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 Chamaecrista (L.) Moench, Methodus: 272. 1794. Figs 22 , 28 , 29 Cassia [infragen. unranked] Chamaecrista L., Sp. Pl. 1: 379. 1753. Type: Cassia chamaecrista L., nom. utique rejic. [≡ Cassia fasciculata Michx. (≡ Chamaecrista fasciculata (Michx.) Greene)] Cassia sect. Chamaecrista (L.) DC., Hist. Nat. Med . Casses 24, 118.1816. Lectotype (designated by Britton and Rose 1930 ): Cassia nictitans L. [≡ Chamaecrista nictitans (L.) Moench] Sooja Siebold, Verh. Batav. Genootsch. Kunst. 12: 56. 1830. Type: Sooja nomame Siebold, nom. inval. (nom. nud.) [≡ Cassia mimosoides var. nomame Makino (≡ Chamaecrista nomame (Makino) H. Ohashi)] Disterepta Raf., Sylva Tellur.: 126. 1838. Type: Disterepta pilosa (L.) Raf. [≡ Cassia pilosa L. (≡ Chamaecrista pilosa (L.) Greene)] Hepteireca Raf., Sylva Tellur.: 126. 1838. Type: Hepteireca glandulosa (L.) Raf. [≡ Cassia glandulosa L. (≡ Chamaecrista glandulosa (L.) Greene)] Dialanthera Raf., Sylva Tellur.: 127. 1838. Type: Dialanthera glandulosa (L.) Raf. [≡ Cassia glandulosa L. (≡ Chamaecrista glandulosa (L.) Greene)] Xamacrista Raf., Sylva Tellur.: 127. 1838. Type: Xamacrista trifolia Raf. [= Cassia chamaecrista L. (≡ Chamaecrista fasciculata (Michx.) Greene)] Nictitella Raf., Sylva Tellur.: 128. 1838. Lectotype (designated by Irwin & Barneby, 1982): Nictitella amena Raf. [= Cassia nictitans L. (≡ Chamaecrista nictitans (L.) Moench)] Ophiocaulon Raf., Sylva Tellur.: 129. 1838. Lectotype (designated by Irwin & Barneby, 1982): Ophiocaulon serpens (L.) Raf. [≡ Cassia serpens L. (≡ Chamaecrista serpens (L.) Greene)] Cassia subg. Lasiorhegma Vogel ex Benth., Fl. Bras. 15(2): 129. 1870. Type not designated. Type . Chamaecrista nictitans (L.) Moench [≡ Cassia nictitans L.] Description. Trees, treelets, shrubs and subshrubs, lacking spines or prickles. Stipules diverse in shape and size, persistent or caducous. Leaves distichous or spiral, bifoliolate or paripinnate; extrafloral nectaries when present on petiole, generally on the rachis between the pairs of leaflets or in the axis of the inflorescence, sessile or stipitate, the secretory surface concave, rarely convex; leaflets 1-65 pairs. Inflorescence a fascicle, raceme or panicle; bract 1, caducous or persistent, bracteoles 2, alternate, located at mid-length or slightly above the pedicels, persistent. Flowers hypogynous, asymmetrical, hypanthium absent; sepals 5, free; petals 5, free, yellow, or yellow with red base, sometimes red, orange or pink; stamens 5-10, homomorphic, filaments glabrous, anthers dehiscent by apical pores, pubescent laterally, rarely with the indumentum covering the entire anther; pollen subprolate to prolate, syncopate, fused at the poles; ovary stipitate. Fruit a compressed legume, valves papyraceous or coriaceous, elastically dehiscent through both margins, becoming twisted after dehiscence. Seeds variable in shape and colour. Chromosome number. Haploid numbers n = 7, 8, 14, 16, 24 ( Goldblatt and Johnson 1979 -; Souza 2004 ). Included species and geographic distribution. 368 species, pantropical with a few species reaching temperate areas (Fig. 28 ). Figure 28. Distribution of Chamaecrista based on quality-controlled digitised herbarium records. See Suppl. material 1 for the source of occurrence data. Ecology. Chamaecrista species typically occur in open environments. Although several species are widespread, such as C. rotundifolia (Pers.) Greene, C. mimosoides (L.) Greene and C. flexuosa (L.) Greene, a high diversity is concentrated in Brazilian savannas and in the "campos rupestres'' vegetation ( Irwin and Barneby 1978 , 1982 ; Rando et al. 2020b ). In these centers of diversity, several species have evolved underground systems that allow survival after fire and during long dry periods ( Rando et al. 2016 ). One clade of arborescent Chamaecrista species is mostly restricted in the Amazon and Atlantic tropical rainforests. Figure 29. Examples of Chamaecrista diversity A