POALES Small Main Tree, Synapomorphies

Poales Small, Fl. S.E. U.S.: 48. 22 Jul 1903; Poanae Takht. ex Reveal et Doweld in Novon 9: 550. 30 Dec 1999

Habit Usually bisexual (sometimes monoecious, andromonoecious, gynomonocious, polygamomonoecious, dioecious, androdioecious, or gynodioecious), usually perennial, biennial or annual herbs (sometimes suffrutices, rarely trees, shrubs, or lianas). Sometimes hygrophytic or aquatic. Often xerophytes.

Vegetative anatomy Mycorrhiza often absent. Phellogen absent. Primary vascular tissue two or more cylinders of vascular bundles or scattered bundles. Secondary lateral growth absent. Vessels present in roots and/or stem and/or leaves. Vessel elements with scalariform or simple (rarely reticulate) perforation plates; lateral pits scalariform or alternating. Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids usually P2c or P2cf Type (rarely P2cs type). Nodes? Secretory cavities absent. Idioblasts with silica bodies (often conical, rounded, saddle-shaped or quadratic) present or absent. Calciumoxalate raphides present or absent. C₄ and CAM physiology occurring.

Trichomes Hairs unicellular or multicellular, uniseriate, simple or branched (sometimes stellate or peltate-lepidote, or T-shaped, and sometimes malpighiaceous, hairs), or absent; glandular hairs sometimes present; microhairs sometimes present.

Leaves Alternate (distichous or tristichous, sometimes tetrastichous), simple, entire, usually linear, sometimes differentiated into pseudopetiole and pseudolamina, sometimes reduced to leaf sheath, with supervolute/convolute, involute, conduplicate, plicate, adplicate, revolute, or plicate (rarely circinate) ptyxis. Stipules absent; leaf sheath usually well developed, open or closed, often with membranous adaxial ligule at distal end of sheath (ligule sometimes modified into hairs or absent). Venation usually parallelodromous, often with transverse veins (sometimes pinnate-parallelodromous). Stomata paracytic or tetracytic (rarely anomocytic), neighbouring cells with oblique or non-oblique divisions. Cuticular wax crystalloids usually as longitudinally aggregated rodlets (Strelitzia type; sometimes scale-like or spherical). Epidermis often with cell walls containing one or several silica bodies. Mesophyll sometimes with sclerenchymatous idioblasts. Mucilaginous idioblasts present or absent. Mesophyll usually without calciumoxalate crystals (sometimes with druses, styloids or single prismatic crystals). Secretory cavities usually absent. Cystoliths absent. Leaf margin usually entire (sometimes serrate or spinose-dentate).

Inflorescence Terminal, panicle, thyrsoid, corymb, simple or compound, spike-, head-, spadix- or raceme-like, or spike, raceme or head (flowers sometimes solitary). Floral prophylls (bracteoles) one, two or absent.

Flowers Usually actinomorphic (rarely more or less zygomorphic). Hypogyny. Tepals usually three, 3+3 (sometimes 2+2, rarely one, four, 2+4, or more than six) or modified into scales or bristles (rarely hairs), or absent; outer tepals with contorted or imbricate (sometimes open, rarely cochlear or conduplicate) aestivation, sepaloid or petaloid; inner tepals with imbricate or contorted aestivation, sepaloid or petaloid. Nectary usually absent (septal nectaries sometimes present). Disc absent.

Androecium Stamens usually three or 3+3 (sometimes one, two, 2+2, five, or more than six). Filaments usually free from each other (sometimes more or less connate), usually free from tepals (sometimes epitepalous). Anthers dorsifixed, subbasifixed or basifixed, versatile or non-versatile, usually tetrasporangiate (rarely disporangiate or trisporangiate), introrse, latrorse or extrorse, longicidal (dehiscing by longitudinal slits) or poricidal (dehiscing by apical pores or short slits). Tapetum secretory. Staminodia usually absent (female flowers sometimes with staminodia).

Pollen grains Microsporogenesis usually successive (rarely simultaneous). Pollen grains usually monoporate to monoulcerate, diporate to tetraporate (to heptaporate) or pantotreme (sometimes spiraperturate or trichotomosulcate, rarely disulculate or inaperturate), often operculate, usually shed as monads (sometimes as cryptotetrads), usually tricellular (sometimes bicellular) at dispersal. Exine tectate or semitectate, usually with columellate (sometimes granular) infratectum, reticulate to punctate, echinulate, echinate, spinulate, finely scabrate, microverrucate, scrobiculate, or smooth.

Gynoecium Pistil composed of (one to) three (or four) connate carpels. Ovary superior, usually unilocular or trilocular (rarely bilocular). Style single, simple, or stylodia two or three, free or more or less connate. Stigma single, usually bilobate (rarely uni-, tri- or quadrilobate), or stigmas two or three (to nine), papillate or non-papillate, Dry type. Pistillodium usually absent (male flowers sometimes with pistillodium).

Ovules Placentation axile, basal, apical or parietal. Ovules one to more than 50 per ovary, anatropous, hemianatropous, amphitropous, orthotropous or campylotropous, ascending, horizontal or pendulous (often orthotropous pendulous), usually bitegmic (rarely unitegmic or almost ategmic), crassinucellar or incompletely tenuinucellar (to pseudocrassinucellar). Micropyle usually endostomal (sometimes bistomal). Funicular obturator occasionally present. Parietal cell formed from archesporial cell, or absent. Nucellar cap often present. Megagametophyte usually monosporous, Polygonum type (rarely disporous, Allium type, or tetrasporous, close to Drusa type). Synergids sometimes haustorial. Antipodal cells persistent, sometimes binucleate, sometimes more than three, sometimes proliferating (rarely absent). Endosperm development ab initio nuclear or helobial. Endosperm haustoria chalazal and micropylar, or absent. Embryogenesis usually asterad (sometimes onagrad).

Fruit A loculicidal and/or septicidal capsule, a nutlet (achene), a follicle, a berry or a nut-like caryopsis with pericarp usually fused with seed coat in adaxial end (sometimes a drupe, rarely a nutlet with membranous or gelatinous pericarp free from seed coat, or a syncarp or coenocarp).

Seeds Aril absent. Caruncular elaiosome occasionally present. Testa usually dry (sometimes fibrous; sometimes developing into a sarcotesta; or degenerating). Exotesta sometimes with silica bodies, sometimes fibrous. Endotesta sometimes persistent, sometimes with silica bodies. Tegmen sometimes fibrous, sometimes tanniniferous. Exotegmen sometimes degenerating. Perisperm not or only slightly developed. Endosperm copious, starchy, usually with proteinaceous tissue (rarely absent), with aleurone and sometimes oils; starch grains simple or compound. Embryo usually minute (sometimes large), straight or curved, usually well differentiated (sometimes rudimentary), with or without chlorophyll. Cotyledon one, not photosynthesizing. Cotyledon hyperphyll compact (sometimes modified into haustorium) and not assimilating, or elongate and assimilating. Hypocotyl internode present or absent. Mesocotyl sometimes present. Coleoptile modified into substrate-penetrating plumule envelope, with or without lamina, or absent. Germination phanerocotylar or cryptocotylar.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin, syringetin), laricitrin, flavonol glycosides (glycosides of myricetin etc.), flavonol sulfate, flavone sulfates, flavone-5-, flavone-6- and C-glycosides, catechin, gossypetin, aurones, luteolin, 8-hydroxyluteolin, luteolin-5-methylether, derivatives of 6-hydroxyluteolin and 6-hydroxymyricetin (hydroxyflavonoids), luteolinidin glycoside, tricine (flavone-3’,5’-dimethyl ether of tricetine), 6-hydroxyflavonoids (quercetagetin, patuletin), flavanones, cyanidin 3-deoxyanthocyanins, triterpenes, chalcones, tannins, cinnamic acids, daphnetin, juncosol (phenol), isoquinoline alkaloids, tryptophane- or tyramine-derived alkaloids (gramine, hordenine, tyramine, etc.), pyrrolizidine alkaloids as 1-aminopyrrolizidine derivatives (loline, lolinine, norline, etc.), indole alkaloids, cyanogenic glycosides (triglochinin), steroidal saponins, tyrosine-derived cyanogenic compounds, chrysazine (anthraquinone), -sitosterol, and S-methylcysteine present. Ferulic, diferulic and p-coumaric acids (esterified) components of non-lignified cell walls. Ellagic acid not found.

Systematics Poales are sister-group to [Commelinales+Cannales] (Commelinanae).

Bromeliaceae and Typhaceae are basal branches in Poales. They were identified as sister-groups in several studies, and Stevens (2001 onwards) cites the following potential synapomorphies for the clade [Bromeliaceae+Typhaceae]: stomatal subsidiary cells with oblique divisions; leaf without distinct sheath; three-nucleotide deletion in the atpA gene. However, these features may be polarized as plesiomorphies instead. Oblique divisions of subsidiary cells also occurs outside Poales (e.g. Iridales). There is a large variation in leaf base sheathing among monocots in general. Finally, the three-base deletion may be interpreted as an insertion in the other clades. Givnish & al. (2010) found Bromeliaceae as sister to the remaining Poales, whereas Typhaceae are successive sister-group to the rest.

Rapateaceae are sister to the remaining Poales except Bromeliaceae and Typhaceae (with various higher or lower support) in many molecular analyses. The remainder of Poales form a somewhat different topology depending on the genes and methods used for the analyses. One possible topology is [Prionium+[Thurnia+[Juncaceae+Cyperaceae]]]+[[Mayaca+[Xyridaceae+ Eriocaulaceae]]+pooids]]. Another plausible topology may be the following (2012 version of Stevens 2001 onwards): [[[Xyridaceae+Eriocaulaceae]+[Mayaca+[Thurniaceae+[Juncaceae+ Cyperaceae]]]]+pooids], where Mayacaceae are sister to the cyperoids. The main clade, representing the vast majority of Poales is characterized by the features (Stevens 2001 onwards): cellulose fibrils in outer epidermal cell walls of root elongation zone arranged parallel to root axis; trichoblast in atrichoblast/trichoblast cell pair adjacent to apical meristem; absence of septal nectaries; pollen grains tricellular at dispersal; absence of parietal tissue in ovule; absence (loss) of the mitochondrial gen sdh4; and presence of (1->3),(1->4)-ß-D-glucans in cell walls. C₄ metabolism is widely distributed here. Moreover, the root hairs in most of those Poales which have been studied develop exclusively from special small cells in the epidermal (piliferous) layer. In most vascular plants the root hairs develop from any cell of the epidermal (piliferous) layer.

Stevens (2001 onwards) lists the following potential synapomorphies of the clade [[Eriocaulaceae+Xyridaceae]+[Mayaca+[Prionium+[Thurnia+[Juncaceae+Cyperaceae]]]]]: leaves spirally arranged; anthers basifixed; outer tepals persistent in fruit; deletions in ORF2280 region; loss of entire plastid gene accD and mitochondrial gene sdh4; and presence of flavonoids.

The xyrid clade, comprising Mayacaceae, Eriocaulaceae and Xyridaceae has, e.g., the following potential synapomorphies (Stevens 2001 onwards): silica bodies absent; spiral leaves; perianth differentiated into outer sepalous and inner petalous tepals; basifixed anthers; pollen grains with spinulate exine; style present; micropyle bistomal; calyx persistent in fruit; seed coat endostomal-tegmic; presence of operculum (’embryostega’) on the seed; undifferentiated embryo; and presence of flavonoids. The position of and support of this clade varies according to analysis methods.

Eriocaulaceae and Xyridaceae (at least Xyris itself) form a well supported monophyletic group in most analyses, characterized by the following potential synapomorphies, according to Stevens (2001 onwards): vessel elements with simple perforation plates; absence of silica bodies; inflorescence scapose, capitate, and provided with involucral bracts; clawed inner tepals; filaments adnate to inner tepals, antepetalous; presence of exothecium; exine provided with spines; seed ridged, with tegmic operculum; and cuticular layer present between testa and tegmen. The last character also occurs in Rapateaceae.

The strongly supported cyperoid clade [Prionium+[Thurnia+[Juncaceae+Cyperaceae]]] has the following potential synapomorphies (Stevens 2001 onwards): absence of mycorrhiza; trichoblasts arising from distal cell in a cell pair; culm angular in cross-section; leaves tristichous, with air canals; leaf sheath usually closed; perianth small, membranous and undifferentiated, or absent; microsporogenesis simultaneous; pollen grains porate, shed in tetrads, and tricellular at dispersal; ovules anatropous; seed coat testal-tegmic; photosynthesizing unifacial cotyledon hyperphyll (phanomer) usually present; presence of hypocotyl; seedling collar minute, with rhizoids; chromosomes with diffuse centromeres; deletions in ORF2280; a 3 bp deletion in the plastid gene atpA; and presence of amylophilic pteridophyte type starch grains, 3-deoxyanthocyanins and luteolin-5-methylether. The clade [Juncaceae+Cyperaceae] alone has endostomal micropyle; absence (loss) of plastid gene rpl23; and presence of luteolin-5’methyl ether.

The clade [[Mayaca+[Eriocaulaceae+Xyridaceae]]+pooids], with fairly weak support in some molecular analyses, is supported by the characters tenuinucellate ovules and minute and little differentiated embryo.

The pooid clade is strongly supported by molecular data and comprises [[Anarthriaceae+ [Centrolepidaceae+Restionaceae]]+[Flagellariaceae+[Joinvilleaceae+Ecdeiocoleaceae+Poaceae]]]]. Stevens (2001 onwards) cites the following potential synapomorphies: sieve tube plastids with cuneate and additional loosely packed protein crystals; leaves usually distichous and sheathing; floral prophylls (bracteoles) often absent; tepals membranous and undifferentiated; endothecial cell wall thickenings girdle-like; pollen grains monoporate, annulate/ulcerate and with scrobiculate exine (with minute pores penetrating tectum and foot layer); stigma plumose, with receptive cells on multicellular branches; placentation apical; ovule one per carpel, orthotropous; micropyle bistomal; embryo minute, undifferentiated, broad; seedling with collar rhizoids; deletions in the ORF2280 region; primary cell wall also containing (1–3, 1–4)-β-D-glucans; and presence of flavones.

The restioid clade of this monophyletic group includes Anarthriaceae, Restionaceae and Centrolepidaceae (the last group possibly being nested inside Restionaceae) and is supported by the synapomorphies: usually dioecious; root hairs developing from any epidermal cell (a reversal); culm with parenchymatous sheath, palisade chlorenchymatous tissue, and sclerenchymatous cylinder with vascular bundles inside; chlorenchyma with peg cells; stamens three, antepetalous; anthers dorsifixed; phanomer (photosynthesizing unifacial cotyledon hyperphyll) present; and loss of plastid gene rpoC1. Restionaceae and Centrolepidaceae share the apomorphies: absence of mycorrhiza; absence of hairs; anthers disporangiate and monothecal; pollen grain pore non-annulate, with irregular margin; cells of megasporangial epidermis anticlinally elongated; megagametophyte with compound starch grains (in particular around polar nuclei); and antipodal cells persistent and proliferating.

The pooids in a strict sense may have the topology [Flagellariaceae+[[Joinvilleaceae+ Ecdeiocoleaceae]+Poaceae]], which is followed here. On the other hand, Ecdeiocoleaceae are sometimes recovered as sister to Poaceae. In any case, the pooids s.str. clade is characterized by the following potential synapomorphies (Stevens 2001 onwards): trichoblasts arising from distal cell of each cell pair; pseudolamina with transverse veins; leaf sheath with distal ligule; inflorescence a panicle, with adaxial swellings on branches; presence of nucellar cap; massive suprachalazal zone; indehiscent fruit; cotyledon non-photosynthesizing; and primary cell walls with mixed-linkage glucans. Finally, the monophyletic group formed by Joinvilleaceae, Ecdeiocoleaceae and Poaceae may have the synapomorphies: silica bodies cuboid; epidermis with microhairs; foliar epidermis with long and silica-containing short cells; guard cells dumbbell-shaped; presence of fusoid cells (large colourless cells in the central mesophyll); hollow stem; presence of nucellar cap; first seedling leaf lacking lamina; presence of one 28 kb and one 6,4 kb inversion in the plastid genome; and loss of the functional plastid gene ycf2.

Hopkinsiaceae B. G. Briggs et L. A. S. Johnson in Telopea 8: 484. Jul 2000; Lyginiaceae B. G. Briggs et L. A. S. Johnson in Telopea 8: 488. Jul 2000

Habit Usually dioecious (rarely monoecious), perennial herbs. Graminids. Rarely with stilt roots. Hygrophytes. Culm branched (Anarthria) or simple (Lyginia), in cross-section terete or flattened.

Vegetative anatomy Root hairs lignified, developed from any epidermal cell. Lateral roots developing from a zone opposite protoxylem poles. Chlorenchymatous layer of peg cells present inside epidermis; inside this layer a cylinder of vascular bundles. Anarthria: common parenchymatic and sclerenchymatic envelope not forming vascular cylinder inside chlorenchyma, instead surrounding each vascular bundle; palisade tissue absent. Hopkinsia and Lyginia: subepidermal chlorenchyma separated from cortex by common sclerenchyma cylinder and parenchyma sheath. Phellogen absent. Secondary lateral growth absent. Vessels present in stem and leaves (probably absent from roots). Vessel elements with scalariform perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids probably P2c type, with cuneate protein crystals. Nodes? Silica bodies absent. Calciumoxalate crystals and druses present in Lyginia.

