FABALES Bromhead Main Tree, Synapomorphies

Habit Usually bisexual (rarely monoecious, andromonoecious or polygamomonoecious), evergreen or deciduous trees, shrubs, lianas or suffrutices, perennial, biennial or annual herbs.

Vegetative anatomy Root nodules containing nitrogen fixing bacteria usually present. Phellogen ab initio superficially or deeply seated. Primary vascular tissue cylinder without separate vascular bundles, or cylinder of bundles. Secondary lateral growth normal or anomalous from concentric cambia, or absent. Vessel elements usually with simple (rarely scalariform) perforation plates; lateral pits alternate, simple or bordered pits. Vestured pits present. Imperforate tracheary xylem elements libriform fibres or tracheids (sometimes fibre tracheids) with usually simple (sometimes bordered) pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, reticulate, unilateral, vasicentric, or banded. Sieve tube plastids Ss, Pc or Pcs type (rarely Pfs type). Nodes 1:1 or 1:3, unilacunar with one or three leaf traces, or 3:3, trilacunar with three traces (sometimes 5:5, pentalacunar with five traces). Secretory cavities often present. Heartwood often with resins or resin-like substances. Silica bodies present or absent. Calciumoxalate as prismatic or acicular crystals, druses, styloids, or crystal sand.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple or branched, furcate or stellate, sometimes dendritic; pearl glands, stalked multicellular glands and/or glandular hairs (sometimes lepidote) often present; stinging hairs rare.

Leaves Usually alternate (spiral or distichous; rarely opposite or verticillate), paripinnate, imparipinnate or trifoliolate (sometimes bipinnate or multifoliolate, rarely unifoliolate), usually with entire (rarely lobate or serrate) opposite conduplicate leaflets (rarely alternate), or scales, simple, entire, with conduplicate ptyxis. Stipules intrapetiolar, often foliaceous or modified into spines or glands, caducous or persistent; leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Venation pinnate, usually brochidodromous, or palmate. Stomata anomocytic, paracytic, anisocytic, tetracytic, parallelocytic or cyclocytic. Cuticular wax crystalloids usually as rosettes of platelets (Fabales type; sometimes as scales or granules). Epidermis with or without mucilaginous idioblasts. Mesophyll often with sclerenchymatous idioblasts (with fibres or branched sclereids). Secretory cavities with ethereal oils, mucilage or resin often present. Leaf margin and leaflet margins serrate or entire.

Inflorescence Terminal or axillary, corymb, panicle, botryoid, raceme, spike, head, or flowers solitary axillary.

Flowers Usually zygomorphic (sometimes actinomorphic). Hypogyny. Sepals (three to) five (or six), with valvate or imbricate aestivation, free or more or less connate, with helical initiation. Petals (one to) three or five (rarely four), with valvate, imbricate or cochlear-descending aestivation, often clawed, free or more or less connate. Nectariferous disc intrastaminal, annular or unilateral, or absent.

Androecium Stamens (two or) five to ten (to numerous), in one or two (or three) whorls. Filaments free or more or less connate into tube, usually free from tepals (sometimes epipetalous). Anthers basifixed or dorsifixed, versatile or non-versatile, usually tetrasporangiate (rarely disporangiate), introrse or latrorse, usually longicidal (dehiscing by longitudinal slits) or poricidal (dehiscing by apical or subapical pores or very short slits). Tapetum usually secretory. Staminodia usually absent (sometimes four or five or more).

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3–23(–33)-colporate (sometimes porate, colporoidate, colpate, rugate or inaperturate, rarely syncolporate), usually shed as monads (sometimes tetrads or polyads), usually bicellular (sometimes tricellular) at dispersal. Exine tectate or semitectate, with usually columellate infratectum, perforate, microreticulate, reticulate, punctate, striate, or rugulate, gemmate, verrucate, granulate, fossulate, foveolate or psilate.

Gynoecium Pistil composed of usually one carpel (sometimes two to 16 free carpels), or two to eight connate (rarely entirely or partially free) carpels. Ovary superior, usually unilocular (monomerous) or bilocular to quinquelocular. Style single, simple or bilobate, often hollow (rarely five free stylodia). Stigma capitate or bilobate (sometimes complex), papillate or non-papillate, Dry or Wet type. Pistillodium usually absent (male flowers sometimes with pistillodium).

Ovules Placentation marginal-ventral, axile or parietal. Ovules one to more than 20 per carpel, usually anatropous, anacampylotropous or campylotropous (sometimes hemianatropous or amphitropous, rarely pleurotropous), pendulous, horizontal or ascending, epitropous, usually bitegmic (rarely unitegmic), usually crassinucellar (rarely tenuinucellar). Micropyle bistomal or endostomal (rarely exostomal). Nucellar cap sometimes present. Megagametophyte usually monosporous, Polygonum type (rarely disporous, Allium type). Synergids often with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustorium usually chalazal (sometimes also micropylar; rarely lateral) or absent. Embryogenesis onagrad, asterad or caryophyllad.

Fruit Usually a loculicidal capsule (often a legume; sometimes a nut, samara, drupe, berry, follicle, schizocarp, or syncarp).

Seeds Aril often present. Elaiosome or carunculus sometimes present. Seed coat exotestal or endotestal. Testa sometimes multiplicative. Exotesta usually palisade (often with lignified malpighian cells, polygonal in cross-section, with linea lucida (light line) separating heavily thickened outer anticlinal cell walls). Mesotesta consisting of stellate cells or crushed. Endotestal cells palisade, crystalliferous, or crushed. Tegmen usually crushed. Perisperm not developed. Endosperm absent or sparse (sometimes copious). Embryo straight or curved, well differentiated, usually with chlorophyll. Cotyledons two, often well developed and nutritious. Germination phanerocotylar or cryptocotylar.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavone-C-glycosides, afzelechin, 5-deoxyflavonoids, isoflavonoids (pterocarpanes, isoflavanes), cyanidin, epigallocatechin-3-gallate, oleanolic acid derivatives, dammarane, surianol, methylated ellagic acids, ellagitannins (e.g. geraniin), proanthocyanidins (prodelphinidins), pyridine alkaloids, quinolizidine alkaloids, indole alkaloids, loline alkaloids, pyrrolizidine alkaloids as macrocyclic diesters, triterpene saponins, tyrosine-, leucine- or phenylalanine-derived cyanogenic compounds, anthraquinones, pinitol (cyclitol), simmondsin-like compounds, lignans (syringaresinol), toxic non-protein amino acids, and lectins (hemagglutinins, especially in seeds) present. Non-methylated ellagic acids not found.

The sister-group relationships within Polygalales are not unambiguously resolved and the bootstrap or bayesian support for any branch is more or less weak. Hence, Fabaceae, Polygalaceae, Quillajaceae or Surianaceae may be identified as sister to the remainder, depending on the characters and analysis methods used.

Lindley in Edwards’s Bot. Reg. 22: ad t. 1845. 1 Apr 1836 [‘Leguminosae, or Fabaceae’], nom. cons.

LeguminosaeJuss., Gen. Plant.: 345. 4 Aug 1789, nom. cons. et nom. alt.; PapilionaceaeGiseke, Prael. Ord. Nat. Plant.: 415. Apr 1792, nom. cons. et nom. alt.; CaesalpiniaceaeR. Br. in M. Flinders, Voy. Terra Austral. 2: 551. 19 Jun 1814 [’Lomentaceaevel Caesalpineae’], nom. cons.; MimosaceaeR. Br. in M. Flinders, Voy. Terra Austral. 2: 551. 19 Jul 1814 [’Mimoseae’], nom. cons.; CassiaceaeVest, Anleit. Stud. Bot.: 270, 291. 1818 [’Cassioideae’]; RobiniaceaeVest, Anleit. Stud. Bot.: 270, 291. 1818 [’Robinioideae’]; AspalathaceaeMartinov, Tekhno-Bot. Slovar: 51. 3 Aug 1820 [’Aspalathoides’]; AstragalaceaeBercht. et J. Presl, Přir. Rostlin: 229. Jan-Apr 1820 [’Astragalides’]; BauhiniaceaeMartinov, Tekhno-Bot. Slovar: 67. 3 Aug 1820 [’Bauhineae’]; CoronillaceaeMartinov, Tekhno-Bot. Slovar: 162. 3 Aug 1820 [’Coronillae’]; CytisaceaeBercht. et J. Presl, Přir. Rostlin: 229. Jan-Apr 1820 [‘Citiseae’]; DaleaceaeBercht. et J. Presl, Přir. Rostlin: 230. Jan-Apr 1820 [‘Daleae’]; GaledupaceaeMartinov, Tekhno-Bot. Slovar: 277. 3 Aug 1820 [’Galedupeae’]; HedysaraceaeBercht. et J. Presl, Přir. Rostlin: 230. Jan-Apr 1820 [‘Hedysareae’]; SophoraceaeBercht. et J. Presl, Přir. Rostlin: 229. Jan-Apr 1820; TamarindaceaeMartinov, Tekhno-Bot. Slovar: 622. 3 Aug 1820 [‘Tamarindi’]; TamarindalesBercht. et J. Presl, Přir. Rostlin: 230. Jan-Apr 1820 [‘Tamaryndeae’]; TrifoliaceaeBercht. et J. Presl, Přir. Rostlin: 230. Jan-Apr 1820 [‘Trifoliae’]; VicialesBercht. et J. Presl, Přir. Rostlin: 229. Jan-Apr 1820 [‘Viciae’]; LotaceaeOken, Lehrb. Naturgesch. 2(2.2): xv, 727. 1826 [’Loteae’]; ViciaceaeOken, Lehrb. Naturgesch. 2(2.2): xv. 1826 [’Vicieae’]; CassialesLink, Handbuch 2: 135. 4-11 Jul 1829 [’Cassiaceae’]; CeratoniaceaeLink, Handbuch 2: 135. 4-11 Jul 1829; MimosalesLink, Handbuch 2: 131. 4-11 Jul 1829 [‘Mimoseae’]; Swartziaceae(DC.) Bartl., Ord. Nat. Plant.: 231, 413. Sep 1830 [’Swartzieae’]; Detariaceae(DC.) J. Hess, Übers. Phan. Nat. Pfl.-Fam.: 46. 1832 [’Detarieae’]; Dalbergiaceae(DC.) Mart., Consp. Regn. Veg.: 34. Sep-Oct 1835; Geoffroeaceae(DC.) Mart., Consp. Regn. Veg.: 34. Sep-Oct 1835 [‘Geoffroeae’]; LathyraceaeBurnett, Outl. Bot.: 660, 1092, 1138. Feb 1835; LotalesBurnett, Outl. Bot.: 1138. Jun 1835 [‘Lotianae’], nom. illeg.; Phaseolaceae(DC.) Mart., Consp. Regn. Veg.: 34. Sep-Oct 1835 [‘Phaseoleae’]; AcaciaceaeE. Mey., Comm. Plant. Afr. Austr. 1: 164. 14 Feb-5 Jun 1836 [‘Acacieae’]; FabalesBromhead in Edinburgh New Philos. J. 25: 126. Jul 1838; CiceraceaeSteele, Handb. Field Bot.: xx, 63. 1847 [’Cicereaevel Leguminosae’]; InocarpaceaeZoll., Syst. Verz. 2: 117. 1854-1855 [’Inocarpeae’]; FabanaeR. Dahlgren ex Reveal in Phytologia 74: 179. 25 Mar 1993

Fossils Pollen grains are known from the Paleocene and the Eocene of Europe, Africa and Texas. Late Cretaceous fossils of Fabaceae are questionable.

Habit Usually bisexual (rarely monoecious, andromonoecious or polygamomonoecious), evergreen or deciduous trees, shrubs, lianas or suffrutices, perennial, biennial or annual herbs. Numerous species are xerophytes, whereas some are aquatic. Petiole or branch sometimes modified into photosynthesizing phyllodia or phyllocladia, respectively. Many species are spiny. Sometimes with large plank buttresses.

Vegetative anatomy Mycorrhiza sometimes absent. Root nodules (usually originating in cortex, sometimes as modified lateral roots) containing nitrogen-fixing bacteria (Azorhizobium, Bradyrhizobium, Rhizobium, and Sinorhizobium) present in most species. Nodules usually with bacterial colonies in centre and with vascular tissue in peripheral parts. Nodule-forming nitrogen-fixing bacteria usually α-2-proteobacteria (in some Mimoseae and Papilionoideae also certain β-proteobacteria). Infection threads permanent or absent, nodules long-lived or short-lived. Nitrogen metabolism usually very specific: free amino acids frequent and nitrogen in xylem juice transported as mixture of different amino acids, amids and sometimes also ureids (very small amount transported as nitrate). Ectomycorrhiza present in many (perhaps especially in non-nodulated?) species (i.a. in Cynometreae and Detarieae). Phellogen ab initio superficially or deeply seated (in Papilionoideae sometimes outer-cortical). Primary vascular tissue cylinder without separate vascular bundles, or ring of bundles. Endodermis in Cercis conspicuous. Secondary lateral growth normal or anomalous from concentric cambia or absent. Cambium and wood elements sometimes storied. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Vestured pits often present. Imperforate tracheary xylem element libriform fibres with usually simple (sometimes bordered) pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma usually abundant, apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, reticulate, vasicentric, unilateral, or banded. Secondary phloem often stratified into hard fibrous layers alternating with soft parenchymatous layers. Sieve tube plastids usually Pcs type (in Mimoseae Pc type) with protein crystalloids (rarely Ss type, with starch and proteins or only starch). Nodes usually 3:3, trilacunar with three leaf traces (sometimes 5:5, pentalacunar with five traces). Secretory cells and cavities present or absent. Heartwood often with resins and resin-like substances. Silica bodies present or absent. Prismatic calciumoxalate crystals and styloids abundant (especially in axial parenchyma), druses, navicular, acicular (organic) and spherical crystals sometimes present as well as crystal sand.

Trichomes Hairs unicellular or multicellular (usually tricellular), uniseriate or multiseriate, simple or branched, furcate or stellate, sometimes dendritic; pearl glands (pearl bodies), stalked multicellular glands and/or glandular hairs (sometimes lepidote) often present; microhairs sometimes present; stinging hairs rare.

Leaves Usually alternate (spiral or distichous; rarely opposite), usually pinnately or palmately compound (paripinnate, imparipinnate or trifoliolate, sometimes bipinnate or multifoliolate, rarely unifoliolate or simple), usually with entire (rarely lobed or serrate) opposite conduplicate leaflets (often pulvinate, rarely alternate) usually with entire margins, sometimes with foliar tendrils, sometimes with stipulules, sometimes coriaceous, sometimes reduced and scale-like, usually with conduplicate ptyxis? Stipules intrapetiolar, often foliaceous or modified into spines, scales or glands (extrafloral nectaries?), caducous or persistent; leaf sheath absent. Colleters often frequent. Petiole vascular bundles of various shape. Leaflets often pulvinate. Stipulules sometimes present. Venation pinnate or palmate. Stomata anomocytic, paracytic, anisocytic, tetracytic, parallelocytic, or cyclocytic. Cuticular wax crystalloids as rosettes of platelets (Fabales type), scales or granules. Epidermis with or without mucilaginous idioblasts. Mesophyll with or without sclerenchymatous idioblasts (with fibres or branched sclereids). Secretory cavities with oils, mucilage or resin present or absent. Silica sometimes frequent. Leaf and leaflet margins serrate or entire. Extrafloral nectaries often present on stipules, petiole or lamina.

Inflorescence Terminal or axillary, racemose, corymb, panicle, fascicle, raceme, spike, or head, or flowers solitary axillary. Lateral inflorescence branches sometimes with extrafloral nectaries. Floral prophylls (bracteoles) in Papilionoideae sometimes suppressed.

Flowers Usually zygomorphic (in Mimosoideae actinomorphic; actinomorphic flowers in Cadia is a reversal, due to dorsalization of flower). Hypanthium often well developed. Hypogyny. Sepals (three to) five (or six), with valvate or imbricate (in Schwartzieae irregular) aestivation, usually persistent, free or more or less connate (rarely strongly reduced); median sepal abaxial. Petals usually five (rarely one, three or four), with valvate (Mimoseae), imbricate, imbricate-ascending or cochlear-descending (adaxial-median petal internal) aestivation, often clawed, persistent or caducous, free or two front petals connate at base into carina (especially in Papilionoideae, sometimes four petals connate) or all petals connate (especially in Mimoseae); median petal adaxial (vexillum in Papilionoideae; lateral petals alae in Papilionoideae; median petal sometimes absent. Nectariferous disc intrastaminal or absent. Development of floral parts largely centripetalous; gynoecium initiated prior to petals.

Androecium Stamens usually ten (sometimes two to nine; in Mimoseae sometimes numerous, in Maniltoa to more than 100), usually in one whorl (at initiation usually in two whorls, diplostemonous; sometimes in three to six? whorls), unidirectionally initiated. Filaments free or more or less connate into tube (often with one adaxial stamen free), usually free from (sometimes adnate to) petals. Anthers basifixed and/or dorsifixed, usually versatile, tetrasporangiate, introrse or latrorse, usually longicidal (dehiscing by longitudinal slits; in some Cassieae poricidal, with apical pores); connective sometimes with caducous apical gland; placentoid sometimes present. Tapetum usually secretory, with uninucleate or quadrinucleate (to septanucleate) cells. Staminodia sometimes five or more.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–6)-colporate (sometimes 2–6-porate, -colpate, -rugate or inaperturate; sometimes with pseudocolpi), usually shed as monads (sometimes tetrads or polyads with eight, 18 or more pollen grains, especially in Mimoseae), usually bicellular (in some Mimoseae tricellular) at dispersal. Exine tectate or semitectate, with columellate or acolumellate? (granular?) infratectum, perforate, microreticulate, reticulate or striate-reticulate to striate, rugulate, verrucate or psilate.

