Campsiandreae Legume Phylogeny Working Group, tribus nov.

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 Campsiandreae Legume Phylogeny Working Group, tribus nov.

Figs 94 , 95 , 96 , 97

Diagnosis.

Campsiandreae is characterised by a combination of flowers with a cupular hypanthium, imbricate petals and stamens long-exserted from the corolla. The genus Dinizia is similar in appearance to genera of tribes Dimorphandreae and Mimoseae by its bipinnate leaves and small flowers in dense spicate racemes but differs by the combination of alternate leaflets, imbricate petal aestivation and all stamens fertile (in Mimoseae petals are valvate and genera of Dimorphandreae with bipinnate leaves and spicate inflorescences have opposite leaflets and an androecium comprising 5 fertile stamens and 5 staminodes). The genus Campsiandra is similar in appearance to Arapatiella Rizzini & A. Mattos (tribe Sclerolobieae ) in having paripinnate leaves and regular perigynous flowers with a turbinate hypanthium and exserted stamens but differs by the variable number of stamens (vs. stamens 10 in Arapatiella ), fruits indehiscent or inertly dehiscent, the valves not strongly twisting after dehiscence (vs. fruits elastically dehiscent from the apex, the valves rolling inwards in Arapatiella ).

Type.

Campsiandra Benth.

Included genera

(2). Campsiandra Benth. (3 to ca. 20 species), Dinizia Ducke (2).

Description.

Medium to large trees. Stipules caducous or lacking. Leaves pinnate with opposite, often gland-dotted leaflets, or bipinnate with alternate, eglandular leaflets; extrafloral nectaries absent. Inflorescence a compound spiciform raceme, or showy multi-flowered panicle. Flowers hermaphrodite or functionally staminate, pedicellate (the pedicels up to 3 cm long in Campsiandra ); calyx with 5 imbricate lobes or a tube with 5 broadly triangular lobes; petals 5, imbricate, whitish (sometimes with rose-reddish markings), cream coloured or yellow; stamens 10-17 (25) per flower, exserted from the corolla, anthers eglandular; pollen in monads or tetrads; ovary stipitate. Fruit laterally compressed, coriaceous or woody, inertly dehiscent or indehiscent. Seeds discoid with a marginal spongy wing, or elliptic to obovate, hard, and wingless.

Distribution.

South America, mainly in the Amazon and Orinoco basins in flooded forests and swamp forests ( Campsiandra ) or in non-flooded Amazonian forests ( Dinizia excelsa Ducke), or semi-deciduous Atlantic rainforest ( Dinizia jueirana-facao G.P. Lewis & G.S. Siqueira).

Clade-based definition.

The most inclusive crown clade containing Campsiandra laurifolia Benth. and Dinizia excelsa Ducke, but not Delonix decaryi (R. Vig.) Capuron, Dimorphandra conjugata (Splitg.) Sandwith or Mimosa sensitiva L. (Fig. 94 ).

Notes.

Phylogenetic analyses of a few molecular markers had suggested that the genus Dinizia could be more closely related to the Dimorphandra group of the old sense subfamily Caesalpinioideae , but with low support ( Luckow et al. 2000, 2003; Wojciechowski et al. 2004; Bruneau et al. 2008; LPWG 2017). The position of Dinizia as sister to the Amazonian genus Campsiandra is more clearly supported in the phylogenomic analyses of Zhang et al. (2020) and Ringelberg et al. (2022), however both genera are subtended by a long branch. Campsiandra comosa Benth. and Dinizia jueirana-facao are confirmed to nodulate with a fixation thread type of nodule anatomy ( Faria et al. 2022). However, Dinizia excelsa is reported to be non-nodulating ( Sprent 2001; Lewis et al. 2017), making Dinizia one of the few Caesalpinioideae genera presently known to contain both nodulating and non-nodulating species.

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 94. Generic membership and phylogenetic position of tribe Campsiandreae. For description of phylogeny and support values, see Fig. 6 caption (page 63).

Figure 95. Flower, fruit and vegetative characters of tribe Campsiandreae A foliage and fruits, Campsiandra comosa Benth., Brazil B compound inflorescence, Campsiandra angustifolia Spruce ex Benth., Peru C fruit and seed, Campsiandra laurifolia Benth., cultivated Rio de Janeiro Botanic Gardens, Brazil D inflorescence and foliage, Campsiandra sp., Amazonas, Brazil E-I Dinizia jueirana facao G. P. Lewis & G. S. Siqueira, Reserva Natural Vale, Espirito Santo, Brazil E foliage and fruits F trunk and crown of mature tree G inflorescences H mature fruits (hand for scale) I rough bark of trunk. Photo credits A D Cardoso B T Pennington D M Falcao E, G, I D Folli C, F, H GP Lewis.

Figure 96. Distribution of Campsiandra based on quality-controlled digitised herbarium records. See Suppl. material 1 for the source of occurrence data.

Figure 97. Distribution of Dinizia based on quality-controlled digitised herbarium records. See Suppl. material 1 for the source of occurrence data.