Moullava Adans., Fam. Pl. 2: 318. 1763.

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

Moullava Adans., Fam. Pl. 2: 318. 1763.

Figs 37 , 38 , 61

Almeloveenia Dennst., Schlüssel Hortus Malab.: 32. 1818. Type: Almeloveenia spinosa Dennst. [= Moullava spicata (Dalzell ex Wight) Nicolson]

Cinclidocarpus Zoll. & Moritzi, Natuur-Geneesk. Arch. Ned.- Indië iii: 81. 1846. Type: Cinclidocarpus nitidus Zoll. & Moritzi [= Moullava tortuosa (Roxb.) Gagnon & G.P. Lewis]

Wagatea Dalzell, Hooker’s J. Bot. Kew Gard. Misc. 3: 90. 1851. Type: Wagatea spicata Dalzell ex Wight [≡ Moullava spicata (Dalzell ex Wight) Nicolson]

Caesalpinia sect. Cinclidocarpus (Zoll. & Moritzi) Benth. & Hook., Gen. Pl. 6.: 565-567. 1865. Type not designated.

Type.

Moullava spicata (Dalzell ex Wight) Nicolson [≡ Wagatea spicata Dalzell ex Wight]

Description.

Lianas and scrambling shrubs, armed with deflexed prickles. Leaves bipinnate, ending with a pair of pinnae; pinnae in 7-20 opposite pairs; leaflets in 5-40 opposite pairs per pinna. Stipules caducous or lacking (not seen). Inflorescence an elongated terminal or axillary raceme, the racemes sometimes aggregated into panicles. Flowers bisexual, sub-actinomorphic or zygomorphic; hypanthium persisting either as a distinct cup or as a wide shallow calyx remnant at the pedicel apex as the fruit matures; sepals 5, caducous, eglandular, glabrous, the lower sepal strongly cucullate, covering the other 4 sepals in bud; petals 5, free, yellow, the median (innermost upper) and lateral petals sometimes streaked red, eglandular; stamens 10, free, barely exserted beyond the corolla, densely pubescent on lower half of filaments; ovary glabrous or pubescent. Fruit fleshy, oblong-elliptic, unarmed, indehiscent, sub-torulose, with thickened sutures, drying black (immature fruits of M. spicata red-tomentose), exocarp and endocarp strongly adnate, glabrous, 1-4-seeded. Seeds sub-globular, olive-brown to black.

Chromosome number.

2 n = 24 [ M. digyna (Rottler) Gagnon & G.P. Lewis, M. spicata ] ( Rice et al. 2015).

Included species and geographic distribution.

Four species, three in southern Asia and one in Africa (Fig. 61 ).

Ecology.

The Asian species are found in seasonally dry tropical semi-evergreen forest margins, secondary thickets, and on mountain slopes, up to 1200 m elevation. The African species occurs mostly in riverine habitats in lowland rainforests.

Etymology.

Derived from the vernacular name of Moullava spicata , “mulu” (Malayalam: spiny), a spiny climber.

Human uses.

Moullava spicata is used for medicine ( Lewis 2005b).

Notes.

First described as a monospecific genus from India, its description was emended in Gagnon et al. (2016) to include three other species, all with similar torulose fruits, not found elsewhere in the Caesalpinieae .

Taxonomic references.

Ansari (1990); Brenan (1963a, 1967); Brummitt et al. (2007, see both Moullava and Mezoneuron welwitschianum Oliv.); Chen et al. (2010a); Gagnon et al. (2016); Hattink (1974); Lewis (2005b); Nicolson (1980); Sanjappa (1992); Vidal and Hul Thol (1976).

Figures

Figure 37. Fruits of the Caesalpinieae A Gelrebia rubra (Engl.) Gagnon & G. P. Lewis, Namibia B Moullava spicata (Dalzell ex Wight) Nicolson, India, Maharashtra C Hultholia mimosoides (Lam.) Gagnon & G. P. Lewis, India D Guilandina bonduc L., Madagascar E Pterolobium stellatum (Forssk.) Brenan, Zimbabwe F Ticanto sinensis (Hemsley) R. Clark & Gagnon, China (Clark 415) G Biancaea decapetala (Roth) O. Deg., Peru, Ancash (Hughes et al. 2227) H Mezoneuron kauaiense (H. Mann) Hillebr. I Mezoneuron andamanicum Prain, Thailand (Clark 251). Photo credits A Dave U, iNaturalist (https: // www. inaturalist. org / photos / 41085094) B P Awale, Flowers of India (http: // www. flowersofindia. net /) C VR Vinayaraj, India Biodiversity Portal (https: // indiabiodiversity. org / observation / show / 335158), the basionym of Hultholia mimosoides D GP Lewis E BT Wursten, Flora of Zimbabwe (https: // www. zimbabweflora. co. zw / speciesdata / image-display. php? species _ id = 127190 & image _ id = 1) F, I P Suksathan G CE Hughes H D Eickhoff, https: // www. flickr. com / photos / dweickhoff / 4822012867 / in / photostream /.

Figure 38. Vegetative traits of the Caesalpinieae A Hoffmannseggia minor (Phil.) Ulibarri, Bolivia B Hererolandia pearsonii (L. Bolus) Gagnon & G. P. Lewis, Namibia, Sesriem Canyon C Moullava spicata (Dalzell ex Wight) Nicolson, India, Maharastra D Cordeauxia edulis Hemsl., undersurface of leaflets showing glands, Somalia E Pomaria austrotexana B. B. Simpson, USA, Texas F Small tree of Erythrostemon nicaraguensis (G. P. Lewis) Gagnon & G. P. Lewis, Nicaragua, Esteli (Hawkins et al. 4) G Leopard bark of Libidibia paraguariensis (D. Parodi) G. P. Lewis, Bolivia, Santa Cruz (Hughes 2475) H Lenticelled bark of Erythrostemon nicaraguensis (G. P. Lewis) Gagnon & G. P. Lewis, Nicaragua, Esteli (Hawkins et al. 4) I Young flush of leaves of Cenostigma pluviosum var. intermedium (G. P. Lewis) Gagnon & G. P. Lewis, Brazil, Bahia (Lima et al. 7901) J Fluted trunk of Haematoxylum brasiletto H. Karst., Mexico, Oaxaca, (Hughes 1947) K Prickles on woody protuberances on a young trunk of Paubrasilia echinata (Lam.) Gagnon, H. C. Lima & G. P. Lewis, Brazil, Bahia (Lima et al. 7909) L Recurved spines of Biancaea decapetala (Roth) O. Deg., Peru, Ancash, (Hughes et al. 3055). Photo credits A GP Lewis B AA Dreyer C Shivaprakash, iNaturalist (https: // inaturalist. ca / observations / 81864757) D M Thulin E WR Carr F-H, J CE Hughes I, K, L E Gagnon.

Figure 61. Distribution of Moullava based on quality-controlled digitised herbarium records. See Suppl. material 1 for the source of occurrence data.