Gynoecium Pistil composed of usually one carpel (in some Mimoseae two to 16 free carpels, [apocarpy]), often on gynophore. Ovary superior, usually unilocular (monomerous; rarely appearing bilocular by secondary septum), sometimes stipitate. Style single, simple, usually curved upwards, often hollow. Stigma capitate (sometimes widened), sometimes hollow, often papillate, Dry or Wet type. Male flowers sometimes with pistillodium? Enantiostyly present in some species (flowers then asymmetric).

Ovules Placentation marginal-ventral (along abaxial suture). Ovules (one or) two to numerous per carpel, usually anatropous, anacampylotropous or campylotropous (sometimes hemianatropous or amphitropous), ascending to pendulous, usually bitegmic (rarely unitegmic), usually crassinucellar (rarely incompletely tenuinucellar). Funicle often long or stout. Micropyle endostomal or bistomal, often Z-shaped (zig-zag). Outer integument two to ten cell layers thick, with vascular strand. Inner integument two or three cell layers thick. Parietal tissue one or two cell layers thick. Nucellar cap approx. three cell layers thick. Archespore often multicellular. Megagametophyte usually monosporous, Polygonum type (rarely disporous, Allium type). Synergids often with a filiform apparatus. Porogamy. Endosperm development ab initio nuclear. Endosperm haustorium usually chalazal (sometimes also micropylar; rarely [Adesmieae] lateral; sometimes elongate). Embryogenesis onagrad, asterad or caryophyllad.

Fruit A legume (in some genera modified into nut, drupe, berry, abaxially dehiscing follicle, or lomentum, in some species of Mimoseae a syncarp), sometimes with accrescent calyx.

Seeds Aril often present. Funicle often thick and fleshy (arilloid). Elaiosome sometimes present. Seed usually without hilar sulcus. Seed coat usually exotestal (sometimes indistinct). Testa sometimes undefined or ruminate. Exotesta often with palisade, more or less lignified Malpighian cells, polygonal in cross-section, often with pleurogram with deeply situated linea lucida (light line, separating very thick outer anticlinal cell walls from thin inner cell walls) and rounded linea fissura demarcating pleurogram. Mesotesta consisting of stellate cells. Endotesta and tegmen collapsed. Perisperm not developed. Endosperm usually absent (sometimes sparse, rarely copious, rarely thick-walled mucilaginous, with galactomannans). Embryo usually large, straight or curved, well differentiated (especially in Papilionoideae), with chlorophyll. Suspensor with very various structure; suspensor haustoria often present. Cotyledons two, well developed and nutritious, fatty, proteinaceous and often starchy, in Amherstieae, Dearieae and Sclerolobieae with amyloid (xyloglucans). Radicula straight, oblique or curved and lateral (Papilionoideae). Germination phanerocotylar or cryptocotylar.

DNA The plastid genome is extensively rearranged in Fabaceae. Examples are as follows. Plastid gene rpl22 transferred from plastid genome to nuclear genome. Plastid inverted repeat lost in six tribes and Wisteria. Plastid genome in most Papilionoideae with inversion of c. 50 kb. Plastid genome in most Phaseoleae with inversion of 78 kb. Robinieae (except Sesbania) with inversion of c. 30 kb. Plastid genome in Medicago with several inversions (62 kb inversion etc.). At least eight inversions present in Pisum (in P. humile inversion of 4 kb). Two or three inversions present in Vicia faba. Plastid gene rps16 lost in Moldenhawera and 16 tribes of Papilionoideae. Plastid gene ycf4 lost in most Papilionoideae. Plastid gene ndhF lost in Hebestigma cubense. Plastid gene accD (zpfA, ORF512) lost in Trifolium. Plastid gene clpP lost in Cicer and other genera. Plastid ORF224 lost in Pisum sativum and ORF84 lost multiple times. Intron lost in plastid gene rpoC1 in Medicago suffruticosa. Intron in plastid gene rpl12 transferred to nucleus in, i.a., Bauhinia and Desmodium (present in, e.g., Brya, Soemmeringia and Mucuna), mitochondrial gene srp12 in, i.a., Bauhinia. Numerous additional structural rearrangements, intron losses, indels etc. present in different clades.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavone-C-glycosides, afzelechin, 5-deoxyflavonoids, isoflavonoids (pterocarpans, genistein, pachyrrhizin, coumestrol, rotenone and other rotenoids, munduserone, angolensin, etc. in Papilionoideae), cyanidin, epigallocatechin-3-gallate, oleanolic acid derivatives, dammarane, phytoalexins, methylated ellagic acids, ellagitannins, geraniin, proanthocyanidins (prodelphinidins), pyrrolizidine alkaloids as macrocyclic diesters (abundant at least in Crotalaria), lupine alkaloids (cytisine, N-methylcytisine, anagyrine, thermopsine, lupanine, sparteine, etc.), loline alkaloids (in Genisteae, but not in Crotalaria), physostigmine and similar alkaloids, pyridine, quinolizidine and indole alkaloids, Erythrina alkaloids, triterpene saponins, tyrosine-, leucine- or phenylalanine-derived cyanogenic compounds, anthraquinones, pinitol (cyclitol), simmondsin-like compounds, lignans (syringaresinol), toxic free non-protein amino acids, and lectins (hemagglutinins, especially in seeds) present. Gums often frequent (especially in seeds). Storage polysaccharides (especially in seeds) present as galactomannans (galactose/mannose relationship possibly phylogenetically informative). Numerous secondary metabolites synthesized by endophytic fungi or bacteria.

Use Ornamental plants, edible fruits and seeds, vegetables, forage plants, spices and flavours, dragant and gums (Astragalus gummifer, Acacia spp.), dyeing substances (Indigofera, Genista tinctoria, etc.), medicinal plants, poisons (rotenonoids from Derris, Lonchocarpus and other genera), timber, tanning, fertilizers.

Systematics There is no undisputable sister-group to the Fabaceae, although there are several candidates.

The presentation below mainly follows that in LPWG (2013, 2017). Duparquetioideae or Cercideae are sister-group to all other Fabaceae.

Duparquetioideae Legume Phylogeny Working Group in Taxon 66(1): 69. Feb. 2017

Duparquetiinae H. S. Irwin et Barneby in R. M. Polhill et P. H. Raven, Adv. Legume System. 1: 102. 1981

1/1. Duparquetia (1; D. orchidacea; tropical West and Central Africa). – Liana, unarmed. Root nodules absent. Vestured pits absent. Stipules lateral, free. Leaves pinnately compound, imparipinnate. Leaflets/pinnae opposite (lamina entire). Extrafloral nectaries absent. Terminal raceme. Flowers not papilionate. Hypanthium absent. Floral development acropetal. Sepals four, free, unequal, abaxial and adaxial sepals cucullate and sepaloid, lateral sepals petaloid. Petals five, free, imbicate, with stalked glands along margins; adaxial petal outermost. Stamens four, antesepalous. Filaments free. Anthers basifixed, poricidal (dehiscing by pores), with pointed appendages, post-genitally fused into synandrium. Pollen grains asymmetrical. Apertures with ectoaperture encircling equator and with two equatorial endoapertures. Gynoecium unicaprellate, stipitate, with free stipe. Gynoecial initiation not advanced (relative to other organs). Ovary bi- to quinque-ovulate. Fruit a woody pod with spirally coiling valves. Seeds without pleurograms. Embryo straight. n = ? Chemistry unknown.

Cercidoideae Legume Phylogeny Working Group in Taxon 66(1): 68. Feb. 2017

Cercideae Bronn, Form. Plant. Legumin.: ad Sect. 134, 131. 1822

14/300–460. Cercis (12; the Mediterranean, eastern Asia, North America to northeastern Mexico); Adenolobus (2; A. garipensis, A. pechuelii; southern Angola, Namibia, northern Botswana, Northern Cape); Bauhinia (150–300; tropical and subtropical regions of both hemispheres, especially tropical South America), Piliostigma (5; P. foveolatum, P. malabaricum, P. reticulatum, P. thonningii, P. tortuosum; Africa incl. South Africa to southern Asia and Australia), Brenierea (1; B. insignis; Madagascar); Griffonia (4; G. physocarpa, G. simplicifolia, G. speciosa, G. tessmannii; tropical West Africa); Gigasiphon (3; G. gossweileri, G. humblotianum, G. macrosiphon; tropical Africa), Tylosema (5; T. angolense, T. argentea, T. esculentum, T. fassoglensis, T. humifusa; Africa incl. South Africa), Barklya (1; B. syringifolia; northeastern Queensland), Schnella (30–35; tropical South America), Lysiphyllum (8; Malesia to tropical Australia), Phanera (60–65; pantropical, East Asia from eastern Himalayas to Japan, Indochina, Java), Lasiobema (12; India to Southeast Asia and Malesia; in Phanera?), Cheniella (10; India to southern China, Southeast Asia, West and Central Malesia). – Mainly tropical and subtropical regions, few species in warm temperate areas. Trees, shrubs or lianas, often with branch tendrils and/or prickles or infrastipular spines. Root nodules absent. Vestured pits absent. Stipules lateral, free. Petiole with a single joined pulvinus. Leaves simple alternatively unifoliolate (or bifoliolate), usually bilobate or entire (rarely with two leaflets). Leaflets/pinnae opposite when lamina bifoliolate. Lamina when unifoliolate entire or bilobate. Stomata usually anomocytic (rarely paracytic). Extrafloral nectaries usually absent. Raceme or pseudoraceme. Flowers sometimes papilionate. Hypanthium present (often very elongated) to almost absent. Sepals five, connate into spathaceous or bi- to quinquelobate calyx, or free. Petals usually five (rarely two, six or absent), free, imbricate, adaxial petal innermost. Stamens usually ten (sometimes fewer) in two whorls of alternate length. Filaments more or less connate or free. Anthers dorsifixed, longicidal or poricidal. Staminodia sometimes present. Pollen grains tricolporate, 3–6-colpate, 3-porate, 3-pororate, 3–4-colporoidate or inaperturate, rarely in tetrads. Exine striate. Zwischenkörper (pectic substances) present below apertures at least in Cercis. Gynoecium unicarpellate, stipitate, with free or adnate stipe. Ovary uni- to multiovulate. Fruit an often explosively dehiscent pod, with twisted valves, or indehiscent and usually samaroid. Seeds with apical usually crescent-shaped (rarely circular) hilum, without pleurograms, pseudopleurograms, wing or aril. Embryo usually straight. x = 7. Introns in mitochondrial genes srp12 and rpl2 absent in Bauhinia. Coumarins and cyanogenic glucosides present. Non-protein aminoacids (5-hydroxy-L-tryptophan) frequent. – Cercis is sister to the remaining Cercidoideae and Adenolobus successive sister to the two major clades.

Fabaceae except Duparquetia and Cercideae

[Detarioideae+[Dialioideae+[Caesalpinioideae+Papilionoideae]]]

Nitrogen fixation sometimes occurring. Vestured pits usually present. Fruit sometimes a drupe, a samara, a schizocarp, etc. Dumbbell-shaped cells present below palisade exotesta. Free amino acids, especially in seeds.

Detarioideae Burmeist., Handb. Naturgesch.: 319. 1837 [‘Detarieae’]

78/760–>805. Trees or sometimes shrubs (rarely suffruticose). Root nodules absent. Ectomycorrhiza often present. Vestured pits present in secondary xylem. Resinous ducts often present in stem. Stipules usually fused partly or entirely, intrapetiolar (between petiole and axillary bud), often caducous. Leaf phloem transfer cells present. Leaves usually paripinnate or bifoliolate (rarely unifoliolate). Leaflets/pinnae opposite or alternate. Leaflets often with translucent crater-like glands on abaxial surface. Extrafloral nectaries often present on lower side of leaves, rarely on margins of leaflets or on leaf rachis. Raceme or panicle. Floral prophylls (bracteoles) often well developed (often petaloid, valvate, imbricate or partially fused with each other or with hypanthium, partially or completely enclosing bud), caducous. Flowers not papilionate. Hypanthium present (often elongated) to almost absent. Sepals often four or five, with two adaxial sepals often connate (rarely some or all absent or up to seven). Petals up to five (to seven), free, imbricate, equal or unequal, adaxial petal outermost. Stamens usually ten (sometimes two to numerous). Filaments free or more or less connate. Anthers dorsifixed or basifixed, longicidal. Pollen grains usually tricolporate. Exine sometimes striate. Gynoecium unicarpellate, stipitate, with free or adnate stipe. Ovary uni- to multiovulate. Style abaxially curved. Fruit usually a woody dehiscent pod (sometimes indehiscent samaroid). Seeds occasionally with pseudopleurograms, sometimes arillate. Embryo straight. Endosperm absent. Cotyledons with thick-walled cells, with amyloid (xyloglucans). x = 12. Coumarins present. Bicyclic diterpenes (resins) and non-protein amino acids often present. – Two major clades, (1) the resin-producing Detarieae and (2) Amherstieae, comprise the major part of the Detarioideae and these two clades form a polytomy together with a number of small groups. – The clade [Barnebydendreae+Schotieae] is sister-group to Detarieae in a wide sense, according to Estrella & al. (2018).

Barnebydendreae Estrella, L. P. Queiroz et Bruneae in Sci. Rep. 8: 6884. 2018

2/2. Barnebydendron (1; B. riedelii; tropical Central and South America), Goniorrhachis (1; G. marginata; southeastern Brazil). – Zwischenkörper (pectic substances) present below apertures.

Schotieae Estrella, L. P. Queiroz et Bruneae in Sci. Rep. 8: 6884. 2018

1/4–5. Schotia (4–5; S. afra, S. brachypetala, S. capitata, S. latifolia; Africa south of the Zambesi River). – Zwischenkörper (pectic substances) present below apertures.

[[Prioria clade+Daniellieae]+Detarieae]

Prioria clade

3/16. Colophospermum (1; C. mopane; tropical southern Africa), Hardwickia (1; H. binata; drier regions of India), Prioria (14; tropical Africa, tropical Asia to islands in the Pacific, Central America, Jamaica, northwestern South America). – Tropical.

Daniellieae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 252. 20 Jul 1943 [’Danielleae’]

2/6. Brandzeia (2; B. filicifolia, B. rubriflora; Madagascar), Daniellia (4; D. klainei, D. oblonga, D. ogea, D. oliveri; tropical Africa). – Tropical Africa, Madagascar. Brandzeia and Daniellia may be sister-groups, although the support is weak.

Detarieae DC., Prodr. 2: 521. Nov (med.) 1825

14/163–170. Peltogyne (23; Central America, tropical South America), Guibourtia (16; tropical Africa, tropical South America), Stemonocoleus (1; S. micranthus; Central Africa), Augouardia (1; A. letestui; Gabon), Hymenaea (14; southern Mexico, Central America, the West Indies, tropical South America, one species, H. verrucosa, in eastern Africa); Neoapaloxylon (3; N. madagascariense, N. mandrarense, N. tuberosum; Madagascar), Baikiaea (7; B. fragrantissima, B. ghesquiereana, B. insignis, B. plurijuga, B. robynsii, B. suzannae, B. zenkeri; tropical to southern Africa in Namibia and Botswana), Copaifera (40–45; tropical Africa, Borneo, Central America, tropical South America), Detarium (3; D. letestui, D. microcarpum, D. senegalense; tropical and subtropical West Africa to Sudan), Gilletiodendron (5; G. glandulosum, G. kisantuense, G. mildbraedii, G. ogoouense, G. pierreanum; tropical Africa), Hylodendron (1; H. gabunense; coasts of the Gulf of Guinea), Sindora (18–20; Southeast Asia, Malesia), Sindoropsis (1; S. letestui; Gabon), Tessmannia (12; tropical Africa), Eperua (c 15; northeastern South America), Eurypetalum (3; E. batesii, E. tessmannii, E. unijugum; Cameroon to Gabon). – Tropical and subtropical. Flowers bisymmetric or actinomorphic. Petals absent or one to five. Stamens three to ten (when fewer, then staminodia as well). Zwischenkörper (pectic substances) present below apertures. Bicyclic diterpenes present in resinous ducts.

