”The Nitrogen Fixing clade”


[Polygalales+[Rosales+[Cucurbitales+Juglandales]]]


POLYGALALES Juss. ex Bercht. et J. Presl

Berchtold et Presl, Přir. Rostlin: 228. Jan-Apr 1820 [‘Polygaleae’]


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.

Cytology x = 5–7 (11)

DNA Plastid gene infA lost/defunct. Mitochondrial intron coxII.i3 lost.

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.

Systematics Polygalales are sister-group to the clade [Rosales+[Cucurbitales+Juglandales]] (Soltis & al. 2011), or the clade [Cucurbitales+Juglandales] (Wang & al. 2009) or to Rosales (Magallón & Castillo 2009).

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.

Bello & al. (2007, 2009) found the following topologies based on matK, rbcL or combined matK and rbcL, respectively: [Polygalaceae+[Fabaceae+[Quillajaceae+Surianaceae]]], [Fabaceae+[Quillajaceae+[Polygalaceae+Surianaceae]]] and [Quillajaceae+[Surianaceae+ [Fabaceae+Polygalaceae]]]. Persson (2001) received the topology [Polygalaceae+[Surianaceae+ [Fabaceae+Quillajaceae]]]. Qiu & al. (2010) found the following topology: [Quillajaceae+ [Fabaceae+[Polygalaceae+Surianaceae]]]. Soltis & al. (2011) received [[Quillajaceae+Polygalaceae]+[Fabaceae+Surianaceae]]. Finally, Moore & al. (2011) found [Surianaceae+ [Quillajaceae+[Fabaceae+Polygalaceae]]].


FABACEAE Lindl.

( Back to Polygalales )

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

Genera/species c 680–710/20.000–20.200

Distribution Cosmopolitan except Antarctica.

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 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 Mimosoideae and Faboideae 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 Faboideae 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 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 Mimosoideae 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).

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 Faboideae 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 (Mimosoideae), imbricate, imbricate-ascending or cochlear-descending (adaxial-median petal internal) aestivation, often clawed, persistent or caducous, free (especially in ’Caesalpinioideae’) or two front petals connate at base into carina (especially in Faboideae, sometimes four petals connate) or all petals connate (especially in Mimosoideae); median petal adaxial (vexillum in Faboideae; lateral petals alae in Faboideae; median petal sometimes absent, especially in Cercidoideae, ’Caesalpinioideae’ and Mimosoideae). 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 Mimosoideae 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 (Mimosoideae, Faboideae) 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 Mimosoideae), usually bicellular (in some Mimosoideae 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 Mimosoideae 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 Mimosoideae 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 Faboideae), 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 (Mimosoideae, ‘Caesalpinioideae’), oblique (‘Caesalpinioideae’) or curved and lateral (Faboideae). Germination phanerocotylar or cryptocotylar.

Cytology x = 7, 8, 12, 14

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 Faboideae with inversion of c. 50 kb. Plastid genome in most Phaseolinae 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 Faboideae. Plastid gene ycf4 lost in most Faboideae. 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, munduserone, angolensin, etc. in Faboideae), 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 (rotenone from Lonchocarpus), 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). Duparquetia may be sister to all other Fabaceae, the clade [Cercis+Bauhinia] or Cercideae possibly being successive sister to the remainder. The traditional subdivision of Fabaceae into the three subfamilies Caesalpinioideae, Mimosoideae and Faboideae (Papilionoideae) has collapsed. Caesalpinioideae form a paraphyletic group with Mimosoideae as an ingroup within it. Faboideae (Papilionoideae) are sister-group to a somewhat reduced although obviously monophyletic Caesalpinioideae.

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 Africa). – Liana. Vestured pits absent. Leaves pinnately compound. Floral development acropetal. Sepals four, petaloid. Median petal adaxial. Stamens four, antesepalous. Filaments more or less connate. Anthers poricidal (dehiscing by pores). Pollen grains asymmetrical. Apertures with ectoaperture encircling equator and with two endoapertures. Gynoecial initiation not advanced (relative to other organs). Fruit a legume.

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

13/310–335. Cercis (10; the Mediterranean, eastern Asia, North America to northeastern Mexico); Adenolobus (2; southern Angola, Namibia, northern Botswana, Northern Cape); Bauhinia (c 100–110; tropical and subtropical regions of both hemispheres, especially tropical South America), Piliostigma (3; Africa to South Africa and eastwards to southern Asia and Australia), Brenierea (1; B. insignis; Madagascar); Griffonia (4; tropical West Africa); Gigasiphon (4–5; tropical Africa), Tylosema (4; Africa southwards to South Africa), Barklya (1; B. syringifolia; northeastern Queensland), Schnella (c 40; tropical South America), Lysiphyllum (8; Malesia and eastwards to tropical Australia), Phanera (c 120–130; pantropical), Lasiobema (15–20; East Asia from eastern Himalaya to Japan, Indochina, Java). – Mainly tropical and subtropical regions, few species in warm temperate areas. Trees, shrubs or lianas with branch tendrils. Vestured pits absent. Petiole with a single joined pulvinus. Leaves simple alternatively unifoliolate, usually bilobate or entire (rarely with two leaflets). Stomata usually anomocytic (rarely paracytic). Sepals five, connate. Median petal often adaxial. Exine striate. Zwischenkörper (pectic substances) present below apertures at least in Cercis. Fruit a legume, often explosively dehiscent. x = 7. Introns in mitochondrial genes srp12 and rpl2 absent in Bauhinia. – Cercis is sister to the remaining Cercideae and Adenolobus successive sister to the two major clades.

Fabaceae except Duparquetia and Cercideae

[Detarioideae+[Poeppigieae+[Caesalpinioideae+Faboideae]]]

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 Kostel. In Allg. Med.-Pharm. Fl. 4: 1315. Jan-Oct 1835

Ectomycorrhiza often present. Vestured pits absent. Resinous ducts often present in stem. Stipules intrapetiolar, often caducous. Leaf phloem transfer cells present. Leaflets often with crater-like glands on abaxial surface. Floral prophylls (bracteoles) often well developed, caducous. Size or number of sepals and/or petals often reduced. Stamen number often increased. Exine sometimes striate. Style abaxially curved. Endosperm absent. Cotyledons with thick-walled cells, with amyloid (xyloglucans). x = 12. Bicyclic diterpenes 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.

Barnebydendron (1; B. riedelii; tropical Central and South America). – Zwischenkörper (pectic substances) present below apertures.

Goniorrhachis (1; G. marginata; southeastern Brazil).

Schotia (4–5; Africa south of the Zambesi River). – Zwischenkörper (pectic substances) present below apertures

[Prioria clade+[Brandzeia+Daniellia]+Detarieae]

Prioria clade

3/3. Colophospermum (1; C. mopane; tropical southern Africa), Hardwickia (1; H. binata; drier regions of India), Prioria (1; P. copaifera; Central and northwestern South America, Jamaica). – Tropical.

