Hymenolobium mesoamericanum, copyright Rolando Pérez.

Belongs within: Leguminosae.
Contains: Dillwynia, Jacksonia, Dipterygeae, Amburaneae, Swartzieae, Exostyleae, Amorpheae, Dalbergieae, Luetzelburgia, Ormosieae, Brongniartieae, Leptolobieae, Sophoreae, Genisteae, Crotalarieae, Baphieae, Mirbelieae, Hologalegina, Indigofera, Millettieae, Diocleae, Kennediinae, Desmodieae, Mucuna, Cajaninae, Erythrininae, Phaseolinae, Glycine, Psoraleeae.

The Papilionoideae are a major clade within the Fabaceae/Leguminosae, alternatively recognised as the Fabaceae sensu stricto or Papilionaceae. Members of a number of genera within the Papilionoideae are significant food crops (such as Glycine, soybean, and Cicer arietinum, chickpea) whereas others are incorporated into pastures to enrich the soil through their nitrogen-fixing nodules (such as Trifolium, clover, and Lotus).

Members of the Papilionoideae are characterised by the presence of ‘pea’ flowers with petals differentiated into a standard, wings and keel. The degree of petal differentiation may differ between taxa and past classifications have differentiated between ‘advanced’ taxa, with higly differentiated flowers, and ‘primitive’ taxa with more radial flowers. However, more recent phylogenetic analyses have indicated that the so-called ‘primitive’ taxa are polyphyletic and do not form a coherent grouping (Cardoso et al. 2013). The majority of members of the group are herbaceous though some are shrubs or trees. These woody taxa include the more basally diverging subgroups, outside the so-called ’50-kb inversion clade’. This clade, including all papilionoids except the Angylocalyceae, Dipterygeae, Amburaneae, Swartzieae and a small clade of taxa related to Cladrastis, is characterised by a large inversion in the plastid genome.

The Angylocalyceae are a disjunctly distributed tropical group characterised by mostly ornithophilous flowers with the calyx and hypanthium enlarged, the petals thickened, the standard often distinctly large, the lower petals undifferentiated or reduced, and the stamens and gynoecium exserted. The Angocalyceae, Dipterygeae and Amburaneae form a morphologically diverse clade that is mostly supported by moecular data though some members of the Dipterygeae and Amburaneae do share features of resin production. The Cladrastis clade are a group of trees and shrubs with papilionate flowers, united by molecular data, that have a disjunct distribution in eastern Asia and southern North America (Cardoso et al. 2013).

Basal relationships within the 50-kb inversion clade are poorly resolved (Cardoso et al. 2013). Amphimas pterocarpoides, bokanga, is a West African species of tree used as a source of wood. Dermatophyllum, mescalbeans, is a genus of highly toxic shrubs and small trees bearing purple flowers in dense racemes, found in southern North America.

Characters (from the Angiosperm Phylogeny Website): N-fixing common; isoflavonoids, prenylated flavonoids, pyrrolizidine, indolizidine, and quinolizidine alkaloids present; sieve tubes with spindle-shaped non-dispersive protein bodies; leaves once compound, palmate or pinnate; androecium with unidirectional [abaxial to adaxial] initiation, adaxial-median corolla external; androecium usually connate [e.g. 9 + 1]; pleurogram absent, linea fissura absent, hour-glass cells present, counter palisade present or absent, raphe shorter than the antiraphe, hilum with a hilar groove, tracheid bar [group of tracheids just below surface of hilum] present; embryo curved, with well-developed suspensor, radicle long, cotyledons accumbent, not investing radicle, cotyledon areole present; duplication of CYC gene.