Leaves Anarthria: alternate (distichous), simple, entire, linear, equitant, isobifacial, laterally flattened, with margin facing culm, with convolute (supervolute) ptyxis; Hopkinsia and Lyginia: leaves reduced to scale-like sheaths. Stipules absent; leaf sheath open, with distal ligule. Venation parallelodromous. Stomata (brachy)paracytic, in furrows, with Poaceae type guard cells. Cuticular waxes? Epidermis without lines of alternating long and short cells. Mesophyll with sclerenchymatous idioblasts; mesophyll without calciumoxalate crystals. Leaf margin entire.

Inflorescence Terminal, few- to many-flowered, usually panicle (not consisting of spikelets, or spikelets one-flowered). Each flower subtended by one or two bracts. Floral prophyll (bracteoles) one, two or absent.

Flowers Actinomorphic, small. Hypogyny. Tepals 3+3, free, membranous to hard. Nectary absent. Disc absent.

Androecium Stamens three, antepetalous (outer staminal whorl absent). Filaments filiform, free or connate (Lyginia), free from tepals. Anthers subbasifixed, versatile, disporangiate (in Anarthria) or tetrasporangiate (in Hopkinsia and Lyginia), latrorse-introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Female flowers often with staminodia.

Pollen grains Microsporogenesis successive? Pollen grains graminoid, monoporate to monoulcerate, operculate? (ulcus with annulus), shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, scrobiculate to smooth (in Hopkinsia and Lyginia microverrucate).

Gynoecium Pistil usually composed of three connate antesepalous carpels (in Hopkinsia of one carpel). Ovary superior, trilocular. Stylodia three, free or connate at base, with adaxial stigmatic areas. Stigmas three, papillate, Dry type? (sometimes extremely elongated). Pistillodium absent or rudimentary in male flowers.

Ovules Placentation apical-axile. Ovule one per carpel, orthotropous, pendulous, bitegmic, tenuinucellar. Micropyle bistomal? Outer integument ? cell layers thick. Inner integument ? cell layers thick, tanniniferous. Hypostase present adjacent to chalazal end. Megagametophyte monosporous, Polygonum type (in Lyginia with compound starch grains). Antipodal cells? Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit Usually a loculicidal capsule (in Hopkinsia a nutlet with fleshy pedicel and persistent tepals).

Seeds Aril absent. Testa? Tegmen? Perisperm not developed. Endosperm copious, starchy. Embryo small?, chlorophyll? Cotyledon one (in Hopkinsia not photosynthesizing). Cotyledon hyperphyll? Hypocotyl internode? Mesocotyl? Coleoptile absent? Plumule? Collar hairs? Germination? Phanomer?

DNA ORF 2280 present. The plastid gene trnL has a 3 bp deletion and a 5 bp insertion. The plastid genome lacks the 28 kb inversion often present in Restionaceae, Joinvilleaceae, Ecdeiocoleaceae, and Poaceae.

Phytochemistry Insufficiently known. Flavonols (kaempferol, quercetin, myricetin) and flavonol glycosides present. Fructans present in Lyginia. Proanthocyanidins and flavonoid sulphates not found.

Bromeliales Link, Handbuch 1: 207. 4-11 Jul 1829 [’Bromelieae’]; Tillandsiaceae Wilbr., Nat. Pflanzenfam.: 44. Jun-Dec 1834 [’Tillandsieae’]; Bromeliopsida Brongn., Enum. Plant. Mus. Paris: xv, 23. 12 Aug 1843 [’Bromelioideae’]; Bromelianae R. Dahlgren ex Reveal in Novon 2: 235. 13 Oct 1992; Bromeliidae C. Y. Wu in Acta Phytotaxon. Sin. 40: 299. 2002

Distribution Tropical and subtropical regions of America from Virginia in the United States to Patagonia in South America; one species of Pitcairnia in tropical West Africa.

Fossils Uncertain. Karatophyllum bromelioides, a fossil from Costa Rica of the mid-Cenozoic, has been attributed to the Bromeliaceae.

Habit Usually bisexual (sometimes functionally unisexual), usually perennial herbs (rarely trees or lianas). Many species are epiphytic or epilithic. Some species are succulent. Two species of Brocchinia and one species of Catopsis are carnivorous.

Vegatative anatomy Majority of species with CAM metabolism (evolved multiple times). Intracauline adventitious roots present in stem cortex in species with a distinct stem. Lateral roots developing from a zone opposite protoxylem poles. Roots in epiphytes and epiliths modified into climbing organs; some species lack roots entirely. Phellogen? Secondary lateral growth absent. Vessels present in roots and often also in stem and leaves. Vessel elements usually with scalariform (in roots sometimes simple) perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays? Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals, or P2c_cl type, with cuneate and several additional loosely packed protein crystals. Nodes? Stem epidermis with silica bodies in inner periclinal cell walls. Mucilaginous idioblasts (raphide sacs) with calciumoxalate raphides.

Trichomes Water absorbing peltate hairs consisting of a proximal foot cell, a multicellular uniseriate stalk of living cells and a stellate to discoid peltate upper part consisting of numerous usually dead cells, often pressed against epidermis; rarely with multicellular simple uniseriate or stellate hairs or glandular hairs. Water absorbing stellate or lepidote peltate hairs present especially at leaf bases.

Leaves Alternate (usually spiral; in some species of Tillandsia distichous), simple, entire, often linear (sometimes filiform), usually hard, coriaceous or succulent (sometimes transformed into spines), with convolute (supervolute), curved or flat ptyxis. Stipules absent; leaf sheath indistinctly delimited from lamina. Venation parallelodromous. Stomata tetracytic (sometimes with six subsidiary cells, sometimes paracytic?), with oblique cell divisions. Cuticular wax crystalloids usually absent (sometimes as aggregated rodlets, often longitudinally orientated, Strelitzia type, or as scales). Epidermis with silica bodies (one per cell, imbedded in inner periclinal cell walls). Mesophyll with water-storing tissue and mucilaginous idioblasts containing calciumoxalate raphides. Envelopes of vascular bundles fibrous. Hypodermal cells often with transparent yellowish tannine drips. Leaf margin serrate (often spinose-dentate) or entire.

Inflorescence Usually terminal, spike or raceme (rarely head), often in panicle or thyrse (flowers in Tillandsia solitary), often with showy colourful bracts (inflorescence in Deuterocohnia lignified, with cambium-like meristem and anthesis persisting for several years). Extrafloral nectaries sometimes present.

Flowers Usually actinomorphic (in Pitcairnia and Tillandsia somewhat zygomorphic). Hypogyny (probably secondary) to epigyny. Tepals 3+3, free or connate; outer tepals usually with contorted (sometimes imbricate, rarely cochleate) aestivation, sepaloid or petaloid, persistent; inner tepals with imbricate or contorted aestivation, petaloid, often connivent into a tube, often considerably longer than outer tepals, often with one or two basal adaxial scale-like outgrowths (nectaries?) and/or longitudinal bulges. Septal nectaries present. Disc absent.

Androecium Stamens 3+3. Filaments flattened, more or less fleshy or plicate, usually free (sometimes connate at base), usually free from tepals (sometimes adnate to inner tepals). Anthers basifixed or dorsifixed, usually non-versatile?, tetrasporangiate, introrse to latrorse, longicidal (dehiscing by longitudinal slits); connective in Androlepis prolonged. Tapetum secretory, with uni- to multinucleate cells, with ab initio secretory cells subsequently invading. Female flowers with staminodia.

Pollen grains Microsporogenesis successive. Pollen grains monosulcate, diporate to tetraporate or pantotreme (in Aechmea, Canistrum and Guzmania inaperturate), usually shed as monads (in Androlepis, Hohenbergiopsis and some Orthophytum as tetrads), usually bicellular (rarely tricellular) at dispersal. Pollen grains sometimes without starch. Exine semitectate, with columellate infratectum, usually reticulate (sometimes scrobiculate or pertectate).

Gynoecium Pistil composed of three connate carpels; median carpel abaxial. Ovary superior (although initiated inferior) to inferior, trilocular. Style simple, long, subulate, straight or curved, trifid at apex (absent in some genera). Stigma usually trilobate (in Aechmea and Werauhia entire), with straight or spirally twisted, sometimes foliaceous, branched or irregularly folded lobes, sometimes cupular, often papillate, Dry or Wet type. Male flowers with pistillodium; sterile flowers sometimes with pistillodium.

Ovules Placentation axile. Ovules two to more than 50 per carpel, usually anatropous (rarely campylotropous or orthotropous), bitegmic, crassinucellar, often with a chalazal appendage (rarely several). Micropyle endostomal. Outer integument up to seven cell layers thick. Inner integument two cell layers thick. Megasporangium one cell layer thick at micropyle; epidermal cells elongate. Parietal cell formed from archesporial cell. Nucellar cap sometimes two cell layers thick. Megagametophyte monosporous, Polygonum type. Antipodal cells sometimes proliferating. Endosperm development ab initio helobial; cell wall formation in small chalazal space preceding wall formation in large micropylar spaces. Endosperm haustoria absent? Embryogenesis asterad.

Fruit A usually septicidal (sometimes septicidal-loculicidal, rarely loculicidal) capsule or a berry with persistent outer tepals (berries in some species fused with each other and with inflorescence axis into a syncarp).

Seeds Aril absent. Seed sometimes winged or plumose (hair tufts in Tillandsioideae formed by splitting of testa). Operculum sometimes? present. Seed coat testal-tegmic. Testa dry or developing into a sarcotesta. Exotegmen sometimes thickened. Endotegmen tanniniferous. Perisperm not developed. Endosperm copious, starchy, periferal cells with aleurone and lipids; starch grains simple or compound. Embryo usually small, straight, often lateral, well differentiated, without chlorophyll, Trillium type. Cotyledon one. Cotyledon hyperphyll elongate and assimilating, or compact and not assimilating. Hypocotyl internode present or absent. Coleoptile absent. Radicula absent in Tillandsioideae. Germination phanerocotylar? or cryptocotylar.

Cytology n = 16, 17, 21, 24, 25, 27, 32, 36, 48 – Chromosomes usually not more than 2.75 μm long.

Phytochemistry Flavonols (kaempferol, quercetin), C-glycosylated/6-oxygenated flavones, derivatives (partially O-methylated) of 6-hydroxyluteolin and 6-hydroxymyricetin, cyaniding?, triterpenes and steroidal saponins present. Ferulic, diferulic and p-coumaric acids (esterified) components of non-lignified cellwalls. Ellagic acid and alkaloids not found. Ananas comosus contains bromelain (a mixture of at least five different proteolytic enzymes); fruit scent largely due to undecatriene and undecatetraene.

Use Ornamental plants, fruits (Ananas comosus, Aechmea, Bromelia, Greigia), fibres (Ananas lucidus, Aechmea magdalenae, Neoglaziovia variegata).

de Jussieu, Gen. Plant.: 26. 4 Aug 1789 [’Cyperoideae’, ’Cyperoïdeae’], nom. cons.

Scirpaceae Batsch ex Borkh., Bot. Wörterb. 2: 340. 1797 [’Scirpeae’]; Scleriaceae Bercht. et J. Presl, Přir. Rostlin: 263. Jan-Apr 1820; Caricaceae Bercht. et J. Presl, Přir. Rostlin: 263. Jan-Apr 1820 [’Caricinae’], nom. illeg. – non Caricaceae Dumort. 1829; Papyraceae Burnett, Outl. Bot.: 761, 1129. Feb 1835; Caricineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1691. 1846; Cyperineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1691, 1702. 1846; Scirpineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1691, 1698. 1846; Kobresiaceae Gilly in Iowa State Coll. J. Sci. 26: 210. Jan 1952; Mapaniaceae Shipunov in Žurn. Obshchei Biol. 64: 505. Dec 2003

Distribution Cosmopolitan, with their largest diversity in temperate and alpine regions.

Fossils Fossil pollen and infructescence of Volkeria messelensis from the mid-Eocene of Germany were assigned to Mapanioideae. Fossil fruits (Caricoidea, Cladiocarya, Polycarpella, Scleriocarya, etc. and also several extant genera) of Cyperaceae (Cyperoideae) have been recorded from the mid-Paleocene onwards in, above all, Europe, Asia and North America, and pollen grains are relatively frequent in Cenozoic layers.

Habit Usually bisexual or monoecious (rarely andromonoecious, gynomonoecious, dioecious, androdioecious or gynodioecious; unisexual flowers sometimes in bisexual pseudanthia), usually perennial (sometimes annual) herbs (rarely shrubs, lianas or epiphytes; Microdracoides consists of small trees). Graminids. Some species have bulb-like or tuberous swollen internodes or stem bases. A few species possess stilt roots. Rhynchospora anomala xeromorphic, with adventitious roots (with well developed velamen) running down along envelope formed by persistent leaf bases. Many species are helophytes (sometimes aquatic). Culm usually medullated (rarely hollow or septate) and terete to sharply triangular in cross-section (rarely flattened, winged or quadrangular to sexangular in cross-section).

Vegetative anatomy Mycorrhiza usually absent (vesicular-arbuscular mycorrhiza present in some species). Dauciform roots (with dense long root hairs, epidermal cells elongate at right angles to long axis of root) sometimes present. Lateral roots developing from zone opposite protophloem or opposite protoxylem poles. Phellogen absent. Secondary lateral growth absent. Vessels present in roots, stem and leaves. Vessel elements with scalariform and/or simple perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals, or P2cf, with cuneate protein crystals and peripheral protein filaments. Nodes? Many species possessing C₄ photosynthesis with a green envelope of Kranz’ cells surrounding vascular bundles (C₄ photosynthesis and Kranz’ anatomy evolved perhaps six times; Kranz’ anatomy present in at least four types: Chlorocyperus, Fimbristylis, Rhynchospora, and Eleocharis types; also reversals to C₃ photosynthesis in, e.g., many species of Cyperus), with or without an envelope of parenchyma cells. Chlorenchyma with lobed cells (peg cells?) present. Idioblasts with frequent tannins and polyphenols. Silica bodies conical, inserted on cell walls, usually inner periclinal epidermal cell walls near veins. Calciumoxalate raphides absent.

Trichomes Hairs usually unicellular (sometimes multicellular, uniseriate) or absent; often with unicellular ‘prickle hairs’, with sharp-pointed apex and usually swollen base.

Leaves Alternate (usually tristichous, sometimes distichous, tetrastichous or spiral), simple, entire, usually linear (rarely ensiform or terete), often reduced to a mere leaf sheath, with conduplicate, plicate, revolute, convolute (supervolute) or involute ptyxis. Stipules absent; leaf sheath usually closed (in Coleochloa and Oreobolus open), often with distal ligule (rarely with distal contraligule). Venation parallelodromous. Stomata usually paracytic (sometimes tetracytic), with dumbbell-shaped guard cells. Cuticular wax crystalloids as aggregated rodlets (Strelitzia type) or non-orientated. Air canals present. Epidermis usually with silica bodies. Mesophyll without mucilaginous idioblasts and calciumoxalate raphides. Leaf margin usually entire (sometimes finely serrate-dentate).

Inflorescence Terminal (sometimes pseudolateral), usually compound (sometimes simple) panicle, corymb, spike-like, anthela and/or head-like, consisting of axillary one- or many-flowered spikelets (sometimes a solitary spikelet), each in axil of often foliaceous bract and with one or more bracts without flowers; bracts with or without sheath; floral bracts usually small and scale-like, persistent or caducous; lowermost spikelet scale often prophyll, usually without flower. Spikelet usually monopodial (indeterminate; sometimes sympodial?). Central axis of spikelet, rhachilla, persistent or caducous (rarely fragmented into one-flowered units); rhachilla internodes short or long, sometimes winged or curved around fruit/fruits. Female spikelet in Carex reduced to pistil and rhachilla, enclosed by urceolate utriculus, perigynium, formed by basal prophyll of one-flowered female spikelet.

Flowers Actinomorphic (zygomorphic through reduction), small. Hypogyny. Tepals usually (one to) three (or four) or 3+3 (sometimes more than six), usually hair-, bristle- or scale-like, usually free (sometimes connate), or absent. Nectary absent. Disc usually absent.

Androecium Stamens (one to) three (to six; in Chrysithrix numerous [in pseudanthium?]), at least sometimes antesepalous. Filaments usually free (rarely more or less connate), usually free from tepals. Anthers basifixed, versatile, tetrasporangiate, introrse or latrorse, longicidal (dehiscing by longitudinal slits); connective often comb-shaped and prolonged. Tapetum secretory, with binucleate to multinucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous (Cyperaceae variation). Pollen grains graminoid, usually monoulcerate or monoporate (in Carex and other Cyperaceae with pseudomonads five or six circular or elongate poroids present: one distal and additional poroids equatorial), shed as pseudomonads (cryptotetrads), three microspores in each tetrahedral tetrad degenerating and incorporated into wall of functioning fourth pollen grain, usually tricellular (sometimes bicellular) at dispersal. Exine tectate, with columellate infratectum, smooth; exine formed from microsporocyte wall. Intine in Carex sometimes thin below aperture, thick in interapertural regions.