Daniellia clade

2/10. Brandzeia (1; B. filicifolia; Madagascar), Daniellia (9; 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

15–17/135–140. Peltogyne (23; tropical America), Guibourtia (14–15; tropical Africa, tropical America), Stemonocoleus (1; S. micranthus; Central Africa), Augouardia (1; A. letestui; Gabon), Hymenaea (14; eastern Africa, tropical America); Neoapaloxylon (3; Madagascar), Baikiaea (5–6; tropical to southern Africa in Namibia and Botswana), Bathiaea (1; B. rubriflora; Madagascar), Copaifera (c 35; tropical Africa, Malesia, tropical America), Detarium (3; tropical and subtropical West Africa and eastwards to Sudan), Gilletiodendron (5; tropical Africa), Hylodendron (1; H. gabunense; along the Gulf of Guinea), Sindora (18–20; Southeast Asia, Malesia), Sindoropsis (1; S. letestui; Gabon), Tessmannia (11–12; tropical Africa), Micklethwaitia (1; M. carvalhoi; Mozambique)?, Normandiodendron (2; western Central Africa)? – 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)

55?/570–575? Endertia (1; E. spectabilis; Borneo), Lysidice (2; southern China, Vietnam), Saraca (11; India, Sri Lanka, southern China, Southeast Asia, Malesia eastwards to Sulawesi); Afzelia (11; tropical regions in the Old World), Brodriguesia (1; B. santosii; eastern Brazil), Intsia (3; tropical Asia eastwards to Pacific coasts); Brownea (12; Costa Rica and southwards to Peru, the West Indies), Browneopsis (6; Panamá to Peru), Ecuadendron (1; E. acosta-solisianum; western Ecuador), Elizabetha (11; tropical South America), Heterostemon (7; tropical South America), Paloue (4; the Guiana Shield in northern South America), Paloveopsis (1; P. emarginata; northeastern South America); Berlinia (17; tropical Africa); Annea (2; tropical West and Central Africa), Brachystegia (c 25; central tropical Africa and southwards to Botswana), Cryptosepalum (11; tropical Africa), Cynometra (c 85; pantropical), Eperua (c 15; northeastern South America), Eurypetalum (3; Cameroun to Gabon), Gabonius (1; G. ngouniensis; Gabon), Gilbertiodendron (c 30; tropical and subtropical Africa), Hymenostegia (16; along the Gulf of Guinea), Lebruniodendron (1; L. leptanthum; along the Gulf of Guinea), Leonardoxa (1–4; L. africana; Central Africa), Leucostegane (2; Malay Peninsula, Borneo), Loesenera (4; tropical West Africa), Macrolobium (c 70; tropical Central and South America), Maniltoa (20–25; India, Sri Lanka, Southeast Asia, Malesia, especially New Guinea), Neochevalierodendron (1; N. stephanii; Gabon), Oddoniodendron (2–3; along the Gulf of Guine), Paramacrolobium (1; P. coeruleum; tropical Africa), Plagiosiphon (5; Central Africa), Scorodophloeus (2; coasts of Guinea and East Africa), Talbotiella (3; tropical West Africa), Tamarindus (1; T. indica; Africa, Asia), Zenkerella (5; coast of Guinea, tropical East Africa); Aphanocalyx (14; tropical West and Central Africa), Bikinia (c 10; western Central Africa), Icuria (1; I. dunensis; Central Mozambique), Julbernardia (11; tropical to southern Africa), Michelsonia (1; M. microphylla; eastern Congo), Tetraberlinia (7; Central Africa); Amherstia (1; A. nobilis; Burma, extinct?), Anthonotha (28; tropical Africa), Brachycylix (1; B. vageleri; northern Central Colombia), Crudia (50–55; tropical Africa, Asia and America), Dicymbe (16; tropical South America), Didelotia (10–11; Central Africa), Englerodendron (1; E. usambarensis; Usambaras in Tanzania), Humboldtia (6; southern India, Sri Lanka), Isoberlinia (5; tropical Africa), Librevillea (1; L. klainei; Gabon), Microberlinia (2; along the Gulf of Guinea), Polystemonanthus (1; P. dinklagei; tropical West Africa), Pseudomacrolobium (1; P. mengei; Congo). – Pantropical. This is a highly provisional constellation of genera, since many of them have not yet been investigated by molecular methods.

[Poeppigieae+[Caesalpinioideae+Faboideae]]

Poeppigieae Britton et Rose in N. Amer. Fl. 23: 201, 218. 18 Nov 1930

4–5/60–65. Poeppigia (1; P. procera; tropical America); Dialium (c 40; tropical to southern Africa, Madagascar, South and Southeast Asia, one species in tropical America); Petalostylis (2–3; drier regions in Australia), Labichea (14; Australia except central parts), Baudouinia (6; Madagascar)? – Tropical. Vestured pits? Inflorescence often cymose, primarily simple or compound cymes or dichasia. Fruit often drupe or samara. – Many representatives have lost one or several floral organs during evolution.

[Caesalpinioideae+Faboideae]

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’] s.str.

Umtiza clade

7/27. Ceratonia (2; southeastern Mediterranean to Somalia and the Arabian Peninsula), Acrocarpus (1; A. fraxinifolius; India, Sri Lanka, Southest Asia, Malesia), Tetrapterocarpon (2; Madagascar); Arcoa (1; A. gonavensis; Hispaniola); Gymnocladus (6; East and Southeast Asia, eastern North America), Umtiza (1; U. listeriana; Eastern Cape), Gleditsia (14; Caspian, China, Japan and southwards 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. – The Umtiza clade may be sister-group to the remainder of Caesalpinioideae.

[[Pterogyne+Caesalpinieae+Cassieae]+[[Dimorphandra clade+Peltophorum clade+ Tachigali clade]+[[Diptychandra+Moldenhawera]+Pachyelasma+Erythrophleum+ Mimosoideae]]]

[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’]

21?/205–210. Cordeauxia (1; C. edulis; northeastern Africa), Stuhlmannia  (1; S. moavi; coast of Tanzania); Erythrostemon  (12–13; warmer regions in America), Poincianella(c 35; tropical America), Pomaria (15–16; tropical Africa and America); Libidibia (6–8; tropical America), Stahlia (1; S. monosperma; Hispaniola, Puerto Rico); Balsamocarpon (1; B. brevifolium; Chile), Zuccagnia (1; Z. punctata; the Andes in Chile and Argentina), Hoffmannseggia (28; tropical to southern Africa, southwestern United States to Chile), Stenodrepanum (1; S. bergii; Argentina); Guilandina (7; warmer regions on both hemispheres), Pterolobium (11; tropical and subtropical regions in the Old World), Mezoneuron (c 25; Africa, Asia and eastwards to the Hawaiian Islands), Moullava (1; M. spicata; western India), Tara (3; tropical America), Coulteria (9–10; Mexico to Colombia), Caesalpinia (c 40; warmer regions on both hemispheres), Lophocarpinia (1; L. aculeatifolia; Paraguay, northern Argentina), Haematoxylum (3; tropical Africa and southwards to Namibia, tropical 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.– Cordeauxiaand Stuhlmanniaform a clade sister to the remaining Caesalpinieae.

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

c 19?/c 700. Cassia (c 30; tropical and subtropical regions on both hemispheres; incl. Latrobea?), Latrobea (6; southwesternmost Western Australia; in Cassia?), Senna (c 300; warm-temperate to tropical regions, especially America); Batesia (1; B. floribunda; Amazonia), Chamaecrista (c 330; Africa, East Asia, temperate and tropical regions in America), Melanoxylum (1; M. brauna; Amazonia), Recordoxylon (3; Amazonia); Apuleia (1–3; A. leiocarpa; northeastern Peru, southeastern Brazil, northern Argentina)?, Dicorynia (1–2; D. guianensis; tropical South America)?, Distemonanthus (1; D. benthamianus; tropical West Africa)?, Eligmocarpus (1; E. cynometroides; southeastern Madagascar)?, Koompassia (3; Malesia)?, Martiodendron (4; tropical South America)?, Mendoravia (1; M. dumaziana; southeastern Madagascar)?, Storckiella (4; northeastern Queensland, New Caledonia, 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)?, Vouacapoua (3)? – 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. – This is a highly provisional constellation of genera, since many of them have not yet been investigated by molecular methods.

[[Dimorphandra clade+Peltophorum clade+Tachigali clade]+[[Diptychandra+Moldenhawera]+Pachyelasma+Erythrophleum+Mimosoideae]]

[Dimorphandra clade+Peltophorum clade+Tachigali clade]

Dimorphandra clade

5–6/c 55. Burkea (1; B. africana; tropical and subtropical Africa southwards to Namibia and northern South Africa), Campsiandra (c 20; tropical South America, especially Amazonia)?, Dimorphandra (c 25; tropical America), Dinizia (1; D. excelsa; Guayana, Brazil), Mora (6; tropical America), Stachyothyrsus (2; tropical West Africa). – Tropical Africa and America. The Dimorphandra clade is weakly supported.