Chamaecrista xinguensis (Ducke) H.S. Irwin & Barneby trunk of mature individual ( Rando et al. 1208 ) B C. compitalis (H.S. Irwin & Barneby) H.S. Irwin & Barneby base of trunk of mature individual ( Rando et al. 1364 ) C C. ensiformis (Vell.) H.S. Irwin & Barneby flowering branch ( Rando & Cota 1366 ) D C. flexuosa (L.) Greene flower, leaves in background E C. desvauxii var. latistipula (Benth.) G.P. Lewis, branch with flowers and fruit F C. ramosa var. curvifolia (Vogel) G.P. Lewis branches and flower G C. distichoclada (Benth.) H.S. Irwin & Barneby flowering branch ( Rando et al. 1230 ) H C. lineata (Sw.) Greene leaves and flower ( Rando 964 ) I C. andromedea (Mart. ex Benth.) H.S. Irwin & Barneby branch with leaves and flowers ( Rando et al. 1251 ) J C. vauthieri (Benth.) H.S. Irwin & Barneby flowering branches ( Cardoso et al. 4096 ) K C. ochnacea var. purpurascens (Benth.) H.S. Irwin & Barneby inflorescence and leaves L C. scabra (Pohl ex Benth.) H.S. Irwin & Barneby leaves and inflorescence ( Rando et al. 1266 ). Photo credits A, C, G-I JG Rando B JG Jardim D-F, K H Moreira J D Cardoso L MF Simon. Human uses. Some species are used in traditional African medicine. For example, C. absus (L.) H.S. Irwin & Barneby is used as a purgative, for treating wounds and sores, and also against syphilis ( Lewis 2005a ). In China and Japan, C. mimosoides is used as a tea, and in Tanzania, against snake bites and scorpion stings ( Lewis 2005a ). In Brazil, dried leaflets and branches of some species [ C. choriophylla (Vogel) H.S. Irwin & Barneby, C. cotinifolia (G.Don) H.S. Irwin & Barneby, C. orbiculata (Benth.) H.S. Irwin & Barneby and C. rotundata (Vogel) H.S. Irwin & Barneby] are used as decorative objects ( Cota et al. 2020 ). Etymology. A composite name from the Greek Chamae (= small, of little growth), and the Latin crista , referring to the crest ( Rizzini and Rizzini 1983 ; Radcliffe-Smith 1998 ). The name was applied in reference to the very short filaments of stamens forming a crest ( Greene 1905 ). Notes. The largest genus of the tribe can be easily recognised by a set of features: the presence of two bracteoles on the pedicels, stamens generally homomorphic, poricidal anthers (Fig. 22 ) and elastically dehiscent fruits. Since the segregation of Chamaecrista from Cassia , the infrageneric classification of the genus remains the subject of intensive studies, changing the rank, expanding, restricting, or combining the names within the genus (e.g., Bentham 1871 ; Irwin 1964 ; Irwin and Rogers 1967 ; Irwin and Barneby 1976a , 1976b , 1977 , 1978 , 1982 ; Rando et al. 2016 ; Souza et al. 2021 ). In short, three highly supported clades with strong correlation in habit and habitat variation are currently well established in Chamaecrista : (i) a clade of arborescent species, with ramiflorous inflorescences and extrafloral nectaries; (ii) a clade of shrubs, with axillary and reduced racemes, distichous phyllotaxy, and with extrafloral nectaries; and (iii) the most diverse clade, embracing shrubs with terminal racemes or panicles, spiral phyllotaxy, without extrafloral nectaries, and commonly with glandular trichomes on branches, leaves and inflorescences ( Conceicao et al. 2009 ; Souza et al. 2021 ). Within these groups there are also some highly supported clades, consistent with morphology, and the trend has been to improve their circumscription to arrive at a stable and practical infrageneric classification. However, more molecular phylogenetic studies are needed to clarify relationships in the most diverse clades (ca. 200 species in savannas and rocky fields), in which recent diversification (ca. 5 Ma; Rando et al. 2016 ; Vasconcelos et al. 2020 ) complicates the understanding of relationships among species. Taxonomic references. Bentham (1871) ; Buril et al. (2011) ; Cota et al. (2020) ; Irwin (1964) ; Irwin and Barneby (1976a , 1976b , 1977 , 1978 , 1981 , 1982 ); Irwin and Rogers (1967) ; Rando et al. (2016 , 2020b ); Souza et al. (2021) .