opposite; leaflets 1-10 (20) pairs per pinna, opposite, rhomboid, sometimes asymmetrically elliptical or lanceolate. Inflorescences cylindrical spikes, white to yellowish or greenish, solitary (rarely 2) in the axils of coevally developing leaves. Flowers 5-merous; calyx cupulate; petals free (sometimes joined in one species); stamens 10, anthers with an apical gland; pollen in polyads with 8 or 16 grains; ovary included or exserted from the corolla; stigma in a terminal pore. Fruit a follicle, flat compressed, margins deeply and regularly constricted, rarely sinuous and shallowly constricted. Seeds flat compressed, dark, narrowly winged, or rarely ovoid or discoid, white and wingless, pleurogram absent from dark seeds, but present in the white ones.

Chromosome number.

2 n = 26 ( Alves and Custódio 1989).

Included species and geographic distribution.

Seven species occurring in three major areas in tropical America: Mexico and Central America, northern South America (Guyana, Venezuela), and eastern Brazil (Fig. 180 ).

Ecology.

Rainforests (Brazil), seasonally dry tropical forests and woodlands (Brazil and Mexico), savannas (Venezuela) and Chaco (Paraguay).

Etymology.

From Greek, pityron (= scurf; husks of bran) and carpus (= fruit), in reference to the leprose fruits of some species (e.g., Pityrocarpa moniliformis ).

Human uses.

Pityrocarpa moniliformis is used as fodder, timber for light constructions, firewood, and for soil enrichment, ecological restoration and honey production; it is also rich in tannins ( Carvalho 2010; Carvalho 2014).

Notes.

Originally described as a section of Piptadenia ( Bentham 1841b), Pityrocarpa included three species with leprose fruits ( Britton and Rose 1928). The limits of the genus were briefly expanded due to nomenclatural mistakes ( Brenan 1955), which, after being corrected, culminated in a return to its original circumscription and rank ( Brenan 1963b). Although phylogenetic evidence provided renewed support for recognition of Pityrocarpa at the generic level ( Jobson and Luckow 2007), more recent analyses recovered the genus as paraphyletic in relation to part of Pseudopiptadenia , including its type species ( Simon et al. 2016; Borges et al. 2022). As Pityrocarpa has publication priority, Pseudopiptadenia was subsumed under it.

The recent updates to the circumscription of Pityrocarpa rendered the genus more variable with respect to seed morphology -a character previously used to distinguish it from Parapiptadenia and Pseudopiptadenia , for example- but also highlighted the taxonomic relevance of particular traits for recognition of the genus, such as extrafloral nectaries between or just below the first pair of pinnae and inflorescences in general solitary and axillary to coevally developing leaves ( Borges et al. 2022). Another distinctive, although not exclusive, feature of Pityrocarpa is the deeply constricted, moniliform fruits ( Bentham 1875; Simon et al. 2016; Borges et al. 2022).

Taxonomic references.

Borges et al. (2022); Jobson and Luckow (2007); Lewis and Lima (1991); Simon et al. (2016).

Figures

Figure 172. Flowering branches in the Stryphnodendron clade A Gwilymia coriacea (Benth.) A. G. Lima, Paula-Souza & Scalon (Simon 3482) B Microlobius foetidus (Jacq.) M. Sousa & G. Andrade C Stryphnodendron adstringens (Mart.) Coville D S. gracile Rizzini & Heringer E Marlimorimia bahiana (G. P. Lewis & M. P. Lima) L. P. Queiroz & L. M. Borges F Pityrocarpa moniliformis (Benth.) Luckow & R. W. Jobson G Parapiptadenia blanchetii (Benth.) Vaz & M. P. Lima H Naiadendron duckeanum (Occhioni) A. G. Lima, Paula-Souza & Scalon (Simon 1457). Photo credits A, H MF Simon B T Iwane C, D H Moreira E LP Queiroz F D Cardoso G PG Ribeiro.

Figure 173. Habit and fruits in the Stryphnodendron clade A Marlimorimia psilostachya (DC.) L. P. Queiroz & Marc. F. Simon showing the tree habit B Stryphnodendron platyspicum Rizzini & Heringer (Simon 2017) showing the geoxyle subshrub habit C Gwilymia coriacea (Benth.) A. G. Lima, Paula-Souza & Scalon (Simon 3730) D Microlobius foetidus (Jacq.) M. Sousa & G. Andrade E Stryphnodendron rotundifolium Mart. fruit, manually opened revealing internal septa and seeds F Marlimorimia contorta (DC.) L. P. Queiroz & P. G. Ribeiro G Naiadendron duckeanum (Occhioni) A. G. Lima, Paula-Souza & Scalon (Pereira-Silva 15692) H Parapiptadenia rigida (Benth.) Brenan showing a yellow inflorescence and dehiscing fruit I Pityrocarpa leptostachya (Benth.) L. P. Queiroz & P. G. Ribeiro. Photo credits A R Aguilar B, C, E MF Simon D H Hulsberg F G Carvalho-Sobrinho G G Pereira-Silva H RT Queiroz https: // rubens-plantasdobrasil. blogspot. com / I LM Borges.

Figure 180. Distribution of Pityrocarpa based on quality-controlled digitised herbarium records. See Suppl. material 1 for the source of occurrence data.