Gynoecium Pistil composed of two or three (or four) connate and at least sometimes antesepalous carpels (median carpel in Carex adaxial, inverted); gynoecium developing from annular primordium. Ovary superior, unilocular (sometimes on gynophore). Stylodia two or three (rarely four), simple or branched, free or connate in lower part; stylar base often enlarged and persistent. Stigmas two or three (to nine), filiform, adaxially decurrent, papillate or non-papillate, Dry type. Pistillodium absent.

Ovules Placentation basal. Ovule one per ovary, anatropous, bitegmic, crassinucellar. Micropyle usually endostomal (at least in Hypolytrum often bistomal, Z-shaped). Outer integument two? cell layers thick. Inner integument two? cell layers thick. Funicular obturator present near micropyle often present. Hypostase present or absent. Parietal cell formed from archesporial cell. Parietal tissue two to four cell layers thick. Megasporangium rarely one cell layer thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria chalazal and micropylar, or absent. Embryogenesis onagrad (Juncus variation).

Fruit Usually a nutlet (achene), often with persistent tepals as bristles, hairs or scales (achene in Carex surrounded by utriculus/perigynium at dispersal; in Cladium, Mapania and Scirpodendron a drupe).

Seeds Aril absent. Seed coat testal-tegmic. Testa thin, not fused with pericarp. Exotesta with silica bodies. Remaining testal layers fibrous. Tegmen thin and fibrous. Perisperm not developed. Endosperm copious, with starch and oils (aleurone?); starch grains simple. Embryo small, capitate, well or little differentiated, without chlorophyll, Xyris, Carex, Schoenus or Scirpus types. Cotyledon one or indistinct, not photosynthesizing. Cotyledon hypophyll elongate, assimilating. Hypocotyl internode absent. Mesocotyl sometimes present. Coleoptile with chlorophyll. Plumule? Collar very small, with rhizoids. Germination cryptocotylar?

Cytology n = 5–>60 (chromosome numbers in Carex extremely variable). – Centromere in many species diffuse (centric activity present over more or less the entire chromosomes; not limited to a specific site; spindle fibrils attached at several sites along chromosomes), easily leading to aberrant chromosome numbers by chromosome fragmentation, agmatoploidy, detached chromosome segments moving against poles and remaining functional. Following meiosis three nuclei move towards one pole and degenerate; finally, they become embedded in the wall of the surviving microspore.

DNA Deletion encompassing three base pairs in plastid gene atpA. Plastid gene rpl23 lost. Plastid gene infA lost/defunct (Cyperus). One three base pairs insertion present in nuclear 5.8S rDNA. Mitochondrial gene rps14 transferred to nucleus (pseudogene ψrps14 present in mitochondrial genome).

Phytochemistry Flavonols (quercetin), flavone-C-glycosides, flavone sulphates, aurones, flavanones, cyanidin, tricin (tricetin 3’,5’-dimethyl ether; frequent), 6-hydroxyluteolin, luteolin-5’-methyl ether, chalcones, simple indole alkaloids, and daphnetin present. Carbohydrates stored as kestose and isokestose oligosaccharides (fructans). Ferulic and p-coumaric acids (esterified) components of non-lignified cell walls. Ellagic acid not found. Aluminium accumulation occurring in some species.

Use Ornamental plants, vegetables (Cyperus esculentus, Eleocharis dulcis), paper (Cyperus papyrus), thatching (Cladium mariscus etc.), rafts, canoes, carpets, textiles, basketry, bioenergy, forage plants, etc.

Habit Monoecious, perennial herbs. Graminids. Xeromorphic. Culm terete, branched, furrowed, with swollen nodes and solid (medullated?) internodes, photosynthesizing.

Vegetative anatomy Culm ridges sclerenchymatous (sclerenchyma ridges in Georgeantha extending through chlorenchyma to subepidermal sclerenchymatous layer; not in Ecdeiocolea). Epidermis in Ecdeiocolea with rows of alternately long and short cells. Stomata paracytic, in longitudinal furrows along culm. Phellogen absent. Chlorenchyma with peg cells. Sclerenchyma not forming a continuous cylinder. Vascular bundles bicollateral, each one surrounded by sclerenchymatous envelope. Secondary lateral growth absent. Vessels present in roots, rhizome and culm. Vessel elements with scalariform or simple perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Secretory cavities absent. Culm chlorenchyma with cuboid silica bodies as sand. Calciumoxalate crystals absent? (raphides absent). Special short idioblasts with large single silica bodies (phytoliths) absent.

Trichomes Hairs absent (Ecdeiocolea), or multicellular, uniseriate or branched (rhizome, culm bases and spiklets in Georgeantha); microhairs?

Leaves Alternate (distichous), on young shoots simple, entire, linear, on older shoots reduced and only consisting of leaf sheath, with convolute (supervolute) ptyxis. Stipules absent; leaf sheath closed, auriculate, in Georgeantha caducous; ligule absent. Venation parallelodromous (also with transverse veins?). Stomata paracytic, with dumbbell-shaped Poaceae type guard cells. Cuticular waxes? Mesophyll with fusoid cells? Secretory cavities absent. Mesophyll without mucilaginous idioblasts or calciumoxalate raphides. Epidermis without rows of long and short cells. Silica absent from leaf epidermis. Leaf margin entire.

Inflorescence Terminal, cymose, usually branched spike-like head, consisting of one (Ecdeiocolea) or two or three (Georgeantha) racemose spikelets with male flowers alternating with female flowers; inflorescence branches not in axils of spatha-like bracts; basal spikelet bracts, glumes, usually with axillary male or female flowers. Each flower in axil of a bract, lemma. Floral prophylls (bracteoles) absent.

Flowers Zygomorphic (due to reduction), dorsiventrally flattened, small. Hypogyny. Tepals 2+2 (Ecdeiocolea) or 3+3 (Georgeantha), bract-like (sepaloid, glumaceous); two adaxial outer tepals with conduplicate aestivation, keeled, laterally compressed, ciliated at apex, abaxial outer tepal (in Georgeantha) and inner tepals flattened, free, surrounded by a stout scale-like bract. Nectary absent. Disc absent.

Androecium Stamens 2+2 (Ecdeiocolea, possibly corresponding to outer staminal whorl and adaxial stamen of inner staminal whorl) or 3+3 (Georgeantha). Filaments filiform, free from each other and from tepals. Anthers basifixed, versatile, tetrasporangiate, latrorse-introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Endothecial thicknesses girdle type. Female flowers with rudimentary staminodia.

Pollen grains Microsporogenes successive. Pollen grains graminoid, monoporate to monoulcerate, operculate (with a plug, Poaceae type), shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, smooth (not scrobiculate).

Gynoecium Pistil composed of two (Ecdeiocolea) or three (Georgeantha) connate carpels. Ovary superior, unilocular (Ecdeiocolea) or trilocular (Georgeantha). Stylodia two (Ecdeiocolea) or three (Georgeantha), free, covered by adaxial stigmatic surfaces. Stigmas plumose, with papillate hairs, Dry type. Male flowers with rudimentary pistillodium.

Ovules Placentation apical-axile. Ovule one per carpel, orthotropous, pendulous, bitegmic, tenuinucellar. Micropyle bistomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Nucellar cap? Megagametophyte tetrasporous, 16-celled, similar to Drusa type (Ecdeiocolea), or monosporous, Polygonum type (Ecdeiocolea?). Antipodal cells absent. Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A nutlet (achene; Ecdeiocolea) or a one- or two-seeded loculicidal capsule (Georgeantha).

Seeds Aril absent. Exotesta with large cells, in Georgeantha with convex and strongly sinuate walls; thickness very different in Ecdeiocolea. Endotesta? Tegmen? Perisperm not developed. Endosperm copious, starchy? Embryo?, chlorophyll? Cotyledon one, not photosynthesizing. Cotyledon hyperphyll? Hypocotyl internode? Mesocotyl? Coleoptile? Plumule? Collar hairs? Germination?

DNA The plastid genome has an inversion of 28 kb and an inversion of 6,4 kb (Michelangeli & al. 2003). Inversion absent from the plastid gene trnT.

Phytochemistry Flavonols (isorhamnetin, in Georgeantha also quercetin) present. Galactose present in Georgeantha.

Eriocaulales Nakai, Hisi-Shokubutsu: 49. 1930; Eriocaulineae Thorne et reveal in Bot. Rev. (Lancaster) 73: 85. 29 Jun 2007

Distribution Tropical and subtropical regions on the Southern and Northern Hemispheres, with their largest diversity in the Guayana Highlands and southeastern Brazil; a few species of Eriocaulon in temperate parts of Europe, East Asia and North America; Mesanthemum: tropical Africa, Madagascar.

Fossils Uncertain. Subfossil pollen grains of Eriocaulon septangulare (E. pellucidum) are known from Pleistocene layers in Ireland and Canada.

Habit Usually monoecious (rarely dioecious or bisexual), perennial or annual herbs. Many species are helophytes and some are aquatic.

Vegetative anatomy Mycorrhiza? Outer part of root cortex in Eriocaulon with specialized aerenchyma formed by alternating transverse layers of stellate parenchyma cells and radiate rows of longitudinally widened and flattened cells without contact with adjacent lateral cells. Root phloem and xylem elements often intermixed in stele. Lateral roots developing from a zone opposite protophloem or opposite protoxylem poles. Phellogen absent. Stem endodermal cell walls often U- or O-shaped in cross-section. Stem vascular bundles alternately on outside and inside. Secondary lateral growth usually absent (cambium occasionally present, producing secondary tissue with separate vascular bundles). Vessels usually present in roots and stem. Vessel elements with scalariform or simple (sometimes reticulate) perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Silica bodies absent or nearly so. Calciumoxalate crystals of various types (raphides absent).

Trichomes Hairs multicellular, uniseriate, with apical cell unbranched, or T-shaped malpighiaceous hairs; hairs on vegetative parts with foot cell and bulb-like persistent usually dark basal cell; often also glandular hairs.

Leaves Alternate (usually spiral, rarely distichous), simple, entire, linear or subulate, with convolute (supervolute) ptyxis? Stipules absent; leaf sheath indistinct. Vascular bundle envelope with large cells without chloroplasts. Palisade tissue absent. Venation parallelodromous. Stomata usually paracytic, with Poaceae type guard cells. Cuticular wax crystalloids as aggregated rodlets. Epidermis without silica bodies; subepidermal cells in Paepalanthus with silica bodies. Mesophyll without mucilaginous idioblasts. Mesophyll with calciumoxalate as druses, styloids or single prismatic crystals (raphides absent). Leaf margin entire.

Inflorescence Terminal, dense head-like spike, or compound head- or umbel-like capitulum consisting of up to 1.000 or more partial inflorescences. Pseudanthium surrounded by scale-like outer bracts and with ten to more than 1.000 flowers in axils of often petaloid separate inner bracts (absent in Syngonanthus). Peduncle (scape) narrow, spirally twisted, often with ridges or edges and ebracteate, although usually with a closed sheathing basal bract; peduncle with central cylinder consisting of two cylinders of concentric or biconcentric vascular bundles separated by a sclerenchymatous envelope.

Flowers Actinomorphic or zygomorphic, very small. Floral receptacle flat or slightly convex. Hypogyny. Tepals 2+2 or 3+3, with usually open (sometimes valvate) aestivation, persistent; outer tepals dry, membranous, sepaloid, usually free (sometimes connate at base; rarely connate into a tube or spathe-like); inner tepals petaloid, membranous, caducous, usually more or less connate (in, e.g., Eriocaulon free), sometimes absent. Tepal nectaries in Eriocaulon inserted on edges or adaxial side of perianth tube. Septal nectaries absent. Disc absent.

Androecium Stamens in Paepalanthoideae two or three (outer staminal whorl absent), antepetalous; stamens in Eriocaulon and Mesanthemum usually 2+2 or 3+3 (in Syngonanthus amazonicus a single stamen). Stamens sometimes inserted on androphore. Filaments free; inner filaments in Eriocaulon and Mesanthemum adnate to inner tepals (epitepalous). Anthers usually dorsifixed (in Leiothrix basifixed), non-versatile?, usually tetrasporangiate (in many species of Paepalanthus disporangiate due to fusion of microsporangia), introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, usually with uninucleate (rarely binucleate) cells. Female flowers often with (antesepalous) inconspicuous (often scale-like) staminodia.

Pollen grains Microsporogenesis usually successive (rarely simultaneous). Pollen grains usually spiraperturate (one aperture spirally winding over entire pollen grain; sometimes inaperturate), shed as monads, bicellular or tricellular at dispersal. Exine tectate, with columellate? infratectum, echinulate to echinate, often spinulate.

Gynoecium Pistil composed of two or three connate carpels. Ovary superior, bilocular or trilocular. Stylodia two or three, free or connate at base, often with stylar appendages. Stigmas two or three, carinal/dorsal (Eriocaulon, Mesanthemum) or commissural and non-vascularized (with stylodia-like nectariferous appendages between stigmas, in Syngonanthus present on same place as vascularized style in Eriocaulon), simple or bifid, non-papillate, Dry type. Male flowers often with pistillodium.

Ovules Placentation axile-apical. Ovule one per carpel, orthotropous, pendulous, bitegmic, tenuinucellar. Micropyle usually endostomal (in Syngonanthus bistomal). Outer integument two cell layers thick. Inner integument two cell layers thick. Hypostase present. Obturator absent. Parietal cell not formed (parietal tissue absent). Megagametophyte monosporous, Polygonum type. Unique antipodal cyst, developed by fusion of antipodal cells, present in megagametophyte. Endosperm development ab initio nuclear, later cellular (or helobial?). Endosperm haustoria little developed or absent. Embryogenesis asterad.

Seeds Aril absent. Seed coat endotestal or tegmic. Exotesta membranous, usually entirely degenerating or persisting as rows of hooks on seed coat. Endotesta consisting of hexagonal brownish red to yellow cells with prominent anticlinal walls forming a striate pattern. Operculum formed from endotegmen. Exotegmen? Endotegmic cells tanniniferous. Perisperm not developed. Endosperm copious, starchy; starch grains usually compound. Embryo small, lens- to bell-shaped, rudimentary (lacks organs but shows differentiation of dermatogen), enclosed by endosperm at micropylar end, Xyris-Scirpus type, chlorophyll? Cotyledon one, not photosynthesizing. Cotyledon hyperphyll compact, not assimilating. Hypocotyl internode absent. Coleoptile absent. Radicula rudimentary or absent. Collar hairs? Germination?

DNA Significant transfers of ribosomal protein genes and succinate dehydrogenase genes have taken place from the mitochondrial genome (to the nuclear genome?) at least in one species of Paepalanthus (‘Lachnocaulon anceps’). ORF 2280 sometimes present.

Phytochemistry Flavonols (kaempferol, myricetin), 6-hydroxyflavonoids quercetagetin and patuletin (6-methyl ether of quercetagetin) present. Ellagic acid, proanthocyanidins, alkaloids, and cyanogenic or phenolic compounds not found. Ferulic, diferulic and p-coumaric acids (esterified) components of non-lignified cell walls.

Flagellariales (Meisn.) Takht. ex Reveal et Doweld in Novon 9: 550. 30 Dec 1999

Distribution Tropical and southern Africa, Madagascar, Sri Lanka and southern India, Southeast Asia to northern Australia and islands in the southwestern Pacific.

Fossils Uncertain. Pollen grains similar to Flagellaria have been found in Miocene layers on Borneo.

Habit Usually bisexual (sometimes unisexual), perennial herbs. Stout, twining and climbing graminids. Dichotomously branched from distinctly sympodial rhizomes.

Vegetative anatomy Lateral roots developing from a zone opposite protoxylem poles. Phellogen absent. Culm solid, with extensive medulla in internodes and compact swollen nodes, without secretory canals. Vascular bundles embedded in sclerotic tissue between cortex and central cylinder of aerial stems. Endodermal cells radially elongate. Chlorenchyma lacking peg cells. Secondary lateral growth absent. Vessels present in roots, stem and leaves. Vessel elements with scalariform or simple perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c_cl type, with cuneate and several additional loosely packed protein crystals. Nodes? Arm cells and fusoid cells absent. Silica bodies spherical, present only in association with vascular strands. Cells with calciumoxalate crystals present.

Leaves Alternate (distichous), simple, entire, linear, with convolute (supervolute) ptyxis below and circinate ptyxis at apex. Stipules absent; leaf sheath usually closed (rarely open), auriculate, with one pair of distal lateral lobes (ligule?). Venation parallelodromous (also with transverse veins). Stomata paracytic, with Poaceae type guard cells; guard cells not reniform or dumbbell-shaped and without perforations between adjacent guard cells; adjacent cells with oblique divisions. Cuticular wax crystalloids as non-orientated platelets (similar to those in Restionaceae), only adjacent to stomata. Epidermal cells without silica bodies and without phytoliths. Mesophyll with secretory canals and fibrous cells with granular silica bodies below and above vascular strands. Mesophyll cells with or without calciumoxalate crystals. Leaf margin entire. Leaf apex developing into tendril when in contact with support, due to presence of sensitive adaxial cells. Extrafloral nectaries present or absent.