Peltophorum clade

6–8/40–45. Bussea (7; tropical Africa, Madagascar), Peltophorum (c 5; tropical and subtropical regions); Schizolobium (1–2; S. parahyba; tropical America), Delonix (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 (1; C. multiflora; Mexico), Cenostigma (3; Brazil, Paraguay), Parkinsonia (c 15; drier warmer regions in America, northeastern Africa, Namibia and Northern Cape). – Tropical and subtropical. Leaves bipinnate. Petals usually yellow. Seeds narrow. – Schizolobium is sister to the Delonix-Parkinsonia clade.

Tachigali clade

3/c 70. Arapatiella (2; southeastern Brazil), Jacqueshuberia (7; Colombia, Brazil), Tachigali (c 60; tropical America, especially Amazonia). – Tropical America.

[[Diptychandra+Moldenhawera]+Pachyelasma+Erythrophleum+Mimosoideae]

Diptychandra (1; D. aurantiaca; Brazil, Bolivia, Paraguay), Moldenhawera (9; 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 and eastwards to northern Australia). – Tropical regions in the Old World. Erythrophleum may be sister to Mimosoideae, although the support is weak.

Mimosoideae DC., Prodr. 2: 424. Nov (med.) 1825 [‘Mimoseae’] (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)

78/3.270–3.320. Adenanthera (13; tropical Asia and Australia, Pacific islands), Tetrapleura (2; tropical Africa), Amblygonocarpus (1; A. andongensis; tropical Africa southwards to Namibia and Botswana), Calpocalyx (11; tropical West Africa), Pseudoprosopis (7; tropical Africa), Xylia (9; tropical to southern Africa to tropical Asia); Pentaclethra (3; tropical Africa, tropical America); Newtonia (14–15; tropical Africa, tropical America), Fillaeopsis (1; F. discophora; western and southwestern Africa); Plathymenia (1; P. reticulata; Brazil, Argentina); ‘Entada’ (c 30; tropical regions; non-monophyletic; incl. Elephantorrhiza?), Elephantorrhiza (9; tropical to southern Africa; in Entada?), Piptadeniastrum (1; P. africanum; Central Africa); Cylicodiscus (1; C. gabunensis; Central Africa); Calliandropsis (1; C. nervosus; Mexico), Dichrostachys (14; Africa, Madagascar, southern and southeastern Asia to Australia), Gagnebina (10; Madagascar, the Comoro Islands, the Mascarene Islands); Desmanthus (c 25; tropical and subtropical regions in South America), Leucaena (22; southern United States to Peru), Schleinitzia (3–4; Malesia, Vanuatu and eastwards to Tahiti, Guam); Prosopidastrum (5; Mexico, southern South America); Piptadeniopsis (1; P. lomentifera; Paraguay); Mimozyganthus (1; M. carinatus; southeastern Bolivia, southwestern Paraguay, Argentina); Neptunia (12; tropical and subtropical regions), ‘Prosopis’ (c 45; Africa, Southwest Asia, southern United States to South America; non-monophyletic), Xerocladia (1; X. viridiramis; Namibia, Northern Cape); Vachellia (c 160; tropical and subtropical regions on both hemispheres), Parkia (c 35; tropical Africa and Asia, Madgascar, tropical Australia, tropical and subtropical America), ’Piptadenia’ (c 25; tropical America; non-monophyletic), Adenopodia (7; tropical Africa, Central America), Anadenanthera (2; tropical America), Microlobius (1; M. foetidus; Mexico to Honduras, southern Brazil, Paraguay, northern Argentina), Parapiptadenia (3; tropical South America), Pityrocarpa (3; tropical America), Stryphnodendron (c 30; tropical America), Pseudopiptadenia (11; tropical America), Mimosa (c 510; tropical and subtropical regions, with their largest diversity in Central and South America), ‘Senegalia’ (c 200; tropical and subtropical regions in Africa, Asia, and Central and South America; non-monophyletic), Mariosousa (13; southwestern United States, Mexico, Central America); Abarema (c 45; tropical America), Acacia (1.050–1.100; Australia, Pacific islands), Acaciella (15; southern and central United States to Argentina), ‘Albizia’ (c 130; pantropical; paraphyletic), ‘Archidendron’ (c 95; tropical Asia, especially Borneo; non-monophyletic), Archidendropsis (14; Southwest Pacific islands), Balizia (2; New Caledonia), Blanchetiodendron (1; B. blanchetii; eastern Brazil)?, Calliandra (c 200?; pantropical; incl. Guinetia?), Guinetia (1; G. tehuantepecensis; Mexico; in Calliandra?), Zapoteca (c 20; southwestern North America to northern Argentina), Cedrelinga (1; C. cateniformis; Brazil), Cojoba (12; tropical America), Ebenopsis (3; southern Texas, Mexico), Enterolobium (11; tropical America), Faidherbia (1; F. albida; eastern Mediterranean, tropical to southern Africa), Falcataria (3; East Malesia to Queensland), Havardia (5; Texas to Central America), Hesperalbizia (1; H. occidentalis; southwestern Mexico), Hydrochorea (3; South America), Inga (c 300; tropical and subtropical regions in South America), ‘Leucochloron’ (4; Brazil; non-monophyletic), Lysiloma (8; Florida, Mexico, the West Indies), Macrosamanea (11; northern South America)?, Pararchidendron (1; P. pruinosum; Java, northern Australia), Pithecellobium (18; tropical and subtropical regions in America), Serianthes (18; Thailand, Southeast Asia, Malesia, New Guinea, Melanesia, especially New Caledonia, Micronesia, western Polynesia)?, Sphinga (3; tropical America), Thailentadopsis (3; Sri Lanka, Thailand, southern Vietnam)?, Viguieranthus (23; Madagascar, tropical Asia)?, Wallaceodendron (1; W. celebicum; the Philippines, Sulawesi), Zygia (c 45; tropical America), Paraserianthes (1; P. lophantha; Malesia). – Unplaced Mimosoideae Aubrevillea (2; western central and western Africa), Dinizia (1; D. excelsa; Guyana, Brazil), Indopiptadenia (1; I. oudhensis; India, Nepal), Lemurodendron (1; L. capuronii; northeastern Madagascar). – 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 Parkieae and Mimozygantheae 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 Mimosoideae are largely unresolved.

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

Faboideae (Giseke) Rudd in Rhodora 70: 496. 31 Dec 1968 (Papilionoideae L. ex DC., Prodr. 2: 94. Mid Nov 1825)

440–467/14.710–14.780. Cosmopolitan except Antarctica. Bisexual evergreen or deciduous trees, shrubs, lianas, suffrutices, or perennial, biennial or annual herbs, often twining with foliar tendrils. Root nodules with nitrogen fixing bacteria frequent. Nodules usually arising in association with lateral roots. Ectomycorrhiza sometimes present. Phellogen superficially or deeply seated. Sieve tube plastids PIVb type or S type, with spindle-shaped non-dispersive protein bodies (forisomes). Leaves pinnately or palmately compound (rarely unifoliolate or simple). Stipules intrapetiolar, often scale-like or modified into spines or glands. Stomata usually anomocytic or paracytic (sometimes anisocytic, tetracytic or cyclocytic). Often with secretory chambers containing mucilage, resins or oils. Inflorescence terminal or axillary, panicle, fascicle, raceme, spike or head. Flowers usually zygomorphic (papilionoid; rarely actinomorphic). Hypanthium? often present (in some species of Amorpheae stemonozone instead of hypanthium). Sepals usually five, with imbricate aestivation, in lower parts usually connate into tube. Petals five (rarely one, three or four; absent in some Amorpheae, in other Amorpheae more or less uniform), with imbricate-ascending (-descending?) 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 (five to) ten (or 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, 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 Medicagineae. 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. Pleurogram and linea fissura usually absent. Endosperm often absent; cell walls sometimes thick. Embryo usually curved, 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 Faboideae (absent in Swartzieae and in Sophoreae). Isoflavonoids (pterocarpans, isoflavanes), prenylated flavonoids, pyrrolizidine, indolizidine and quinolizidine alkaloids, canavanine and other free (non-protein) amino acids present. – Forisomes providing a regulatory mechanism of phloem transport, are a synapomorphy of Faboideae, which has been lost several times (i.a. in the astragalean clade).

