Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification Author Bruneau, Anne https://orcid.org/0000-0001-5547-0796 Institut de recherche en biologie vegetale and Departement de Sciences biologiques, Universite de Montreal, 4101 Sherbrooke E., Montreal (QC) H 1 X 2 B 2, Canada anne.bruneau@umontreal.ca Author de Queiroz, Luciano Paganucci https://orcid.org/0000-0001-7436-0939 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Ringelberg, Jens J. https://orcid.org/0000-0003-0567-5210 Department of Systematic and Evolutionary Botany, University of Zurich, Zollikerstrasse 107, 8008 Zurich, Switzerland & School of Geosciences, University of Edinburgh, Old College, South Bridge, Edinburgh EH 8 9 YL, UK Author Borges, Leonardo M. https://orcid.org/0000-0001-9269-7316 Universidade Federal de Sao Carlos, Departamento de Botanica, Rodovia Washington Luis, Km 235, 13565 - 905, Sao Carlos, SP, Brazil Author Bortoluzzi, Roseli Lopes da Costa https://orcid.org/0000-0002-7445-7244 Programa de Pos-graduacao em Producao Vegetal, Universidade do Estado de Santa Catarina, Centro de Ciencias Agroveterinarias, Avenida Luiz de Camoes 2090, 88520 - 000, Lages, Santa Catarina, Brazil Author Brown, Gillian K. https://orcid.org/0000-0002-7940-5435 Queensland Herbarium and Biodiversity Science, Department of Environment and Science, Toowong, Queensland, 4066, Australia Author Cardoso, Domingos B. O. S. https://orcid.org/0000-0001-7072-2656 Instituto de Pesquisas Jardim Botanico do Rio de Janeiro, Pacheco Leao 915, 22460 - 030, Rio de Janeiro, RJ, Brazil & Programa de Pos-Graduacao em Biodiversidade e Evolucao (PPGBioEvo), Instituto de Biologia, Universidade Federal de Bahia (UFBA), Rua Barao de Jeremoabo, s. n., Ondina, 40170 - 115, Salvador, BA, Brazil Author Clark, Ruth P. https://orcid.org/0000-0001-9974-2933 Accelerated Taxonomy Department, Royal Botanic Gardens, Kew, Richmond, TW 9 3 AE, UK Author Conceicao, Adilva de Souza https://orcid.org/0000-0002-8800-422X Programa de Pos-graduacao em Diversidade Vegetal, Universidade do Estado da Bahia, Herbario HUNEB, Campus VIII, Rua do Gangorra 503, 48608 - 240, Paulo Afonso, Bahia, Brazil Author Cota, Matheus Martins Teixeira https://orcid.org/0000-0003-0654-7501 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Demeulenaere, Else https://orcid.org/0000-0002-1815-3051 Center for Island Sustainability and Sea Grant, University of Guam, UOG Station, Mangilao, 96923, Guam Author de Stefano, Rodrigo Duno https://orcid.org/0000-0003-1707-4121 Centro de Investigacion Cientifica de Yucatan, A. C. (CICY), Calle 43 No. 130 x 32 y 34, Chuburna de Hidalgo; CP 97205, Merida, Yucatan, Mexico Author Ebinger, John E. Eastern Illinois University, Charleston, IL 61920, USA Author Ferm, Julia https://orcid.org/0000-0002-8762-3942 Department of Ecology, Environment and Plant Sciences, 10691, Stockholm University, Stockholm, Sweden Author Fonseca-Cortes, Andres https://orcid.org/0000-0001-7207-9940 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Gagnon, Edeline https://orcid.org/0000-0003-3212-9688 Department of Integrative Biology, University of Guelph, 50 Stone Road, Guelph (ON) N 1 G 2 W 1, Canada & Chair of Phytopathology, Technical University Munich, 85354 Freising, Germany & Royal Botanic Garden Edinburgh, 20 A Inverleith Row, Edinburgh, EH 3 5 LR, UK Author Grether, Rosaura https://orcid.org/0000-0003-2673-665X Departamento de Biologia, Universidad Autonoma Metropolitana-Iztapalapa, Apdo. Postal 55 - 535, 09340 Ciudad de Mexico, Mexico Author Guerra, Ethiene https://orcid.org/0000-0002-9495-1717 Universidade Federal do Rio Grande do Sul, Programa de Pos-Graduacao em Botanica, Av. Bento Goncalves 9500, Bloco IV - Predio 43433, Porto Alegre, RS, 91501 - 970, Brazil Author Haston, Elspeth https://orcid.org/0000-0001-9144-2848 Royal Botanic Garden Edinburgh, 20 A Inverleith Row, Edinburgh, EH 3 5 LR, UK Author Herendeen, Patrick S. https://orcid.org/0000-0003-2657-8671 Chicago Botanic Garden, 1000 Lake Cook Road, Glencoe, IL 