Inflorescence Terminal, compound panicle, with spike- or raceme-like partial inflorescences terminating branches. Lateral branches often with distinct adaxial swellings at base, and with first two bracts usually inserted transversely at almost same level. Floral prophylls (bracteoles) absent.

Flowers Actinomorphic, small. Hypogyny. Tepals 3+3, pseudouniseriate, sepaloid, membranous, persistent, connate at base. Nectary absent. Disc absent.

Androecium Stamens usually 3+3 (sometimes fewer, with fertile stamens replaced by staminodia?). Filaments free from each other and from tepals. Anthers basifixed, versatile, usually tetrasporangiate (rarely tri-, hexa- or heptasporangiate), latrorse or introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Endothecial thickenings with complete basal plate. Female flowers with staminodia.

Pollen grains Microsporogenesis successive. Pollen grains monoporate to monoulcerate, with or without rudimentary operculum, shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, microperforate, verrucate to scrobiculate.

Gynoecium Pistil composed of usually three (sometimes two) connate carpels. Ovary superior, trilocular. Stylodia three, free or somewhat connate at base. Stigmas three, adaxially papillate (papillae multicellular), Dry type. Male flowers with pistillodium.

Ovules Placentation apical-axile. Ovule one per carpel, almost orthotropous, pendulous, bitegmic, crassinucellar. Micropyle endostomal; endostoma formed by fast growth of inner integument. Outer integument apporox. four cell layers thick. Inner integument two? cell layers thick. Parietal cell formed from non-dividing archesporial cell. Parietal tissue one cell layer thick. Epidermal cells of megasporangium dividing periclinally. Megagametophyte at least sometimes disporous, 8-nucleate, Allium type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad or asterad.

Seeds Aril absent. Seed coat consisting of exotesta fused with endocarp. Operculum present. Exotesta with persistent outer periclinal wall. Endotesta? Tegmen? Perisperm not developed. Endosperm copious, starchy; starch grains simple or compound. Embryo small, lens-shaped, little differentiated, Xyris-Scirpus type, chlorophyll? Cotyledon one, not photosynthesizing. Cotyledon hyperphyll? Hypocotyl internode? Mesocotyl? Coleoptile absent or minute. Plumule? Radicula with collar hairs. First leaf absent. Germination?

DNA The 6,4 kb and 28 kb inversions are absent from the plastid genome. ORF2280?

Phytochemistry Flavonols (kaempferol), alkaloids, cyanogenic glycosides (triglochinin), saponins, and -sitosterol present. Ellagic acid and proanthocyanidins not found. Ferulic, diferulic and p-coumaric acids (esterified) components of non-lignified cell walls.

Distribution West Malesia, Melanesia on islands in the Pacific to Samoa and the Hawaiian Islands (probably absent from East Malesia and Australia).

Fossils Uncertain. Ambiguous fossils from New Zealand have sometimes been assigned to Joinvillea.

Habit Bisexual, perennial herbs. Graminids. Culm simple, usually terete, with hollow internodes and compact swollen nodes.

Vegetative anatomy Phellogen absent. Secondary lateral growth absent. Vessels present in roots, stem and leaves. Vessel elements with scalariform or simple (in leaves sometimes also reticulate) perforation plates; lateral pits? small, numerous. Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c_cl type, with cuneate and several additional loosely packed protein crystals. Nodes? Chlorenchyma without peg cells. Arm cells probably absent. Fusoid cells present. Silica bodies cuboid, especially frequent in epidermal cell walls and in association with vascular strands. Special short idioblasts with large single silica bodies (phytoliths) present.

Trichomes Hairs unicellular or multicellular, branched or uniseriate, often pointed (‘prickle hairs’); multicellular microhairs often present.

Leaves Alternate (distichous), simple, entire, linear, with plicate ptyxis. Stipules absent; leaf sheath open, auriculate, with distal ligule. Venation parallelodromous to palmate-parallelodromous; veins anastomosing; main veins connected by transverse veins. Stomata paracytic, with dumbbell-shaped Poaceae type guard cells. Cuticular wax crystalloids absent. Mesophyll with fusoid cells? Epidermis with rows of long cells alternating with short cells containing silica bodies and cuboidal unlobed phytoliths. Mesophyll without mucilaginous idioblasts and without calciumoxalate raphides. Mesophyll cells with calciumoxalate crystals, tannins and silica. Foliar epidermis with microhairs? Leaf margin finely serrate.

Inflorescence Terminal, compound panicle with spike- or raceme-like partial inflorescences. Lateral branches with distinct adaxial swellings at base. Prophylls caducous.

Flowers Actinomorphic, small. Hypogyny. Tepals 3+3, dry, bract-like, sepaloid (outer tepals cucullate), persistent, free or connate at base. Nectary absent. Disc absent.

Androecium Stamens 3+3. Filaments free, sometimes adnate at base to tepals. Anthers subbasifixed, versatile, tetrasporangiate, latrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Endothecial thickenings girdle type. Staminodia absent.

Pollen grains Microsporogenesis successive. Pollen grains graminoid, monoporate to monoulcerate, with or without rudimentary operculum, shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, microperforate, scrobiculate.

Gynoecium Pistil composed of three connate carpels. Ovary superior, trilocular. Stylodia three, free or somewhat connate at base, sometimes persistent. Stigmas three, with plumose lobes, and with scattered detached papillate surfaces on multiseriate branches, Dry? type. Pistillodium absent.

Ovules Placentation apical-axile. Ovule one per carpel, orthotropous, pendulous, bitegmic, tenuinucellar? Micropyle bistomal. Outer integument two cell layers thick. Inner integument two cell layers thick. Parietal cell not formed (parietal tissue absent). Nucellar cap? Megagametophyte disporous, Allium? type. Antipodal cells three, binucleate. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis asterad?

Fruit A one- to three-seeded drupe with persistent tepals (and sometimes stylar branches).

Seeds Aril absent. Seed coat tegmic. Testa thin. Exotegmen? Endotegmen tanniniferous. Perisperm not developed. Endosperm copious, starchy; starch grains compound. Embryo small, disc-shaped, little differentiated to rudimentary, Xyris-Scirpus type, chlorophyll? Cotyledon one, not photosynthesizing. Cotyledon hyperphyll? Hypocotyl internode? Mesocotyl? Coleoptile? Plumule? Collar hairs? Germination? First seedling leaf only as leaf sheath.

DNA The plastid genome has an inversion of 28 kb and one inversion of 6,4 kb. The mitochondrial gene rps14 is transferred to the nuclear genome, with pseudogene ψrps14 in mtDNA. The trnT inversion is absent.

Phytochemistry Very insufficiently known. Ferulic, diferulic and p-coumaric acids (esterified) components of non-lignified cell walls.

Juncales Bercht. et J. Presl, Přir. Rostlin: 266. Jan-Apr 1820 [‘Junceae’]; Juncopsida Bartl., Ord. Nat. Plant.: 23, 34. Sep 1830 [’Juncinae’]; Juncanae Takht., Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 510. 4 Feb 1967; Juncidae Doweld, Tent. Syst. Plant. Vasc.: lxii. 23 Dec 2001

Distribution Cosmopolitan, with their highest diversity in cold and temperate regions in the Northern and Southern Hemispheres.

Fossils Fossil seeds have been found in Late Eocene to Early Oligocene layers in England and from the Miocene onwards in several places in Europe. Pollen grains, at least of Juncus and Luzula, are known from the Cenozoic of Russia, Germany and North America.

Habit Usually bisexual (rarely monoecious, dioecious or gynodioecious), usually perennial (sometimes annual) herbs. Graminids. Many species are aquatic or hygrophytic. Culm terete or flattened in cross-section, smooth or with longitudinal ridges and furrows.

Vegetative anatomy Mycorrhiza at least usually absent. Dauciform roots (with dense long root hairs, epidermal cells elongate at right angles to long axis of root) sometimes present. Root hairs sometimes developing from short cells in root epidermis. Root endodermis one cell layer thick, with cell wall thickenings U-shaped in cross-section. Lateral roots developing from a zone opposite protophloem or opposite protoxylem poles. Central rhizome parenchyma with several scattered vascular bundles. Culm with one or few peripheral concentric cylinders of vascular bundles, without endodermis. Phellogen absent. Medulla hollow or consisting of aerenchyma with stellate cells. Secondary lateral growth absent. Vessels present in roots, stem and leaves. Vessel elements in Juncus and Luzula with scalariform and/or simple (in remaining genera usually entirely scalariform) perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Silica bodies usually absent (in Juncus rarely as silica sand). Calciumoxalate raphides absent.

Trichomes Hairs usually absent (multicellular, uniseriate hairs present on leaf margins in Luzula).

Leaves Alternate (usually tristichous, rarely distichous), simple, entire, linear, usually unifacial (subulate or filiform; rarely laterally flattened and equitant; in some species consisting of mere leaf sheath), often with convolute (supervolute) ptyxis. Stipules absent; leaf sheath usually open (in Luzula closed), often auriculate (sometimes with ligule formed by fusion of auriculae). Venation parallelodromous. Stomata paracytic (in xeromorphic species Poaceae type paracytic). Cuticular waxes? Air canals present. Mesophyll without calciumoxalate raphides. Leaf margin usually entire (in, e.g., Juncus trifidus serrate).

Inflorescence Usually terminal or pseudolateral, racemose, panicle, corymb, spike- or head-like, in anthela, drepania or rhipidia (flowers in Marsippospermum and Rostkovia solitary terminal; in Distichia, Oxychloe and Patosia solitary lateral). Basal inflorescence bract in some species of Juncus elongate (appearing as prolongation of culm). Floral prophylls (bracteoles) one adaxial, two transverse or absent.

Flowers Actinomorphic, usually small (in Marsippospermum relatively large). Hypogyny. Tepals (2–)3+(2–)3, sepaloid, caducous or persistent, free. Nectary absent. Disc absent.

Androecium Stamens usually 3+3 (rarely 2+2, two or three [inner staminal whorl absent]), usually alternitepalous (in Luzula antetepalous). Filaments filiform or flattened, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse or latrorse, longicidal (dehiscing by longitudinal slits); connective sometimes slightly prolonged at apex. Tapetum secretory, with uninucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous (sometimes with ephemeral cell plate following meiosisI). Pollen grains monoulcerate with indistinct aperture, shed as tetrahedral tetrads with common exine, tricellular at dispersal. Exine tectate, thin, with columellate? infratectum, smooth.

Gynoecium Pistil composed of usually three (rarely two) connate antesepalous carpels; median carpel abaxial. Ovary superior, usually trilocular (sometimes unilocular and incompletely septate; rarely bilocular). Stylodia usually three (rarely two), free or connate below. Stigmas usually three (rarely two), adaxial, papillate (with unicellular papillae), Dry type, sometimes twisted. Pistillodium absent.

Ovules Placentation usually axile (when ovary unilocular then parietal placentation; in Luzula basal). Ovules one (Luzula) or three to c. 50 per carpel, anatropous, ascending, bitegmic, weakly crassinucellar. Micropyle endostomal or bistomal. Outer integument two (Juncus), or three or four (Distichia, Luzula) cell layers thick. Inner integument two cell layers thick. Funicular obturator (as hair) present. Hypostase present or absent. Parietal cell formed from archespore. Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus. Antipodal cells persistent. Endosperm development ab initio helobial. Endosperm haustoria? Embryogenesis onagrad (Juncus or Luzula variation).

Fruit A usually many-seeded (in Luzula three-seeded) loculicidal, usually trilocular capsule (in Distichia, Oxychloe and Patosia more irregularly dehiscing or sometimes a pyxidium).

Seeds Seeds in many species of Luzula with a chalazal (apical) or micropylar (basal) fleshy caruncular elaiosome. Seed coat in Juncus largely exotestal-endotegmic. Exotesta sometimes mucilaginous; in Luzula membranous. Endotesta? Exotegmen? Endotegmen in Luzula thickened. Perisperm not developed. Endosperm copious, starchy, outer layers with aleurone and oils. Suspensor few-celled, little developed. Embryo small, straight, without chlorophyll, enclosed by endosperm, Xyris-Scirpus type. Cotyledon one, usually photosynthesizing. Cotyledon hyperphyll elongate, assimilating. Hypocotyl rudimentary or absent. Hypocotyl internode present or absent. Coleoptile absent. Collar very small, with rhizoids. Phanomer (photosynthesizing unifacial cotyledon hyperphyll) usually present. Germination?

Cytology n = 9–65 (Juncus); n = 3–42 (Luzula); n = 8 (Oxychloe) – Centromeres at least in Luzula diffuse (centric activity principally over the entire chromosomes, not situated at a specific site; spindle fibrils attaching to several sites along the chromosomes), easily resulting in anomalous chromosome numbers by chromosome fragmentation, agmatoploidy; detached chromosome segments move against the poles and remain functional.

DNA A deletion of three base pairs present in the plastid gene atpA. The plastid gene rpl23 is absent.

Phytochemistry Flavonols (quercetin), flavones, flavone sulphates, luteolin-5’-methyl ether, luteolinidin based glycosides (replacing anthocyanins), cyanidin, tannins, juncosol (a phytotoxic phenol), 7,8-dihydroxy coumarin, daphnetin including its 8-methyl-ether, and tyrosine-derived cyanogenic compounds present. Flavone-C-glycosides, tricin, anthocyanins, ellagic acid, alkaloids, and saponins not found.

Use Ornamental plants, carpets, baskets, candle wicks (Juncus effusus etc.).

Kunth in Abh. Königl. Akad. Wiss. Berlin, Phys. Abh. 1840: 93. 1842 [’Mayaceae’], nom. cons.

Mayacales Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov]: 214. 20 Jul 1943

Distribution Central Africa, southeastern United States, the West Indies, Central and tropical South America.

Vegetative anatomy Root hairs developing from specialized rhizodermal cells. Lateral roots developing from a zone opposite protophloem poles. Phellogen absent. Cortex aerenchymatous, consisting of septate air canals. Central stem stele consisting of three to five collateral vascular bundles in a single cylinder separated by endodermis from cortex; vascular bundles outside endodermis from that layer. Secondary lateral growth absent. Vessels present in roots, stem and leaves. Vessel elements with scalariform (sometimes reticulate?) perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Uniseriate colleters present. Silica bodies absent. Calciumoxalate raphides absent.

Trichomes Hairs almost absent (present in leaf axils); uniseriate glandular hairs present in leaf axils.

Leaves Alternate (spiral), simple, entire, linear to filiform, with adplicate (flat) ptyxis. Stipules and leaf sheath absent. Leaf axils with caducous uniseriate glandular hairs (colleters?). Leaf one-veined. Stomata paracytic, with subsidiary cells with intersecting oblique cell divisions. Cuticular waxes? Mesophyll without mucilaginous idioblasts and calciumoxalate crystals. Leaf margin entire. Leaf apex usually bifid to bidentate.

Inflorescence Flowers axillary (appearing terminal), solitary, open above water surface. Flowers associated with a wide adaxial prophyll-like structure (bracteole?).

Flowers Actinomorphic, small. Hypogyny. Tepals 3+3, free; outer tepals with valvate or subvalvate aestivation, sepaloid; inner tepals with imbricate aestivation, petaloid, shortly clawed. Nectary absent. Disc absent.

Androecium Stamens three (inner staminal whorl absent), antesepalous. Filaments filiform, free from each other and from tepals. Anthers basifixed, non-versatile, usually tetrasporangiate (sometimes disporangiate, rarely trisporangiate), extrorse, poricidal (dehiscing by an apical pore, a short slit or a pore at apex of a tubular outgrowth). Tapetum secretory, with uninucleate cells. Staminodia absent.

Pollen grains Microsporogenesis successive. Pollen grains monosulcate, shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, finely reticulate, spinulate.

Gynoecium Pistil composed of three connate antepetalous carpels. Ovary superior, unilocular. Style single, simple, with stylar canal. Stigma short, capitate or somewhat trilobate, type? Pistillodium absent.

Ovules Placentation parietal. Ovules two to c. 30 per ovary, orthotropous, ascending or horizontal, bitegmic, tenuinucellar to weakly crassinucellar. Micropyle bistomal (endostomal?). Outer integument ? cell layers thick. Inner integument ? cell layers thick. Hypostase well developed, tanniniferous. Obturator present. Megasporangial epidermis basally thickened. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad.

Seeds Aril absent. Seeds operculate. Seed coat tegmic. Testa? Exotegmic cells with U-shaped lignifications. Operculum (somewhat similar to embryostega in Commelinaceae) mostly developed from endotegmen. Perisperm not developed. Endosperm copious, with starch and aleurone, and with a proteinaceous outer layer. Embryo small, undifferentiated, micropylar, Xyris-Scirpus type. Cotyledon one, consisting of a closed leaf sheath and an apical haustorium. Radicula absent. Germination?

Phytochemistry Very insufficiently known. Flavonols (quercetin) and phenolic compounds present. Ellagic acid, proanthocyanidins, and cyanogenic compounds not found.