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

8/c 215. Swartzia (c 180; tropical America), Bobgunnia (2; tropical and southern Africa), Bocoa (3; tropical South America), Candolleodendron (1; C. brachystachyum; northeastern South America); Fairchildia (1; F. panamensis; Central and northwestern South America), Trischidium (5; South America), Ateleia (c 20; tropical America), Cyathostegia (1; C. mathewsii; 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 Faboideae. – Some molecular analyses (using the trnL intron) suggested that Swartzioideae are sister to the remaining Faboideae (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/c 80. Angylocalyceae clade Angylocalyx (7; tropical Africa), Xanthocercis (3; Gabon, southern Africa, Madagascar), Castanospermum (9; tropical America, one species, C. australe, in western New Britain, eastern Queensland and New South Wales, New Caledonia and Vanuatu). – Dipterygeae clade Monopteryx (3–4; northern South America), Taralea (7; tropical America), Dipteryx (1; D. odorata; tropical America), Pterodon (3; Brazil, Bolivia). – Amburaneae clade Dussia (c 12; tropical America), Petaladenium (1; P. urceoliferum; Rio Negro in Brazil), Myrospermum (2–3; tropical America), Myrocarpus (5; tropical South America), Myroxylon (2; tropical America), Dupuya (2; Madagascar), Aldina (15; tropical South America), Amburana (3; tropical South America), Cordyla (8; tropical Africa to South Africa, Madagascar). – Tropical America and Africa, Madagascar, Papuasia, Melanesia. Adaxial two sepals in Dipteryx, Pterodon and Taralea enlarged petaloid; abaxial three sepals forming tridentate lobe. – Some molecular analyses (using matK with low bootstrap support) suggest that the clade [Swartzioideae+Dipterygeae] is sister-group to the remaining Faboideae.

[cladrastidoids+50 kb inversion clade]

The bootstrap support of this clade is weak.

Cladrastidoids

1–3/16. ’Cladrastis’ (6; East Asia, North America; paraphyletic; incl. Pickeringia and Styphnolobium?), Pickeringia (1; P. montana; southwestern North America; in Cladrastis?), Styphnolobium (9; Central America; in Cladrastis?). – East Asia, America. – The cladrastidoids may be sister-group to the remaining Faboideae, but the bootstrap support is weak.

50 kb inversion clade

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.

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

6/21. Holocalyx (1; H. balansae; tropical South America), Lecointea (4; tropical America), Zollernia (10; tropical America); Harleyodendron (1; H. unifoliolatum; northeastern Brazil), Exostyles (4; southeastern Brazil); Uribea (1; U.tamarindoides; Central America). – Tropical America.

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

Pyrrolizidine or quinolizidine alkaloids present. x = 9.

Dermatophyllum clade

1/3–4. Dermatophyllum (3–4; southwestern North America). – Trees or shrubs.

Ormosieae Yakovlev in Bobovye Zemnogo Shara: 72. 1991

9/c 150. Ormosia (c 130; East Asia to Queensland, Central and eastern South America), Haplormosia (1; H. monophylla; western central and western Africa), Pericopsis (4; tropical Africa, Sri Lanka to Micronesia), Dicraeopetalum (3; southeastern Ethiopia, southern Somalia, northeastern Kenya, Madagascar), Platycelyphium (1; P. voense; drier regions in northeastern and eastern Africa), Bolusanthus (1; B. speciosus; Northern Province, Mpumalanga, Swaziland), Neoharmsia (2; western and northwestern Madagascar), Sakoanala (2; northwesterna and eastern Madagascar), Clathrotropis (6; tropical South America). – Pantropical, few species in temperate East Asia.

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

62–70/2.265–2.280. – Brongniartieae and Podalyrieae form well supported clades, whereas the Sophoreae/Crotalarieae/Thermopsideae/Genisteae clade is still largely unresolved. 50 kb inversion absent in Sophoreae.

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

13/c 160. Poecilanthe (7; tropical South America), Harpalyce (c 25; Mexico, Cuba, Brazil), Amphiodon (1; A. effusus; tropical South America), Tabaroa (1; T. caatingicola; southwestern Bahia in Brazil), Brongniartia (c 65; tropical and subtropical America), Hovea (c 40; Australia), Templetonia (11; Australia), Cyclolobium (1; C. brasiliense; tropical South America), Limadendron (2; Colombia, southern Venezuela, Guayanas, Amazonian Brazil), Plagiocarpus (1; P. axillaris; northern Australia), Cristonia (1; C. biloba; western Western Australia), Lamprolobium (2; northeastern Queensland), Thinicola (1; T. incana; northern Western Australia). – Tropical and subtropical America, Australia.

Bowdichia clade

5/27. Diplotropis (12; Amazonia), Guianodendron (1; G. praeclarum; Guyana, Brazil), Staminodianthus (3; Amazonas), Bowdichia (2; tropical South America), Leptolobium (10; Mexico to northern South America). – Tropical America. – The Bowdichia clade may be sister-group to the remaining Genistoideae, although the bootstrap support is low.

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

[Sophoreae+Euchresteae+Thermopsideae]

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

6–10/65–75. Camoensia (2; along the Gulf of Guinea and southwards to Angola), Ammodendron (4–5; West and Central Asia to northwestern China), Ammothamnus (2; Central Asia), Maackia (8; East Asia), Salweenia (1; S. wardii; southeastern Tibet), Sophora (c 50; warmer regions on both hemispheres). – Unplaced ‘Sophoreae’ s.lat. Amphimas (3–4; western Central Africa), Panurea (2; Colombia, northern Brazil), Spirotropis (2–3; northeastern South America), Uleanthus (1; U. erythrinoides; Amazonian Brazil). – Tropical Africa, Asia, South America. – Panurea and Bowdichia (the latter genus in the Bowdichia clade) have similar pollen morphology.

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

5/45–47. Anagyris (1–2; the Canary Islands, the Mediterranean, the Middle East to Iran), Piptanthus (2; Himalayas), Ammopiptanthus (1–2; Central Asia), Baptisia (17; eastern United States), Thermopsis (23; East Asia, North America). – Macaronesia and the Mediterranean to Central and East Asia, North America (‘Tethys distribution’).

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

1/4. Euchresta (4; India, Bhutan, China, the Korean Peninsula, Japan, Southeast Asia, Java).

[Podalyrieae+[Crotalarieae+Genisteae]]

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

8–10/c 145. Cyclopia (23; Western and Eastern Cape), ’Amphithalea’ (c 20; 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; Western Cape), Podalyria (c 25; Western Cape to KwaZulu-Natal), Liparia (c 20; Western and Eastern Cape), Virgilia (2; Western Cape, western Eastern Cape), ’Calpurnia’ (16?; Africa southwards to South Africa; paraphyletic), Cadia (7; Madagascar, one species in northeastern tropical Africa to southern Arabian Peninsula). – South Africa, few species to the northeast. Methylated anthocyanins, carboxylic acid esters of quinolizidine alkaloids, 3’-hydroxydaidzein, and canavanine (in seeds) present. Alkaloids not found.

[Crotalarieae+Genisteae]

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

19/1.320–1.325. ’Wiborgiella’ (9; Northern and Western Cape; polyphyletic), Ezoloba (1; E. macrocarpa; Western Cape), Calobota (13; North and South Africa), ’Lebeckia’ (c 35; 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 280; Western Cape to KwaZulu-Natal), ’Lotononis’ (c 150; the Mediterranean, Africa, southwestern Asia to India, with their largest diversity in South Africa; paraphyletic), Bolusia (5–6; arid and semiarid regions of Africa south of the equator), Crotalaria (c 700; tropical and subtropical regions on both hemispheres, with their largest diversity in Africa and Madagascar), Listia (7; southeastern Africa), Pearsonia (7; tropical to southern Africa), Rothia (2; Africa southwards to South Africa and eastwards to India and Australia), Robynsiophyton (1; R. vanderystii; southern Central Africa), Leobordea (51; tropical and southern Africa, northeastern Africa, southwestern Asia), Euchlora (1; E. hirsuta; Northern and Western Cape), Dichilus (5; southern Africa), Polhillia (7; Western Cape), Melolobium (c 20; southern Africa). – Mainly South Africa, few species in eastern and northeastern Africa and southwestern Asia.