Amherstieae Benth. in J. Bot. (Hooker) 2: 73. Mar 1840

Cynometreae Benth. in J. Bot. (Hooker) 2: 74. Mar 1840

c 56/570–>605. Saraceae Estrella, L. P. Queiroz et Bruneae in Sci. Rep. 8: 6884. 2018. Endertia (1; E. spectabilis; Borneo), Lysidice (2; L. brevicalyx, L. rhodostegia; southern China, Vietnam), Saraca (11; India, Sri Lanka, southern China, Southeast Asia, Malesia to Sulawesi), Leucostegane (2; L. grandis, L. latistipulata; the Malay Peninsula, Borneo). – Afzelieae Estrella, L. P. Queiroz et Bruneae in Sci. Rep. 8: 6884. 2018. Afzelia (13; tropical Africa, tropical Asia), Brodriguesia (1; B. santosii; eastern Brazil), Intsia (4; I. acuminata, I. bijuga, I. moeleri, I. palembanica; Southeast Asia to Pacific coasts). – Amherstieae sensu stricto. Ecuadendron (1; E. acostasolisianum; western Ecuador), Brownea (c 40; Costa Rica to Peru, the West Indies), Macrolobium (75–80; Central America, tropical South America), ‘Heterostemon’ (1–7; H. mimosoides; tropical South America; diphyletic), ‘Elizabetha’ (6; E. bicolor, E. coccinea, E. durissima, E. paraensis, E. princeps, E. speciosa; tropical South America; polyphyletic), Paloue (4; P. brasiliensis, P. guianensis, P. induta, P. riparia; the Guayana Shield in northern South America), Paloveopsis (1; P. emarginata; northeastern South America); Crudia (50–55; tropical Africa, tropical Asia, tropical America), Lebruniodendron (1; L. leptanthum; coasts of the Gulf of Guinea), Normandiodendron (2; N. becquaertii, N. romii; western Central Africa), ‘Cynometra’ pro parte (c 85 + 20–25; pantropical; paraphyletic), Plagiosiphon (5; P. discifer, P. emarginatus, P. gabonensis, P. longitubus, P. multijugus; Central Africa), Zenkerella (6; Z. capparidacea, Z. citrina, Z. egregia, Z. grotei, Z. perplexa, Z. schliebenii; coast of Guinea, tropical East Africa), Neochevalierodendron (1; N. stephanii; Gabon), Gabonius (1; G. ngouniensis; Gabon), Micklethwaitia (1; M. carvalhoi; Mozambique), Annea (2; A. afzelii, A. laxiflora; tropical West and Central Africa), Scorodophloeus (2; S. fischeri, S. zenkeri; coasts of Guinea and East Africa), ‘Hymenostegia’ pro parte (7–13?; along the Gulf of Guinea; diphyletic); Librevillea (1; L. klainei; Gabon), ‘Gilbertiodendron’ (c 30; tropical and subtropical Africa; non-monophyletic), Didelotia (10–11; Central Africa), ‘Anthonotha’ (c 30; tropical Africa; polyphyletic), Oddoniodendron (6; O. gambanum, O. gilletii, O. micranthum, O. normandii, O. reitsmarum, O. romeroi; along the Gulf of Guinea), Englerodendron (1?; E. usambarensis; Usambara Mountains in Tanzania), ‘Isoberlinia’ (5; I. angolensis, I. dalzielii, I. doka, I. scheffleri, I. stolzii; tropical Africa; polyphyletic), ‘Berlinia’ (17; tropical Africa; polyphyletic), Microberlinia (2; M. bisulcata, M. brazzavillensis; coasts of the Gulf of Guinea), Brachystegia (25–30; central tropical Africa to Botswana), Icuria (1; I. dunensis; Central Mozambique), Aphanocalyx (14; tropical West and Central Africa), Julbernardia (11; tropical to southern Africa), ‘Bikinia’ (c 10; western Central Africa; paraphyletic; incl. Tetraberlinia?), Tetraberlinia (7; T. baregarum, T. bifoliata, T. korupensis, T. longiracemosa, T. moreliana, T. polyphylla, T. tubmaniana; Central Africa; in Bikinia?); Cryptosepalum (12; tropical Africa), Humboldtia (7; H. bourdillonii, H. brunonis, H. decurrens, H. laurifolia, H. trijuga, H. unijuga, H. vahliana; southern India, Sri Lanka), Paramacrolobium (1; P. coeruleum; tropical Africa), Cynometra sensu stricto (tropical Africa), Dicymbe (c 20; the Guayana Shield to western Amazonia in Brazil), Polystemonanthus (1; P. dinklagei; tropical West Africa); Tamarindus (1; T. indica; tropical Africa), Loesenera (2; L. kalantha, L. talbotii; tropical West Africa), Talbotiella (5–8; T. bakossiensis, T. batesii, T. breteleri, T. ebo, T. eketensis, T. gentii, T. korupensis, T. velutina; tropical West Africa), Leonardoxa (1; L. africana; Cameroon), Hymenostegia sensu stricto (3?; H. floribunda; coast of the Gulf of Guinea); unplaced: Amherstia (1; A. nobilis; Burma, extinct?), Brachycylix (1; B. vageleri; northern Central Colombia), Michelsonia (1; M. microphylla; eastern Congo), Pseudomacrolobium (1; P. mengei; Congo). – Pantropical. – The sister-group of Amherstieae is not yet clarified.

[Dialioideae+[Caesalpinioideae+Papilionoideae]]

Dialioideae Legume Phylogeny Working Group in Taxon 66(1): 69. Feb. 2017

Dialiinae H. S. Irwin et Barneby in Polhill et Raven, Adv. Legume Syst. 1: 100. 1981

16/c 83. Poeppigia (1; P. procera; southern Mexico, Central America, tropical South America); Dialium (c 40; tropical to southern Africa, Madagascar, South and Southeast Asia, one species, one species, D. guianense, in tropical America); Petalostylis (2; P. cassioides, P. labicheoides; drier regions in Australia), Labichea (14; Australia except central parts), Baudouinia (6; B. capuronii, B. fluggeiformis, B. louvelii, B. orientalis, B. rouxevillei, B. sollyaeformis; Madagascar), Distemonanthus (1; D. benthamianus; tropical West Africa), Storckiella (4; S. australiensis: northeastern Queensland; S. neocaledonica, S. pancheri: New Caledonia; S. vitiensis: Fiji), Zenia (1; Z. insignis; southern China, Thailand, Vietnam), Androcalymma (1; A. glabrifolium; upper Amazon Basin in Brazil), Kalappia (1; K. celebica; Malili in Sulawesi), Apuleia (1; A. leiocarpa; northeastern Peru, southeastern Brazil, northern Argentina), Dicorynia (2; D. guianensis, D. paraensis; tropical South America), Eligmocarpus (1; E. cynometroides; southeastern Madagascar), Koompassia (3; K. excelsa, K. grandiflora, K. malaccensis; Malesia), Martiodendron (4; M. elatum, M. excelsum, M. mediterraneum, M. parviflorum; tropical South America), Mendoravia (1; M. dumaziana; southeastern Madagascar). – Tropical. Trees or shrubs (rarely suffruticose), unarmed. Root nodules absent. Vestured pits rarely present in secondary xylem. Stipules lateral, free, or absent. Leaves imparipinnate (rarely paripinnate, unifoliolate or palmately compound). Leaflets/pinnae usually alternate (rarely opposite). Extrafloral nectaries absent. Inflorescence usually cymose, primarily simple or compound cymes, thyrsoid or dichasia (sometimes racemes with distichous floral arrangement or flowers solitary). Flowers sometimes papilionate. Hypanthium usually absent (rarely present). Sepals (3–)5(–6), free. Petals usually five or fewer (rarely six), free, imbricate, adaxial petal innermost. Stamens usually five or fewer (rarely six to ten). Filaments free. Anthers usually basifixed (rarely dorsifixed), longicidal to poricidal. Staminodia often present. Pollen grains tricolporate. Exine punctate or finely reticulate (rarely striate). Gynoecium usually unicarpellate (sometimes bicarpellate), stipitate with free stipe, or sessile. Ovary usually biovulate (rarely uni- to multiovulate). Fruit usually drupe or samara (rarely dehiscent or drupe with endocarp indurating into one-seeded segments). Seeds without pleurograms. n = 14. Chemistry unknown. – Many representatives have lost one or several floral organs during evolution.

[Caesalpinioideae+Papilionoideae]

Vestured pits usually present (absent in Cassieae). Non-protein amino acids sometimes present (especially in seeds).

Caesalpinioideae DC., Prodr. 2: 473. Nov (med.) 1825 [‘Caesalpineae’]

148/4.100–>4.200. Trees, shrubs, lianas, suffruticose or functionally herbaceous, unarmed or often armed with prickles or spines. Root nodules often present. Vestured pits present in secondary xylem. Stipules lateral, free, or absent. Leaves bipinnate or pinnate (then usually paripinnate, rarely imparipinnate or bifoliolate, modified into phyllodes or absent). Leaflets/pinnae usually opposite (rarely alternate). Extrafloral nectaries often present on petiole and/or on primary and secondary rachises, usually between pinnae or leaflet pairs (sometimes on stipules or bracts). Inflorescence globose, spike, panicle, raceme or flowers in fascicles. Flowers sometimes papilionate or asymmetric. Hypanthium absent or cupular (rarely tubular). Sepals (3–)5(–6), free. Petals (3–)5(–6), free or connate, with valvate or imbricate aestivation, then adaxial petal innermost. Stamens diplo- or haplostemonous (sometimes 3–5), often numerous (up to more than 100). Filaments free or connate. Anthers basifixed or dorsifixed, longicidal or poricidal (apically or basally), often with stipitate or sessile apical gland. Staminodia present or absent. Pollen grains shed as monads, tetrads, bitetrads or polyads, tricolporate or porate. Exine never striate. Gynoecium usually unicarpellate (rarely polycarpellate), stipitate, with free stipe, or sessile. Ovary uni- to multiovulate. Fruit a one- to many-seeded pod, dehiscent along one or two sutures, often a lomentum, a craspedium, or thick, woody and indehiscent or explosively dehiscent. Seeds usually with open or closed pleurogram on both sides (sometimes with aril och sarcotesta, sometimes winged), hilum usually apical. Embryo straight. x = 7, 12. Non-protein amino acids, coumarins, cyanogenic glucosids, phenylethylamine, traptamines and β-carboline alkaloids present.

Ceratonieae Reichb., Flora Germ. Excurs. 2(2): 544. 1832

7/24–27. Ceratonia (2; C. oreothauma, C. siliqua; southeastern Mediterranean to Somalia and the Arabian Peninsula), Acrocarpus (1; A. fraxinifolius; India, Sri Lanka, Southest Asia, Malesia), Tetrapterocarpon (2; T. geayi, T. septentrionalis; Madagascar); Arcoa (1; A. gonavensis; Hispaniola); Gymnocladus (5–6; G. angustifolius, G. assamicus, G. burmanicus, G. chinensis, G. dioicus; East and Southeast Asia, eastern North America), Umtiza (1; U. listeriana; Eastern Cape), Gleditsia (14; the Caspian area, China, Japan to New Guinea, eastern North America, South America). – Subtropical and tropical regions. Most species are usually dioecious (not so in Acrocarpus and Umtiza). The perianth is often greenish and more or less reduced. – Ceratonieae may be sister-group to the remainder of Caesalpinioideae.

[[Pterogyne+Caesalpinieae+Cassieae]+[[Dimorphandra clade A+Peltophoreae+Tachigaleae]+Dimorphandra clade B]]

[Pterogyne+Caesalpinieae+Cassieae]

Pterogyne (1; P. nitens; tropical South America). – Pterogyne is sister to either Caesalpinieae or Cassieae.

Caesalpinieae Rchb., Fl. Germ. Excurs. 2(2): 544. 1832 [‘Caesalpineae s. Cassieae’]

27/224. Cordeauxia (1; C. edulis; Somalia, Ethiopia), Stuhlmannia (1; S. moavi; coast of Kenya and Tanzania, northern Madagascar), Lophocarpinia (1; L. aculeatifolia; Paraguay, northern Argentina, southern Brazil?), Haematoxylum (5; H. brasiletto, H. calakmulense, H. campechianum, H. dinteri, H. sousanum; Mexico, Central America, the West Indies, northwestern South America), Hererolandia (1; H. pearsonii; Ababes in Namibia), Denisophytum (8; the Arabian Peninsula, Somalia, Kenya, northern Madagascar, Florida, Mexico, the West Indies, Paraguay, northern Argentina), Coulteria (7; C. cubensis, C. glabra, C. mollis, C. platyloba, C. pringlei, C. pumila, C. velutina; Mexico, Central America, the West Indies, Colombia), Tara (3; T. cacalaco, T. spinosa, T. vesicaria; Mexico, Central America, the West Indies, northwestern South America), Gelrebia (8; northern Kenya, Ethiopia, Somalia, Congo, southern Africa), Hultholia (1; H. mimosoides; India, Bangladesh, Burma, Yunnan, Thailand, Indochina), Guilandina (19; warmer regions on both hemispheres), Moullava (4; M. digyna, M. spicata, M. tortuosa, M. welwitschiana; India, Sri Lanka, Burma, Yunnan, Hainan, Southeast Asia, Malesia, one species, M. welwitschiana, in Central Africa), Biancaea (6; B. decapetala, B. godefroyana, B. millettii, B. oppositifolia, B. parviflora, B. sappan; India, Burma, Southeast Asia, Malesia to the Philippines and Borneo), Ticanto (15; southern China, Southeast Asia, Malesia), Pterolobium (10; India, southern China, Thailand, Indochina, Malesia, one species, P. stellatum, in southern and eastern tropical Africa and the Arabian Peninsula), Mezoneuron (c 25; tropical Africa, Madagascar, tropical and subtropical Asia to tropical Australia and the Hawaiian Islands), Paubrasilia (1; P. echinata; eastern Brazil), Caesalpinia (9; Mexico, Central America, the West Indies, tropical South America), Cenostigma (14; southern Mexico, Central America, the West Indies, tropical South America), Stenodrepanum (1; S. bergii; central and western Argentina), Hoffmanseggia (23; tropical to southern Africa, southwestern United States to Chile and Argentina), Balsamocarpon (1; B. brevifolium; northern Chile), Zuccagnia (1; Z. punctata; the Andes in northern and central Chile and western central Argentina), Libidibia (7; L. coriaria, L. ferrea, L. glabrata, L. monosperma, L. paraguariensis, L. punctata, L. sclerocarpa; Mexico, Central America, the West Indies, tropical South America), Pomaria (16; southern Africa, southern United States, Mexico, southeastern South America), Arquita (5; A. ancashiana, A. celendiniana, A. grandiflora, A. mimosifolia, A. trichocarpa; the Andes in Ecuador, Peru, Bolivia and northwestern Argentina), Erythrostemon (31; southern United States, Mexico, Central America, the West Indies, South America). – Tropical and subtropical regions, few species in warm-temperate parts of the world. Trees or shrubs. Sieve tube plastids also with fibres. Leaves often bicompound. Ovules usually campylotropous. Outer integument usually with vascular bundle. Aril often present. – Cordeauxia and Stuhlmannia form a clade sister to the remaining Caesalpinieae.

Cassieae Bronn, Form. Plant. Legumin.: 78, 127, 130. 1822

8/600–780. Cassia (30–90 or more?; tropical and subtropical regions on both hemispheres; incl. Latrobea?), Latrobea (4–6; L. colophona, L. diosmifolia, L. genistoides, L. hirtella; southwesternmost Western Australia; in Cassia?), Senna (260–350; warm-temperate to tropical regions, especially America); Batesia (1; B. floribunda; Amazonas), Chamaecrista (300–330?; Africa, East Asia, temperate and tropical regions in America), Melanoxylum (1; M. brauna; Amazonas), Recordoxylon (1; R. speciosum; Amazonas); Vouacapoua (3; V. americana, V. macropetala, V. pallidior; tropical South America). – Mainly tropical and subtropical regions, few species in warm-temperate regions. Trees or shrubs (rarely herbs). Vestured pits absent. – Chamaecrista, Melanoxylon and Recordoxylon have the ability to form bacterial nodules. Rhizobia in Chamaecrista remain in infection threads.

[[Dimorphandra clade A+Schizolobieae+Tachigaleae]+Diptychandra clade B]

[Dimorphandra clade A+Schizolobieae+Tachigalieae]

Dimorphandra clade A

6/43–63. Burkea (1–2; B. africana, B. caperangau; tropical and subtropical Africa to Namibia and northern South Africa), Campsiandra (c 20; tropical South America, with their highest diversity in Amazonas), Dimorphandra pro parte (c 25; tropical America), Dinizia (1; D. excelsa; Guayana, Brazil), Mora (7–10; M. abbottii, M. ekmanii, M. excelsa, M. gonggrijpii, M. megistosperma, M. oleifera, M. paraensis; Central America, the West Indies, northern tropical South America), Stachyothyrsus (3; S. stapfiana, S. staudtii, S. tessmannii; tropical West Africa). – Tropical Africa and America. The Dimorphandra clade is weakly supported.

Schizolobieae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 251. 20 Jul 1943

8/45–49. Bussea (7; B. eggelingii, B. gossweileri, B. massaiensis, B. occidentalis, B. perrieri, B. sakalava, B. xylocarpa; tropical Africa, Madagascar), Peltophorum (11; tropical and subtropical regions); Schizolobium (2; S. amazonicum, S. parahyba; tropical South America), Delonix (10–11; eastern Africa, Madagascar, the Arabian Peninsula, India; incl. Colvillea and Lemuropisum?), Colvillea (1; C. racemosa; Madagascar; in Delonix?), Lemuropisum (1; L. edule; Madagascar; in Delonix?), Conzattia (2; C. multiflora, C. sericea; Mexico), Parkinsonia (12–15; drier warmer regions in America, northeastern Africa, Namibia and Northern Cape, and from Mexico to Argentina). – Tropical and subtropical. Leaves bipinnate. Petals usually yellow. Seeds narrow. – Schizolobium is sister to the Delonix-Parkinsonia clade.

Tachigalieae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 252. 20 Jul 1943

3/100–105. Arapatiella (2; A. emarginata, A. psilophylla; southeastern Brazil), Jacqueshuberia (7; J. amplifolia, J. brevipes, J. loretensis, J. purpurea, J. pustulata, J. quinquangulata, J. splendens; Colombia, Brazil), Tachigali (c 95; tropical America, especially Amazonas). – Tropical America.

Dimorphandra clade B

[Dimorphandra pro parte+[Diptychandra+Moldenhawera]+Pachyelasma+Erythrophleum+Sympetalandra+Mimoseae]

Dimorphandra pro parte

Diptychandra (1; D. aurantiaca; Brazil, Bolivia, Paraguay), Moldenhawera (10; Venezuela, eastern Brazil). – Tropical South America. Diptychandra and Moldenhawera may be sister-groups, but the support is weak.

Pachyelasma (1; P. tessmannii; tropical West Africa).

Erythrophleum (10; tropical to southern Africa, Madagascar, southern and eastern Asia to northern Australia). – Tropical regions in the Old World. Erythrophleum may be sister to Mimoseae, although the support is weak.