Gramineae Juss., Gen. Plant.: 28. 4 Aug 1789, nom. cons. et nom. alt.; Aegilopaceae Martinov, Tekhno-Bot. Slovar: 221. 3 Aug 1820 [‘Egilopeae’, ‘Aegilopeae’]; Agrostidaceae Bercht. et J. Presl, Přir. Rostlin: 264. Jan-Apr 1820 [‘Agrostideae’]; Alopecuraceae Martinov, Tekhno-Bot. Slovar: 19. 3 Aug 1820 [‘Alopecurinae’]; Andropogonaceae Martinov, Tekhno-Bot. Slovar: 28. 3 Aug 1820 [‘Andropogones’]; Avenaceae Martinov, Tekhno-Bot. Slovar: 60. 3 Aug 1820; Bambusaceae Bercht. et J. Presl, Přir. Rostlin: 265. Jan-Apr 1820; Bromaceae Bercht. et J. Presl, Přir. Rostlin: 264. Jan-Apr 1820; Chloridaceae Bercht. et J. Presl, Přir. Rostlin: 265. Jan-Apr 1820 [‘Chlorideae’]; Hordeaceae Bercht. et J. Presl, Přir. Rostlin: 265. Jan-Apr 1820; Maydaceae Martinov, Tekhno-Bot Slovar: 387. 1820 [‘Mayseae’], nom. illeg.; Melicaceae Martinov, Tekhno-Bot. Slovar: 390. 3 Aug 1820 [‘Meliceae’]; Nardaceae Martinov, Tekhno-Bot. Slovar: 413. 3 Aug 1820 [‘Nardinae’]; Olyraceae Bercht. et J. Presl, Přir. Rostlin: 265. Jan-Apr 1820 [‘Olyreae’]; Oryzaceae Bercht. et J. Presl, Přir. Rostlin: 265. Jan-Apr 1820 [‘Oryzeae’]; Panicaceae Bercht. et J. Presl, Přir. Rostlin: 264. Jan-Apr 1820 [‘Paniceae’]; Saccharaceae Bercht. et J. Presl, Přir. Rostlin: 265. Jan-Apr 1820 [‘Saccharinae’]; Stipaceae Bercht. et J. Presl, Přir. Rostlin: 264. Jan-Apr 1820; Festucaceae Spreng. in J. Sadler, Fl. Comit. Pest. 1: 28. 27 Jun 1825; Anthoxanthaceae Link, Hort. Berol. 1: 271. 1 Oct-27 Nov 1827; Asperellaceae Link, Hort. Berol. 1: 106, 270. 1 Oct-27 Nov 1827 [’Asperellinae’]; Cenchrinaceae Link, Hort. Berol. 1: 9, 268. 1 Oct-27 Nov 1827 [‘Cenchrinae’]; Chaeturaceae Link, Hort. Berol. 1: 106, 269. 1 Oct-27 Nov 1827 [‘Chaeturinae’]; Chamagrostidaceae Link, Hort. Berol. 1: 268. 1 Oct-27 Nov 1827 [‘Chamagrostideae’], nom. illeg; Chondrosaceae Link, Hort. Berol. 1: 269. 1 Oct-27 Nov 1827 [‘Chondrosiaceae’]; Cynodontaceae Link, Hort. Berol. 1: 51, 269. 1 Oct-27 Nov 1827 [‘Cynodonteae’]; Cynosuraceae Link, Hort. Berol. 1: 271. 1 Oct-27 Nov 1827 [‘Cynosurinae’]; Echinariaceae Link, Hort. Berol. 1: 271. 1 Oct-27 Nov 1827; Ehrhartaceae Link, Hort. Berol. 1: 271. 1 Oct-27 Nov 1827 [‘Ehrhartinae’]; Glyceriaceae Link, Hort. Berol. 1: 271. 1 Oct-27 Nov 1827 [‘Glycerinae’]; Lappaginaceae Link, Hort. Bot. 1: 11, 268. 1 Oct-27 Nov 1827 [‘Lappagineae’], nom. illeg.; Loliaceae Link, Hort. Berol. 1: 6, 267. 1 Oct-27 Nov 1827; Miliaceae Link, Hort. Berol. 1: 91, 270. 1 Oct-27 Nov 1827; Ophiuraceae Link, Hort. Berol. 1: 3, 268. 1 Oct-27 Nov 1827 [‘Ophiurinae’]; Paspalaceae Link, Hort. Berol. 1: 47, 269. 1 Oct-27 Nov 1827 [‘Paspalinae’, ‘Paspalaceae’]; Phleaceae Link, Hort. Berol. 1: 269. 1 Oct-27 Nov 1827 [‘Phleodeae’]; Spartinaceae Link, Hort. Berol. 1: 46, 268. 1 Oct-27 Nov 1827; Triticaceae Link, Hort. Berol. 1: 22, 268. 1 Oct-27 Nov 1827 [’Triticeae’]; Zoysiaceae Link, Hort. Berol. 1: 8, 268. 1 Oct-27 Nov 1827 [’Zoysinae’]; Holcaceae Link, Hort. Berol. 2: 252. Jul-Dec 1833 [‘Holcoideae’]; Laguraceae Link, Hort. Berol. 2: 250. Jul-Dec 1833 [‘Lagurinae’]; Phalaridaceae Link, Hort. Berol. 2: 250. Jul-Dec 1833 [‘Phalarideae’]; Tristeginaceae Link, Hort. Berol. 2: 220. Jul-Dec 1833 [’Tristeginae’], nom. illeg.; Avenales Bromhead in Edinburgh New Philos. J. 24: 417. Apr 1838; Arundinaceae Döll, Rhein. Fl.: 60. 24-27 Mai 1843; Sesleriaceae Döll, Rhein. Fl.: 60. 24-27 Mai 1843; Coleanthaceae (Link) Pfeiffer, Nomencl. Bot. 1(2): 818. 3 Oct 1873 [‘Coleanthinae’]; Zeaceae A. Kern., Pflanzenleben 2: 651. 8 Aug 1891; Arundinellaceae (Stapf) Herter in Revista Sudamer. Bot. 6: 136. Jun 1940; Eragrostidaceae (Stapf) Herter in Revista Sudamer. Bot. 6: 145. Jun 1940; Lepturaceae (Dumort.) Herter in Revista Sudamer. Bot. 6: 147. Jun 1940; Pappophoraceae (Kunth) Herter in Revista Sudamer. Bot. 6: 145. 1940; Pharaceae (Stapf) Herter in Revista Sudamer. Bot. 6: 139. Jun 1940; Anomochloaceae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 222. 20 Jul 1943; Parianaceae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 222. 20 Jul 1943; Streptochaetaceae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov]: 222. 20 Jul 1943

Fossils Well preserved fossil grass phytoliths have been found in a dinosaur coprolith from the Maastrichtian Deccan Intertrappean Beds of India, and phytoliths (sometimes associated with epidermal fragments) are known from Cenozoic layers in many places. Pollen grains resembling those in Poaceae have been found in Maastrichtian layers, whereas leaves and flowers are known from the Early Eocene onwards. Paleocene grass pollen have been recorded from Brazil and tropical West Africa. Inflorescence parts have been detected in Oligocene layers in Germany and a spikelet was found in the Early Eocene of Tennessee.

Habit Usually bisexual (sometimes monoecious, andromonoecious, gynomonocious, polygamomonoecious, dioecious, androdioecious, or gynodioecious), usually perennial, biennial or annual herbs (sometimes woody, up to c. 40 m tall). Graminids. Sometimes helophytes, rarely aquatic. Numerous species are xerophytes. Culm terete to elliptic in cross-section, usually with hollow (fistulose; sometimes medullated solid) internodes and solid swollen nodes.

Vegetative anatomy Adventitious roots developing directly from hypocotyl. Endophytic fungi (i.a. Class I endophytes, Clavicipitaceae) frequent. Vesicular-arbuscular mycorrhiza often present. Roots producing siderophores that chelate ferric ions subsequently taken up by plant. Lateral roots developing from zone opposite protophloem or opposite protoxylem poles. Phellogen absent. Chlorenchyma without peg cells. Primary vascular tissue two or more cylinders of vascular bundles or scattered bundles. Secondary lateral growth absent. Vessels present in roots, stem and leaves. Vessel elements with scalariform or simple perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c_c type (with cuneate protein crystals) or P2c_cl type (with cuneate and several additional loosely packed protein crystals); P proteins absent from sieve elements. Nodes often swollen, with pulvinus; sclerenchyma usually absent; vascular bundle extending into leaf, with trace two nodes below leaf and connecting through next node above; nodes containing diffuse vascular bundles, transverse vascular bundles and expanded vascular bundles. Secretory cavities absent. Arm cells and fusoid cells present or absent. Special short idioblasts with large single silica bodies (phytoliths) of various shape (cuboidal, rounded, quadratic, cross- or saddle-shaped, etc.). Calciumoxalate absent or scarce. C₄ and CAM physiologies frequently present (at least 50% of species; evolved several times).

Trichomes Hairs unicellular or multicellular, uniseriate, or absent; glandular hairs sometimes present; usually bicellular (sometimes multicellular) microhairs present in most Poaceae.

Leaves Alternate (usually distichous; in Micraira tristichous), simple, entire, usually linear, with supervolute(-plicate), conduplicate (involute, revolute, convolute, or plicate) ptyxis (sometimes differentiated into pseudopetiole and pseudolamina). Stipules absent; leaf sheath well developed, usually open (rarely closed), long, usually with membranous adaxial distal ligule (sometimes modified into hairs or absent); sometimes with abaxial contraligule. Venation usually parallelodromous, often with transverse veins (in Leptaspis and Pharus pinnate-parallel); midvein usually distinct. Stomata paracytic, usually with dumbbell-shaped Poaceae type guard cells (not in, e.g., Neostapfia) and conical to dome-shaped subsidiary cells. Cuticular wax crystalloids as longitudinally aggregated or non-orientated rodlets and platelets, chemically dominated by polymeric aldehydes, or as tubuli, chemically characterized by hydroxy-β-diketones (sometimes Strelitzia type). Central mesophyll with large uncoloured fusoid cells. Epidermis usually with rows of long cells alternating with short cells containing silica (with longitudinal axis parallel to lamina); short ones usually containing silica crystals. Most Poaceae forming pairs of silica-cork short cells. Mucilaginous idioblasts absent. Secretory cavities usually absent. Cystoliths absent. Leaf margin usually entire (sometimes serrate). Epidermis usually with microhairs. Extrafloral nectaries rarely present.

Inflorescence Terminal, compound panicle, or simple or branched spike-, head-, spadix- or raceme-like (sometimes surrounded by spatha), consisting of spikelets subtended by one basal pair of bracts, glumes (glumae), without axillary flowers; above these one, several or numerous flowers each subtended and often surrounded by a bract (or outer tepal?), lemma, and opposite this a two-keeled and usually two-veined palea (possibly corresponding to a floral prophyll [bracteole] or two connate outer tepals). Glumes and/or lemmas (rarely paleas) often provided with long and narrow projections, awns. Lateral branches often bearing distinct adaxial swellings at base. Spikelet bracts usually suppressed. Some genera have modified sterile flowers or sterile branchlets serving as dispersal mechanisms.

Flowers Zygomorphic (due to reduction), small. Hypogyny. Each flower with usually two (rarely one, three or more than three) very small scale-like lodiculae, probably corresponding to reduced tepals of inner whorl (when more than three, then supernumerary lodiculae possibly corresponding to reduced modified sterile stamens; sometimes connate; often absent in female flowers). Nectary absent. Disc absent.

Androecium Stamens usually three (often 3+3, rarely one, two, four, five, or more than six, in Ochlandra up to c. 120 [to c. 170]). Filaments long, filiform, usually free from each other, free from lodiculae. Anthers dorsifixed, centrifixed or subbasifixed, versatile, tetrasporangiate, introrse, latrorse or extrorse, longicidal (dehiscing by longitudinal slits) or poricidal (dehiscing by apical pores or short slits). Tapetum secretory, with uni-, bi- or multinucleate cells. Staminodia absent.

Pollen grains Microsporogenesis usually successive (in Streptochaeta simultaneous). Pollen grains graminoid, usually monoporate to monoulcerate, operculate, shed as monads, usually tricellular at dispersal. Exine tectate, with columellate infratectum, smooth, echinulate, spinulate, or finely scabrate, with thin intraexinous channels (not scrobiculate).

Gynoecium Pistil composed of probably three connate carpels (three vascular bundles reaching stigma), with abaxial carpel fertile leading to pseudomonomery. Ovary superior, unilocular (pseudomonomerous). Style single, simple, solid or hollow, or stylodia two (rarely three; third style often represented by a reduced basal tissue), apical or lateral, free or more or less connate. Stigma single and usually bilobate (rarely uni-, tri- or quadrilobate), or stigmas two (rarely three), with often plumose lobes, papillate or non-papillate, Dry type, usually with scattered discontinuous receptive surfaces on multiseriate branches (sometimes on distinct zones) (pollen tubes growing between elongate transfer cells in multicellular stigmatic lobes). Pistillodium absent.

Ovules Placentation basal to parietal. Ovule one per ovary, usually hemicampylotropous (sometimes amphitropous or campylotropous, rarely orthotropous, pendulous), usually bitegmic (rarely unitegmic or almost ategmic), tenuinucellar or pseudocrassinucellar. Funicle short. Micropyle usually endostomal (sometimes bistomal). Outer integument two? cell layers thick, usually degenerating following fertilization. Inner integument at least two cell layers thick. Parietal cell not formed (parietal tissue absent). Megasporangium multilayered. Nucellar cap usually (not in Pooideae) formed through periclinal cell divisions in megasporangial epidermis. Megagametophyte monosporous, Polygonum type (Poaceae variation). Synergids sometimes haustorial (in ’Cortaderia’ and other Danthonioideae). Antipodal cells binucleate, more than three, often proliferating (rarely up to c. 300 cells). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis asterad (special variation). Polyembryony frequent.

Fruit Usually a nut-like caryopsis with adaxial end of seed coat usually fused with endocarp (sometimes an achene with membranous or gelatinous pericarp not adherent to seed coat; rarely a berry or a drupe).

Seeds Aril absent. Seed-coat usually tegmic. Testa usually not persistent. Tegmen often crushed. Perisperm not or only slightly developed. Endosperm copious, starchy, usually with proteinaceous tissue (rarely absent); peripheral cell layers of mature endosperm showing meristematic activity, with aleurone and sometimes oils; starch grains simple or compound. Embryo small to large, straight or curved, lateral-abaxial (sometimes inserted adjacent to seed coat on one side), well differentiated, with special and unique shape, usually almost enclosed by cylindrical coleoptile, without chlorophyll. Cotyledon one, lateral, not photosynthesizing. Haustorial cotyledon hyperphyll compact, not assimilating, modified into absorptive scutellum appressed to endosperm. Hypocotyl internode absent. Collar modified to epiblast, cotyledon ligule, often stout, or absent (alternatively, epiblast representing projection from first lateral root). Plumule terminal. Coleoptile first leaf, modified into substrate penetrating plumular envelope, with or without lamina. Alternatively, coleoptile and scutellum being different parts of cotyledon. Mesocotyl present as part of axis between coleoptile and scutellum, or absent. Radicula surrounded by coleorrhiza and scutellum, or absent. Germination cryptocotylar.

Cytology n = 11, 18 (Anomochlooideae); n = (2) (4) 5–12 (14) (other Poaceae) (n = 2–90) x = 5 or 11? – Polyploidy and aneuploidy frequently occurring. Agamospermy present in many genera. – ADP-glucose pyrophosphorylase present in cytosol.

DNA The plastid genome has one inversion of 28 kb, a second inversion of 6,4 kb, and a third inversion of 0,2–2 kb, and only 17 introns. The plastid gene accD, ORF244 and the introns 1 and 2 in clpP are often absent (at least in Bambusa, Oryza and Zea). ORF2280 is absent in many Poaceae. The intron is absent from the plastid gene rpoC1 (at least in Oryza and Zea). An insertion is present in the plastid gene rpoC2. A trnT inversion is present in the single copy region of the plastid genome. The IR region of the plastid DNA is expanded. A translocation has taken place in the plastid genome, thus creating the pseudogene ψrpl23.

Duplication of the nuclear genes AP1/FUL (FUL1 and FUL2) has taken place in the crown clade (this duplication may be involved in the evolution of the spikelet): one copy coding for degradation of starch into fermentable sugars in endosperm, the second copy with wider expression. Vast gene duplications have taken place in the API, AG and SEP gene families, although not in the AP3 evolutionary line. A duplication of the gene LOFSEP may have taken place in Poaceae (Malcomber & Kellogg 2005), the entire genome being duplicated in a clade comprising at least Zea, Oryza, Hordeum, and Sorghum. Loss of nuclear genes ycf1 and ycf2, the c. 200 bp ψycf1 being left in the genome. The mitochondrial gene rps14 has been transferred to the nuclear genome, the pseudogene ψrps14 being left in mtDNA.