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

c 6/c 500. 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 (c 200; 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). – Eurasia, Africa, North and South America. α-pyridone-type quinolizidine alkaloids present.

[Vataireoid clade+[Andirinae+Dalbergioideae]]

Vataireoid clade

4/21. Luetzelburgia (13; northeastern South America to Bolivia, with their highest diversity in Brazil); Sweetia (1; S. fruticosa; tropical South America), Vataireopsis (4; 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’]

2/c 47. Hymenolobium (17; tropical South America), Andira (c 30; tropical Central and South America, one species in tropical West Africa). – Tropical Central and South America, tropical West Africa.

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

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

c 8/240–245. Amorpha (16; North America, with their highest diversity in southeastern United States), Parryella (1; P. filifolia; Mexico), Apoplanesia (1; A. paniculata; drier regions in Central America), Errazurizia (4; arid regions in southwestern United States, coastal Chile), Eysenhardtia (12–15; Central America), ’Psorothamnus’ (9; 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

45–46/1.280. Riedeliella (3; Brazil, Paraguay); Adesmia (c 240; South America), Poiretia (11; tropical America), Amicia (7; tropical America), Zornia (c 80; warmer regions on both hemispheres, with their largest diversity in Brazil), Nissolia (14; Mexico, Central America, the West Indies, tropical South America), Chaetocalyx (13; tropical America); Platypodium (1–2; P. elegans; tropical America), Paramachaerium (5; tropical America), Pterocarpus (c 35; tropical and subtropical regions), Etaballia (1; E. dubia; Amazonia), Ramorinoa (1; R. girolae; western Argentina), Centrolobium (7; tropical America), Tipuana (1; T. tipu; Brazil, Bolivia, Argentina), Inocarpus (3; Malesia, Pacific islands), Discolobium (8; South America), Platymiscium (c 20; tropical America), Acosmium (3; tropical America), Maraniona (1; M. lavinii; northern Peru), Chapmannia (4; Somalia, Socotra), Arachis (c 70; South America), Stylosanthes (c 25; tropical and subtropical regions), Grazielodendron (1; G. riodocensis; Brazil), Cranocarpus (3; Brazil), Brya (4; the West Indies), Cascaronia (1; C. astragalina; Bolivia, northern Argentina), Geoffroea (2; South America), Fissicalyx (1; F. fendleri; Venezuela, Guyana), Fiebrigiella (1; F. gracilis; Bolivia), Machaerium (c 155; tropical America, one species also in coastal tropical West Africa), Steinbachiella (1; S. leptoclada; Bolivia), Dalbergia (c 250; tropical and subtropical regions), ’Aeschynomene’ (c 175; tropical and subtropical regions; polyphyletic), Cyclocarpa (1; C. stellaris; tropical regions in the Old World), Kotschya (c 30; tropical to southern Africa, Madagascar), Soemmeringia (1; S. semperflorens; tropical South America), Smithia (c 20; tropical and subtropical regions in the Old World), Humularia (c 35; tropical Africa), Bryaspis (2; tropical West Africa), Geissaspis (3; tropical and subtropical regions in Asia), Pictetia (8; the West Indies), Diphysa (15; tropical America), Zygocarpum (6; Somalia, southern Arabian Peninsula, Socotra), Peltiera (2; central Madagascar, extinct?), Ormocarpum (c 20; tropical and subtropical regions in the Old World), Ormocarpopsis (6; Madgascar; in Ormocarpum?), Weberbauerella (2; 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

5/55–60. Baphia (45–50; tropical to southern Africa, Madagascar), Baphiopsis (1; B. parviflora; tropical Africa), Leucomphalos (6; Africa, Madagascar, East Asia), Airyantha (1; A. schweinfurthii; Central Africa), Dalhousiea (3; tropical West Africa, northeastern India, Bangladesh). – Tropical Africa to East Asia. Trees, shrubs, lianas. – Baphia is sister to the remaining Faboideae, the Canavanine-accumulating clade.

Canavanine-accumulating clade

Canavanine (non-protein amino acid) present. – The peptide pea albumin is present in Millettieae 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; Western and Eastern Cape).

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

c 33/790–800. 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 (22; Australia, with their highest diversity in southwestern Australia), Goodia (2; southwestern and southeastern Australia), Bossiaea (c 60; temperate Australia, with their highest diversity in southwestern Australia), Platylobium (4; southeastern South Australia to southeasternmost Queensland, Tasmania), Muelleranthus (3; Australia), Paragoodia (1; P. crenulata; southwestern Western Australia), Ptychosema (2; Central and Western Australia), Aenictophyton (1; A. reconditum; northwestern Australia), Paragoodia (1; P. crenulata; southwestern Western Australia). – Multiple embryo-sacs group Isotropis (14; Australia), Chorizema (27; southwestern and eastern Australia), ‘Mirbelia’ (32; Australia; polyphyletic), Oxylobium (15; Australia, with their highest diversity in southwestern Australia), ‘Podolobium’ (6; southwestern and southeastern Australia; polyphyletic), Callistachys (1; C. lanceolata; southwestern Western Australia), Gastrolobium (c 110; Australia), Euchilopsis (1; E. linearis; southwestern Western Australia), Phyllota (11; southwestern and eastern Australia), Latrobea (6; southwestern Western Australia), Jacksonia (c 75; Australia), Leptosema (13; northern, central and southwestern Australia), Urodon (4; southwestern Western Australia), Otion (8; northern, central and southwestern Australia), Aotus (18; Australia), Stonesiella (1; S. selaginoides; Tasmania), Almaleea (5; eastern New South Wales, northeasternmost Victoria), Eutaxia (8–9; Australia, with their highest diversity in southwestern Australia), Dillwynia (c 40; Australia), ‘Pultenaea’ (c 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 etc.]+Hologalegina]

[Indigofereae+Phaseoleae etc.]

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

7/785–790. Disynstemon (1; D. paullinioides; southwestern Madagascar); Indigofera (c 730; tropical and subtropical regions), Cyamopsis (4; drier regions in Africa southwards to South Africa and eastwards to India), Indigastrum (9; tropical and southern Africa, one pantropical species), Phylloxylon (7; Madagascar), Microcharis (c 35; Africa southwards to South Africa, Madagascar, the Arabian Peninsula), Rhynchotropis (2; southern Central Africa). – Pantropical, with their largest diversity in Africa and Madagascar. Inflorescence racemose. – Disynstemon is sister to the remaining Indigofereae.

The Xeroderris, Dalbergiella etc. basal grade’

Austrosteenisia (4; New Guinea, northern Australia), Burkilliodendron (1; B. album; Malay Peninsula)?, Craibia (10; tropical to southern Africa), Craspedolobium (1; C. schochii; western China), Dalbergiella (3; tropical Africa), ‘Fordia’ (18; Southeast Asia, West Malesia; non-monophyletic), Kunstleria (8; Kerala in southwestern India, West and Central Malesia), Leptoderris (20; tropical Africa), Ostryocarpus (1–2; tropical western central Africa; incl. Xeroderris?)?, Xeroderris (1; X. stuhlmannii; tropical to southern Africa; in Ostryocarpus?)?, Platycyamus (2; Brazil, Peru), Platysepalum (7–8; tropical Africa), Schefflerodendron (3–4; tropical Africa), Sylvichadsia (4; Madagascar).