60022, USA Author Hernandez, Hector M. https://orcid.org/0000-0002-1741-5515 Departamento de Botanica, Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Cd. Universitaria, 04510 Ciudad de Mexico, Mexico Author Hopkins, Helen C. F. https://orcid.org/0000-0003-4984-8224 Accelerated Taxonomy Department, Royal Botanic Gardens, Kew, Richmond, TW 9 3 AE, UK Author Huamantupa-Chuquimaco, Isau https://orcid.org/0000-0002-4153-5875 Herbario Alwyn Gentry (HAG), Universidad Nacional Amazonica de Madre de Dios (UNAMAD), AV. Jorge Chavez N ° 1160, Madre de Dios, Peru Author Hughes, Colin E. https://orcid.org/0000-0002-9701-0699 Department of Systematic and Evolutionary Botany, University of Zurich, Zollikerstrasse 107, 8008 Zurich, Switzerland Author Ickert-Bond, Stefanie M. https://orcid.org/0000-0001-8198-8898 Department of Biology & Wildlife & Herbarium (ALA) at the University of Alaska Museum of the North, University of Alaska Fairbanks, P. O. Box 756960, Fairbanks AK 99775 - 6960, USA Author Iganci, Joao https://orcid.org/0000-0002-5740-3666 Universidade Federal do Rio Grande do Sul, Programa de Pos-Graduacao em Botanica, Av. Bento Goncalves 9500, Bloco IV - Predio 43433, Porto Alegre, RS, 91501 - 970, Brazil & Programa de Pos-Graduacao em Fisiologia Vegetal, Universidade Federal de Pelotas, Instituto de Biologia, Campus Universitario Capao do Leao, Passeio Andre Dreyfus, Departamento de Botanica, Predio 21, Pelotas, Rio Grande do Sul, 96010 - 900, Brazil Author Koenen, Erik J. M. https://orcid.org/0000-0002-4825-4339 Evolutionary Biology & Ecology, Universite Libre de Bruxelles, Faculte des Sciences, Campus du Solbosch - CP 160 / 12, Avenue F. D. Roosevelt, 50, 1050 Bruxelles, Belgium Author Lewis, Gwilym P. https://orcid.org/0000-0003-2599-4577 Accelerated Taxonomy Department, Royal Botanic Gardens, Kew, Richmond, TW 9 3 AE, UK Author de Lima, Haroldo Cavalcante https://orcid.org/0000-0003-2154-670X Instituto de Pesquisas Jardim Botanico do Rio de Janeiro, Pacheco Leao 915, 22460 - 030, Rio de Janeiro, RJ, Brazil & Instituto Nacional da Mata Atlantica / INMA-MCTI, Av. Jose Ruschi, 4, Centro, 29650 - 000, Santa Teresa, Espirito Santo, Brazil Author de Lima, Alexandre Gibau https://orcid.org/0000-0002-9168-2507 Instituto de Pesquisas Jardim Botanico do Rio de Janeiro, Pacheco Leao 915, 22460 - 030, Rio de Janeiro, RJ, Brazil & Department of Biological and Environmental Sciences, University of Gothenburg, Gothenburg, Sweden Author Luckow, Melissa https://orcid.org/0009-0007-2543-0516 School of Integrative Plant Science, Plant Biology Section, Cornell University, 215 Garden Avenue, Roberts Hall 260, Ithaca, NY 14853, USA Author Marazzi, Brigitte https://orcid.org/0000-0003-3252-5816 Natural History Museum of Canton Ticino, Viale C. Cattaneo 4, 6900 Lugano, Switzerland Author Maslin, Bruce R. https://orcid.org/0000-0002-3039-0973 Western Australian Herbarium, Department of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia, 6983, Australia & Singapore Herbarium, 1 Cluny Road, Singapore, Singapore Author Morales, Matias https://orcid.org/0000-0001-5540-9725 Instituto de Recursos Biologicos, CIRN-CNIA, INTA. N. Repetto & Los Reseros s. n., Hurlingham, Buenos Aires, Argentina & Consejo Nacional de Investigaciones Cientificas y Tecnicas (CONICET), Godoy Cruz 2290 (C 1425 FQB), Ciudad Autonoma de Buenos Aires, Argentina Author Morim, Marli Pires https://orcid.org/0000-0003-0872-8429 Instituto de Pesquisas Jardim Botanico do Rio de Janeiro, Pacheco Leao 915, 22460 - 030, Rio de Janeiro, RJ, Brazil Author Murphy, Daniel J. https://orcid.org/0000-0002-8358-363X Royal Botanic Gardens Victoria, Melbourne, Victoria, 3004, Australia Author O'Donnell, Shawn A. https://orcid.org/0000-0003-0731-7425 Geography and Environmental Sciences, Northumbria University, Ellison Place, Newcastle upon Tyne, NE 1 8 ST, UK Author Oliveira, Filipe Gomes https://orcid.org/0000-0003-0244-3262 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Oliveira, Ana Carla da Silva