Phytochemistry Flavonol sulphates, flavone sulphates, flavone-5- and C-glycosides, luteolin, tricin (tricetin 3’,5’-dimethyl ether, very frequent), 3-deoxyanthocyanins, free triterpenes, isoquinoline alkaloids, tryptophane- or tyramine-derived alkaloids (gramine, hordenine, tyramine, etc.), pyrrolizidine alkaloids (1-aminopyrrolizidine derivatives; loline, lolinine, norline, etc.), indole alkaloids, tyrosine-derived cyanogenic compounds, and chelidonic acid present. Flavonols (kaempferol, quercetin), cyanidin, and saponins rare. Ellagic acid not found. Primary cell wall with abundant arabinoxylans. Xyloglucans without fucose. Lignins with p-coumarylic alcohol (lignins acylated with p-coumarates), and coniferyl and sinapyl monomers. Hemicellulose/pectin composition different from other seed plants. Aluminium accumulation occurring in some species.

Use Ornamental plants, starch sources (cereals), sugar (Saccharum), beer and other alcohol beverages (Hordeum, Triticum, Saccharum, etc.), vegetables, spices and perfumes (Cymbopogon), necklaces (Coix), forage plants, thatching, basketry, textiles, carpets, brushes, bamboo species for timber, carpentries, tools, ornaments, weapons, musical instruments, vegetables, paper, etc.

Systematics Poaceae are sister-group either to Ecdeiocoleaceae or (perhaps more probable) to the clade [Joinvilleaceae+Ecdeiocoleaceae]. Multicellular microhairs are present in Joinvilleaceae and many Poaceae (also in Ecdeiocoleaceae?). - Poaceae incertae sedis: Schmidiella (1; S. maxwellii; Laos).

The system below largely follows: Soreng RJ, Davidse G, Peterson PM, Zuloaga FO, Judziewicz EJ, Filgueiras TS, Morrone O, Romaschenko. 2012 (and continuously revised). A world-wide phylogenetic classification of Poaceae (Gramineae). Published online. Another important source is Peterson & al. (2011).The potential synapomorphies of the clades are mainly according to Stevens (2001 onwards).

A possible topology is the following:

[Anomochloa+[Streptochaeta+[Pharoideae+[Puelioideae+[[Aristidoideae+[Panicoideae+Arundinoideae]]]+[Oryzoideae+[Bambusoideae+Pooideae]]]]]]]

Sometimes, the following two alternative topologies of the “crown clade” are recovered:

[[Pooideae+[Bambusoideae+Oryzoideae]]+[Micrairoideae+[Panicoideae+[[Aristidoideae+Danthonioideae]+[Arundinoideae+Chloridoideae]]]]] or [Bambusoideae+[Oryzoideae+[Pooideae+[Panicoideae+[[Aristidoideae+Danthonioideae]+[Arundinoideae+Chloridoideae]]]]]]

Anomochlooideae Pilg. ex Potzdal in Willdenowia 1: 772. 1 Mar 1957 [‘Anomochloideae’] (paraphyletic?)

2/4. Anomochloa (1; A. marantoidea; Bahia in southeastern Brazil); Streptochaeta (3; S. angustifolia [extinct?], S. sodiroana, S. spicata; Mexico, Central America, the West Indies, tropical South America). – Central America to tropical South America, forests. Pseudopetiole with apical (and sometimes basal) pulvinus. Ligule modified into fringe of hairs. Arm cells absent in Streptochaeta. Inflorescence branches cymose, with two ’bracts’ along each branch unit and two additional ’bracts’ subtending each flower (Anomochloa), or flowers spirally arranged along racemose axis, with several spiral ’bracts’ subtending each flower (Streptochaeta). Tepals 2(–3)+3 or absent. Anomochloa with four stamens and one carpel. Filaments connate at base. Anthers subbasifixed or centrifixed (Anomochloa), latrorse. Anther wall development Reduced type, without endothecium thickenings. Microsporogenesis simultaneous. Stigma in Streptochaeta not plumose. Antipodal cells not proliferating (Streptochaeta). Epiblasts present in Anomochloa, absent in Streptochaeta. First leaf of seedling without pseudolamina. n = 11, 18. Plastid gene rpoC2 insertion with 21 bp subrepeats. A c. 850 bp pseudogene ψycf1 present in nuclear genome in Anomochloa. – Streptochaeta and Anomochloa are sister-groups in some analyses. In others, Streptochaeta is recovered as successive sister to the remaining Poaceae (except Anomochloa). The bicarinate palea in Streptochaeta is sometimes interpreted as originated from two adaxial outer tepals. Likewise, the lodiculae have been interpreted as three modified inner tepals. In a way, morphology indicates that Streptochaeta may be sister to all other Poaceae (including Anomochloa).

[Pharoideae+[Puelioideae+[SAPA+EBP]]] ("The spikelet clade")

Leaves with membranous ligules, sometimes also ciliate. Inflorescence consisting of laterally compressed, racemose, pedunculate spikelets. Spikelet units with two sterile basal glumes (spikelet bract + prophyll). Flowers distichous, each with lemma and palea (possibly equalling one bract and two connate adaxial outer tepals), inverted. Lodiculae two or three (possibly equalling inner tepal whorl; median lodicula adaxial). n = 12. 3’ end of the plastid gene matK with 1 bp deletion. Plastid gene rpoC1 absent (lost). Plastid gene rpoC2 insertion with 39 bp subrepeats.

Pharoideae L. G. Clark et Judz. in Taxon 45: 643. 1996

3/13. Leptaspis (3; L. angustifolia, L. banksia, L. zeylanica; tropical Africa, Madagascar, the Comoros, tropical Asia from India and Sri Lanka to Queensland and Melanesia), Scrotochloa (2; S. urceolata, S. tararaensis; India, Sri Lanka, Burma, Southeast Asia, Malesia to New Guinea, the Bismarck Archipelago, Solomon Islands, Queensland), Pharus (7; P. ecuadoricus, P. lappulaceus, P. latifolius, P. mezii, P. parvifolius, P. virescens, P. vittatus; Florida, southern Mexico, Central America, the West Indies, tropical South America). – Pantropical, forests. Microhairs absent. Leaves resupinate. Lateral veins oblique. Spikelets one-flowered. Stamens 3+3. Anthers centrifixed, latrorse. Anther wall development in Pharus Reduced type, without endothecium thickenings. Style hollow (Pharus) or solid. Micropyle bistomal (Pharus). Antipodal cells not proliferating (Pharus). Coleoptile with pseudolamina. Scutellus tail absent in Pharus.

[Puelioideae+[SAPA+IEBP]] ("The bistigmatic clade")

Phytoliths saddle-shaped. Spikelets disarticulating above glumes. Anthers usually versatile. Stigmas two, with stigmatic branching in two orders. Plastid gene ndhF with 15 bp insertion.

Puelioideae (Soderstr. et R. P. Ellis) L. G. Clark, M. Kobay., S. Mathews, Spangler et E. A. Kellogg in Syst. Bot. 25: 184. Apr-Jun 2000

2/11. Guaduella (6; G. densiflora, G. dichroa, G. humilis, G. macrostachys, G. marantifolia, G. oblonga; tropical West and Central Africa), Puelia (5; P. ciliata, P. coriacea, P. dewevrei, P. olyriformis, P. schumanniana; tropical West and Central Africa). – Tropical West and Central Africa. Flowers bisexual or unisexual, polygamous. Perennial with sympodial rhizome. Kranz’ anatomy absent. Arm cells poorly developed. Fusoid cells well developed. Microhairs multicellular uniseriate (Guaduella) or absent (Puelia). Pseudopetiole present. Contraligule present (Puelia) or absent (Guaduella). Lodiculae three. Stamens 3+3. Filaments free (Guaduella) or connate (Puelia). Stigmas two or three. Embryo small. Seedling leaves of unknown shape. n = 12 (Puelia).

[[Aristidoideae+[Panicoideae+[Arundinoideae+[Danthonioideae+Chloridoidae]]]]+[Ehrhartoideae+[Bambusoideae+Pooideae]]]

Arm cells absent. Fusoid cells absent. No differentiation of leaf into pseudopetiole and pseudolamina. Leaf transverse veins absent. Lodiculae usually two. Stamens usually three. Stylodia two, not connate. Antipodal cells usually proliferating. x = 12. Benzoxazinoids (a type of defense molecules) sometimes present. Genome duplication present. Plastid gene ndhF with 15 bp insertion. Disease resistance through nuclear gene Hm1.

[Aristidoideae+[Panicoideae+[Arundinoideae+[Danthonioideae+Chloridoideae]]]]

Phytoliths axial-bilobate or axial-polylobate.

Aristidoideae Caro in Dominuezia 4: 16. Feb 1982

3/350–360. Aristida (c 290; tropical and subtropical regions on both hemispheres), Sartidia (6; S. angolensis, S. dewinteri, S. isaloensis, S. jucunda, S. perrieri, S. vanderystii; Central to southern Africa, Madagascar), Stipagrostis (c 55; arid and semi-arid subtropical regions in the Mediterranean, Africa and Asia to China). – Tropical and subtropical regions. Often with C₄ photosynthesis. Ligule lined with hairs. Spikelet elongated-cylindrical, disarticulating above glume. Lemma awn trifid, or (one or) three awns, with basal columella. Germination flap present. n = 11, 12. Insertion of 6 bp absent from 3’ end of plastid gene matK. – Aristidoideae are possibly sister-group to the clade [Panicoideae+Arundinoideae+[Danthonioideae+ Chloridoideae]]. However, Aristidoideae are sometimes revealed as sister-group to Danthonioideae, and the analyses by Peterson & al. (2011) place Aristideae as sister to the clade [Hakonechloa+[Danthonieae+[Chlorideae+Centropodieae]]].

[Panicoideae+[Arundinoideae+[Danthonioideae+Chloridoideae]]] (The "PACC clade", the "PACMAD clade", or the "PACCMAD" clade)

C₄ photosynthesis present in approximately 50% of the species. Phytoliths dumbbell-shaped. Ligule often consisting of hairs. Hilum non-linear. Mesocotyl internode elongated. Epiblast absent. Plastid gene ndhF extended from short single copy region (SSC) into inverted repeat (IR). 3’ end of the plastid gene matK usually with 6 bp insertion.

Panicoideae Link, Hort. Berol. 1: 202. 1 Oct-27 Nov 1827 [‘Paniceae’]

217/3.075–3.105. Tropical, subtropical and temperate regions. Culm usually solid. Fusoid cells often absent. C₄ photosynthesis frequent. Elongate microhairs usually present, Panicoid type (with slender thin-walled cap cells). Stomata with triangular or dome-shaped subsidiary cells. Spikelet development basipetal. Spikelet dorsally compressed, without rachilla, two-flowered, lower flower male or sterile, female floret one. Spikelet shed as one-seeded unit by disarticulation below glumes. Style sometimes present. Hilum non-linear. Starch grains simple. Embryo with overlapping leaf margins. Epiblast usually present in ’centothecoids’. Germination flap present. n = 5, 7, 9, 10 (11–14). Plastid gene rpl16 intron with 5 bp insertion. Deletion of 4 bp in rpl16 intron in "centothecoids" Pseudogene ψrps14 absent (lost). – Panicoideae incertae sedis: Alloeochaete (6; A. andongensis, A. geniculata, A. gracillima, A. namuliensis, A. oreogena, A. ulugurensis; southern tropical Africa), Dichaetaria (1; D. wightii; India, Sri Lanka).

Gynerieae Sánchez-Ken et L. G. Clark in Novon 11: 350. 21 Sep 2001

1/1. Gynerium (1; G. sagittatum; Central America, South America south to Paraguay). – Gynerieae may be sister-group to the clade [[Thysanolaeneae+Centotheceae+Cyperochloeae]+[Chasmanthieae+Zeugiteae]]. According to Morrone & al. (2012), they are instead sister to the clade [Paniceae+[Paspaleae+[Arundinelleae+Andropogoneae]]].

[[Thysanolaeneae+Centotheceae+Cyperochloeae]+[Chasmanthieae+Zeugiteae]]

[Thysanolaeneae+Centotheceae+Cyperochloeae]

Thysanolaeneae C. E. Hubb. in J. Hutchinson, Fam. Fl. Plants 2: 222. 20 Jul 1934

1/1. Thysanolaena (1; T. latifolia; southern and southeastern Asia, southern China). – Thysanolaena may be sister to the clade [Centotheceae+Cyperochloeae] or should be included in Centotheceae.

Centotheceae Ridl., Mat. Fl. Malay. Pen. 3: 122. 1907

2/4. Centotheca (3; C. lappacea, C. philippinensis, C. uniflora; tropical West and Central Africa, Madagascar, China, Southeast Asia, Malesia to New Guinea, islands in the Pacific), Megastachya (1; M. mucronata; tropical and southern Africa, Madagascar). – Tropical regions in the Old World. – Centotheceae may be sister to Cyperochloeae.

Cyperochloeae L. Watson et Dallwitz ex Sánchez-Ken et L. G. Clark in Amer. J. Bot. 97: 1744. 1 Oct 2010

2/2. Cyperochloa (1; C. hirsuta; southwestern Western Australia), Spartochloa (1; S. scirpoidea; southwestern Western Australia). – Southwestern Western Australia. – Cyperochloeae should perhaps be included in Centotheceae.

[Chasmanthieae+Zeugiteae]

Chasmanthieae W. V. Br. et B. N. Sm. ex Sánchez-Ken et L. G. Clark in Amer. J. Bot. 97: 1744. 1 Oct 2010

1/6. Chasmanthium (6; C. curvifolium, C. latifolium, C. laxum, C. nitidum, C. ornithorhynchum: southern Canada, eastern United States, northern Mexico; C. gossweileri: Tanzania to Angola). – Eastern and southern tropical Africa, eastern North America. – Chasmanthieae are sister-group to Zeugiteae.

Zeugiteae Sánchez-Ken et L. G. Clark in Amer. J. Bot. 97: 1744. 1 Oct 2010

4/18. Chevalierella (1; C. dewildemanii; Congo), Lophatherum (2; L. gracile, L. sinense; India, Sri Lanka, the Himalayas, China, the Korean Peninsula, Japan, Taiwan, Southeast Asia, Malesia to New Guinea and Queensland), Orthoclada (2; O. africana: southeastern tropical Africa; O. laxa: southern Mexico, Central America, tropical South America), Zeugites (13; Mexico, Central America, the West Indies, tropical South America). – Pantropical.

[[Steyermarkochloeae+Tristachyideae]+[Andropogonodae+Panicodae]]

[Steyermarkochloeae+Tristachyideae]

Steyermarkochloeae Davidse et R. P. Ellis in Ann. Missouri Bot. Gard. 71: 994. 1985

2/2. Arundoclaytonia (1; A. dissimilis; Amazonas, Brazil), Steyermarkochloa (1; S. angustifolia; Colombia, Venezuela). – Tropical South America. – Steyermarkochloeae are sister to Tristachyideae, according to Morrone & al. (2012).

Tristachyideae Sánchez-Ken et L. G. Clark in Amer. J. Bot. 97: 1743. 1 Oct 2010

5/c 70. Danthoniopsis (16; Africa, the Arabian Peninsula to Pakistan), Gilgiochloa (1; G. indurata; tropical Africa), Loudetia (c 25; tropical and southern Africa, Madagascar, tropical South America), Trichopteryx (5; T. dregeana, T. elegantula, T. fruticulosa, T. marungensis, T. stolziana; tropical and southern Africa, Madagascar), Tristachya (22; tropical and southern Africa, Madagascar, tropical South America). – Africa and Madagascar to Pakistan, tropical South America. – Tristachyideae may be sister to Steyermarkochloeae.

[Andropogonodae+Panicodae]

Lecomtelleae Pilg. ex Potztal in Willdenowia 1: 771. 1957

1/1. Lecomtella (1; L. madagascariensis; the Andringitra mountain range in southern Madagascar). – The position of Lecomtella madagascariensis is uncertain. It is sister-group to [Sacchareae+Paspaleae] or [Sacchareae+Paniceae] or [Paniceae+Paspaleae] or [Paniceae+[Paspaleae+Andropogoneae]] or to Paniceae (Besnard & al. 2013).