Phaseoleae DC., Prodr. 2: 381. mid Nov 1825

145–155?/3.040–3.050. Abrus (17; tropical and subtropical regions); ’Dioclea’ (c 40; tropical America, few species in the Old World; non-monophyletic), Bionia (5; Brazil, especially Minas Gerais), Luzonia (1; L. purpurea; the Philippines)?, Macropsychanthus (c 2; New Guinea, Micronesia, the Philippines?)?, Canavalia (c 60; warmer regions of both hemispheres incl. the Hawaiian Islands, with their highest diversity in tropical and subtropical America), Cymbosema (1; C. roseum; tropical America), Cleobulia (3–5; Mexico, Brazil), Camptosema (c 10; South America), Cratylia (7; South America), ’Galactia’ (c 55; tropical and subtropical regions, with their largest diversity in South America; non-monophyletic), Rhodopis (2; Hispaniola), Neorudolphia (1; N. volubilis; Puerto Rico)?; Ophrestia (16; Africa, southern Asia), Pseudoeriosema (4–5; tropical Africa)?; ’Millettia’ (c 150; tropical and subtropical regions in the Old World, especially Africa and Madagascar; polyphyletic), Philenoptera (12; tropical Africa, Madagascar), Hesperothamnus (5; Mexico), Piscidia (7; Florida, Central America, the West Indies), Dahlstedtia (2; southern Brazil, northern Argentina), Deguelia (15–17; Panamá to Amazonia), ‘Lonchocarpus’ (c 120; tropical America, one species, L. sericeus, also in tropical West Africa; paraphyletic), Muellera (7; tropical America), Behaimia (1; B. cubensis; Cuba), ’Derris’ (c 65; tropical regions of the Old World eastwards to New Guinea, with their largest diversity in Southeast Asia; non-monophyletic), Solori (9?; Southeast Asia), Brachypterum (9?; Southeast Asia), Aganope (6; tropical Africa, Southeast Asia), Pongamiopsis (3; Madagascar), Pyranthus (6; Madagascar)?, Chadsia (9; Madagascar), Mundulea (c 12; southern Africa, Madagascar), Tephrosia (>350; tropical and subtropical regions on both hemispheres, especially Africa), Apurimacia (2; drier regions in South America), Requienia (3; western to northeastern Africa, southern Africa)?, Ptycholobium (3; drier regions in northeastern and southern Africa, the Arabian Peninsula); ’Otholobium’ (c 60; eastern and southeastern Africa, South Africa, South America; non-monophyletic), Psoralea (c 20; southern Africa, with their largest diversity in Western Cape), Orbexilum (8; the United States, Mesico), Rupertia (3; western North America), Ladeania (2; western United States), Pediomelum (c 20; North America), Cullen (c 35; the Mediterranean, Africa, southern Asia and eastwards to New Guinea and Australia), Bituminaria (3; the Mediterranean); Cologania (12; tropical America, with their highest diversity in Mexico), Calopogonium (5–6; tropical America, one species, C. mucunoides, also in the Old World tropics), Pachyrhizus (5; tropical America), Herpyza (1; H. grandiflora; western Cuba)?, Neorautanenia (3; southern tropical Africa), Teyleria (3; Southeast Asia, West Malesia), Dumasia (10; Africa and eastwards to southern Asia), ’Pueraria’ (17; southern and eastern Asia; polyphyletic), Nogra (3; Asia; non-monophyletic?), Eminia (4; Zambesian region in Africa), Sinodolichos (2; Burma, southern Chine; in Glycine?), Pseudeminia (4; tropical Africa), Pseudovigna (2; tropical Africa), Amphicarpaea (4–5; Africa, East Asia, North America), Teramnus (9; tropical and subtropical regions on both hemispheres), ’Glycine’ (c 20; Africa, southern Asia to Australia; non-monophyletic?), Phylacium (2; Southeast Asia to Queensland), Neocollettia (1; N. wallichii; Burma, Java)?; ’Vigna’ (c 100; tropical and subtropical regions on both hemispheres; non-monophyletic), Ancistrotropis (6; Mexico to Bolivia), Cochliasanthus (1; C. caracalla; northern South America, Bolivia, Argentina), Condylostylis (4; Mexico to Argentina), Helicotropis (3; Mexico to Bolivia), Leptospron (2; Mexico to Colombia), Sigmoidotropis (9; Mexico to Peru, Hispaniola), Ramirezella (7; Mexico, Central America), Sphenostylis (7–8; Africa, India), Wajira (5; Africa, India, Sri Lanka), Nesphostylis (4; tropical Africa, tropical Asia), Oxyrhynchus (4; East Malesia, Central America), Physostigma (4; tropical Africa), Alistilus (3; southern tropical Africa, Madagascar), Strophostyles (3; North America), Dolichopsis (1; D. paraguariensis; southern Brazil, Paraguay, northern Argentina), Mysanthus (1; M. uleanus; Brazil), Dipogon (1; D. lignosus; Western and Eastern Cape), Lablab (1; L. purpureus; tropical? Africa), Oryxis (1; O. monticola; Minas Gerais in Brazil), Austrodolichos (1; A. errabundus; northern Australia), Spathionema (1; S. kilimandscharicum; tropical Africa), Vatovaea (1; V. pseudolablab; tropical East Africa to Oman), Phaseolus (c 60; tropical and subtropical regions in America), Dolichos (c 60; tropical and subtropical Africa to South Africa and eastwards to India and East Asia), Macrotyloma (c 25; tropical and subtropical Africa and Asia), Macroptilium (17; tropical America); Erythrina (c 120; tropical and subtropical regions on both hemispheres), Psophocarpus (c 10; tropical and subtropical regions in the Old World), Dysolobium (4; eastern India, southwestern China, Indochina, Malesia)?, Otoptera (2; tropical Africa, Madagascar), Decorsea (4; tropical to southern Africa, Madagascar), Strongylodon (12; Madagascar, tropical Asia and eastwards to Polynesia), Adenodolichos (15; tropical Africa), Paracalyx (6; Ethiopia, Somalia, Socotra, India, Indochina), Bolusafra (1; B. bituminosa; Western Cape), Carrissoa (1; C. angolensis; Angola)?, Chrysoscias (3–4; Western Cape)?, Rhynchosia (c 230; tropical and subtropical regions), Eriosema (c 150; tropical and subtropical regions), Dunbaria (c 20; India, Southeast Asia to China, Malesia, New Guinea, northern Australia), Cajanus (c 35; tropical regions in the Old World eastwards to Australia), Flemingia (c 30; tropical regions in the Old World), Spatholobus (c 30; Southeast Asia to Central Malesia), Butea (2; tropical Asia), Meizotropis (2; India, Himalayas, western Indochina)?, Apios (10; East Asia, North America), Cochlianthus (2; Nepal, western China), Shuteria (4–5; tropical Asia), Mastersia (2; Assam, Central Malesia), Diphyllarium (1; D. mekongense; Indochina)?, Mucuna (c 105; tropical and subtropical regions on both hemispheres), Campylotropis (c 37; Himalayas, China, the Korean Peninsula, Taiwan), Kummerowia (2; East Asia, North America), Lespedeza (c 35; tropical and eastern Asia, Australia, temperate North and South America), Dendrolobium (18; tropical Asia, Indian Ocean islands, Australia), Phyllodium (8; tropical Asia to northern Australia), Ougeinia (1; O. oojeinensis; India, western Nepal)?, Aphyllodium (7; India, Sri Lanka, Hainan, Indochina, Malesia, New Guinea, northern Australia)?, Ohwia (2; India, China, Japan, Indochina, Malesia)?, Hanslia (2; Malesia, New Guinea, northern Queensland, Vanuatu)?, Arthroclianthus (c 30; New Caledonia, one species also in Vanuatu)?, Nephrodesmus (6; New Caledonia)?, Droogmansia (5; tropical Africa; incl. Tadehagi?)?, Tadehagi (5; tropical Asia; in Droogmansia?)?, Trifidacanthus (1; T. unifoliolatus; Hainan, southern Vietnam, Lombok, Flores, the Philippines)?, ‘Desmodium’ (c 275; tropical and subtropical regions; non-monophyletic), Hylodesmum (14; tropical Asia), Verdesmum (1; V. hentyi; Yunnan), Ototropis (1; O. sambuensis; tropical Asia, southern China, Taiwan), Codariocalyx (2; India, Sri Lanka, China, Taiwan, Indochina, Malesia to tropical Australia), Pseudarthria (3–4; southern Africa, Madagascar, Mauritius, Réunion, tropical Asia), Pycnospora (1; P. lutescens; tropical East Africa to Somalia, India, East and Southeast Asia to northern Australia), Mecopus (1; M. nidulans; India, Hainan, Indochina, Java)?, Uraria (c 20; tropical and subtropical regions in the Old World), Christia (c 10; India to China and Indochina, Malesia, northern Australia)?, Alysicarpus (25–30; tropical and subtropical Africa, southern Asia), Melliniella (1; M. micrantha; western central Africa)?, Leptodesmia (3; Madagascar, India)?, Eleiotis (2; India and Sri Lanka to Burma)?; Kennedia (16; New Guinea, Australia), Hardenbergia (3; Australia), Vandasina (1; V. retusa; New Guinea, northeastern Queensland); Clitoria (c 60; tropical and subtropical regions, with their largest diversity in South America), Centrosema (c 35; tropical and subtropical regions in America), Periandra (6; Hispaniola, Brazil, Bolivia)?, Clitoriopsis (1; C. mollis; Congo, Sudan)?; unplaced: Dewevrea (1–2; tropical West Africa). – Tropical to warm-temperate regions. Inflorescence often pseudoracemose. – The relationships among the many lineages in Millettieae are far from understood and the clades have low bootstrap support.