https://orcid.org/0000-0001-7042-5360 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Rando, Juliana Gastaldello https://orcid.org/0000-0002-3714-8231 Programa de Pos-graduacao em Ciencias Ambientais, Universidade Federal do Oeste da Bahia, Rua Professor Jose Seabra Lemos 316, 47800 - 021, Barreiras, Bahia, Brazil Author Ribeiro, Petala Gomes https://orcid.org/0000-0002-0070-9971 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Ribeiro, Carolina Lima https://orcid.org/0000-0001-9508-2894 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Santos, Felipe da Silva https://orcid.org/0000-0002-1068-0578 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Seigler, David S. https://orcid.org/0009-0003-5177-5893 Department of Plant Biology, University of Illinois, Urbana, IL 61801, USA Author da Silva, Guilherme Sousa https://orcid.org/0000-0002-4250-0017 Instituto de Biologia, Universidade Estadual de Campinas, Campinas, 13083 - 876, Sao Paulo / SP, Brazil Author Simon, Marcelo F. https://orcid.org/0000-0002-5732-1716 Empresa Brasileira de Pesquisa Agropecuaria (Embrapa) Recursos Geneticos e Biotecnologia, Parque Estacao Biologica, Caixa Postal 02372, 70770 - 917, Brasilia / DF, Brazil Author Soares, Marcos Vinicius Batista https://orcid.org/0000-0003-2660-1771 Universidade Federal do Rio Grande do Sul, Programa de Pos-Graduacao em Botanica, Av. Bento Goncalves 9500, Bloco IV - Predio 43433, Porto Alegre, RS, 91501 - 970, Brazil Author Terra, Vanessa https://orcid.org/0000-0001-5669-1304 Instituto de Biologia, Universidade Federal de Santa Maria, 97105 - 900, Santa Maria / RS, Brazil text PhytoKeys 2024 2024-04-03 240 1 552 http://dx.doi.org/10.3897/phytokeys.240.101716 journal article http://dx.doi.org/10.3897/phytokeys.240.101716 1314-2003-240-1 B699D9DE2B435B1093DE3C38C703D430 Inga Mill., Gard. Dict. Abr. 2. 1754. Figs 266 , 268 , 274 , 276 Affonsea A. St.-Hil., Voyage Distr. Diamans Bres , 1: 385. 1833. Type: Affonsea juglandifolia A. St.-Hil. [≡ Inga globularis T.D. Penn.] Feuilleea Kuntze, Revis. Gen. Pl. 1: 182. 1891. Type: Feuilleea inga (L.) Kuntze [≡ Mimosa inga L. (≡ Inga vera Willd.)] Type . Inga vera Willd. [≡ Mimosa inga L.] Description. Unarmed trees or treelets to ca. 40 m high, with a smooth cylindrical bole (Fig. 268D ), bark usually smooth, numerous lenticels arranged in horizontal or vertical rows. Stipules present, up to 8.5 cm diam. Leaves paripinnate (Fig. 274C ), rarely bifoliolate; petiole and leaf rhachis cylindrical, narrowly margined or winged; foliar glands nearly always present between each pair of leaflets, occasionally present on the leaflet midrib, rarely absent; leaflets 1-11 pairs, often strongly asymmetrical at the base and with eucamptodromus or brochidodromus venation, secondary veins oblique, perpendicular or reticulate. Inflorescences umbellate, capitate, racemose or spicate, usually axillary, rarely ramiflorus or Inga cauliflorus (Fig. 274D-G ); floral bracts often conspicuous and persistent, occasionally partly fused to form an involucre. Flowers 5-merous and regular; calyx gamosepalous, the tube cup-shaped, funnel-shaped or tubular, nearly always exceeding the lobes, open in bud and then (4) 5 (6)-lobed, or closed in bud and then splitting irregularly, the apex rostrate and then opening by a single lateral slit; corolla gamopetalous, greenish, yellowish-white or bright yellow, rarely pink or red, tubular or rarely funnel-shaped, tube exceeding the lobes; stamens ca. 20-350, united in the lower half into a tube, tube shorter than, equaling or exceeding the corolla, free portion of filaments long-exserted; pollen in 16, 20, 24, 28, 30 or 32-celled polyads; intrastaminal disc small, cup-shaped; gynoecium 1-9-carpellate, ovules 10-32 per ovary. Fruits indehiscent, woody, leathery or fleshy, usually flat or convex, less frequently quadrangular or cylindrical, straight, curved or spirally twisted or coiled (Fig. 266L-O ), faces usually much broader than the margins, or margins sometimes raised or winged, when cylindrical then margin highly developed and partially or completely covering the faces. Seeds fleshy, thin-walled, the testa developing a thick white sugary sarcotesta (Fig. 266M, N ); pleurogram absent. Chromosome number. 