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 Subfamily Caesalpinioideae DC., Prodr. [A. P. de Candolle] 2: 473. 1825. Caesalpiniaceae R. Br., in M. Flinders, Voy. Terra Austral. 2: 551. 1814. Type: Caesalpinia L. Type . Caesalpinia L. Description. Trees, shrubs, lianas, suffruticose or functionally herbaceous, occasionally aquatic, either unarmed or commonly armed with prickles, spines, or thorns; specialised extrafloral nectaries often present on the petiole and/or on the primary and secondary leaf rachides, usually between pinnae or leaflet pairs, more rarely stipular or bracteal. Stipules in lateral position and free or absent, usually entire, less frequently divided or spinescent. Leaves usually pulvinate, bipinnate, otherwise pinnate (sometimes both types on the same plant) and then mostly paripinnate, rarely imparipinnate, less often bifoliolate, modified into phyllodes or lacking, arrangement of the pinnae and leaflets mostly opposite, rarely alternate; stipels rare and not to be confused with the more commonly present paraphyllidia. Inflorescences globose or ellipsoid capitula, spicate, paniculate, racemose or in fascicles; bracteoles commonly small or absent. Flowers usually bisexual, rarely unisexual (species dioecious or monoecious), or bisexual flowers combined with unisexual and/or sterile flowers in heteromorphic inflorescences ( Mimoseae ), radially, less frequently bilaterally symmetrical, or asymmetrical; hypanthium lacking or cupular, rarely tubular; sepals (3) 5 (6-8), free or fused; petals (3) 5 (6-8), free or fused, the sepals or petals or both sometimes lacking, aestivation valvate ( Mimoseae ) or imbricate and then the adaxial petal innermost; stamens commonly diplostemonous or haplostemonous, sometimes reduced to 3 or 4 (in some Mimosa species), frequently polystemonous (to 100+ in some Mimoseae ), free or fused, sometimes heteromorphic, some or all sometimes modified or staminodial, anthers basifixed or dorsifixed, often with a stipitate or sessile apical gland, dehiscing via longitudinal slits or apical or basal poricidal slits or pores; pollen in tricolporate monads, or commonly in tetrads, bitetrads or polyads (most Mimoseae ); gynoecium uni- or rarely polycarpellate, 1-many-ovulate. Fruit typically dry and dehiscent, either a legume (dehiscent along both sutures) or a follicle (dehiscent along the adaxial suture only), or dry and segmented into one-seeded articles, either without a persistent margin (a lomentum) or with a persistent margins forming a replum like a frame (a craspedium), sometimes indehiscent and somewhat fleshy (an indehiscent legume), or dry and winged (a samara), when dehiscent with papery, leathery, or woody valves, dehiscence passive (inert valves), elastic, or explosive (the valves becoming curved, spirally coiled, or arched backwards), less frequently with entire valves, but with the endocarp segmented into one-seeded articles (a cryptolomentum). Seeds usually with an open (U-shaped) or closed (O-shaped) pleurogram on both faces or lacking a pleurogram, sometimes with a fleshy aril or sarcotesta, sometimes winged, in cross section terete (with a 1:1 ratio), compressed (with more or less 2:1 ratio) or flattened (with> 4:1 ratio; Gunn 1991 ); hilum usually apical, lens usually inconspicuous; embryo straight. Chromosome number. 2 n mostly 24, 26, 28, but also reported 2 n = 14, 16, 20, 22, 36, 52, 54, 56, 72, 78, 104, 112. Included taxa. Caesalpinioideae in its emended circumscription contains eleven tribes, 163 genera and ca. 4680 species. Tribes: Caesalpinieae Rchb. (27 genera / ca. 223 species), Campsiandreae LPWG (2 / 5-22), Cassieae Bronn (7 / 695), Ceratonieae Rchb. (4 / 6), Dimorphandreae Benth. (4 / 35), Erythrophleeae LPWG (2 / 13), Gleditsieae Nakai (3 / 20), Mimoseae Bronn (100 / ca. 3510), Pterogyneae LPWG (1 / 1), Schizolobieae Nakai (8 / 42-43), Sclerolobieae Benth. & Hook. f. (5 / ca. 113) (Fig. 4 ). Clade-based definition. The most inclusive crown clade containing Arcoa gonavensis Urb. and Mimosa pudica L., but not Bobgunnia fistuloides (Harms) J.H. Kirkbr. & Wiersema, Duparquetia orchidacea Baill., or Poeppigia procera C. Presl. Distribution and ecology. Pantropical, common in both wet and dry regions, with a handful of species extending to the temperate zone, less frequently frost-tolerant ( Gleditsia J. Clayton, Gymnocladus Lam. and some species of Acacia Mill., Desmanthus Willd. and Senna Mill.). Caesalpinioideae species are infrequent above 2500 m in the tropics and are largely absent from mid- and high-elevation tropical montane forests. Generic diversity is highest in the Neotropics, and there are important centres of high species diversity in Mexico and Central America, central-east South America, Africa, Madagascar, parts of South East Asia and Australia (Fig. 3 , Table 1 ). Notes. This clade was referred to as the Mimosoideae - Caesalpinieae - Cassieae (MCC clade) ( Doyle 2011 , 2012 ) or the Gleditsia - Chamaecrista - Mimosoideae (GCM clade) ( Marazzi et al. 2012 ). In LPWG (2017) , Caesalpinioideae was chosen over Mimosoideae as the preferred name for the MCC clade, leaving open the option for naming the morphologically distinct mimosoid clade at the tribal level, as is done here (Fig. 5 ; see Mimoseae , page 201). In the following treatments, the type species of different taxa are presented. Their homotypic (nomenclatural) synonyms are indicated by the identity symbol (≡) and heterotypic (taxonomic) synonyms by the equality symbol (=), as specified in the International Code of Nomenclature for algae, fungi, and plants ( Turland et al. 2018 ).