Sympetalandra (1; S. schmutzii; Flores).

Mimoseae Bronn, Form. Plant. Legumin.: 78, 127, 130. 1822 (incl. Acacieae Dumort., Anal. Fam. Plant.: 40. 1829 and Ingeae Benth. et Hook. f., Gen. Plant. 1: 437. 19 Oct 1865)

82/3.035–3.150. Adenanthera (13; tropical Asia and Australia, islands in the Pacific), Tetrapleura (1–2; T. chevalieri, T. tetraptera; tropical Africa), Amblygonocarpus (1; A. andongensis; tropical Africa to Namibia and Botswana), Calpocalyx (11; tropical West Africa), Pseudoprosopis (7; P. bampsiana, P. claessensii, P. euryphylla, P. fischeri, P. gilletii, P. sericea, P. uncinata; tropical Africa), Xylia (9–10; tropical to southern Africa, tropical Asia); Pentaclethra (3; P. eetveldeana, P. macroloba, P. macrophylla; tropical West and Central Africa, Central America, tropical South America); ‘Entada’ (c 30; tropical regions; non-monophyletic; incl. Elephantorrhiza?), Elephantorrhiza (9; tropical to southern Africa; in Entada?), Piptadeniastrum (1; P. africanum; Central Africa); Newtonia (14–15; tropical Africa, tropical America), Fillaeopsis (1; F. discophora; western and southwestern Africa), Indopiptadenia (1; I. oudhensis; India, Nepal), Lemurodendron (1; L. capuronii; northeastern Madagascar); Plathymenia (1; P. reticulata; Brazil, Argentina); ‘Prosopis’ (26; Africa, Southwest Asia, southern United States to South America; non-monophyletic), Xerocladia (1; X. viridiramis; Namibia, Northern Cape); Cylicodiscus (1; C. gabunensis; Central Africa); Prosopidastrum (5; P. angusticarpum, P. globosum, P. gracile, P. mexicanum, P. striatum; Mexico to southern South America); Piptadeniopsis (1; P. lomentifera; Paraguay), Mimozyganthus (1; M. carinatus; southeastern Bolivia, southwestern Paraguay, Argentina); Desmanthus (c 25; southern United States, Mexico, Central America, the West Indies, tropical South America), Leucaena (c 25; Texas to Peru), Schleinitzia (2; S. fosbergii, S. insularum; Malesia, Vanuatu to Tahiti, Guam); Calliandropsis (1; C. nervosus; Mexico), Dichrostachys (c 25; tropical Africa, Madagascar, southern and southeastern Asia to Australia), Gagnebina (5; G. bakoliae, G. calcicola, G. commersoniana, G. icrocephala, G. pterocarpa; Madagascar, the Comoros, the Mascarene Islands); Neptunia (8; tropical and subtropical regions); Aubrevillea (2; A. kerstingii, A. platycarpa; tropical West and Central Africa); Chidlowia (1; C. sanguinea; western tropical Africa); Parkia (c 35; tropical Africa and Asia, Madagascar, tropical Australia, tropical and subtropical America), Anadenanthera (2; A. colubrina, A. peregrina; Hispaniola, Puerto Rico, tropical South America); Adenopodia (7; A. gymnantha, A. oaxacana, A. patens, A. rotundifolia, A. scelerata, A. schlechteri, A. spicata; tropical Africa, Mexico, Central America), Mimosa (c 400?; tropical and subtropical regions, with their largest diversity in Central and tropical South America), ’Piptadenia’ (c 30; southern Mexico, Central America, tropical South America; non-monophyletic), Pityrocarpa (4; P. leucoxylon, P. moniliformis, P. obliqua, ‘Pseudopiptadenia’ brenanii; southern Mexico, Central America, tropical South America), Stryphnodendron duckeanum (Brazil), Parapiptadenia (3; P. excelsa, P. pterosperma, P. rigida; tropical South America), Pseudopiptadenia (5–10; P. contorta, P. leptostachya, P. psilostachya, P. suaveolens, P. warmingii; Central America, tropical South America), ‘Stryphnodendron’ (c 12; Central America, tropical South America; polyphyletic), Vachellia (163; tropical Africa, Madagascar, the Mascarene Islands, tropical Asia, tropical Australia, islands in the Pacific), ‘Senegalia’ (c 225; tropical and subtropical regions in Africa and Asia, Mexico, Central America, tropical South America; polyphyletic), Pseudosenegalia (2; P. feddeana, P. riograndensis; Bolivia), Parasenegalia (7; P. lundellii, P. muricata, P. rurrenabaqueana, P. santosii, P. skleroxyla, P. visco, P. vogeliana; Central America, the West Indies, tropical South America), Mariosousa (13; southwestern United States, Mexico, Central America), Lachesiodendron (1; L. viridiflorum; Mexico, Guatemala, Colombia and Venezuela to eastern Brazil, Bolivia, Paraguay and northern Argentina), Acaciella (15; southern and central United States to Argentina), Acacia (900–1.000; Madagascar, tropical Asia, Australia, islands in the Pacific); Pseudosamanea (1; P. cubana; Cuba), Cedrelinga (1; C. cateniformis; Brazil), Hesperalbizia (1; H. occidentalis; southwestern Mexico), ‘Enterolobium’ (12; tropical America; polyphyletic), ‘Archidendron’ (95–100; tropical Asia, with their highest diversity on Borneo; non-monophyletic), Archidendropsis (14; islands in southwestern Pacific), Wallaceodendron (1; W. celebicum; the Philippines, Sulawesi), Samanea (5; S. dinklagei, S. guineensis, S. inopinata, S. leptophylla, S. tubulosa; Central America, the West Indies, tropical South America), ‘Abarema’ (c 45; New Caledonia, Mexico, Central America, tropical South America; polyphyletic), Balizia (2; B. elegans, B. leucocalyx; southern Mexico, Central America, tropical South America), Blanchetiodendron (1; B. blanchetii; eastern Brazil), Leucochloron (2; L. foederale, L. minarum; Brazil), Macrosamanea (8; tropical South America), Zygia (14; southern Mexico, Central America, tropical South America), Inga (c 300; tropical and subtropical regions in South America), Ebenopsis (2; E. confinis, E. ebano; southern Texas, Mexico), Sphinga (3; S. acatlensis, S. platyloba, S. prehensilis; southern Mexico, Central America, the West Indies, tropical South America), Pithecellobium (c 75; tropical and subtropical regions), Havardia (6; H. albicans, H. campylacanthus, H. elachistophylla, H. mexicana, H. pallens, H. sonorae; Texas to Central America), Thailentadopsis (3; T. nitida, T. tenuis, T. vietnamensis; Sri Lanka, Thailand, southern Vietnam), Viguieranthus (18; Madagascar, tropical Asia), Zapoteca (c 20; southwestern North America to northern Argentina), Cojoba (c 20; southern Mexico, Central America, the West Indies, tropical South America), Afrocalliandra (2; A. gilbertii, A. redacta; tropical to southern Africa), Calliandra (c 140; pantropical), Pararchidendron (1; P. pruinosum; Java, northern Australia), Falcataria (3; F. moluccana: East Malesia to New Guinea, the Bismarck Archipelago, Solomon Islands; F. pullenii: Papua New Guinea; F. toona: Queensland), Paraserianthes (1; P. lophantha; Malesia), ‘Acacia’ pro parte, Lysiloma (11; Florida, Mexico, the West Indies), Faidherbia (1; F. albida; eastern Mediterranean, tropical to southern Africa), ‘Albizia’ (c 150; pantropical; paraphyletic), Cathormion (4; C. altissimum, C. berteroanum, C. rhombifolium, C. umbellatum; tropical West and Central Africa), Serianthes (18; Thailand, Southeast Asia, Malesia to New Guinea, Melanesia, especially New Caledonia, Micronesia, western Polynesia). – Tropical, subtropical and warm-temperate. Bisexual, monoecious, andromonoecious or polygamomonoecious, evergreen or deciduous trees or shrubs (rarely lianas or perennial, biennial or annual herbs). Often with root nodules containing nitrogen fixing bacteria. Ectomycorrhiza sometimes present. Fibres usually non-septate. Axial parenchyma sometimes aliform. Wood rays usually 20 or more cells high. Sieve tube plastids PIVa type, also with fibres. Leaves usually bipinnately compound (sometimes pinnately compound or modified into phyllodia). Stipules intrapetiolar (often scale-like or modified into spines or glands). Petiolar extrafloral nectaries frequent (sometimes present on petiolules). Stomata usually anomocytic or paracytic (sometimes anisocytic, tetracytic or cyclocytic). With or without mucilaginous idioblasts and secretory cavities containing mucilage, resins or oils. Inflorescence terminal or axillary, usually capitate. Flowers often synchronously developing and opening simultaneously. Floral prophylls (bracteoles) absent. At least corolla actinomorphic, usually small. Hypanthium often absent. Sepals (three to) five (or six), usually with valvate (in the Parkia and Mimozyganthus clades imbricate; in Calliandra cochlear-descending) aestivation, usually more or less connate. Petals (three or) four or five (or six), sessile, usually with valvate (in Dinizia imbricate) aestivation, not clawed, in lower part usually connate into tube; median petal usually abaxial. Stamens as many or twice as many as petals or numerous (in Calliandra spirally initiated), often showy. Filaments free or connate at base, free or adnate to petals, often much exserted. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective often with caducous apical gland. Tapetal cells uninucleate. Pollen grains usually shed in tetrads or polyads (sometimes monads), usually bicellular (sometimes tricellular) at dispersal. Pistil composed of usually one carpel (in Affonsea, Archidendron and Klugiodendron two to 16 free carpels, sometimes antesepalous), usually stipitate (gynophore). Ovary unilocular. Stigma usually cup-shaped (sometimes peltate), Dry type. Placentation marginal (along abaxial suture). Ovules usually anatropous, ascending or pendulous. Micropyle bistomal or endostomal. Synergids often with a filiform apparatus. Arillus sometimes present. Funicle filiform. Testa with vascular bundle. Pleurogram usually present; linea fissura U-shaped (sometimes absent). Endosperm haustorium usually chalazal (rarely lateral). Embryogenesis onagrad, asterad or caryophyllad? Fruit usually a legume (rarely a follicle, a samaroid or a lomentum). Funicular aril sometimes present. Testa with vascular strand. Pleurogram usually present, as well as a linea fissura (a thin line delimitating pleurogram). Funicle often long and thin. Seed usually without hilum fissure. Endosperm thin, with thickened cell walls, with or without starch, or absent. Embryo straight, with chlorophyll. Radicula short and thick, straight. Cotyledons usually without starch grains, without amyloid. Germination phanerocotylar or cryptocotylar. Often with x = 13, 14. Seed often with rare free (non-protein) amino acids (e.g. albizziine; canavanine absent). – The relationships among the different lineages within Mimoseae are largely unresolved.

Phylogeny (somewhat simplified) of Mimosoideae based on DNA sequence data (LPWG 2013).

Papilionoideae L. ex DC., Prodr. 2: 94. Nov (mid) 1825

Faboideae (Giseke) Rudd in Rhodora 70: 496. 31 Dec 1968

460/14.315–14.700. Cosmopolitan except Antarctica. Bisexual evergreen or deciduous trees, shrubs, lianas, suffrutices, or perennial, biennial or annual herbs, often twining with foliar tendrils, usually unarmed. Root nodules with nitrogen fixing bacteria usually present. Nodules usually arising in association with lateral roots. Ectomycorrhiza sometimes present. Phellogen superficially or deeply seated. Vestured pits present in secondary xylem. Sieve tube plastids PIVb type or S type, with spindle-shaped non-dispersive protein bodies (forisomes). Stipules usually intrapetiolar, often scale-like or modified into spines or glands, free or absent (very rarely interpetiolar). Leaves usually paripinnate, imparipinnate or palmately compound, often unifoliolate or trifoliolate (rarely bifoliolate or tetrafoliolate). Leaflets/pinnae opposite or alternate, sometimes modified into tendrils (rarely in phyllodes). Stomata usually anomocytic or paracytic (sometimes anisocytic, tetracytic or cyclocytic). Often with secretory chambers containing mucilage, resins or oils. Extrafloral nectaries absent from petiole and leaf rachis (occasionally present on stipules, stipels, bracts, or swollen and nectar-secreting peduncles or sepals). Inflorescence terminal or axillary, panicle, fascicle, raceme or pseudoraceme (sometimes cyme, spicate or capitate, or flowers solitary). Flowers usually zygomorphic and papilionate (rarely actinomorphic or asymmetrical). Hypanthium present or absent (in some species of Amorpheae stemonozone instead of hypanthium). Sepals (three to) five, with imbricate aestivation, in lower parts usually connate into tube (sometimes entire and splitting into irregular lobes or lobes dimorphic and some petaloid). Petals (1–)5(–6) (absent in some Amorpheae, in other Amorpheae more or less uniform), with imbricate-ascending aestivation; two lower petals usually connate into carina; median adaxial (upper) petal vexillum, enclosing remaining petals in bud; lateral petals alae, enclosing carina in bud (in some genera only adaxial petal free). Petals in Mucuna holtoni functioning as reflectors for ultrasounds (cheiropterophily with nectar guide). Sepals, corolla and stamens with unidirectional (abaxial to adaxial) initiation. Stamens (nine or) ten (to numerous). All filaments connate into tube surrounding pistil or adaxial (posterior) stamen free (rarely all filaments free), usually free from petals (in, e.g., Dalbergieae, Genisteae, Mirbelieae, and Trifolieae often adnate to petals). Anthers basifixed and/or dorsifixed, often versatile, tetrasporangiate, introrse or latrorse, longicidal; connective sometimes with apical gland. Tapetal cells uninucleate. When androecium monadelphic, then pollinator reward often pollen grains liberated by secondary pollen presentation (pollen pump). When androecium diadelphic, then pollinator reward often nectar from glands present between filament tube and gynoecium. Pollen grains shed as monads, usually tricolporate, tricolpate or triporate, bicellular at dispersal. Pollen grains in Lupinus shed as threads. Pistil composed of usually one carpel (monomery; in Swartzia often more than one), often stipitate (gynophore). Ovary unilocular (rarely bilocular due to secondary septum). Pollen grains in Vicia presented on stigmatic brush. Secondary pollen display explosive in theMedicago clade. Placentation marginal (along abaxial suture). Ovules usually campylotropous, ascending or pendulous. Micropyle bistomal (often Z-shaped) or endostomal. Megagametophyte sometimes protruding into micropyle. Synergids often with a filiform apparatus. Endosperm haustorium usually chalazal (rarely lateral). Funicle short. Embryogenesis onagrad, asterad or caryophyllad. Fruit usually a legume, dehiscing along dorsal and ventral sutures simultaneously, valves twisting (often explosively dehiscent; rarely follicle, nut, samaroid, lomentum, or drupe); two layers of lignified fibres present, one or two oblique to long axis of legume. Aril often present. Seed with hilar furrow; hilum often with group of tracheids immediately below hilar surface. Raphe shorter than antiraphe. Testa sometimes multiplicative. Exotesta palisade, cells beneath exotesta hour-glass. Pleurograms and linea fissura usually absent. Endosperm often absent; cell walls sometimes thick. Embryo usually curved (rarely straight), with chlorophyll, sometimes starchy. Suspensor well developed. Cotyledons accumbent, with margins against but not investing radicula, without amyloid (xyloglucans); cotyledon areoles frequent. Radicula usually long, curved (in Sophoreae short and straight). Germination phanerocotylar or cryptocotylar. Duplication of gene CYC. 25 kb plastid IR absent. Inversion of 50 kb in trnL intron in plastid LSC region present in numerous Papilionoideae (absent in Swartzieae and in Sophoreae). Isoflavonoids (pterocarpans, isoflavanes), prenylated flavonoids, pyrrolizidine, indolizidine and quinolizidine alkaloids, canavanine and other non-protein amino acids present. – Forisomes providing a regulatory mechanism of phloem transport, are a synapomorphy of Papilionoideae, which has been lost several times (i.a. in the astragalean clade).

Swartzioideae DC., Prodr. 2: 422. Nov (med.) 1825 [‘Svarzieae’]

8/210–230. Fairchildia (1; F. panamensis; Central America, northwestern South America), Swartzia (180–200; southern Mexico, Central America, the West Indies, tropical South America), Bobgunnia (2; B. fistuloides: tropical and southern Africa; B. madagascariensis: Madagascar), Bocoa (4; B. marionii, B. prouacensis, B. ratteri, B. viridiflora; tropical South America, especially the Guayana Shield and northern Brazil), Candolleodendron (1; C. brachystachyum; northeastern South America); Trischidium (5; T. alternum, T. decipiens, T. limae, T. molle, T. racemulosum; South America), Ateleia (18; Central America, the West Indies, tropical South America), Cyathostegia (1–2; C. mathewsii, C. weberbaueri; Ecuador, Peru). – Tropical and subtropical America and Africa. Trees or shrubs. Root nodules often formed. Floral prophylls (bracteoles) absent. Sepals in Swartzia connate, early caducous. Petal one. Nectary in at least Swartzia absent. Stamens numerous, free, developing from annular meristem. Pistil composed of usually one carpel (sometimes several carpels). Ovary in Swartzia stipitate. Arillus usually present. Embryo in Swartzia straight. – Swartzioideae (at least Swartzia) may be sister-group to the remaining Papilionoideae except Dipterygeae. Some molecular analyses (using the trnL intron) suggested that Swartzioideae are instead sister to the remaining Papilionoideae (including the Dipterygeae), although the support is weak.