Papilionoideae [Faboideae, Papilionaceae, Papilionatae, Phaseoleae]
    |  |  `--AmburaneaeCP13
    |  `--AngylocalyceaeCP13
    |       |  i. s.: UleanthusCP13
    |       |--+--Xanthocercis zambesiacaCP13
    |       |  `--Angylocalyx talbotiiCP13
    |       `--+--Castanospermum australeCP13
    |          `--AlexaCP13
    |               |--A. bauhiniifloraCP13
    |               |--A. canaracunensisCP13
    |               |--A. grandifloraCP13
    |               `--A. wachenheimiiCP13
       `--+--+--‘Cladrastis’ platycarpaCP13
          |  `--+--Pickeringia montanaCP13
          |     |    |--P. m. var. montanaCP13
          |     |    `--P. m. var. tomentosaCP13
          |     `--+--CladrastisCP13
          |        |    |--C. delavayiCP13
          |        |    |--C. kentukeaCP13
          |        |    `--C. luteaCP13
          |        `--StyphnolobiumCP13
          |             |--S. affineCP13
          |             |--S. japonicumCP13
          |             `--+--S. conzattiiCP13
          |                `--+--S. burseroidesCP13
          |                   `--S. monteviridisCP13
          `--+--Amphimas pterocarpoidesCP13
             |  `--DalbergieaeCP13
             |    |--D. arizonicumCP13
             |    `--D. secundiflorumCP13
             |    |--A. insignisCP13
             |    `--+--A. heterophyllaCP13
             |       `--A. latifoliaCP13
             |--+--Sweetia fruticosaCP13
             |  |--VataireopsisCP13
             |  |    |--V. ararobaCP13
             |  |    |--V. speciosaCP13
             |  |    `--V. surinamensisCP13
             |  `--+--LuetzelburgiaCP13
             |     `--VataireaCP13
             |          |--+--V. heteropteraCP13
             |          |  `--V. macrocarpaCP13
             |          `--+--+--V. fuscaCP13
             |             |  `--V. lundeliiCP13
             |             `--+--V. erythrocarpaCP13
             |                |--V. guianensisCP13
             |                |--V. paraensisCP13
             |                `--V. sericeaCP13
             |  |    |  i. s.: A. fraxinifoliaCP13
             |  |    |--A. carvalhoiCP13
             |  |    |--A. humilisCP13
             |  |    |--A. legalisCP13
             |  |    |--A. marauensisCP13
             |  |    `--+--A. inermisCP13
             |  |       `--+--A. galeottianaCP13
             |  |          `--A. ormosioidesCP13
             |  `--HymenolobiumCP13
             |       |--H. alagoanumCP13
             |       |--H. grazielanumCP13
             |       |--H. heringerianumCP13
             |       |--H. janeirenseCP13
             |       |--H. mesoamericanumCP13
             |       |--H. petraeumCP13
             |       `--+--H. heterocarpumCP13
             |          `--H. sericeumCP13
             |  `--+--‘Clathrotropis’ macrocarpaCP13
             |     `--+--BrongniartieaeCP13
             |        |--LeptolobieaeCP13
             |        `--+--Camoensia [Camoensieae]CP13
             |           |    |--C. brevicalyxCP13
             |           |    `--C. scandensCP13
             |           `--+--SophoreaeCP13
             |              `--+--+--GenisteaeCP13
             |                 |  `--CrotalarieaeCP13
             |                 `--Podalyrieae [Liparieae]CP13
             |                      |--Cadia purpureaCP13
             |                      |--Calpurnia aureaCP13
             |                      |--AmphithaleaCP13
             |                      |--CyclopiaCP13
             |                      |--LipariaCP13
             |                      |--Podalyria albaCP13, J23
             |                      |--StirtonanthusCP13
             |                      |--Virgilia oroboidesCP13, B00
             |                      `--XiphotecaCP13
                   |  `--Hypocalyptus [Hypocalypteae]CP13
                   |       `--H. coluteoidesCP13
                         |    |--Phylloxylon spinosaCP13
                         |    `--+--IndigoferaCP13
                         |       `--+--Microcharis karinensisCP13
                         |          `--CyamopsisCP13
                         |               |--C. senegalensisCP13
                         |               `--C. tetragonolobaZ94
                         `--+--+--Ostryocarpus ripariusCP13
                            |  `--+--Aganope thyrsifloraCP13
                            |     `--Xeroderris stuhlmanniiCP13
                               |    |--Dalbergiella nyasaeCP13
                               |    `--+--CentrosemaCP13
                               |       |    |--C. pubescensM99
                               |       |    `--C. sagittatumCP13
                               |       `--ClitoriaCP13
                               |            |--C. australisLK14
                               |            |--C. racemosaA96
                               |            |--C. rubiginosaBMM99
