Tetrapterocarpon Humbert, Compt. Rend. Hebd. Seances Acad. Sci. 208: 374. 1939.

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

Tetrapterocarpon Humbert, Compt. Rend. Hebd. Seances Acad. Sci. 208: 374. 1939.

Figs 7 , 9

Type.

Tetrapterocarpon geayi Humbert

Description.

Unarmed trees or shrubs, dioecious; brachyblasts absent. Stipules inconspicuous, minute and caducous. Leaves bipinnate, ending in a terminal pinna; leaflets alternate. Inflorescences axillary, spike-like racemes of small, subsessile flowers, usually aggregated into panicles. Flowers unisexual, actinomorphic, 4-merous (sepals and petals 4 per flower), greenish; sepals equal, petals equal, both whorls imbricate in young bud; androecium diplostemonous in staminate flowers, with one whorl of 4 fertile stamens, their filaments with an apical tuft of hairs behind the anthers, and one whorl of 4 hairy staminodes, lacking anthers; pollen with a scabrate-punctate sculpture pattern; pistillate flowers with a stipitate, compressed-fusiform ovary, stigma capitate and bilobed. Fruits membranous, indehiscent, compressed, 4-winged (in two unequal pairs), 1-seeded (Fig. 7B ). Seeds trapezoid, subterete, club-shaped, pleurogram lacking.

Chromosome number.

Unknown.

Included species and geographic distribution.

Two species, both endemic to Madagascar (Fig. 9 ).

Ecology.

Seasonally dry tropical to xerophytic forest and thicket, on limestone, basalt, or sand.

Etymology.

From Greek, tetra - (= four), ptero - (= winged) and carpos (= fruit), the fruits have four dry, papery wings in unequal pairs.

Human uses.

Tetrapterocarpon geayi is used locally for carpentry, cart construction and to make charcoal (Du Puy and Rabevohitra 2002).

Notes.

Although placed in the Dimorphandra group of the Caesalpinieae by Polhill and Vidal (1981), the genus was resolved as sister to an Acrocarpus - Ceratonia clade in a study of the " Umtiza clade" by Herendeen et al. (2003b), sharing with these two genera a bipinnate leaf with an unequal leaflet base, and this relationship is supported in the phylogenomic analysis of Ringelberg et al. (2022) (Fig. 6 ).

Taxonomic references.

Du Puy and Rabevohitra (2002); Lewis (2005b).

Figures

Figure 6. Generic relationships in tribe Ceratonieae. Left part of figure shows complete genus-level Caesalpinioideae phylogeny with the Ceratonieae indicated with a red rectangle. Branch lengths are expressed in coalescent units and terminal branches were assigned an arbitrary uniform length for visual clarity. Support for relationships is based on fractions of supporting and conflicting gene trees: pie charts show gene tree support and conflict per node (blue representing supporting gene trees, green gene trees supporting the most common alternative topology, red gene trees supporting further alternative topologies, grey gene trees uninformative for this node), and numbers above pie charts are Internode Certainty All support values [both calculated with PhyParts (Smith et al. 2015)]. If present, red numbers below pie charts are non-significant (i. e.> 0.05) outcomes of ASTRAL's polytomy test (Sayyari and Mirarab 2018), which tests for each node whether the polytomy null model can be rejected. Monophyletic genera are represented by single branches; see Suppl. material 2 for a phylogeny with all accessions. See Suppl. material 3 for gene tree support across the phylogeny. The phylogeny is a pruned version of the backbone phylogeny of Ringelberg et al. (2023), where full details of the data and phylogenomic analysis methods are presented.

Figure 7. Flower, fruit, and vegetative characters of tribe Ceratonieae A Arcoa gonavensis Urb., foliage and fruits, Dominican Republic B Tetrapterocarpon geayi Humbert, fruits and part of bipinnate leaf, Madagascar (Du Puy M 410) C-E Acrocarpus fraxinifolius Wight & Arn., India, cultivated tree C inflorescence D dehisced fruits and new flush foliage E part of a bipinnate leaf F-J Ceratonia siliqua L. F tree, Jordan, semi-desert near Petra G fruits, Israel, Mt. Scopus Botanical Gardens, Jerusalem H pistillate flowers, Crete I staminate flowers, Crete J staminate inflorescences, Israel, Jerusalem Botanical Gardens. Photo credits A F Jimenez R. B D Du Puy C-E https: // efloraofindia. com / 2011 / 02 / 01 / acrocarpus-fraxinifolius / E D Valke F, G, J O Fragman-Sapir H, I G Lewis.

Figure 9. Distribution of Tetrapterocarpon based on quality-controlled digitised herbarium records. See Suppl. material 1 for the source of occurrence data.