Hologalegina

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

[Robinioid clade+IRLC]

Robinioid clade

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

11/75–80. Robinia (4–5; North America), Genistidium (1; G. dumosum; Texas, Mexico), Peteria (4; southwestern North America), Coursetia (38; southwestern United States, Mexico, Central America, the West Indies, tropical South America to Brazil and Peru), Olneya (1; O. tesota; southwestern North America), Sphinctospermum (1; S. constrictum; southwestern North America), Poissonia (4–5; Peru, Bolivia, Argentina), Lennea (5; Central America), Hebestigma (1; H. cubense; Cuba), Gliricidia (5; tropical America), Poitea (12; the West Indies). – Warm-temperate to tropical regions in America. Loss of plastid inverted repeat.

[Sesbanieae+Loteae]

The support of this clade is 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.

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

15–20/c 255. 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; western Mediterranean, northeastern Africa, the Arabian Peninsula), Ornithopus (5; Europe, the Mediterranean, western Asia, Atlantic islands, temperate South America), Hosackia (11; southwestern Canada to Guatemala), Acmispon (8; southern Canada to Mexico, one species, A. subpinnatus, in Chile; incl. Kebirita, Ottleya and Syrmatium?), Ottleya (13; southwestern North America; in Acmispon?), Syrmatium (14; western North America; in Acmispon?), Kebirita (1; K. roudairei; northwestern Africa; in Acmispon?), Hippocrepis (34; Europe, the Mediterranean, western Asia), Scorpiurus (2; Macaronesia, the Mediterranean, the Middle East to Iran), Coronilla (9; Europe, the Mediterranean, Atlantic islands; incl. Securigera?), Securigera (13; Europe, the Mediterranean, northeastern Africa to Somalia; in Coronilla?), Anthyllis (22; Europe, Macaronesia, the Mediterranean, northeastern Africa), Hammatolobium (2; the Mediterranean), Tripodion (1; T. tetraphyllum; the Mediterranean), Cytisopsis (2; eastern Mediterranean, North Africa), Lotus (c 100; temperate regions in the Old World; incl. Dorycnium?), Dorycnium (8; Macaronesia, the Mediterranean, southwestern Asia; in Lotus?). – Temperate regions on the Northern Hemisphere, temperate South America.

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

1/18. Glycyrrhiza (18; Europe, Asia, Australia, North America, temperate South America; incl. ‘Calleryaatropurpurea).

[Wisterieae+[Astragaleae+Vicieae]]

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

5/30–35. Wisteria (6; China, the Korean Peninsula, Japan, eastern United States), ’Callerya’ (c 20; East and Southeast Asia and eastwards to northeastern Australia and New Caledonia; polyphyletic), Endosamara (1–2; India, Sri Lanka, Southeast Asia, Malesia), Afgekia (3; southern China, Burma, Thailand), Antheroporum (1 or 4; southwestern China, Indochina). – Eastern North America, East, South and Southeast Asia, New Guinea, Australia, New Caledonia. n = 8. rps12 intron present.

[Hedysareae+Fabeae]

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

25–27/>4.300. Erophaca (1; E. baetica; the Mediterranean), Oxytropis (c 350; temperate regions on the Northern Hemisphere, especially Central Asia), Astragalus (>3.270; mainly temperate and subtropical regions on the Northern Hemisphere, tropical African mountains, northern India, the Andes southwards to Chile, one species in southern Africa; incl. Calophaca?), Calophaca (5–8; Siberia, Central Asia; in Astragalus?); Colutea (28; the Mediterranean, Asia eastwards to China, Himalayas, northestern and eastern Africa), Oreophysa (1; O. microphylla; Iran), Eremosparton (3; southeastern Russia to Central Asia), Sphaerophysa (2; eastern Mediterranean and eastwards to Central Asia), Smirnowia (1; S. turkestana; Turkestan), Lessertia (c 55; tropical eastern and southern Africa), Swainsona (c 85; drier regions in Australia), Clianthus (2; northeastern New Zealand), Carmichaelia (23; New Zealand, Lord Howe Island), Streblorrhiza (1; S. speciosa; Philip Island near Norfolk Island, extinct); Chesneya (c 30; Southwest and Central Asia to Mongolia), Spongiocarpella (7; southern Central Asia and Himalaya to western China)?, Gueldenstaedtia (14; Siberia to Himalaya and western China; incl. Tibetia?), Tibetia (4; Central Asia, Himalaya, western China; in Gueldenstaedtia?); Caragana (c 80; eastern Europe, Siberia, West and Central Asia to China), Halimodendron (1; H. halodendron; Europe, Turkey to Central Asia); Alhagi (1; A. maurorum; the Mediterranean and eastwards to Nepal), Onobrychis (c 130; Europe, the Mediterranean, Asia, Ethiopia), Sartoria (1; S. hedysaroides; southern Turkey)?, Ebenus (18; the Mediterranean and eastwards to Baluchistan), Hedysarum (c 160; temperate regions on the Northern Hemisphere), Taverniera (15; northeastern Africa, southwestern Asia), Eversmannia (4; southeastern Russia, northern Iran, Central Asia). – Temperate regions on the Northern Hemisphere, southern Africa, Australia, New Zealand, few species in East Africa and temperate South America.

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

10/c 870. Parochetinae Yakovlev, Bobovye Zemnogo Shara: 117. 1991. Parochetus (1; P. communis; mountains of tropical Africa and Asia to Java). – Fabinae (Viciinae Bronn, Form. Plant. Legumin.: ad Sect. 134, 133. 1822 [‘Vicieae’]). Galega (6; eastern Europe, East African mountains, temperate Asia), Cicer (c 45; Greece, the Canary Islands, Morocco, Ethiopia, West and Central Asia), Trifolium (c 240; temperate and subtropical regions on both hemispheres except Australia), Lathyrus (c 160; temperate regions on the Northern Hemisphere, tropical East African mountains, temperate South America); Ononis (c 75; Europe, the Canary Islands, the Mediterranean, Ethiopia, Iran), Medicago (c 85; Europe, the Mediterranean, Ethiopia, southern Africa, Asia), Trigonella (c 55; Macaronesia, the Mediterranean, southern Africa, Australia), Melilotus (c 20; Europe, the Mediterranean, North Africa, Ethiopia, temperate and subtropical Asia); Vicia (c 160; temperate regions on the Northern Hemisphere, tropical East Africa, South America, the Hawaiian Islands). – Subcosmopolitan. – Parochetus is sister to Fabinae. A peculiar type of microhairs is a synapomorphy of Fabinae. Vicia has secondary pollen display by stigmatic pollen brush.