Dipterygeae Polhill in R. M. Polhill et P. H. Raven, Adv. Legume System. 1: 231. 1981 [‘Dipteryxeae’]

16/59. Angylocalyceae clade Angylocalyx (6; A. boutiqueanus, A. braunii, A. oligophyllus, A. pynaertii, A. schumannianus, A. talbotii; tropical Africa), Xanthocercis (3; X. madagascariensis, X. rabiensis, X. zambesiaca; Gabon and Zambia to southern Africa, Madagascar), Castanospermum (1; C. australe; western New Britain, eastern Queensland, eastern New South Wales, New Caledonia, Vanuatu), Uleanthus (1; U. erythrinoides; Amazonian Brazil). – Dipterygeae clade Monopteryx (3; M. angustifolia, M. inpae, M. uaucu; tropical South America), Taralea (5; T. cordata, T. crassifolia, T. oppositifolia, T. phaeophylla, T. reticulata; southern Central America, Hispaniola, tropical South America), Dipteryx (9; Central America, the West Indies, tropical South America), Pterodon (2; P. abruptus, P. emarginatus; Brazil, Bolivia). – Amburaneae clade Dussia (10; southern Mexico, Central America, the West Indies, tropical South America), Petaladenium (1; P. urceoliferum; Rio Negro in Brazil), Myrospermum (1; M. frutescens; Mexico, Central America, the West Indies, northern tropical South America), Myrocarpus (4; M. fastigiatus, M. frondosus, M. leprosus, M. venezuelensis; tropical South America), Myroxylon (2; M. balsamum, M. peruiferum; Central America, tropical South America), Dupuya (2; D. haraka, D. madagascariensis; Madagascar), Amburana (3; A. acreana, A. cearensis, A. erythrosperma; tropical South America), Cordyla (6; C. africana, C. densiflora, C. excelsa, C. pinnata, C. richardii, C. somalensis; tropical Africa to South Africa, Madagascar). – Tropical America and Africa, Madagascar, Papuasia to Melanesia. Adaxial two sepals in Dipteryx, Pterodon and Taralea enlarged petaloid; abaxial three sepals forming tridentate lobe. – Dipterygeae are possibly sister-group to the remaining Papilionoideae. Angylocalyceae seem to be sister to [Dipterygeae+Amburaneae]. Some molecular analyses (using matK with low bootstrap support) suggest that the clade [Swartzioideae+Dipterygeae] is sister-group to the remaining Papilionoideae.

[cladrastidoids+50 kb inversion clade]

The bootstrap support of this clade is weak.

Cladrastidoids

3/18. ’Cladrastis’ (9; East Asia, one species, C. kentukea, in southeastern United States; paraphyletic; incl. Pickeringia and Styphnolobium?), Pickeringia (1; P. montana; California; in Cladrastis?), Styphnolobium (8; southern United States, Mexico, Central America; in Cladrastis?). – East Asia, North to Central America. – The cladrastidoids may be sister-group to the remaining Papilionoideae, but the bootstrap support is weak.

50 kb inversion clade (meso-papilionoids)

Inversion of 50 kb usually present in the plastid Large Single Copy Region (situated in the intron between the genes accD and trnL). Root nodules usually formed. – Unplaced: Amphimas (4; A. ferrugineus, A. klaineanus, A. pterocarpoides, A. tessmannii; western Central Africa).

Exostyleae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 254. 20 Jul 1943

6/17–19. Holocalyx (1; H. balansae; tropical South America), Lecointea (4; L. amazonica, L. ovalifolia, L. peruviana, L. tango; tropical South America), Zollernia (8; Central America, tropical South America); Harleyodendron (1; H. unifoliolatum; northeastern Brazil), Exostyles (2–4; E. glabra, E. venusta; southeastern Brazil); Uribea (1; U. tamarindoides; Central America). – Tropical America.

[Dermatophyllum+Ormosieae+[Brongniartieae+Bowdichia clade+[[Sophoreae +Euchresteae+Thermopsideae]+[Podalyrieae+[Crotalarieae+Genisteae s.str.]]]]]

Pyrrolizidine or quinolizidine alkaloids present. x = 9.

Dermatophyllum clade

1/6. Dermatophyllum (6; D. arizonicum, D. gypsophilum, D. guadalupense, D. juanhintonianum, D. secundiflorum, D. purpusii; southwestern United States, northern Mexico). – Trees or shrubs.

Ormosieae Yakovlev in Bobovye Zemnogo Shara: 72. 1991

4/c 140. Ormosia (c 130; East Asia to Queensland, Central and eastern South America), Clathrotropis (6; C. brachypetala, C. brunnea, C. glaucophylla, C. macrocarpa, C. nitida, C. paradoxa; tropical South America), Panurea (1; P. longifolia; Colombia, northern Brazil), Spirotropis (2; S. candollei, S. longifolia; northeastern South America). – Pantropical, few species in temperate East Asia.

Genistoideae F. Schwarz, Forstl. Bot.: 347. Oct 1891 [‘Genisteae’]

80/2.075–2.130. – Brongniartieae and Podalyrieae form well supported clades, whereas the Sophoreae/Crotalarieae/Thermopsideae/Genisteae clade is still largely unresolved. 50 kb inversion absent in Sophoreae. – Unplaced Genistoideae: Pericopsis (4; P. angolensis, P. elata, P. laxiflora, P. mooniana; tropical Africa, Sri Lanka to Micronesia). – Pericopsis may be closely allied to Camoensia (see below).

Brongniartieae Hutch., Gen. Fl. Plants 1: 393. 3 Dec 1964

c 16/150–155. Haplormosia (1; H. monophylla; tropical West and Central Africa); Poecilanthe (7; P. falcata, P. grandiflora, P. itapuana, P. ovalifolia, P. parviflora, P. subcordata, P. ulei; tropical South America); Harpalyce (c 30; Mexico, Cuba, Brazil), Tabaroa (1; T. caatingicola; southwestern Bahia in Brazil), Amphiodon (1; A. effusus; tropical South America), Limadendron (2; L. amazonicum, L. hostmannii; Colombia, southern Venezuela, the Guianas, Amazonian Brazil), Behaimia (1; B. cubensis; Cuba), Cyclolobium (1; C. brasiliense; Brazil, Paraguay, Bolivia), ‘Brongniartia’ ulbrichiana (Bolivia), Brongniartia (50–55; Mexico, Central America, Ecuador, Peru), ‘Templetonia’ (13; Australia; diphyletic), Hovea (35–40; southwestern, southeastern and eastern Australia, Tasmania, Arnhem Land), Cristonia (2; C. biloba, C. stenophylla; western Western Australia), Plagiocarpus (1; P. axillaris; northeastern Western Australia, northwestern Northern Australia), Lamprolobium (2; L. fruticosum, L. grandiflorum; northeastern Queensland), Thinicola (1; T. incana; northern Western Australia). – Tropical and subtropical America, Australia. – Haplormosia is sister to the remaining Brongniartieae and Poecilanthe successive sister to the rest (Cardoso & al. 2017).

Leptolobieae (Benth.) Cardoso et al. in South Afr. J. Bot. 89: 70. Nov 2013

6/27–32. Orphanodendron (1–2; O. bernalii, O. grandiflorum; northwestern Colombia); Diplotropis (10–12; Amazonas to northern Argentina), Guianodendron (1; G. praeclarum; the Guayana Shield in Guyana and northern Brazil), Staminodianthus (3; S. duckei, S. racemosus, S. rosae; Amazonas), Bowdichia (2; B. nitida, B. virgilioides; tropical South America), Leptolobium (10–12; Mexico, Central America, tropical South America). – Tropical America. – The position of Orphanodendron is uncertain. Leptolobieae may be sister-group to the remaining Genistoideae, although the branch support is low.

[Camoensieae+[[Sophoreae+Euchresteae+Thermopsideae]+[Podalyrieae+[Crotalarieae+Genisteae s.str.]]]]

Camoensieae Yakovlev in Bot. Zhurn. (Moscow & Leningrad) 53: 1477. 12 Jul 1968

1/2. Camoensia (2; C. brevicalyx, C. scandens; coasts of the Gulf of Guinea and south to Angola).

[[Sophoreae+Euchresteae+Thermopsideae]+[Podalyrieae+[Crotalarieae+Genisteae]]]

[Sophoreae+Euchresteae+Thermopsideae]

Sophoreae Spreng. ex DC., Prodr. 2: 94. Nov (med.) 1825

6–11/64–76. Ammodendron (5–10; A. bifolium, A. conollyi, A. eichwaldii, A. karelinii, A. maxima; West and Central Asia to northwestern China), Ammothamnus (3; A. gibbosus, A. lehmannii, A. songoricus; Central Asia), Maackia (10–11; East Asia), Salweenia (1–2; S. bouffordiana, S. wardii; southeastern Tibet), Sophora (45–50; southeastern Europe, South Asia to tropical Australia and islands in the Pacific incl. the Hawaiian Islands, western United States, Florida, Puerto Rico, western South America), Ammopiptanthus (2; A. mongolicus, A. nanus; Central Asia to Mongolia and northwestern China). – Tropical Africa, Asia, South America. – Panurea (Ormosieae) and Bowdichia (Leptolobieae) have similar pollen morphology. – Cardoso & al. (2013b) separated Camoensia as a tribe (Camoensieae) sister to [Sophoreae s.lat.+[Podalyrieae+[Crotalarieae+Genisteae s.str.]]]. Sophoreae in a wider sense include Thermopsideae and Euchresteae. – Unplaced Sophoreae s.lat.: Bolusanthus (1; B. speciosus; Northern Province, Mpumalanga, Swaziland), Dicraeopetalum (3; D. stipulare: southeastern Ethiopia, southern Somalia, northeastern Kenya; D. capuronianum, D. mahafaliense: Madagascar), Neoharmsia (2; N. baronii, N. madagascariensis; western and northwestern Madagascar), Platycelyphium (1; P. voense; drier regions in northeastern and eastern Africa), Sakoanala (2; S. madagascariensis, S. villosa; northwestern and eastern Madagascar).

Thermopsideae Yakovlev in Bot. Žurn. (Moscow et Leningrad) 57: 592. 26 Jun 1972

5/61–62. Anagyris (1–2; A. foetida, A. latifolia; the Canary Islands, the Mediterranean, the Middle East to Iran), Piptanthus (2; P. nepalensis, P. tomentosus; the Himalayas), Baptisia (27; eastern and southern United States), Thermopsis (c 30; East Asia, southern Canada, United States; incl. Vuralia?), Vuralia (1; V. turcica; near Lake Aksehir in Turkey; in Thermopsis?). – Macaronesia and the Mediterranean to Central and East Asia, North America.

Euchresteae (Nakai) Ohashi in J. Jap. Bot. 48: 229. 1973

1/4. Euchresta (4; E. formosana, E. horsfieldii, E. japonica, E. tubulosa; India, the Himalayas, China, the Korean Peninsula, Japan, Southeast Asia, Java, Taiwan).

[Podalyrieae+[Crotalarieae+Genisteae]]

Podalyrieae Benth. in S. F. L. Endlicher, E. Fenzl, G. Bentham et H. W. Schott, Enum. Plant.: 27. Apr 1837

9/138–152. Cyclopia (23; Western and Eastern Cape), ’Amphithalea’ (c 42; Western and Eastern Cape; paraphyletic?), Coelidium (c 20; Northern, Western and Eastern Cape; in Amphithalea?), Xiphotheca (9; Western and Eastern Cape; in Amphithalea?), Stirtonanthus (3; S. chrysanthus, S. insignis, S. taylorianus; Western Cape), Podalyria (25–30; Western Cape to KwaZulu-Natal), Liparia (c 20; Western and Eastern Cape), Virgilia (2; V. divaricata, V. oroboides; Western Cape, western Eastern Cape), ’Calpurnia’ (7 or 16?; C. aurea, C. capensis, C. glabrata, C. robinioides, C. sericea, C. villosa, C. woodii; Africa incl. South Africa; paraphyletic), Cadia (7; C. commersoniana, C. ellisiana, C. emarginatior, C. pedicellata, C. pubescens, C. purpurea, C. rubra; Madagascar, one species in northeastern tropical Africa to southern Arabian Peninsula). – South Africa, few species to northeastern Africa and the Arabian Peninsula. Methylated anthocyanins, carboxylic acid esters of quinolizidine alkaloids, 3’-hydroxydaidzein, and canavanine (in seeds) present. Alkaloids not found. – The position of Cadia is uncertain. Its species possess radially symmetrical flowers, showing complete expression of the Cycloidea genes (Citerne & al. 2006).

Oberholzeria (1; O. etendekaensis; northwestern Namibia). – Oberholzeria is sister to [Crotalarieae+Genisteae], according to Swanepoel & al. (2015).

[Crotalarieae+Genisteae]

Crotalarieae Hutch., Gen. Fl. Plants 1: 364. 3 Dec 1964

16/1.095? Euchlora (1; E. hirsuta; Northern and Western Cape), Bolusia (5; B. acuminata, B. amboensis, B. ervoides, B. grandis, B. resupinata; arid and semiarid regions of Africa south of the equator), Crotalaria (c 500?; tropical and subtropical regions on both hemispheres, with their largest diversity in Africa and Madagascar), ’Wiborgiella’ (9; Northern, Western and Eastern Cape; polyphyletic), Ezoloba (1; E. macrocarpa; Western Cape), Calobota (15; North and South Africa), ’Lebeckia’ (22; Namibia, Northern, Western and Eastern Cape, KwaZulu-Natal; paraphyletic), Wiborgia (10; Northern and Western Cape), Rafnia (19; Western and Eastern Cape, KwaZulu-Natal), Aspalathus (c 290; Western Cape to KwaZulu-Natal), ’Lotononis’ (c 150; the Mediterranean, Africa, southwestern Asia to India, with their largest diversity in South Africa; paraphyletic), Listia (7; L. angolensis, L. bainesii, L. heterophylla, L. marlothii, L. minima, L. solitudinis, L. subulata; southeastern Africa), Pearsonia (c 12; tropical to southern Africa), Rothia (2; R. hirsuta, R. indica; Africa to South Africa, India and Australia), Robynsiophyton (1; R. vanderystii; southern Central Africa), Leobordea (51; tropical and southern Africa, northeastern Africa, southwestern Asia). – Mainly South Africa, few species in eastern and northeastern Africa and southwestern Asia.

Genisteae Bronn in B. C. J. Dumortier, Fl. Belg.: 98. 1827

9/525–545. Cytisus (c 140; Europe, Africa, Asia), Genista (c 130; Europe, Macaronesia, the Mediterranean, Asia), ’Adenocarpus’ (c 15; the Canary Islands, the Mediterranean, tropical African mountains; paraphyletic), Lupinus (200–220; the Mediterranean, North and East Africa, America, with their largest diversity in western North and western South America), Anarthrophyllum (15; the Andes), Sellocharis (1; S. paradoxa; southeastern Brazil), Dichilus (5; D. gracilis, D. lebeckioides, D. pilosus, D. reflexus, D. strictus; southern Africa), Polhillia (6; P. brevicalyx, P. canescens, P. connata, P. ignota, P. obsoleta, P. pallens; Western Cape), Melolobium (15; southern Africa). – Eurasia, Africa, North and South America. α-pyridone-type quinolizidine alkaloids present.

[Vataireoid clade+[Andirinae+Dalbergioideae]]

Vataireoid clade

4/26. Luetzelburgia (13; northeastern South America to Bolivia, with their highest diversity in Brazil); Sweetia (1; S. fruticosa; tropical South America), Vataireopsis (4; V. araroba, V. iglesiasii, V. speciosa, V. surinamensis; northern South America), Vatairea (8; tropical America). – Tropical America. Root nodules not formed. – Luetzelburgia is sister to [Sweetia+[Vataireopsis+Vatairea]], according to Cardoso & al. (2013).

[Andirinae+Dalbergioideae]

Andirinae Baill. in M. A. Hartog [trans.], Nat. Hist. Plant. 2: 217. Sep-Dec 1872 [‘Andireae’]

3/c 70. Aldina (17; the Guayana Shield to northern Amazonia in Brazil), Hymenolobium (13; tropical South America), Andira (c 40; Central America, tropical South America, one species, A. inermis, also in tropical West Africa). – Tropical Central and South America, tropical West Africa. – Aldina has ectomycorrhiza.

Dalbergioideae Burnett, Outlines Bot.: 661. Feb 1835 [‘Dalbergidae’]

Amorpheae Boriss. in Novit. Syst. Plant. Vasc. 1964: 224. 14 Oct 1964

8/245–250. Amorpha (16; southern Canada, United States, northern Mexico, with their highest diversity in southeastern United States), Parryella (1; P. filifolia; Mexico), Apoplanesia (2; A. cryptopetala, A. paniculata; drier regions in Central America), Errazurizia (4; E. benthamii, E. megacarpa, E. multifoliolata, E. rotundata; arid regions in southwestern United States, coastal Chile), Eysenhardtia (13; Central America), ’Psorothamnus’ (c 10; arid regions in southwestern North America; paraphyletic), Marina (c 40; southwestern United States, Mexico, Central America, Venezuela), Dalea (c 160; arid and semiarid regions in Canada to Argentina, especially Mexico and the Andes). – America, with their highest diversity in drier regions. Petals often absent. Filaments often adnate to petals (epipetalous) forming tubular stemonozone (hypanthium absent).