Andropogonodae L. Liu in Acta Phytotaxon. Sin. 18: 325. Aug 1980 (‘Andropogodae’]

Paspaleae J. Presl in C. B. Presl, Reliq. Haenk. 1: 208. Jan-Jun 1830 [’Paspalineae’]

38/595–600. Reynaudia (1; R. filiformis; Cuba, Hispaniola). – Paspalinae Griseb. in C. F. von Ledebour, Flor. Ross. 4: 466. Jun 1853 [’Paspaleae’]. Aakia (1; A. tuerckheimii; southern Mexico, Central America), Acostia (1; A. gracilis; Ecuador), Anthaenantiopsis (5; A. fiebrigii, A. perforata, A. racemosa, A. rojasiana, A. trachystachya; tropical and subtropical South America), Axonopus (c 70; southern United States, Mexico, Central America, the West Indies, tropical South America, one species, A. compressus, in Africa, and one species, A. foliacea, in Mozambique), Gerritea (1; G. pseudopetiolata; Bolivia), Hopia (1; H. obtusa; southwestern United States, northwestern Mexico), ‘Ichnanthus‘ pro parte (2), Hildaea (6; H. breviscrobs, H. nemorosa, H. pallens, H. parvispiculata, H. ruprechtii, H. tenuis; pantropical), Streptostachys (6; S. acuminata, S. asperifolia, S. macrantha, S. ramosa, S. rigidifolia, S. robusta; Mexico, Central America, the West Indies, tropical South America), Renvoizea (10; Brazil), Oedochloa (9; tropical America), Ocellochloa (12; tropical America), Echinolaena (2; E. gracilis, E. inflexa; southern Mexico, Central America, tropical South America), Ichnanthus (c 30; Mexico, Central America, the West Indies, tropical South America), Lecomtella (1; L. madagascariensis; Madagascar), Osvaldoa (1; O. valida; Brazil, Uruguay, Argentina), Paspalum (c 330; tropical and subtropical regions on both hemispheres, with their largest diversity in tropical America). – Otachyriinae Butzin in Willdenowia 6: 182. 23 Nov 1970. Anthenantia (4; A. lanata, A. rufa, A. texana, A. villosa; southeastern United States), Rugoloa (3; R. hylaeica, R. pilosa, R. polygonata; Jamaica, Brazil), Hymenachne (4; H. amplexicaulis, H. aurita, H. felliana, H. pseudointerrupta; southern Mexico, Central America, the West Indies, tropical South America), ’Hymenachne’ pro parte (10; H. assamica, H. donacifolia, H. grumosa, H. patens, H. pernambucensis, H. wombaliensis, ’Panicum’ harleyi, P. hemitomon, P. leptachne, P. stagnatile; tropical Africa, tropical Asia, Central America, the West Indies, tropical South America), Plagiantha (1; P. tenella; eastern Brazil), Otachyrium (15; southern United States to tropical South America). – Arthropogoninae Butzin in Willdenowia 6: 515. Nov 1972. Apochloa (15; tropical South America), Arthropogon (7; A. chapadense, A. filifolius, A. piptostachyus, A. scaber, A. sorengii, A. villosus, A. xerachne; the Greater Antilles, Colombia, Brazil, Bolivia), Canastra (2; C. aristella, C. lanceolata; Brazil), Coleataenia (7; C. anceps, C. longifolia, C. rigidula, C. scabrida, C. stenodes, C. stipitata, C. tenera; eastern United States, Mexico, Central America, tropical South America), Cyphonanthus (1; C. discrepans; Central America, the West Indies, tropical South America), Homolepis (5; H. aturensis, H. glutinosa, H. isocalycia, H. longispicula, H. villaricensis; Mexico, Central America, the West Indies, tropical South America), Keratochlaena (1; K. rigidifolia; Guyana, northern Brazil), Mesosetum (c 25; Mexico, Central America, the West Indies, tropical South America), Oncorachis (2; O. macrantha, O. ramosa; Brazil), Oplismenopsis (1; O. najada; Uruguay, Argentina), Phanopyrum (1; P. gymnocarpon; southeastern United States), Stephostachys (1; S. mertensii; Mexico, Central America, tropical South America), Tatianyx (1; T. arnacites; Brazil), Triscenia (1; T. ovina; Cuba). – Pantropical, with their largest diversity in tropical America.

[Andropogoneae+Arundinelleae]

Andropogoneae Dumort., Observ. Gramin. Belg.: 84. Jul-Sep 1824 [‘Andropogineae‘]

82/1.110–1.130. Chrysopogon (45–50; warmer regions on both hemispheres, especially in tropical region in the Old World), Thelepogon (1; T. elegans; Ethiopia to Namibia and east to India). – Arthraxoninae Benth. in J. Linn. Soc. London, Bot. 19: 67. 1881 [‘Arthraxeae‘]. Arthraxon (27; tropical and subtropical Africa, Madagascar, islands in the Indan Ocean, India and the Himalayas to southern China and Southeast Asia, Malesia to New Guinea and northern Australia). – Tripsacinae Dumort., Anal. Fam. Plant.: 64. 1829 [’Tripsaceae’]. Elionurus (c 15; Macaronesia, tropical and subtropical Africa, Madagascar, southwestern Asia to Sind in northwestern India, New Guinea, Solomon Islands, Australia, southern United States, Mexico, Central America, the West Indies, tropical South America), Oxyrhachis (1; O. gracillima; tropical East and southern Africa, Madagascar), Rhytachne (12; tropical and southern Africa, Madagascar, tropical South America), Tripsacum (16; southern United States, Mexico, Central America, South America to Paraguay), Urelytrum (7; U. agropyroides, U. annuum, U. auriculatum, U. digitatum, U. giganteum, U. henrardii, U. muricatum; tropical and southern Africa, Madagascar), Vossia (1; V. cuspidata; tropical Africa to Namibia and northern Botswana, eastern India to Burma), Zea (6; Z. diploperennis, Z. luxurians, Z. mays, Z. mexicana, Z. nicaraguensis, Z. perennis; Mexico, Central America). – Chionachninae Clayton in Kew Bull. 35: 813. 1981. Chionachne (12; India to Burma, southern China and Southeast Asia, Malesia to New Guinea, Solomon Islands and northern Australia; C. cookei (Trilobachne cookei): the West Indies). – Coicinae Reichb. ex Clayton et Renvoize in Kew Bull., Addit. Ser. 13: 372. 1986. Coix (4; C. aquatica, C. gasteenii, C. lacryma-jobi, C. puellarum; India to southern China and Southeast Asia, northern Queensland). – Rottboelliinae J. Presl in C. Presl, Reliq. Haenk. 1: 329. Jan-Jun 1830 ['Rottboelliaceae']. Chasmopodium (3; C. afzelii, C. caudatum, C. purpurascens; tropical West and Central Africa), Eremochloa (14; India, Sri Lanka, Southeast Asia, Malesia to New Guinea and tropical Australia), Hemarthria (12; tropical and subtropical regions in the Old World), Ophiuros (4; O. bombaiensis, O. exaltatus, O. megaphyllus, O. papillosus; the Horn of Africa, India and eastern Himalayas to Burma, southern China and Southeast Asia, the Ryukyu Islands, Malesia to New Guinea and Australia), Mnesithea (6; M. annua, M. formosa, M. laevis, M. mollicoma, M. pilosa, M. veldkampii; the Indian Subcontinent, Sri Lanka, China, Japan, Taiwan, Southeast Asia, Malesia to New Guinea and tropical Australia, islands in the Pacific), Hackelochloa (2; H. granularis, H. porifera; sub-Saharan Africa, the Arabian Peninsula, India, eastern Himalayas to Japan and Southeast Asia, Malesia to New Guinea, Micronesia), ’Phacelurus’ (10; tropical and subtropical Africa, Southwest and East Asia, tropical Asia to Indochina and Japan, one species, P. digitatus, on the Balkan Peninsula to Syria; polyphyletic), Rottboellia (6; R. cochinchinensis, R. coelorachis, R. goalparensis, R. laevispica, R. paradoxa, R. purpurascens; tropical Africa, Madagascar, tropical and subtropical Asia and Australia, Melanesia), Glyphochloa (12; central and southwestern India), Heteropholis (3; H. sulcata: Central and tropical East Africa; H. benoistii: Madagascar; H. nigrescens: Sri Lanka), Lasiurus (1; L. scindicus; Morocco and Mali to the Middle East and northeastern India), Loxodera (5; L. bovonei, L. caesitosa, L. ledermannii, L. rhytachnoides, L. strigosa; tropical Africa), Thaumastochloa (9; New Guinea, Australia). – Ischaeminae J. Presl in C. Presl, Reliq. Haenk. 1: 328. Jan-Jun 1830 [’Ischaemeae’]. Andropterum (1; A. stolzii; tropical Africa), Dimeria (c 60; Madagascar, India and Sri Lanka to China, the Korean Peninsula, Japan and Southeast Asia, Malesia to New Guinea and tropical Australia, Micronesia), Ischaemum (c 80; tropical and subtropical regions on both hemispheres, especially Asia), Kerriochloa (1; K. siamensis; Southeast Asia), Pogonachne (1; P. racemosa; the Mumbay area in India), Triplopogon (1; T. ramosissima; western India). – Germainiinae Clayton in Kew Bull. 27: 465. 1972. Apocopis (16; India and Sri Lanka to southern China, Southeast Asia and Sulawesi), Germainia (9; Assam, Southeast Asia, Malesia to Queensland), Lophopogon (2; L. kingii, L. tridentatus; India), Pogonatherum (4; P. biaristatum, P. crinitum, P. paniceum, P. rufobarbatum; Madagascar, the Arabian Peninsula, India to China and Japan, Southeast Asia, Malesia to New Guinea). – Saccharinae Griseb., Spic. Flor. Rumel. 2: 472. Jan 1846 [’Sacchareae’]. Asthenochloa (1; A. tenera; Central Malesia), Euclasta (2; E. clarkei, E. condylotricha; tropical Africa, Madagascar, the Comoros, Oman India, Burma, Mexico to tropical South America), Pseudodichanthium (1; P. serrafalcoides; western India), Cleistachne (1; C. sorghoides; tropical East and southeastern Africa, Oman, India), Sorghastrum (20; tropical and southern Africa, North America, Mexico, Central America, the West Indies, tropical South America), Sorghum (28; tropical and subtropical regions in the Old World, India, Sri Lanka, Burma, Southeast Asia Australia, one species, S. trichocladum, in Mexico and Central America), Trachypogon (4; T. chevalieri, T. macroglossus, T. spicatus, T. vestitusa; tropical and southern Africa, Madagascar, southern United States and Mexico to tropical South America), ’Eulalia’ (34; tropical and subtropical regions in Africa, Madagascar, Asia and Australia; polyphyletic), Homozeugos (6; H. conciliatum, H. eylesii, H. fragile, H. gossweileri, H. huillense, H. katakton; tropical Africa), Miscanthus (16; tropical and subtropical regions in the Old World, southern Africa, East Asia), Polytrias (1; P. indica; Southeast Asia), Pseudosorghum (2; P. fasciculare, P. zollingeri; the Himalayas, Burma, Yunnan, Southeast Asia, Java, the Philippines; in Eulalia?), Saccharum (35–40; tropical and subtropical regions on both hemispheres), Tripidium (>3; T. bengalense, T. ravennae, T. strictum; southeastern Europe, the Mediterranean, southwestern and southern Asia), Imperata (12; tropical and subtropical regions on both hemispheres), Veldkampia (1; V. sagaingensis; Southeast Asia), Agenium (3; A. leptocladum, A. majus, A. villosum; Brazil, Uruguay, Paraguay, Bolivia, Argentina), Eriochrysis (10; tropical and southern Africa, India, tropical America). – Andropogoninae J. Presl in C. Presl, Reliq. Haenk. 1: 331. Jan-Jun 1830 [’Andropogoneae’]. Andropogon (c 100; tropical and subtropical regions on both hemispheres), Bhidea (3; B. borii, B. burnsiana, B. fischeri; India), Diheteropogon (4; D. amplectans, D. filifolius, D. hagerupii, D. microterus; tropical and southern Africa), Hyparrhenia (c 55; Macaronesia, the Mediterranean, Africa, Madagascar, southwestern and southern Asia to Pakistan and India, tropical America, with their highest diversity in Africa), Schizachyrium (c 65; tropical and subtropical regions on both hemispheres), Anadelphia (21; western to southeastern tropical Africa), Bothriochloa (c 38; tropical and subtropical regions on both hemispheres), Capillipedium (18; East Africa, tropical Asia to Australia and New Caledonia), Clausospicula (1; C. extensa; Northern Territory), Cymbopogon (55–60; tropical and subtropical regions in Africa, Madagascar, Asia and Australia), Dichantium (19–23; tropical and subtropical regions in the Old World), Elymandra (6; E. androphila, E. archaelymandra, E. gossweileri, E. grallata, E. lithophila, E. subulata; tropical and southern Africa, one species, E. lithophila, also in Brazil), Eremopogon (1; E. foveolatus; tropical Africa, the Middle East to India and Sri Lanka), Exotheca (1; E. abyssinica; tropical East Africa, Vietnam), Heteropogon (5; H. contortus, H. fischerianus, H. melanocarpus, H. ritchiei, H. triticeus; tropical and subtropical regions on both hemispheres), Hyperthelia (7; H. colobantha, H. cornucopiae, H. dissoluta, H. edulis, H. kottoensis, H. macrolepis, H. polychaeta; tropical and southern Africa), Iseilema (c 30; India, Sri Lanka, Southeast Asia, Malesia to New Guinea and tropical Australia), Monocymbium (3; M. ceresiiforme, M. deightonii, M. lanceolatum; tropical and southern Africa), Parahyparrhenia (6; P. annua, P. bellariensis, P. laegaardii, P. perennis, P. siamensis, P. tridentata; tropical West and Central Africa, India, Thailand), Pseudanthistiria (4; P. burmanica, P. emeinica, P. heteroclita, P. umbellata; India, Thailand, southern China), Spathia (1; S. neurosa; tropical Australia), Spodiopogon (16; Turkey, Southwest, Central, East and tropical Asia to the Russian Far East, Japan, Taiwan and the Malay Peninsula), Themeda (26; tropical and subtropical regions in Africa, Asia and Australia); unplaced Andropogoninae: Lakshmia (1; L. venusta; Sri Lanka, Western Ghats). – Andropogoneae incertae sedis: Apluda (1; A. mutica; Madagascar, Oman, Socotra, the Mascarene Islands, Central Asia, the Indian Subcontinent, Tibet, China, Japan, the Ryukyu Islands, Taiwan, Southeast Asia, Malesia to New Guinea and Melanesia), Eulaliopsis (2; E. binata, E. sykesii; Afghanistan, India, China, Taiwan, the Philippines), Microstegium (c 25; tropical and subtropical Africa and Asia), Sehima (5; S. galpinii, S. ischaemoides, S. nervosum, S. notatum, S. sulcatum; the Cape Verde Islands, tropical and southern Africa, Southwest Asia and India to southern China and Burma, Southeast Asia to Java, New Guinea). – Coix is sister to Rottboelliinae (Soreng & al. 2017). – Tropical and subtropical regions on both hemispheres. Spikelets paired. C4 photosynthetic pathway (NADP-ME subtype) present.

Arundinelleae Stapf in W. H. Harvey et O. W. Sonder (ed. W. T. Thiselton-Dyer), Fl. Cap. 7: 314. Jul 1898

2–4/75–80. Arundinella (45–50; tropical and subtropical regions on both hemispheres), Garnotia (c 30; southern and eastern Asia to islands in the Pacific). – Plausible Arundinelleae: Chandrasekharania (1; C. keralensis; Kerala in India), Jansenella (1; J. griffithiana; India, Sri Lanka). – Tropical and subtropical regions, especially Asia. – Arundinelleae are sister to Andropogoneae (Sánchez-Ken & Clark 2010). The positions of Chandrasekharania and Jansenella are uncertain.