2 n = 26, and in some tetraploid species 2 n = 52 ( Pennington 1997 ). Included species and geographic distribution. ca. 300 species, restricted to tropical America. The species are distributed in Mexico, southern Central America, western South America, Venezuela and the Guianas, the coastal states of Brazil and the West Indies. The highest species diversity is concentrated in the Andean foothills of Peru, Ecuador, Colombia and in southern Central America, occupying a wide variety of habitats from sea level to 3000 m ( Pennington 1997 ) (Fig. 276 ). Figure 276. Distribution of Inga based on quality-controlled digitised herbarium records. See Suppl. material 1 for the source of occurrence data. Ecology. The genus is largely confined to wet forests, having a strong relationship with the germination process of the seeds, as they need shade and high humidity for immediate germination ( Pennington 1997 ). Some species occur in semi-arid areas, but then are mostly confined to gallery forests along temporary or permanent watercourses. Etymology. Derived from "inga" , the Brazilian name of several species of mimosoid legumes, especially of the genus Inga . Originated from the Tupi word in-ga , which probably means "soaked, stewed", due to the consistency of the pulp that surrounds the seeds ( Ferreira 1986 ). Human uses. The genus has many uses. The fruits are edible, and the fleshy sarcotesta surrounding the seed can be eaten fresh or in recipes (Fig. 266O ). The earliest records of this usage are from coastal Peru, where remnants of fruits and seeds of Inga feuillei DC. have been recorded from tombs dating back to the Chimu and Mochica periods, ca. 2000 years ago. Ceramics from the same period made in the form of the fruit of I. feuillei have also been found in archaeological remains ( Pennington 1997 ). Some species have edible embryos, which are used in soups or prepared by boiling in water or by roasting. Species of Inga are widely used by farmers as shade trees, mainly in coffee, cocoa and tea plantations, due to their architecture, which produces a flat or umbrella-shaped evergreen crown ( Pennington 1997 ). In addition, they are widely used as fuelwood and to produce high volumes of timber in a short period of time ( Pennington 1997 ). The species are also commonly used in soil restoration and agroforestry ( Pennington 1997 ). Notes. Inga has been supported as monophyletic in several phylogenetic studies of the mimosoid legumes although only about one third of the species have been sampled ( Nicholls et al. 2015 ; Dexter et al. 2017 ). Barneby and Grimes (1996) included Inga in the informal Inga alliance together with Macrosamanea , Cojoba Britton & Rose, Zygia , Calliandra Benth., Zapoteca H.M. Hern. and Archidendron F. Muell. Subsequent studies demonstrated that this alliance is polyphyletic and that Inga is probably sister to the genus Zygia ( Koenen et al. 2020a ; Ringelberg et al. 2022 ). The infrageneric taxonomy has changed importantly over the years. Bentham (1845 , 1865 , 1875 ) divided Inga into five sections: Leptinga Benth., Diadema Benth., Bourgonia Benth., Pseudinga Benth. and Euinga . Pennington (1997) remarked that the boundaries of sections and series are blurred, but that the series might be more natural than the sections. Pennington (1997) classified the genus into 14 sections, within which only informal species groups were recognised. The sections are largely based on several overlapping and quantitative characters and are difficult to separate with a simple key, probably reflecting the relatively recent origin and explosive radiation of the genus in Neotropical rainforests ( Pennington and Dick 2009 ; Dexter et al. 2017 ). Taxonomic references. Bentham (1845 , 1875 , 1876 ); Pennington (1997) ; Poncy (1985) ; Sousa (1993) .