Unplaced Fabaceae

Chidlowia (1; C. sanguinea; western tropical Africa), Orphanodendron (1–2; O. bernalii; northwestern Colombia), Sympetalandra (5; Malesia including the Philippines).

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.

POLYGALACEAE Hoffmanns. et Link

( Back to Polygalales )

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

Genera/species c 28/900–1.000

Distribution Cosmopolitan except polar regions, Polynesia and New Zealand.

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 comprises achlorophyllous root holoparasites and Epirixanthes holomycotrophs). Some species are xerophytes. Often with paired crateriform glands (extrafloral nectaries) or spines at nodes (sometimes elsewhere).

Vegetative anatomy 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.

Trichomes Hairs unicellular, simple, or absent.

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).

Systematics Xanthophyllum is sister to the remaining Polygalaceae.

Xanthophyllum

1/c 95. Xanthophyllum (c 95; southern India, Sri Lanka, Southeast Asia, Malesia, New Guinea, the Solomon Islands, Queensland). – Trees or shrubs. Axial parenchyma apotracheal, diffuse. Glands present at nodes. Hypogyny. Sepals unequal, caducous. Petals five, with contorted aestivation. Stamens (seven or) eight (to ten), diplostemonous. Filaments expanded and hairy below, not adnate to petals. Anthers dorsifixed, introrse, longicidal. Pollen grains 5–11-colporate. Pistil composed of two connate carpels. Ovary unilocular, stipitate. Style single. Stigma small, usually bilobate (sometimes capitate). Placentation parietal. Ovules two to 16 per carpel, apotropous, inserted in two rows. Outer integument four to twelve cell layers thick. Fruit sometimes an irregularly loculicidal capsule. Testa multiplicative, often indistinct. Hypostase massive. Aluminium accumulated. n = ?

Polygaloideae Eaton, Bot. Dict., ed. 4: 46. Apr-Mai 1836 [‘Polygaleae’]

c 17/830–845. Hypogyny. Petals three or five, large. Filaments connate, often adnate to petals, often monadelphous. Anthers dehiscing by short apical slits. Pistil composed of two to five (to eight) carpels. Ovule one per carpel, epitropous. Funicular or exostomal aril often present.

Moutabeeae Chodat in Engler et Prantl, Nat. Pflanzenfam. III, 4: 329. Jul 1896

c 5/14–15. ’Moutabea’ (8; tropical America; paraphyletic), Balgoya (1; B. pacifica; New Caledonia), Eriandra (1; E. fragrans; New Guinea), Barnhartia (1; B. floribunda; tropical South America), Diclidanthera (3–4; tropical South America). – Tropical America, New Guinea to New Caledonia. Axial parenchyma apotracheal banded (Moutabea). Leaves (and sometimes nodes) with glands. Sepals adnate to petals. Petals five, abaxial petal not keeled. Stamens six to ten. Anthers dehiscing by confluent short apical slits. Pistil composed of three to eight connate carpels. Stigma capitate. Funicular aril present. n = 14. Aluminium accumulated.

[Carpolobieae+Polygaleae]

Carpolobieae B. Eriksen in Plant Syst. Evol. 186: 47. 1993

2/6. Atroxima (2; tropical West and Central Africa), Carpolobia (4; tropical Africa, Madagascar). – Tropical Africa, Madagascar. Petals five, sometimes with contorted aestivation; abaxial petal keeled. Stamens (four or) five. Anthers dehiscing by confluent short apical slits. Pistil composed of three connate carpels. Stigma capitate. n = 9–11.

Polygaleae Fr., Fl. Scan.: 35, 38. 1835

c 20/800–900. ’Bredemeyera’ (c 60; New Guinea, tropical Australia, tropical America; polyphyletic), Acanthocladus (8; Central America, tropical South America), Gymnospora (2; South America), Badiera (11; the West Indies), Hebecarpa (c 60; southern United States to South America), Securidaca (c 80?; tropical regions on both hemispheres), Comesperma (c 40; Australia), Ancylotropis (2; Brazil), Monnina (c 150; New Mexico and southwards to Chile, with their highest diversity in the Andes; incl. Pteromonnina?), Pteromonnina (c 30; the Andes; in Monnina?), Phlebotaenia (3; Cuba, Puerto Rico), Rhinotropis (17?; southwestern United States, Mexico), Asemeia (28; southern United States, Mexico, Central America, the West India, South America), Caamembeca (11; tropical South America), Muraltia (c 120; tropical and southern Africa, with their largest diversity in the Cape Provinces), Epirixanthes (5; tropical Asia), Polygala (300–400; almost cosmopolitan), Salomonia (3; tropical Asia, tropical Australia to eastern Queensland), Heterosamara (5?; tropical and southern Africa, tropical and subtropical Asia), Polygaloides (?; Europe, the Mediterranean, North Africa, western Asia, eastern North America). – Cosmopolitan except polar areas, Polynesia and New Zealand. Vestured pits present. Axial parenchyma paratracheal banded. Two abaxial lateral sepals minute, two adaxial lateral sepals alae. Two abaxial lateral petals minute (sometimes absent), two connate adaxial petals velum, abaxial petal carina (often fimbriate). Stamens (two to) four or six to eight (in Polygala myrtifolia, with eight stamens, two median stamens on opposite sides of flower probably lost). Anthers dehiscing by apical pores. Pistil composed of two connate carpels (adaxial carpel sometimes reduced; gynoecium sometimes pseudomonomerous). Stylar canal present. Stigma bilobate, asymmetric. Carunculus often present. Chalazal aril sometimes present. Fruit a capsule (often flattened), a samara or a drupe. n = 6 or more. – Epirixanthes consists of holomycoheterotrophs, whereasSalomonia comprises root holoparasites.

Cladogram of Polygalaceae based on DNA sequence data (Persson 2001).

Cladogram of Polygalaceae based on DNA sequence data (Forest & al. 2007; Banks & al. 2008). The generic nomenclature is updated.

Phylogeny (Bayesian inference) of Polygalaceae based on DNA sequence data (Pastore 2012; Epirixanthes added here; Nylandtia synonymized with Muraltia).

QUILLAJACEAE D. Don

( Back to Polygalales )

Don in Edinburgh New Philos. J. 10: 299. 1831 [‘Quillajeae’]

Quillajales Doweld, Tent. Syst. Plant. Vasc.: xxxviii. 23 Dec 2001

Genera/species 1/3–4

Distribution Temperate and subtropical South America.

Fossils Unknown.

Habit Bisexual or unisexual, evergreen small trees.

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.

Trichomes Hairs verrucose.

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?

Cytology n = 14, 17

DNA Gene rpl22 present in nuclear or plastid genome?

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

Use Medicinal plants, detergents, food industry.

Systematics Quillaja (3–4; Q. saponaria: central Chile from 31S to 38S; Q. brasiliensis: Peru?, southeastern Brazil to Uruguay and northern Argentina).

SURIANACEAE Arn.

( Back to Polygalales )

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

Genera/species 5/8

Distribution Tropical beaches, Australia, Mexico.

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.

Cytology n = ?

DNA Gene rpl22 present in nuclear or plastid genome?

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

Use Unknown.

Systematics Recchia (3; Mexico), Cadellia (1; C. pentastylis; southeastern Queensland, northeastern New South Wales), Suriana (1; S. maritima; pantropical, sea-shores), Guilfoylia (1; G. monostylis; eastern Queensland, northeastern New South Wales), Stylobasium (2; mainly western and central Australia).

Analyses of matK (Bello & al. 2009) resulted in the topology [[Recchia+Cadellia]+[Suriana+[Guilfoylia+Stylobasium]]].

Nelsen strict consensus tree of Surianaceae based on DNA sequence data (Bello & al. 2009).


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