Dalbergieae DC., Prodr. 2: 415. Med Nov 1825

45/1.290–1.335. Adesmia (210–220; South America), Poiretia (11; tropical America), Amicia (8; Mexico, Central America, tropical South America), Zornia (80–85; warmer regions on both hemispheres, with their largest diversity in Brazil), Nissolia (14; Mexico, Central America, the West Indies, tropical South America), Chaetocalyx (c 15; southern Mexico, Central America, Hispaniola, tropical South America); Riedeliella (3; R. graciliflora, R. magalhaesii, R. sessiliflora; Brazil, Paraguay); Platypodium (2; P. elegans, P. viride; Central America, tropical South America), Paramachaerium (5; P. gruberi, P. krukovii, P. ormosioides, P. schomburgkii, P. schunkei; Central America, tropical South America), Pterocarpus (c 35; tropical and subtropical regions), Ramorinoa (1; R. girolae; western Argentina), Centrolobium (7; C. microchaete, C. ochroxylum, C. paraense, C. robustum, C. sclerophyllum, C. tomentosum, C. yavizanum; southern Central America, tropical South America), Tipuana (1; T. tipu; Brazil, Bolivia, Argentina), Inocarpus (3; I. fagifer, I. glabellus, I. papuanus; Malesia to New Guinea, islands in the Pacific), Discolobium (8; eastern South America), Platymiscium (c 20; Mexico, Central America, tropical South America), Acosmium (3; A. cardenasii: southwestern Brazil, Bolivia; A. diffusissimum, A. lentiscifolium: southeastern Brazil), Maraniona (1; M. lavinii; northern Peru), Chapmannia (7; C. floridana, C. gracilis, C. prismatica, C. reghidensis, C. sericea, C. somalensis, C. tinireana; Somalia, Yemen, Socotra, one species, C. floridana, in Florida, southern Mexico, Guatemala, northern Venezuela), Arachis (c 70; South America), Stylosanthes (40–45; tropical and subtropical regions), Grazielodendron (1; G. riodocensis; Brazil), Cranocarpus (3; C. gracilis, C. martii, C. mezii; Brazil), Brya (6; B. buxifolia, B. chrysogonii, B. ebenus, B. hirsuta, B. microphylla, B. subinermis; the West Indies), Cascaronia (1; C. astragalina; Bolivia, northern Argentina), Geoffroea (2; G. decorticans, G. spinosa; South America), Fissicalyx (1; F. fendleri; Venezuela, Guyana), Fiebrigiella (1; F. gracilis; Bolivia); Machaerium (c 155; southern Mexico, Central America, the West Indies, tropical South America, one species, M. lunatum, also in coastal tropical West Africa), Steinbachiella (1; S. leptoclada; Bolivia), Dalbergia (250–270; tropical Africa, Madagascar, tropical Asia, Central America, tropical South America), ’Aeschynomene’ (c 175; tropical and subtropical regions; polyphyletic), Cyclocarpa (1; C. stellaris; tropical Africa, tropical Australia), Kotschya (c 30; tropical to southern Africa, Madagascar), Soemmeringia (1; S. semperflorens; tropical South America), Smithia (c 22; tropical and subtropical regions in the Old World), Humularia (30–35; Central Africa), Bryaspis (2; B. humularioides, B. lupulina; tropical West Africa), Geissaspis (4; G. cristata, G. keilii, G. psittacorhyncha, G. tenella; tropical and subtropical Asia), Pictetia (10; the West Indies), Diphysa (c 20; Mexico, Central America, the West Indies, tropical South America), Zygocarpum (6; Z. coeruleum, Z. dhofarense, Z. gillettii, Z. rectangulare, Z. somalense, Z. yemenense; Somalia, southern Arabian Peninsula, Socotra), Ormocarpum (16; tropical and subtropical regions in the Old World), Ormocarpopsis (7–8; O. aspera, O. calcicola, O. itremoensis, O. mandrarensis, O. nitida, O. parvifolia, O. tulearensis; Madagascar; in Ormocarpum?), Weberbauerella (1–2; W. brongniartioides, W. raimondiana; coast of Peru). – Tropical and subtropical with their largest diversity in tropical South America. Desmodioid (aeschynomenoid) root nodules with determinate growth and associated with lateral roots and glandular-based trichomes are synapomorphies of Dalbergieae (Chaetocalyx and Nissolia produce no nodules; glandular-based trichomes are present in most species).

Old World clade

[Baphieae+Canavanine-accumulating clade]

Baphieae Yakovlev in Bot. Žurn. (Moscow et Leningrad) 53: 1477. 1968

5/58–63. Baphia (45–50; tropical to southern Africa, Madagascar), Baphiopsis (1; B. parviflora; tropical Africa), Leucomphalos (6; L. brachycarpus, L. callicarpa, L. capparideus, L. discolor, L. insignis, L. mildbraedii; tropical Africa, Madagascar), Airyantha (3; A. borneensis, A. confusum, A. schweinfurthii; Central Africa), Dalhousiea (3; D. africana, D. bracteata, D. paucisperma; tropical West Africa, northeastern India, Bangladesh). – Tropical Africa to East Asia. Trees, shrubs, lianas. – Baphia is sister to the remaining Papilionoideae, the Canavanine-accumulating clade.

Canavanine-accumulating clade (non-protein amino acid-accumulating clade, NPAAA clade)

Canavanine (non-protein amino acid) present. – The peptide pea albumin is present in Phaseoleae and Hologalegina (absent in Robinieae).

Hypocalypteae (Yakovlev) A. L. Schutte in A. L. Schutte et B.-E. van Wyk, Novon 8: 180. 1998

1/3. Hypocalyptus (3; H. coluteoides, H. oxalidifolius, H. sophoroides; Western and Eastern Cape).

Mirbelieae (Benth.) Polhill in R. M. Polhill et P. H. Raven, Adv. Legume System. 1: 391. 1981

32/715–>745. Giant antipodals group 1 Daviesia (c 135; Australia, with their highest diversity in southwestern Australia), Erichsenia (1; E. uncinata; southwestern Western Australia), Viminaria (1; V. juncea; southwestern Western Australia, coastal areas of southeastern Australia). – Giant antipodals group 2 Gompholobium (44; Australia, with their highest diversity in southwestern Australia), Sphaerolobium (18–22; Australia, with their highest diversity in southwestern Australia), Goodia (2; G. lotifolia, G. medicaginea; southwestern and southeastern Australia), Bossiaea (c 60; temperate Australia, with their highest diversity in southwestern Australia), Platylobium (4; P. alternifolium, P. formosum, P. obtusangulum, P. triangulare; southeastern South Australia to southeasternmost Queensland, Tasmania), Muelleranthus (2; M. stipularis, M. trifoliolatus; Australia), Paragoodia (1; P. crenulata; southwestern Western Australia), Ptychosema (2; P. anomalum, P. pusillum; Central and Western Australia), Aenictophyton (1; A. reconditum; northwestern Australia). – Multiple embryo-sacs group Isotropis (14–16; Australia); Jacksonia (35–40; Australia); Leptosema (6; L. aculeatum, L. anomalum, L. bossiaeoides, L. chambersii, L. daviesioides, L. uniflorum; northern, central and southwestern Australia); Chorizema (c 20; southwestern and eastern Australia), ‘Mirbelia’ (25–30; Australia; polyphyletic), Oxylobium (17; Australia, with their highest diversity in southwestern Australia), ‘Podolobium’ (6?; southwestern and southeastern Australia; polyphyletic); Callistachys (6; C. aciculifera, C. cordifolia, C. hamulosa, C. lanceolata, C. microphylla, C. procumbens; southwestern Western Australia), Gastrolobium (100–110; Australia); Euchilopsis (1; E. linearis; southwestern Western Australia), Phyllota (10–11; southwestern and eastern Australia), Latrobea (5; L. brunonis, L. diosmifolia, L. genistoides, L. hirtella, L. tenella; southwestern Western Australia), Urodon (2; U. capitatus, U. dasyphyllus; southwestern Western Australia), Otion (8; northern, central and southwestern Australia)?, Aotus (14–18; Australia), Stonesiella (1; S. selaginoides; Tasmania), Almaleea (5; A. cambagei, A. capitata, A. incurvata, A. paludosa, A. subumbellata; eastern New South Wales, northeasternmost Victoria), Eutaxia (c 25; Australia, with their highest diversity in southwestern Australia), Dillwynia (c 22; Australia), ‘Pultenaea’ (>110; southwestern and southeastern Australia; non-monophyletic). – Australia, with their highest diversity in southwestern and southern parts. – Mirbelieae are an exclusively Australian clade. Aberrant types of megagametophyte development or antipodal cells of unusual size are peculiar features characterizing subclades in this group.

[[Indigofereae+Phaseoleae]+Hologalegina]

[Indigofereae+Phaseoleae]

Floral development with early expression of monosymmetry.

Indigofereae Benth. in C. F. von Martius, S. Endlicher et I. Urban, Fl. Bras. 15(1): 5-6, 35-36. 30 Jul 1859

6/>800. Phylloxylon (7; P. arenicola, P. decipiens, P. perrieri, P. phillipsonii, P. spinosa, P. xiphoclada, P. xylophylloides; Madagascar); Indigofera (>750; tropical and subtropical regions), Cyamopsis (4; C. dentata, C. senegalensis, C. serrata, C. tetragonoloba; drier regions in Africa south to South Africa and east to India), Indigastrum (9; tropical and southern Africa, one pantropical species), Microcharis (c 35; Africa south to South Africa, Madagascar, the Arabian Peninsula), Rhynchotropis (2; R. marginata, R. poggei; southern Central Africa). – Pantropical, with their largest diversity in Africa and Madagascar. Inflorescence racemose.

Disynstemon (1; D. paullinioides; southwestern Madagascar) may be sister to the millettioids (Phaseoleae).

Phaseoleae DC., Prodr. 2: 381. mid Nov 1825 (the millettioids)