                               |            `--C. ternateaCP13
                                  |    |--A. blackiiCP13
                                  |    `--A. glabristaWO95
                                  `--+--+--Disynstemon paullenoidesCP13
                                     |  `--+--Abrus [Abreae]CP13
                                     |     |    |--A. precatoriusCP13
                                     |     |    `--A. pulchellusSR07
                                     |     `--+--MillettieaeCP13
                                     |        `--DiocleaeCP13
                                     `--+--Platycyamus regnelliiCP13
                                        `--+--Kunstleria ridleyiCP13
                                              |    |--A. americanaCP13
                                              |    `--A. tuberosaC55
                                              `--+--+--Shuteria vestitaCP13
                                                 |  `--KennediinaeCP13
                                                 |  `--+--Craspedolobium schochiiCP13
                                                 |     `--MucunaCP13
                                                    |  `--ButeaCP13
                                                    |       |--B. frondosaZ94
                                                    |       |--B. monospermaCP13
                                                    |       |    |--B. m. var. monospermaS02
                                                    |       |    `--B. m. var. luteaS02
                                                    |       |--B. parvifloraP03
                                                    |       `--B. superbaP03
                                                       `--+--+--Cologania pallidaCP13
                                                          |  `--PhaseolinaeCP13
                                                          `--+--+--Calopogonium mucunoidesCP13, BMM99
                                                             |  `--Pseudovigna argenteaCP13
                                                             |--+--Neorautanenia mitisCP13
                                                             |  `--Pachyrhizus erosusCP13
                                                             `--+--Neonotonia wightiiCP13
                                                                `--+--+--Amphicarpaea bracteataCP13
                                                                   |  `--+--Teramnus uncinatusCP13
                                                                   |     `--+--GlycineCP13
                                                                   |        `--PsoraleeaeCP13
                                                                        |--P. hirsutaAP80
                                                                        |--P. lobata (Willd.) Ohwi 1947SY72 (see below for synonymy)
                                                                        |--P. montanaCP13
                                                                        |--P. trilobaBMM99
                                                                        `--P. tuberosaSR07
Papilionoideae incertae sedis:
    |--E. microphyllaB00
    |--E. parvifoliaG04
    `--E. virgataGK00
    |--C. aciculareOS04
    |--C. ericifoliumK90
    |--C. ilicifoliumGK00
    `--C. nanumGK00
  Nemcia obovataOS04
    |--B. aphyllaK90
    |--B. daviesioidesK90
    |--B. macrocarpumK90
    |--B. melanopetalumGK00
    `--B. praemorsumGK00
  Callistachys lanceolataGK00
    |--O. atropurpureumD84
    |--O. cordifoliumM89
    |--O. ellipticumKC83
    |--O. ilicifoliumM65
    |--O. lineareGK00
    `--O. trilobatumH87
    |--S. drummondiiGK00
    |--S. linophyllumGK00
    |--S. macranthumGK00
    |--S. mediumGK00
    `--S. vimineumGK00
  Goodia lotifoliaGK00
  Viminaria junceaGK00
  Latrobea tenellaGK00
  Atylosia scarabaeoidesP03
  Maughania bracteataP03
  Pseudathria viscidaP03
  Sarothamnus scopariusH59
  Psoralidium lanceolatum [incl. Psoralea lanceolata ssp. scabra]H93
  Corallodendron Mill. 1754KC01
  Genistella Ortega 1773KC01
  Sterbeckia Schreb. 1789KC01
  Xerocarpus Guill., Perr. & Rich. 1832KC01
  Goniogyna DC. 1825 [incl. Heylandia DC. 1825]SY72
    `--G. hirta (Willd.) Ali 1967 [=Hallia hirta Willd. 1803; incl. Heylandia latebrosa DC. 1825]SY72
  Eleotis monophylla [incl. E. sororia]VS73
  Moghania paniculataVS73, M72
    |--S. bracteolatusFJ99
    |--S. gyrocarpusK03
    `--S. littoralisA78
Nomen nudum: Nemcia aspera Crisp in Obbens & Sage 2004OS04

Pueraria lobata (Willd.) Ohwi 1947SY72 [=Dolichos lobatus Willd. 1803SY72, P. montana var. lobataH06; incl. Pachyrrhizus thunbergianus Sieb. & Zucc. 1846SY72, Pueraria thunbergiana (Sieb. & Zucc.) Benth. 1865SY72]

*Type species of generic name indicated


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