Gleditsieae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 253. 1943.

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

Tribe Gleditsieae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 253. 1943.

Figs 12 , 13 , 14 , 15 , 16 , 17

Type.

Gleditsia J. Clayton

Included genera

(3). Gleditsia J. Clayton (13 species), Gymnocladus Lam. (6), Umtiza Sim (1).

Description.

Deciduous or evergreen trees, with simple or branched thorns or unarmed; branches with or without lateral brachyblasts. Stipules inconspicuous ( Gleditsia ), foliaceous, caducous ( Gymnocladus ), or absent ( Umtiza ). Leaves bipinnate, pinnate, or intermediate with one or more pinnae replaced by a leaflet. Inflorescences panicles or racemes. Flowers regular, bisexual ( Umtiza ), or unisexual and then androdioecious or dioecious ( Gleditsia , Gymnocladus ), small, white or greenish to violet, mostly 5-merous; hypanthium short to elongate; calyx gamosepalous; petals small; stamens (6) 10, free; pollen in tricolporate monads, exine perforate, reticulate; ovary sessile or stipitate. Fruit compressed or turgid, papery, leathery or woody, dehiscent, tardily dehiscent or indehiscent, sometimes with pulpy interior, with few to 25 (40) seeds. Seeds compressed to subterete, orbicular or ovoid-elliptic.

Distribution.

Temperate to subtropical North America, South America, Asia, and South Africa.

Clade-based definition.

The most inclusive crown clade containing Umtiza listeriana Sim and Gymnocladus dioicus (L.) K. Koch, but not Ceratonia siliqua L., Dimorphandra conjugata (Splitg.) Sandwith or Mimosa sensitiva L. (Fig. 12 ).

Notes.

Tribe Gleditsieae was first named by Nakai (1943) to accommodate Gleditsia and Gymnocladus . A close relationship between the two genera has long been known ( Polhill and Vidal 1981; Polhill 1994; Lewis 2005b). Gleditsia and Gymnocladus were resolved together with Umtiza , as well as with Tetrapterocarpon Humbert, Arcoa Urb., Ceratonia L. and Acrocarpus Wight ex Arn., in a clade that was informally named the Umtiza clade ( Herendeen et al. 2003a, 2003b). The close relationship of Gleditsia , Gymnocladus and Umtiza has been confirmed by Manzanilla and Bruneau (2012), LPWG (2013, 2017) and Ringelberg et al. (2022), but the monophyly of the larger Umtiza clade is not supported (Fig. 12 ). Umtiza was previously placed in the Cynometra group of tribe Detarieae by Cowan and Polhill (1981) with some uncertainty. The small regular flowers and thorns in Umtiza are notable morphological similarities to Gleditsia . The three genera differ most in fruit structure and dehiscence.

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 12. Generic relationships in tribe Gleditsieae. For description of phylogeny and support values, see Fig. 6 caption (page 63).

Figure 13. Gleditsieae, diversity of vegetative morphology and fruits A-C Umtiza listeriana Sim A small tree B branch with pinnately compound leaves and inflorescence C branch with thorn at node D Gymnocladus assamicus Kanjilal ex P. C. Kanjilal branch with bipinnate leaves and inflorescence E Gleditsia japonica Miq., bipinnate leaf with several pinnae replaced by single leaflets, cultivated at the Royal Botanic Gardens, Kew F Gleditsia amorphoides (Griseb.) Taub., branched thorns G Umtiza listeriana dehisced fruits H Gymnocladus dioicus L., fruit I Gleditsia triacanthos L., fruits. Photo credits A-C, G SAplants, Wikimedia Commons (CC-BY-SA 4.0) D B Choudhury E, F GP Lewis H Chicago Botanic Garden (photographer unknown) I A Schnabel.

Figure 14. Gleditsieae, floral diversity A, B Umtiza listeriana Sim A branch with inflorescence of hermaphrodite flowers B close-up of flowers C, D Gymnocladus assamicus Kanjilal ex P. C. Kanjilal C inflorescence of hermaphrodite flowers D dissected hermaphrodite flower E Gymnocladus chinensis Baill., inflorescence, cultivated at the US National Arboretum F Gymnocladus dioicus L., cultivated at Chicago Botanic Garden, upright inflorescence of hermaphrodite flowers G branch with leaves and inflorescence H Gleditsia triacanthos L., inflorescence of staminate flowers I Gleditsia caspica Desf., inflorescence, cultivated at the Royal Botanic Gardens, Kew. Photo credits A, B SAplants, Wikimedia Commons (CC-BY-SA 4.0) C, D Murata E PS Herendeen F, G R Carlson H Royal Botanic Gardens, Kew (photographer unknown) I M Svanderlik.

Figure 15. Distribution of Umtiza based on quality-controlled digitised herbarium records. See Suppl. material 1 for the source of occurrence data.

Figure 16. Distribution of Gymnocladus based on quality-controlled digitised herbarium records. See Suppl. material 1 for the source of occurrence data.

Figure 17. Distribution of Gleditsia based on quality-controlled digitised herbarium records. See Suppl. material 1 for the source of occurrence data.