164/3.210–3.300. Clitoriinae DC., Prodr. 2: 216. Nov (med.) 1825 [‘Clitoriae’]. Clitoria (c 45; tropical and subtropical regions, with their largest diversity in South America), Centrosema (c 35; tropical and subtropical regions in America), Periandra (7; P. berteriana, P. coccinea, P. densiflora, P. gracilis, P. heterophylla, P. mediterranea, P. pujalu; Hispaniola, Brazil, Bolivia), Clitoriopsis (1; C. mollis; Congo, Sudan). – Austrosteenisia (4; A. blackii, A. glabristyla, A. mollitricha, A. stipularis; New Guinea, northern Australia). – Abrinae Wight et Arn. ex Endl., Gen. Plant.: 1301. Oct 1840 [‘Abrineae’]. Abrus (13–18; tropical and southern Africa, Madagascar, the Arabian Peninsula, tropical Asia to China and tropical Australia). – Diocleinae Benth., Comm. Legum. Gen.: 113. Jun 1837 [‘Diocleae’]. Canavalia (c 50; warmer regions of both hemispheres incl. the Hawaiian Islands, with their highest diversity in tropical and subtropical America); ’Dioclea’ (c 50; tropical America, few species in the Old World; non-monophyletic; incl. Cleobulia, Cymbosema, Macropsychanthus), Macropsychanthus (3; M. dolichobotrys, M. lauterbachii, M. mindanaensis; New Guinea, New Britain, Solomon Islands; in Dioclea), Cymbosema (2; C. apurense, C. roseum; tropical America; in Dioclea), Cleobulia (3–5; C. multiflora; Mexico, Brazil; in Dioclea); Rhodopis (2; R. lowdenii, R. planisiliqua; Hispaniola), Neorudolphia (1; N. volubilis; Puerto Rico); Bionia (5; B. bella, B. coccinea, B. coriacea, B. pedicellata, B. tomentosa; Brazil, especially Minas Gerais), Cratylia (4; C. argentea, C. bahiensis, C. hypargyraea, C. mollis; South America), Lackeya (1; L. multiflora; southeastern United States), ‘Camptosema’ (5–10; C. coriaceum, C. paraguariense, C. pedicellatum, C. rubicundum, C. scarlatinum; South America; polyphyletic), ’Galactia’ (c 55; tropical and subtropical regions, with their largest diversity in South America; non-monophyletic), Luzonia (1; L. purpurea; the Philippines). – Millettieae Miq., Fl. Ned. Ind. 1(1): 137. 2 Aug 1855. Dalbergiella (2; D. nyasae, D. welwitschii; tropical Africa), ’Millettia’ (c 150; tropical and subtropical regions in the Old World, especially Africa and Madagascar; polyphyletic), Deguelia (15–17; Panamá to Amazonas), Leptoderris (c 40; tropical Africa), Philenoptera (13; tropical Africa, Madagascar), Ophrestia (15–20; Africa, southern Asia), Apurimacia (2; A. boliviana, A. dolichocarpa; drier regions in South America), Mundulea (13–14; southern Africa, Madagascar), Ptycholobium (3; P. biflorum, P. contortum, P. plicatum; drier regions in northeastern and southern Africa, the Arabian Peninsula), Tephrosia (>350; tropical and subtropical regions on both hemispheres, especially Africa), Piscidia (4; P. carthagenensis, P. grandifolia, P. mollis, P. piscipula; Florida, Central America, the West Indies), Lonchocarpus (120–130; tropical America, one species, L. sericeus, also in tropical West Africa; paraphyletic?), Pongamiopsis (3; P. amygdalina, P. pervilleana, P. viguieri; Madagascar), ‘Fordia’ (18; Southeast Asia, West Malesia; non-monophyletic), Burkilliodendron (1; B. album; the Malay Peninsula), ‘Solori’ (9?; Southeast Asia; non-monophyletic), ’Derris’ (c 65; tropical regions of the Old World to New Guinea, with their largest diversity in Southeast Asia; non-monophyletic); Platycyamus (1; P. regnellii; Brazil, Peru); Kunstleria (11; Kerala in southwestern India, West and Central Malesia, tropical Australia); Apios (8–10; East Asia, southern Canada, United States); Craibia (2; C. atlantica, C. brevicaudata; tropical to southern Africa), Ostryocarpus (2; O. riparius, O. zenkerianus; tropical West and Central Africa; incl. Xeroderris?), Xeroderris (1; X. stuhlmannii; tropical to southern Africa; in Ostryocarpus?), Platysepalum (7–8; tropical Africa), Schefflerodendron (3–4; S. usambarense; tropical Africa), Sylvichadsia (4; S. grandidieri, S. grandifolia, S. macrophylla, S. perrieri; Madagascar), Hesperothamnus (5; H. ehrenbergii, H. grandis, H. littoralis, H. pentaphyllus, H. purpusii; Mexico), Dahlstedtia (1; D. pinnata; southern Brazil, northern Argentina), Aganope (3–6; A. gabonica, A. heptaphylla, A. thyrsiflora; tropical Africa, Southeast Asia), Pyranthus (6; P. alasoa, P. ambatoana, P. lucens, P. monantha, P. pauciflora, P. tulearensis; Madagascar), Chadsia (c 17?; Madagascar), Requienia (3; R. obcordata, R. pseudosphaerosperma, R. sphaerosperma; western to northeastern Africa, southern Africa), Dewevrea (1; D. bilabiata; tropical West Africa). – Kennediinae Benth., Comm. Legum. Gen.: 112. Jun 1837 [‘Kennedyeae’]. Shuteria (3; S. densiflora, S. hirsuta, S. involucrata; tropical Asia), Kennedia (14–16; New Guinea, Australia), Hardenbergia (3; H. comptoniana, H. perbrevidens, H. violacea; Australia), Vandasina (1; V. retusa; New Guinea, northeastern Queensland). – Desmodiinae Benth. et Hook. f., Gen. Plant. 1: 449. 19 Oct 1865 [‘Desmodieae’]. Craspedolobium (2; C. schochii, C. unijugum; western China); Campylotropis (c 37; the Himalayas, China, the Korean Peninsula, Taiwan), ‘Lespedeza’ (c 40; tropical and eastern Asia, Australia, temperate North and South America; paraphyletic), Kummerowia (2; K. stipulacea, K. striata; East Asia; in Lespedeza?); Hylodesmum (16; tropical Asia), Pseudarthria (6; P. confertiflora, P. crenata, P. fagifolia, P. hookeri, P. macrophylla, P. viscida; southern Africa, Madagascar, Mauritius, Réunion, tropical Asia), ‘Desmodium’ (c 275; tropical and subtropical regions; polyphyletic), Grona (41; tropical and subtropical regions), ‘Alysicarpus’ (30–35; tropical and subtropical Africa, southern Asia; paraphyletic), Melliniella (1; M. micrantha; western central Africa; in Alysicarpus?), Leptodesmia (3; L. bojeriana, L. congesta, L. perrieri; Madagascar, India?), Codariocalyx (2; C. gyroides, C. motorius; India, Sri Lanka, China, Indochina, Malesia to tropical Australia, Taiwan), ‘Uraria’ (c 20; tropical and subtropical regions in the Old World; non-monophyletic), Mecopus (1; M. nidulans; India, Hainan, Indochina, Java), Christia (2; C. obcordata, C. vespertilionis; India to China and Indochina, Malesia to northern Australia), Eleiotis (2; E. monophyllos, E. rottleri; India and Sri Lanka to Burma), Ototropis (13?; tropical Asia, southern China, Taiwan), Pycnospora (1; P. lutescens; tropical East Africa to Somalia, India, East and Southeast Asia to northern Australia), Trifidacanthus (1; T. unifoliolatus; Hainan, southern Vietnam, the Philippines, Lombok, Flores); Ougeinia (1; O. oojeinensis; India, western Nepal), Aphyllodium (4; A. australiense, A. biarticulatum, A. hispidum, A. novoguineense; India, Sri Lanka, Hainan, Indochina, Malesia to New Guinea, northern Australia), Tadehagi (4; T. pseudotriquetrum, T. robustum, T. rodgei, T. triquetrum; tropical Asia), Akschindlium (1; A. godefroyanum; Southeast Asia), Droogmansia (6; D. chevalieri, D. megalantha, D. munamensis, D. pteropus, D. scaettaiana, D. wheptei; tropical Africa), Dendrolobium (c 20; tropical Asia, Indian Ocean islands, Australia), Phyllodium (8; tropical and subtropical Asia to northern Australia), Ohwia (1; O. caudata; India, China, Japan, Indochina, Malesia, Taiwan), Hanslia (2; H. hentyi, H. ormocarpoides; Malesia to New Guinea, northern Queensland, Vanuatu), ‘Nephrodesmus’ (5–7; N. albus, N. ferrugineus, N. francii, N. hochreutineri, N. macrobotryosus, N. parvifolius, N. sericeus; New Caledonia; polyphyletic), Arthroclianthus (c 20; New Caledonia, one species also in Vanuatu), Verdesmum (1; V. hentyi; Yunnan). – Cajaninae Benth., Comm. Legum. Gen.: 113. Jun 1837 [‘Cajaneae’]. Butea (2; B. monosperma, B. superba; tropical Asia), Cajanus (37; tropical regions in the Old World to Australia), Rhynchosia (c 250; tropical and subtropical regions), Eriosema (>150; tropical and subtropical regions), Adenodolichos (22; tropical Africa), Paracalyx (6; P. balfourii, P. microphyllus, P. nogalensis, P. scariosus, P. schweinfurthii, P. somalorum; Ethiopia, Somalia, Socotra, India, Indochina), Bolusafra (1; B. bituminosa; Western Cape), Carrissoa (1; C. angolensis; Angola), Dunbaria (20–25; India, Southeast Asia to China, Malesia to New Guinea, northern Australia), Flemingia (c 30; tropical regions in the Old World). – Erythrininae Benth., Comm. Legum. Gen.: 113. Jun 1837 [‘Erythrineae’]. Otoptera (2; O. burchelli: tropical Africa; O. madagascariensis: Madagascar), Erythrina (c 130; tropical and subtropical regions on both hemispheres), Psophocarpus (7; P. grandiflorus, P. lancifolius, P. lukafuensis, P. monophyllus, P. palustris, P. scandens, P. tetragonolobus; tropical and subtropical regions in the Old World), Spatholobus (c 30; Southeast Asia to Central Malesia), Meizotropis (2; M. buteiformis, M. pellita; India, the Himalayas, western Indochina), Cochlianthus (2; C. gracilis, C. montanus; Nepal, western China). – Psoraleeae Benth. in C. F. von Martius, S. Endlicher et I. Urban, Fl. Bras. 15(1): 3–4, 31–34. 1859 [‘Psoraleae’]. Calopogonium (4; C. caeruleum, C. galactioides, C. mucunoides, C. velutinum; Mexico, Central America, the West Indies, tropical South America, one species, C. mucunoides, also in the Old World tropics), Pseudovigna (1; P. argentea; tropical Africa), Neorautanenia (3; N. amboensis, N. ficifolius, N. mitis; southern tropical Africa), Pachyrhizus (5; P. ahipa, P. erosus, P. ferrugineus, P. panamensis, P. tuberosus; Mexico, Central America, the West Indies, tropical South America); Neonotonia (2; N. verdcourtii, N. wightii; tropical and southern Africa, tropical Asia from southern Arabian Peninsula to Malesia); ’Pueraria’ (17–20; southern and eastern Asia; non-monophyletic); Amphicarpaea (5; A. africana, A. bracteata, A. edgeworthii, A. ferruginea, A. linearis; Africa, East Asia, North America), Teramnus (7; T. flexilis, T. labialis, T. micans, T. mollis, T. repens, T. uncinatus, T. volubilis; tropical and subtropical regions on both hemispheres), ’Glycine’ (c 25; Africa, southern Asia to Australia; non-monophyletic?), Sinodolichos (2; S. lagopus, S. oxyphyllus; Burma, southern China; in Glycine?), Bituminaria (4; B. acaulis, B. bituminosa, B. flaccida, B. morisiana; the Mediterranean to the Caucasus and the Middle East), ’Otholobium’ (50–55; eastern and southeastern Africa, South Africa, South America; non-monophyletic), Orbexilum (11; the United States, Mexico), Psoralea (30–35; southern Africa, with their largest diversity in Western Cape), Cullen (c 35; the Mediterranean, Africa, southern Asia to New Guinea and Australia), Pediomelum (20–25; North America), Ladeania (2; L. juncea, L. lanceolata; western United States), Rupertia (3; R. hallii, R. physodes, R. rigida; southwestern Canada, western United States), Herpyza (1; H. grandiflora; western Cuba), Teyleria (3; T. koordersii; Southeast Asia, West Malesia), Dumasia (c 10; Africa to southern Asia), Nogra (4; N. dalzellii, N. filicaulis, N. grahamii, N. guangxiensis; India to southern China and Thailand; non-monophyletic?), Eminia (4; E. antennulifera, E. benguellensis, E. harmsiana, E. holubii; Zambesian region in Africa), Pseudeminia (4; P. benguellensis, P. comosa, P. mendoncae, P. muxiria; tropical Africa), Phylacium (2; P. bracteosum, P. majus; Southeast Asia to Queensland), Neocollettia (1; N. wallichii; Burma, Java), Mastersia (1; M. bakeri; Assam, Central Malesia), Diphyllarium (1; D. mekongense; Indochina). – Phaseolinae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 346, 402. 1846 [‘Phaseolaceae’]. Cologania (12; tropical America, with their highest diversity in Mexico); Wajira (5; W. albescens, W. danissana, W. grahamiana, W. praecox, W. virescens; Africa, southwestern Arabian Peninsula, India, Sri Lanka); Sphenostylis (5; S. angustifolia, S. erecta, S. marginata, S. schweinfurthii, S. stenocarpa; Africa, India), Decorsea (6; D. dinteri, D. galpinii, D. grandidieri, D. livida, D. meridionalis, D. schlechteri; tropical to southern Africa, Madagascar), Macrotyloma (c 25; tropical and subtropical Africa and Asia), Alistilus (3; A. bechuanicus, A. jumellei, A. magnificus; southern tropical Africa, Madagascar), Dolichos (c 60; tropical and subtropical Africa to South Africa to India and East Asia), Nesphostylis (4; N. bracteata, N. holosericea, N. junodii, N. lanceolata; tropical Africa, tropical Asia); Dipogon (1; D. lignosus; Western and Eastern Cape), Lablab (1; L. purpureus; tropical and subtropical Africa), Spathionema (1; S. kilimandscharicum; tropical Africa), Vatovaea (1; V. pseudolablab; tropical East Africa to Oman), Physostigma (2–4; P. mesoponticum, P. venenosum; tropical Africa), ’Vigna’ (90–100; tropical and subtropical regions on both hemispheres; non-monophyletic), Phaseolus (c 70; tropical and subtropical regions in America), Oxyrhynchus (4; O. papuanus, O. populneus, O. trinervus, O. volubilis; East Malesia, Central America), Ramirezella (7; R. calcoma, R. crassa, R. lozanii, R. micrantha, R. nitida, R. penduliflora, R. strobilophora; Mexico, Central America), Cochliasanthus (1; C. caracalla; northern South America, Bolivia, Argentina), Leptospron (2; L. adenanthum, L. gentryi; Mexico, Central America, Colombia), Sigmoidotropis (9; Mexico, Central America, Hispaniola, tropical South America to Peru), Helicotropis (4; H. hookeri, H. linearis, H. spectabilis, H. stenoloba; Mexico, Central America, tropical South America to Bolivia), Ancistrotropis (6; A. arrabidae, A. clitorioides, A. formula, A. peduncularis, A. robusta, A. subhastata; Mexico, Central America, tropical South America to Bolivia), Condylostylis (4; C. candida, C. latidenticulata, C. venusta, C. vignoides; Mexico, Central America, tropical South America to Argentina), Macroptilium (19; Mexico, Central America, the West Indies, tropical South America), Mysanthus (1; M. uleanus; Brazil), Dolichopsis (2; D. ligulata, D. paraguariensis; southern Brazil, Paraguay, northern Argentina), Strophostyles (3; S. helvola, S. leiosperma, S. umbellata; southern Canada, United States), Mucuna (100–105; tropical and subtropical regions on both hemispheres), Pseudoeriosema (5; P. andongense, P. borianii, P. homblei, P. longipes, P. moeroense; tropical Africa), Oryxis (1; O. monticola; Minas Gerais in Brazil), Austrodolichos (1; A. errabundus; northern Australia), Dysolobium (6; D. apioides, D. dolichoides, D. grande, D. lucens, D. pilosum, D. tetragonum; eastern India, southwestern China, Indochina, Malesia), Strongylodon (14; Madagascar, tropical Asia to Polynesia). – Tropical to warm-temperate regions. Inflorescence often pseudoracemose. – Phaseoleae are still in a state of flux. The relationships among the many lineages are far from understood and the clades have low bootstrap support. However, Clitoriinae may be sister-group to the remaining Phaseoleae, and the clade [Abrinae+[Dioclinae+Millettieae]] (the core millettioids) sister to the rest (the phaseoloids).

Hologalegina (galegoids)

Hologalegina consist of two mainly northern hemispherical major lineages, the Robinioid clade and the Inverted Repeat-Lacking Clade.

[Robinioid clade+IRLC]

Robinioid clade

Loteae DC., Prodr. 2: 115. mid Nov 1825

19/290–300. Antopetitia (1; A. abyssinica; mountains in tropical East Africa), Pseudolotus (1; P. villosus; Oman to Pakistan), Podolotus (1; P. hosackioides; Oman to western India), Dorycnopsis (2; D. abyssinica, D. gerardii; western Mediterranean, northeastern Africa, the Arabian Peninsula), Ornithopus (5; O. compressus, O. micranthus, O. perpusillus, O. pinnatus, O. sativus; Europe, the Mediterranean, western Asia, Atlantic islands, temperate South America), Hosackia (14; southwestern Canada to Guatemala), Acmispon (17; southern Canada to Mexico, one species, A. subpinnatus, in Chile; incl. Kebirita, Ottleya and Syrmatium?), Kebirita (1; K. roudairei; northwestern Africa; in Acmispon?), Ottleya (10; southwestern North America; in Acmispon?), Syrmatium (12; western North America; in Acmispon?), Hippocrepis (34; Europe, the Mediterranean, western Asia), Scorpiurus (2; S. muricatus, S. vermiculatus; Macaronesia, the Mediterranean, the Near East to Iran), Coronilla (c 20; Europe, the Mediterranean, Atlantic islands; incl. Securigera?), Securigera (12; Europe, the Mediterranean, northeastern Africa to Somalia; in Coronilla?), Anthyllis (22; Europe, Macaronesia, the Mediterranean, northeastern Africa), Hammatolobium (2; H. kremerianum, H. lotoides; the Mediterranean), Tripodion (1; T. tetraphyllum; the Mediterranean), Cytisopsis (2; C. dorycniifolia, C. pseudocytisus; eastern Mediterranean, North Africa), Lotus (130–140; temperate regions in the Old World, Macaronesia, the Mediterranean, southwestern Asia). – Temperate regions on the Northern Hemisphere, temperate South America.

[Sesbanieae+Robinieae]

The support of this clade is fairly weak.

Sesbanieae Hutch., Gen. Flow. Pl. 1: 401. 3 Dec 1964

1/c 60. Sesbania (c 60; tropical and subtropical regions on both hemispheres). – Sesbania is sister to either Robinieae or Loteae.

Robinieae Hutch., Gen. Fl. Plants 1: 366. 3 Dec 1964

11/64–76. Robinia (5–10; southern Canada, United States), Genistidium (1; G. dumosum; Texas, Mexico), Peteria (3; P. glandulosa, P. scoparia, P. thompsoniae; southwestern North America), Coursetia (35; southwestern United States, Mexico, Central America, the West Indies, tropical South America to Brazil and Peru), Olneya (1; O. tesota; southwestern United States, northwestern Mexico), Sphinctospermum (1; S. constrictum; western Mexico), Poissonia (4; P. eriantha, P. heterantha, P. orbicularis, P. weberbaueri; Peru, Bolivia, Argentina), Lennea (3–5; L. melanocarpa, L. modesta, L. viridiflora; Central America), Hebestigma (1; H. cubense; Cuba), Gliricidia (5; G. brenningii, G. ehrenbergii, G. maculata, G. robustum, G. sepium; tropical America), Poitea (5–10; P. carinalis, P. florida, P. galegoides, P. paucifolia, P. punicea; the West Indies). – Warm-temperate to tropical regions in America. Loss of plastid inverted repeat.

IRLC (inverted repeat-lacking clade)

Leaves without pulvini, with somewhat different development. Flower with CA primordia. Stamens with bidirectional initiation. Initiation of petals, stamens and carpels overlapping. Plastid inverted repeat absent (lost). Introns in plastid genes clpP and rps12 absent (lost).

Glycyrrhizeae Rydb., Fl. Rocky Mts.: 454. 31 Dec 1917

1/19. Glycyrrhiza (19; Europe, Asia, Australia, North America, temperate South America; incl. ‘Callerya’ atropurpurea).

[Wisterieae+[[Hedysareae+Galegeae]+Fabeae]]

Wisterieae X. Y. Zhu in Cathaya 6: 121. 1994

3/14. Wisteria (8; East and Southeast Asia to northeastern Australia and New Caledonia, eastern United States), Endosamara (1; E. racemosa; India, Sri Lanka, Southeast Asia, Malesia), Antheroporum (5; A. banaense, A. glaucum, A. harmandii, A. pierrei, A. vidalii; southwestern China, Indochina). – Eastern North America, East, South and Southeast Asia, New Guinea, Australia, New Caledonia. n = 8. rps12 intron present.

[[Hedysareae+Galegeae]+Fabeae]

Hedysareae DC., Prodr. 2: 307. mid Nov 1825

14/475–520. Chesneya clade Chesneya (c 35; Southwest and Central Asia to Mongolia and the Himalayas), Chesniella (5; C. ferganensis, C. gansuensis, C. gracilis, C. mongolica, C. villosa; Central Asia and Mongolia to northwestern China), Gueldenstaedtia (5–10; G. himalaica, G. henryi, G. monophylla, G. taihangensis, G. verna; Siberia and Mongolia to the Himalayas and western China), Tibetia (5; T. forrestii, T. himalaica, T. tongolensis, T. yadongensis, T. yunnanensis; Central Asia, the Himalayas, Tibet, western China). – Caragana clade Caragana (80–100; eastern Europe, Turkey and the Caucasus to Siberia, Mongolia, West and Central Asia to China). – Hedysarum clade Alhagi (6; A. canescens, A. graecorum, A. kirghisorum, A. maurorum, A. nepalensis, A. sparsifolia; the Mediterranean to Nepal), Sulla (6; S. aculeolata, S. capitata, S. carnosa, S. coronaria, S. pallida, S. spinosissima; the Mediterranean and North Africa to Siberia), Ebenus (c 20; the Mediterranean to Baluchistan), Taverniera (15; northeastern Africa, southwestern Asia), Hedysarum (c 160; temperate regions on the Northern Hemisphere), Onobrychis (130–150; Europe, the Mediterranean, Asia, Ethiopia), Greuteria (1; G. argyrea; Morocco, West Sahara), Eversmannia (4; E. botschantzevii, E. sarytavica, E. sogdiana, E. subspinosa; southeastern Russia, northern Iran, Central Asia), Corethrodendron (5; C. fruticosum, C. krasnowii, C. lignosum, C. multijugum, C. scoparium; Russia, Kazakhstan, Mongolia, China). – Temperate regions in Eurasia, North Africa to Ethiopia.

Galegeae Bronn in B. C. J. Dumortier, Fl. Belg.: 101. 1827

14/3.685–3.740. Carmichaelia clade Carmichaelia (c 25; New Zealand, Lord Howe), Streblorrhiza (1; S. speciosa; Philip Island near Norfolk Island, extinct), Swainsona (c 85; drier regions in Australia), Clianthus (2; C. maximus, C. puniceus; northeastern New Zealand). – Colutea clade Erophaca (1; E. baetica; the Mediterranean), Oxytropis (300–350; temperate regions on the Northern Hemisphere, especially Central Asia), Astragalus (>3.280; mainly temperate and subtropical regions on the Northern Hemisphere, tropical African mountains, northern India, the Andes to Chile, one species in southern Africa), Colutea (25–28; the Mediterranean, northeastern and eastern Africa, Asia to China and the Himalayas), Oreophysa (1; O. microphylla; Iran), Eremosparton (3; E. aphyllum, E. flaccidum, E. songoricum; southeastern Russia to Central Asia), Sphaerophysa (2; S. kotschyana: Central Anatolia; S. salsula: eastern Mediterranean to Central Asia), Smirnowia (1; S. turkestana; Turkestan), Lessertia (c 55; tropical eastern and southern Africa). – Galega clade Galega (6–8; G. albiflora, G. battiscombei, G. lindblomii, G. officinalis, G. orientalis; eastern Europe, East African mountains, temperate Asia). – Temperate and subtropical regions on the northern hemisphere, the Andes south to Chile, tropical African mountains, South Africa, Australia, Norfolk Island, Lord Howe, New Zealand.

Fabeae Reichenb., Fl. Germ. Excurs. 2(2): 525. 1832

9/755–825. Trifoliinae Bronn, Form. Plant. Legumin.: 127, 132. 1822 [‘Trifolieae’]. Trifolium (240–300; temperate and subtropical regions on both hemispheres except Australia), Ononis (c 30; Europe, the Canary Islands, the Mediterranean, Ethiopia, Iran), Medicago (85–90; Europe, the Mediterranean, Ethiopia, southern Africa, Asia), Trigonella (35–40; Macaronesia, the Mediterranean, southern Africa, Australia), Melilotus (19; Europe, the Mediterranean, North Africa, Ethiopia, temperate and subtropical Asia). – Parochetinae Yakovlev, Bobovye Zemnogo Shara: 117. 1991. Parochetus (1; P. communis; mountains of tropical Africa and Asia to Java). – Viciinae Bronn, Form. Plant. Legumin.: ad Sect. 134, 133. 1822 [‘Vicieae’]. Cicer (c 45; Greece, the Canary Islands, Morocco, Ethiopia, West and Central Asia), Lathyrus (c 160; temperate regions on the Northern Hemisphere, tropical East African mountains, temperate South America); Vicia (c 140; temperate regions on the Northern Hemisphere, tropical East Africa, South America, the Hawaiian Islands). – Subcosmopolitan. – Parochetus may be sister to the remaining Fabeae. A peculiar type of microhairs is a synapomorphy of Fabeae. Vicia has secondary pollen display by stigmatic pollen brush.

Phylogeny (somewhat simplified) of Fabaceae based on DNA sequence data (LPWG 2013). Dotted branches are weakly supported.

Phylogeny (simplified) of Fabaceae based on DNA sequence data (LPWG 2013). Dotted branches are weakly supported.

Hoffmannsegg et Link, Fl. Portug. 1: 62. 1 Sep 1809 [‘Polygalinae’], nom. cons.

Polygalopsida Endl., Gen. Plant.: 1076. Apr 1840 [’Polygalineae’]; Diclidantheraceae J. Agardh, Theoria Syst. Plant.: 195. Apr-Sep 1858 [‘Diclidanthereae’], nom. cons.; Moutabeaceae (Endl.) Endl. in H. Pfeiffer, Nomencl. Bot. 2(1): 364. 24 Jan 1873; Polygalineae Bessey in C. K. Adams, Johnson’s Universal Cyclop. 8: 461. 15 Nov 1895; Xanthophyllaceae (Baill. ex Engl.) Gagnep. ex Reveal et Hoogland in Bull. Mus. Natl. Hist. Nat., sér. 4, sect. B, Adansonia, 12: 206. 24 Nov 1990; Polygalanae Doweld, Tent. Syst. Plant. Vasc.: xxxviii. 23 Dec 2001

Fossils Polycolporopollenites is stephanocolpate pollen from the Palaeogene and Neogene of New Zealand and the Falkland Islands. Paleosecuridaca curtisii comprises fossilized fruits (composed of two-seeded carpels) of Polygalaceae from the Paleocene of North Dakota.

Habit Usually bisexual (in Balgoya functionally unisexual), perennial or annual herbs, evergreen or deciduous? shrubs, trees (rarely lianas; Salomonia and Epirixanthes comprise chlorophyllous hemimycotrophs versus achlorophyllous holomycotrophs). Some species are xerophytes. Often with paired crateriform glands (extrafloral nectaries) or spines at nodes (sometimes elsewhere).

Vegetative anatomy Arbuscular mycorrhiza present in at least some genera (e.g. Polygala, Polygaloides, Salomonia, and Epirixanthes). Phellogen ab initio superficial. Medulla with lignified and unlignified cells. Primary vascular tissue cylinder, without separate vascular bundles. Secondary lateral growth anomalous (from concentric cambia) present in lianas, or absent. Vessel elements (usually solitary) usually with simple perforation plates; lateral pits usually alternate, simple or bordered pits. Imperforate tracheary xylem elements usually tracheids (in Securidaca fibre tracheids) with bordered pits, non-septate (also vasicentric tracheids). Wood rays usually uniseriate (sometimes biseriate or multiseriate), heterocellular. Axial parenchyma usually paratracheal aliform, lozenge-aliform, winged-aliform confluent, vasicentric, reticulate, or banded (sometimes apotracheal diffuse or diffuse-in-aggregates). Tyloses present in some species. Intraxylary (concentric) phloem present in lianas. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Calciumoxalate as druses, crystal sand or prismatic or acicular crystals.

Leaves Usually alternate (rarely opposite or verticillate), simple, entire, often coriaceous, sometimes ericoid or scale-like or absent, with ? ptyxis. Stipules intrapetiolar (sometimes scale-like or modified into spines or paired glands) or absent; leaf sheath absent. Leaf base or abaxial side of lamina in Xanthophyllum with nectariferous glands (extrafloral nectaries; rarely on petiole). Petiole vascular bundle transection? Venation pinnate, usually brochidodromous. Stomata usually anomocytic or paracytic (in Xanthophyllum anisocytic or paracytic; in Balgoya cyclocytic). Cuticular wax crystalloids as rosettes of platelets (Fabales type). Lamina sometimes with lysigenous secretory cavities containing ethereal oils, often with glands or domatia. Leaf margin usually entire (rarely with nectariferous glandular teeth).

Inflorescence Terminal or axillary, usually spike or raceme (sometimes panicle; flowers rarely solitary axillary).

Flowers Usually zygomorphic (rarely actinomorphic). Pedicel often articulated. Hypogyny. Sepals five, with imbricate-quincuncial aestivation, persistent or caducous, free or partially or entirely connate (two lower sepals connate at base; sepals sometimes connate into tube); median sepal adaxial; two adaxial lateral sepals often enlarged into wing-like alae and petaloid, two abaxial lateral sepals small. Petals three or five (in Xanthophyllum rarely four), with imbricate or contorted aestivation, free or connate at base; vexillum consisting of two connate adaxial petals, carina consisting of abaxial petal (often fimbriate), two abaxial-lateral petals small. Alternatively: sepals almost uniform, carina consisting of abaxial petal. Alternatively: flowers more or less actinomorphic, with sepals and petals almost uniform, without distinct carina. Nectariferous disc intrastaminal, annular or unilateral (sometimes excentric), or absent. Some species of Polygala with pollination mechanisms (also secondary pollen display) similar to Faboideae.

Androecium Stamens (two to) five to eight (to ten); median stamen in each whorl usually absent. Filaments usually connate in lower part (often into tube) around pistil, usually more or less adnate at base to petals (epipetalous). Anthers usually basifixed (in Xanthophyllum dorsifixed), usually non-versatile, disporangiate or tetrasporangiate, usually dehiscing by apical or subapical pores or very short slits (rarely by longitudinal introrse? slits). Tapetum secretory. Female flowers with staminodia?

Pollen grains Microsporogenesis simultaneous. Pollen grains (5–)7–23(–33)-polycolporate (sometimes heteropolar and asymmetrical; rarely syncolporate; in Balgoya tricolporate), shed as monads, bicellular or tricellular at dispersal. Exine tectate, with columellate infratectum, perforate (to microreticulate), punctate, or finely rugulate, verrucate, granulate, fossulate, foveolate, or psilate.

Gynoecium Pistil composed of two to eight connate carpels; when two carpels then adaxial carpel reduced. Ovary superior, usually bilocular to quinquelocular (sometimes unilocular, pseudomonomerous, with one carpel reduced; in Xanthophyllum stipitate). Style single, long, erect or curved, simple or bifid (one branch with apical stigma, second branch with hair tuft). Stigma capitate or bilobate, often complex, asymmetrical, non-papillate, Dry type. Male flowers with pistillodium?

Ovules Placentation axile or parietal. Ovule usually one per carpel (in Xanthophyllum four to more than 40 per unilocular ovary composed of two carpels), anatropous or hemianatropous, pendulous, usually epitropous (in Xanthophyllum apotropous), bitegmic, crassinucellar. Micropyle usually bistomal, Z-shaped (zig-zag) (sometimes exostomal or endostomal). Outer integument usually two to five (in Xanthophyllum four to twelve) cell layers thick. Inner integument (one or) two to three (in Securidaca up to nine) cell layers thick. Hypostase enlarged. Parietal tissue approx. two cell layers thick. Nucellar cap present. Megagametophyte 8-nucleate (probably monosporous, Polygonum type; rarely tetrasporous). Synergids sometimes with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis asterad.

Fruit Usually a loculicidal capsule with membranous or fleshy pericarp (often flattened; rarely a berry, a drupe or a nut, often a samara; in, e.g., Polygala with persistent green calyx).

Seeds Hairs (sometimes containing juice, sometimes as coma near hilum), exostomal elaiosome or carunculus, or other arilloid structures often present (funicular aril present in Moutabeeae). Seed coat endotestal. Testa multiplicative. Exotesta subsclerotic. Endotestal cells isodiametric or palisade with narrowly elongate cells, with U-shaped thickenings, usually with crystals. Tegmen? Perisperm not developed. Endosperm copious or sparse, starchy (oily?), or absent. Embryo straight, at least in some species with chlorophyll. Cotyledons two, planoconvex. Germination phanerocotylar or cryptocotylar.

Cytology n = 5–14, 16, 17, 19, 20, 22, 23, 28–30 – Polyploidy and aneuploidy frequently occurring.

DNA Gene rpl22 present in nuclear genome (not in plastid genome; at least in Polygala). Plastid gene rps16 absent (lost) in Polygala (P. lindheimeri investigated) and Securidaca (S. diversifolia investigated).

Phytochemistry Flavonols (kaempferol, quercetin), indole and ergoline alkaloids and triterpene saponins present. Ellagic acid, tannins, proanthocyanidins, and cyanogenic compounds not found. Starch often replaced by saccharose or polygalite (1,5-anhydrosorbite). Aluminium accumulated in Xanthophyllum and Moutabeeae.

Use Ornamental plants (Polygala), medicinal plants (e.g. Polygala senega).

Distribution Central Chile to southern Brazil, Uruguay and northern Argentina.

Vegetative anatomy Phellogen ab initio subepidermal. Endodermis absent. Vessel elements (solitary) with simple or scalariform perforation plates; lateral pits?, simple pits. Vestured pits absent. Imperforate tracheary xylem elements? Wood rays ?-seriate, heterocellular. Axial parenchyma apotracheal. Secondary phloem diffusely expanded (intraxylary?). Sieve tube plastids S type? Nodes 1:3, unilacunar with three leaf traces. Mucilage cells present. Calciumoxalate styloids present.

Leaves Leaves alternate (spiral), simple, entire, coriaceous, with conduplicate ptyxis. Stipules small, petiolar, caducous; leaf sheath absent. Petiole vascular bundle transection arcuate; pericyclic fibres absent. Venation pinnate. Stomata anomocytic? Cuticular wax crystalloids as rosettes of platelets? Leaf margin usually serrate, with hydathodes? (sometimes entire).

Inflorescence Terminal or axillary, botryoid, cymose. Terminal flower bisexual, lateral flowers male.

Flowers Actinomorphic. Hypogyny. Sepals five, with valvate aestivation, persistent and accrescent in fruit, with nectary on lower half, free. Petals five, with contorted aestivation, clawed, free. Disc intrastaminal, wide, thick and fleshy, quinquelobate, with lobes adnate to stamens.

Androecium Stamens ten, diplostemonous; five antesepalous stamens inserted on abaxial margin of disc lobes a distance up sepals (above nectary), and five antepetalous stamens inserted near ovary base (below nectary), unidirectionally developing. Filaments subulate, free from each other and from tepals. Anthers dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, striate; exine protruding at apertures.

Gynoecium Pistil composed of five antesepalous carpels connate in lower part (laterally largely free). Ovary superior, quinquelocular. Stylodia five, free. Stigmas decurrent, type? Pistillodium absent.

Ovules Placentation axile. Ovules numerous per carpel (inserted in two marginal rows), pleurotropous, horizontal, bitegmic, crassinucellar? Micropyle bistomal? Outer integument approx. three cell layers thick. Inner integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A strongly asymmetrical follicular capsule with stellately radiating lobes and accrescent calyx, dehiscing downwards dorsally and ventrally by two coriaceous valves.

Seeds Aril absent. Seed with long apical wing. Seed coat exotestal. Exotesta three-layered; layers thickened, sclerotic. Endotesta? Tegmen degenerated. Perisperm not developed. Endosperm one cell layer thick. Embryo? Chlorophyll? Cotyledons two, conduplicate. Germination?

Phytochemistry Insufficiently known. Flavone-C-glycosides, prodelphinidins (in leaves), and triterpene saponins (high concentrations in bark) present.

Arnott in Wight et Arnott, Prodr. Fl. Ind. Orient. 1: 360. 22 Sep 1834 [’Surianeae’], nom. cons.

Stylobasiaceae J. Agardh, Theoria Syst. Plant.: 169. Apr-Sep 1858 [’Stylobasieae’]); Surianales Doweld, Tent. Syst. Plant. Vasc.: xxxviii. 23 Dec 2001

Fossils Wood fossil with the name of Suriana inordinata have been described from the Eocene of Wyoming.

Habit Usually bisexual (in Stylobasium polygamomonoecious), evergreen small trees or shrubs. Some species are xerophytes.

Vegetative anatomy Phellogen ab initio subepidermal or inner-cortical. Medullary vascular bundles present in Recchia. Vessels in radial multiples. Vessel elements with simple perforation plates; lateral pits alternate, simple or bordered pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres with simple pits, non-septate (in Stylobasium also vasicentric tracheids). Wood rays usually uniseriate (rarely biseriate), homocellular or heterocellular. Axial parenchyma usually absent (in Suriana apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty vasicentric, or usually with crystalliferous strands). Wood elements in Suriana and Stylobasium storied. Sieve tube plastids S type (Suriana) or Pfs type (in Stylobasium). Nodes usually 3:3, trilacunar with three leaf traces (in Suriana 1:1, unilacunar with one trace). Secretory cavities present or absent. Sclereids present. Heartwood often with red or brown resin. Parenchyma cells often with acicular calciumoxalate crystals, styloids, crystal sand, and other types of crystals.

Trichomes Hairs unicellular or multicellular, simple or absent; glandular hairs, with multicellular head and stalk, often present.

Leaves Alternate (spiral or distichous), usually simple, entire (in some species of Recchia pinnately compound, with alternate leaflets, with articulated petiolules), usually coriaceous, with ? ptyxis. Stipules usually replaced by pseudostipules and/or metastipules (absent in Suriana); leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Petiole or mid-vein in Cadellia with extrafloral nectaries. Venation pinnate-reticulate. Stomata anomocytic or anisocytic (in Stylobasium paracytic?). Cuticular wax crystalloids usually as rosettes of platelets? (sometimes as scales). Epidermis without mucilage cells. Mesophyll and epidermis usually with druses or single crystals of calciumoxalate. Leaf margin entire.

Inflorescence Terminal or axillary, cymose, raceme-like or panicle, or flowers solitary axillary.

Flowers Actinomorphic. Pedicel articulated. Hypogyny. Sepals five (to seven), with imbricate-quincuncial aestivation, usually persistent, free or connate at base. Petals usually five (absent in Stylobasium), with contorted aestivation, caducous, in Recchia and Suriana shortly clawed, free (rarely absent). Nectary usually absent. Disc usually absent (present in Recchia).

Androecium Stamens 5+5 or five (and up to five outer staminodia; in Suriana antepetalous inner stamens shorter or staminodial), obdiplostemonous, as many as sepals, antesepalous, persistent or caducous, sometimes articulated, antesepalous stamens usually longer than antepetalous stamens. Filaments filiform or subulate, free from each other and from tepals. Anthers basifixed or dorsifixed, sometimes versatile, tetrasporangiate, usually introrse (rarely latrorse; in Stylobasium extrorse?), longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Staminodia in bisexual and male flowers four or five, intrastaminal or extrastaminal, or absent; female flowers in Stylobasium with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpor(oid)ate, shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate, microreticulate, striate or finely rugulate, sometimes gemmate to verrucate; exine sometimes protruding at apertures.

Gynoecium Carpels one to five (in Stylobasium, Guilfoylia and Recchia usually one carpel [monomery], in Cadellia and Suriana five antepetalous carpels), secondarily free (apocarpy), antepetalous (in Recchia on short nectariferous gynophore). Ovaries superior. Stylodia one to five, gynobasic (ventrobasic). Stigmas peltate to capitate or clavate, papillate, type? Male flowers in Stylobasium with pistillodium.

Ovules Placentation (supra)basal (basal-marginal or subbasal?). Ovules (one or) two (to five) per carpel, collateral, anatropous to campylotropous, ascending, unitegmic, crassinucellar, pachychalazal. Integument three to seven cell layers thick. Obturator absent. Hypostase present. Parietal tissue four or five cell layers thick. Nucellar cap sometimes present. Archespore often multicellular. Megagametophyte monosporous, Polygonum type. Antipodal cells ephemeral, degenerating. Endosperm development ab initio nuclear. Endosperm haustorium chalazal. Embryogenesis onagrad.

Fruit A nut, a drupe or a berry, in Cadellia and Suriana an assemblage of nutlets or drupelets, usually with persistent and accrescent calyx. Exocarp in Cadellia and Recchia with thickened cell walls. Mesocarp usually hard (in Stylobasium fleshy), parenchymatic, with innermost cell layer crystalliferous. Endocarp usually hard, usually three-layered, with outer layer lignified, palisade-sclerenchymatous.

Seeds Aril absent. Seed coat exotestal. Exotesta in Suriana green, with cuboid, enlarged, tanniniferous cells. Mesotestal and endotestal cells crushed. Perisperm not developed. Endosperm usually absent (in Stylobasium sparse). Embryo curved (often hippocrepomorphic) or plicate (conduplicate, transversely induplicate or globular), with chlorophyll. Cotyledons two, usually thick, incumbent, oily or starchy. Germination in Stylobasium phanerocotylar.

Phytochemistry Insufficiently known. Surianol (triterpene diol) present in Suriana. Proanthocyanidins sometimes present. Ellagic acids, saponins and cyanogenic compounds not found.

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”The Nitrogen Fixing clade”

POLYGALALES Juss. ex Bercht. et J. Presl

FABACEAE Lindl.

POLYGALACEAE Hoffmanns. et Link

QUILLAJACEAE D. Don

SURIANACEAE Arn.