Panicodae L. Liu in Acta Phytotaxon. Sin. 18: 324. Aug 1980

Paniceae R. Br. in M. Flinders, Voy. Terra Austral. 2: 582. 19 Jul 1814

74/1.190–1.210. Hylebates (2; H. chlorochloe, H. cordatus; tropical Africa). – Anthephorinae Benth. in J. Linn. Soc. London, Bot. 19: 30. 24 Dec 1881 ['Anthephoreae']. ’Digitaria‘ (270–280; cosmopolitan; non-monophyletic), Chlorocalymma (1; C. cryptacanthum; Tanzania), Anthephora (11; tropical and southern Africa, the Arabian Peninsula, one species, A. hermaphrodita, also in tropical America), Chaetopoa (2; C. pilosa, C. taylorii; Tanzania); unplaced Anthephorinae: Taeniorhachis (1; T. repens; Somalia), Tarigidia (1; T. aequiglumis; Namibia, North-West, Free State), Thyridachne (1; T. tisserantii; tropical Africa), Trachys (1; T. muricata; coastal areas in southern India, Sri Lanka, Bangladesh and Burma). – Dichantheliinae Zuloaga in Plant Syst. Evol. 301: 1697. 1 Jun 2015. Adenochloa (14; tropical and subtropical Africa, Madagascar), Dichanthelium (70–75; North America, Mexico, Central America, the West Indies, tropical and subtropical South America). – Boivinellinae Pilg. in H. G. A. Engler et K. A. E. Prantl, Nat. Pflanzenfam., ed. 2, 14e: 101. 1940. Acroceras (c 25; tropical and southern Africa, Madagascar, Southeast Asia, Malesia), Setiacis (1; S. diffusa; Hainan; in Acroceras?), Alloteropsis (5; A. angusta, A. cimicina, A. paniculata, A. papillosa, A. semialata; tropical and subtropical regions in the Old World), Amphicarpum (2; A. amphicarpon, A. muehlenbergianum; southeastern United States), Cyphochlaena (2; C. madagascariensis, C. sclerioides; Madagascar, the Comoros), Cyrtococcum (14; tropical regions in Africa, Asia and Australia), Echinochloa (c 40; warm-temperate to tropical regions on both hemispheres), Entolasia (6; E. imbricata, E. marginata, E. minutifolia, E. olivacea, E. stricta, E. whiteana; tropical and southern Africa, tropical Asia to eastern Australia, New Caledonia), Lasiacis (16; Florida, Mexico, Central America, the West Indies, tropical South America), Mayariochloa (1; M. amphistemon; Cuba), Microcalamus (1; M. barbinodis; tropical West Africa), Morronea (6; M. arundinariae, M. cayoensis, M. guatemalensis, M. incumbens, M. parviglumis, M. trichidiachnis; Mexico, Central America, the West Indies, tropical South America), Oplismenus (7; O. burmannii, O. compositus, O. flavicomus, O. fujianensis, O. hirtellus, O. thwaitesii, O. undulatifolius; tropical and subtropical regions on both hemispheres), Ottochloa (3; O. gracillima, O. grandiflora, O. nodosa; India, southern China, Southeast Asia, Malesia to northern Australia), Parodiophyllochloa (6; P. cordovensis, P. missiona, P. ovulifera, P. pantricha, P. penicillata, P. rhizogona; Mexico, Central America, tropical South America), Poecilostachys (c 20; tropical Africa, Madagascar), Pseudechinolaena (6; P. camusiana, P. madagascariensis, P. moratii, P. perrieri, P. polystachya, P. tenuis; Madagascar, one species, P. polystachya, pantropical). – Clade. Sacciolepis (c 25; tropical and subtropical regions on both hemispheres, with their largest diversity in tropical Africa); Trichanthecium (38; tropical Africa, tropical America). – Neurachninae Clayton et Renvoize in Kew Bull., Addit. Ser. 13: 377. 1986. Cleistochloa (3; C. rigida, C. sclerachne, C. subjuncea; eastern Queensland, eastern New South Wales), Neurachne (7; N. alopecuroides, N. annularis, N. lanigera, N. minor, N. munroi, N. queenslandica, N. tenuifolia; Australia), Thyridolepis (3; T. mitchelliana, T. multiculmis, T. xerophila; arid regions in Australia). – Homopholis (1; H. belsonii; Australia). – Melinidinae Stapf in D. Oliver (ed. D. Prain), Flora Trop. Afr. 9: 13. 1 Jul 1917 [’Meliniastrae’]. Thuarea (2; T. perrieri: Madagascar; T. involuta: Madagascar, tropical and East Asia to tropical Australia, Melanesia, Micronesia and the Hawaiian Islands), Rupichloa (2; R. acuminata, R. decidua; Bahia, Minas Gerais), Moorochloa (3; M. eruciformis, M. malacodes, M. schoenfelderi; tropical and subtropical America), Tricholaena (4; T. capensis, T. monachne, T. teneriffae, T. vestita; the Canary Islands, the Mediterranean, Africa, Madagascar), Leucophrys (1; L. mesocoma; southern Namibia, Northern Cape), Melinis (22; tropical and southern Africa, Madagascar), Urochloa (18; tropical and subtropical regions on both hemispheres; incl. Chaetium, Eriochloa?), Chaetium (3; C. bromoides, C. cubanum, C. festucoides; southern Mexico, Central America, Cuba, Colombia, Venezuela, Brazil; in Urochloa?), Eriochloa (c 35; tropical and subtropical regions on both hemispheres; in Urochloa?); unplaced Melinidinae: Eccoptocarpha (1; E. obconiciventris; southern tropical Africa), Yvesia (1; Y. madagascariensis; Madagascar). – Panicinae Fr., Flora Scan.: 195. 1835 ['Paniceae']. Louisiella (2; L. fluitans: Sudan, Congo; L. elephantipes: Mexico, Central America, tropical South America), ’Panicum’ (c 300; warm-temperate to tropical regions on both hemispheres; polyphyletic). – Cenchrinae Dumort., Anal. Fam. Plant.: 64. 1829 ['Cenchreae']. Acritochaete (1; A. volkensii; mountains in tropical Africa), Alexfloydia (1; A. repens; Coffs Harbour in New South Wales), Cenchrus (27; warm-temperate to tropical regions in Africa, India and America), Chamaeraphis (1; C. hordeacea; northern Australia), Dissochondrus (1; D. biflorus; the Hawaiian Islands), Holcolemma (3; H. canaliculatum, H. dispar, H. inaequale; tropical East Africa, India, Australia), Hygrochloa (2; H. aquatica, H. cravenii; northern Australia), Ixophorus (1; I. unisetus; Mexico), Paractaenum (2; P. novae-hollandiae, P. refractum; Australia), Paratheria (2; P. glaberrima, P. prostrata; tropical West and Central Africa, Madagascar, Cuba, Brazil), Pseudochaetochloa (1; P. australiensis; northern Australia), Pseudoraphis (7; P. balansae, P. brunoniana, P. jagonis, P. minuta, P. paradoxa, P. sordida, P. spinescens; India to China, Japan and Australia), ’Setaria’ (100–105; warm-temperate to tropical regions on both hemispheres; non-monophyletic?), Setariopsis (2; S. auriculata, S. latiglumis; Arizona, Mexico, Central America, Colombia, Venezuela), Spinifex (4; S. hirsutus, S. littoreus, S. longifolius, S. sericeus; India to East Asia and islands in the Pacific), Stenotaphrum (7; S. clavigerum, S. dimidiatum, S. heiferi, S. micranthum, S. oostachyum, S. secundatum, S. unilaterale; tropical and subtropical regions on both hemispheres), Stereochlaena (4; S. annua, S. caespitosa, S. cameronii, S. tridentata; tropical East to South Africa), Streptolophus (1; S. sagittifolius; Angola), Uranthoecium (1; U. truncatum; tropical Australia), Whiteochloa (6; W. airoides, W. biciliata, W. capillipes, W. cymbiformis, W. multiciliata, W. semitonsa; Aru Islands, tropical Australia), Xerochloa (3; X. barbata, X. imberbis, X. laniflora; Australia), Zuloagaea (1; Z. bulbosa; Arizona, Mexico, Central America, northern South America), Zygochloa (1; Z. paradoxa; arid regions of central Australia). – Unplaced Paniceae: Hydrothauma (1; H. manicatum; Zambia), Kellochloa (2; K. brachyantha, K. verrucosa; eastern and southeastern United States), Oryzidium (1; O. barnardii; Namibia, northern Botswana to Zambia), Thedachloa (1; T. nana; northwestern Australia). – Cosmopolitan, with their highest diversity in tropical and subtropical regions. – Cenchrinae are highly provisional. Most of their subgroups are nested inside ’Setaria’ (Silva & al. 2015).

[Arundinoideae+[Danthonioideae+Chloridoideae]]

Arundinoideae Burmeist., Handb. Naturgesch.: 204. 1837 [‘Arundinaceae’]

20–22/218–220. Ligule hairy. Lemma awned. Starch grains compound. – The clade [Arundineae+Molinieae] may be sister to Micraireae.

[Micrairoideae+[Arundineae+[Crinipedeae+Molinieae]]]

Micrairoideae Pilg. in H. G. A. Engler et K. A. E. Prantl, Nat. Pflanzenfam., ed. 2, 14d: 167. 16 Feb 1956

9/c 185. Micraireae Pilg. in Engler et Prantl, Nat. Pflanzenfam., ed. 2, 14d: 167. 16 Feb 1956. Micraira (16; tropical Australia). – Eriachneae Eck-Boorsb. in Blumea 26: 2. 128. 1980. Eriachne (c 50; Sri Lanka and southern China to southern Malesia and Australia). – Isachneae Benth. in J. Linn. Soc. London, Bot. 19: 30. 24 Dec 1881. Isachne (c 100; tropical and subtropical regions on both hemispheres, especially southern and eastern Asia), Coelachne (11; tropical Africa, Madagascar, tropical Asia from India to New Guinea), Heteranthoecia (1; H. guineensis; tropical Africa), Sphaerocaryum (1; S. malaccense; India to China, Taiwan, Southeast Asia and West Malesia), Limnopoa (1; L. meeboldii; southern India), Hubbardia (2; H. diandra, H. heptaneuron; western India). – Zenkeria (5; Z. elegans, Z. jainii, Z. obtusiflora, Z. sebastinei, Z. stapfii; India, Sri Lanka)? – Pantropical, with their highest diversity in the Old World tropics. Sometimes xerophytic. C4 photosynthesis present in Eriachne. Leaves tristichous. Stomatal subsidiary cells dome-shaped. Ligule with hair fringe. Lemma with or without awn. Embryo small. Starch grains simple. Germination flap present. n = 10. – Micraireae may be sister-group to [Arundineae+[Crinipedeae+Molinieae]] (Hardion & al. 2017). In other analyses they are sister-group to [Panicoideae+[[Aristidoideae+Danthonioideae]+[Arundinoideae+Chloridoideae]]]. According to Soreng & al. (2017), Micrairoideae are sister to Arundinoideae.

Arundineae Dumort., Observ. Gramin. Belg.: 82. Jul-Sep 1824 [’Arundinaceae’]

4/18. Amphipogon (9; Australia), Arundo (6; A. collina, A. donax, A. formosana, A. mediterranea, A. micrantha, A. plinii; the Mediterranean, Southwest Asia to India, the Ryukyu Islands, Taiwan, the Philippines), Monachather (1; M. paradoxa; Australia), Dregeochloa (2; D. calviniensis, D. pumila; southern Namibia, Northern Cape). – Tropical and subtropical regions of the Old World.

Crinipedeae Hardion in Plant Syst. Evol. 303: 1337. Dec 2017

4/8. Crinipes (2; C. abyssinicus, C. longifolius; Sudan, Ethiopia, Uganda), Styppeiochloa (3; S. catherineana, S. gynoglossa, S. hitchcockii; mountains in southern to southeastern tropical Africa, Madagascar), Elytrophorus (2; E. globularis, E. spicatus; tropical and southern Africa, the Indian Subcontinent, Yunnan, Hainan, Indochina, Lesser Sunda Islands, tropical Australia), Pratochloa (1; P. walteri; Namibia). – Tropical and subtropical regions of the Old World, with their largest diversity in southern and eastern Africa.

Molinieae Jirásek in Preslia 38: 33. Jan-Mar 1966

3–5/7–9. Hakonechloa (1; H. macra; Japan), Leptagrostis (1; L. schimperiana; Ethiopia)?, Molinia (2; M. caerulea, M. japonica; Europe, temperate Asia), Phragmites (4; P. australis, P. japonicus, P. karka, P. mauritianus; cosmopolitan), Piptophyllum (1; P. welwitschii; Angola)? – Cosmopolitan, with their largest diversity in tropical and subtropical regions in the Old World. Elongate microhairs Panicoid type (with slender thin-walled cap cells). Hilum short. Epiblasts absent. n = 6, 9, 12.

[Danthonioideae+Chloridoideae]

Hilum short, punctate.

Danthonioideae N. P. Baker et P. H. Linder in Ann. Missouri Bot. Gard. 88: 421. 17 Sep 2001

19/c 285. Merxmuellera (7; M. ambalavaoensis, M. davyi, M. drakensbergensis, M. grandiflora, M. macowanii, M. stereophylla, M. tsaratananensis; tropical and southern Africa, Madagascar), Capeochloa (3; C. arundinacea, C. cincta, C. setacea; southern Africa), Geochloa (3; G. decora, G. lupulina, G. rufa; tropical and southern Africa), Pentameris (c 80; tropical and subtropical Africa, Madagascar, southern and eastern Asia), Chionochloa (26; 23 spp. in New Zealand; C. antarctica: Auckland and Campbell Islands; C. howensis: Lord Howe; C. archboldii: New Guinea), Cortaderia (20; South America; non-monophyletic?), Chaetobromus (1; C. involucratus; southern Namibia, Northern and Western Cape), Pseudopentameris (3; P. brachyphylla, P. caespitosa, P. macrantha; mountains in Western Cape), Austroderia (5; A. fulvida, A. richardi, A. splendens, A. toetoe: New Zealand; A. turbaria: Chatham Islands), Notochloe (1; N. microdon; Blue Mountains in New South Wales), Plinthanthesis (3; P. paradoxa, P. rodwayi, P. urvillei; southeastern Australia), Chimaerochloa (1; C. archboldii; tropical Asia to New Guinea), Danthonia (27; Europe, the Mediterranean to Ukraine and the Caucasus, North Africa, the Russian Far East, Greenland, North, Central and South America, the West Indies), Tenaxia (8; Africa, temperate to tropical Asia), Schismus (5; S. arabicus, S. barbatus, S. inermis, S. scaberrimus, S. schismoides; the Mediterranean, Africa, southwestern Asia to northwestern India and China, especially southern Africa), Tribolium (14; southern Namibia, Northern, Western and Eastern Cape), Rytidosperma (c 75; New Guinea, Australia, New Zealand, South America)); unplaced: Phaenanthoecium (1; P. koestlinii; mountains in northeastern tropical Africa). – Danthonioideae incertae sedis: Danthonidium (1; D. gammiei; India). – Cosmopolitan, with their highest diversity in the Southern Hemisphere, especially in southern and eastern Africa and Australia (few species in Southeast Asia and Malesia). Leaves usually symmetrical. Ligules ciliated. Foliar anatomy festucoid. Prophylls bifid. Spikelets festucoid. Lemma awn trifid or with three awns. Stylar bases much separated. Synergids haustorial, growing out through micropyle. n = 6, 7, 9. – In some analyses Danthonioideae are sister-group to Aristidoideae.

Chloridoideae Burmeist., Handb. Naturgesch.: 205. 1837 [’Chlorideae’]

125–126/1.405–1.420. Centropodieae are sister-group to the remaining Chloridoideae, according to Peterson & al. (2011). – Unplaced Chloridoideae Gossweilerochloa (1; G. delicatula; Angola), Indopoa (1; I. paupercula; India), Lepturopetium (1; L. kuniensis; New Caledonia, Marshall Islands, Cocos Island), Myriostachya (1; M. wightiana; southern India, Sri Lanka, Southeast Asia), Pogonochloa (1; P. greenwayi; southern tropical Africa), Pseudozoysia (1; P. sessilis; Somalia), Silentvalleya (1; S. nairii; India). – Tropical, subtropical and warm-temperate regions, dry habitats (especially in Africa and Australia). C₄ photosynthesis usually present (phosphoenolpyruvate carboxykinase subtype). Microhairs usually with inflated hemispherical thick-walled distal cells and elongate secretory basal cell with internal membranes (’Chloridoid type’; in Neyraudia ‘Panicoid type’). Leaves usually symmetrical. Spikelets disarticulating above glumes. Hilum short. Epiblast usually present. Mesocotyl present. Leaf margins of embryo not overlapping. Plastid gene rpl16 intron with 4 bp insertion. n = (6–8) 9, 10.

[Centropodieae+[Triraphideae+[Eragrostideae+[Zoysieae+Chlorideae]]]]

Centropodieae P. M. Peterson, N. P. Barker et H. P. Linder in Taxon 60(4): 1118. Aug 2011

2/6. Centropodia (4; C. forskalii, C. fragilis, C. glauca, C. mossamedensis; North Africa, the Middle East to India), Ellisochloa (2; E. papposa, E. rangei; southern Africa). – Southern Africa and one species in North Africa eastwards to India. C₃ (Ellisochloa) and C₄ (Centropodia) photosynthesis present. Microhairs probably absent. Haustorial synergids absent. x = 9, 12.

[Triraphideae+[Eragrostideae+[Zoysieae+Chlorideae]]]

Triraphideae P. M. Peterson in Mol. Phylogen. Evol. 55(2): 591. May 2010

4/14. Habrochloa (1; H. bullockii; Central Africa), Neyraudia (4; N. arundinacea, N. curvipes, N. montana, N. reynaudiana; tropical and subtropical regions in Africa and Asia), Nematopoa (1; N. longipes; Zambia, Zimbabwe; in Triraphis?), Triraphis (8; tropical Africa, the Arabian Peninsula, Australia, one species, T. devia, in central Brazil). – Tropical and subtropical regions in the Old World, central Brazil.

[Eragrostideae+[Zoysieae+Chlorideae]]

Eragrostideae Stapf in W. H. Harvey et O. W. Sonder (ed. W. T. Thiselton-Dyer), Fl. Cap. 7: 316. Jul 1898 [’Eragrosteae’]

13/410–415. Cotteinae Reeder in Madroño 18: 25. 10 Feb 1965. Cottea (1; C. pappophorides; southern United States to central Mexico, Ecuador to Argentina), Enneapogon (c 25; dry warmer regions on both hemispheres), Kaokochloa (1; K. nigrirostris; northwestern Namibia), Schmidtia (2; S. kalahariensis, S. pappophoroides; Africa, Cape Verde Islands, Pakistan). – Unioliinae Clayton in Kew Bull. 37: 417. 30 Dec 1982. Entoplocamia (1; E. aristulata; Namibia), Fingerhuthia (2; F. africana, F. sesleriiformis; southern tropical and southern Africa, the Arabian Peninsula, Afghanistan to India), Tetrachaete (1; T. elionuroides; Ethiopia, Tanzania, the Arabian Peninsula), Tetrachne (1; T. dregei; Eastern Cape, Free State, Lesotho, Pakistan), Uniola (5; U. condensata, U. paniculata, U. peruviana, U. pittieri, U. virgata; southern United States, Mexico, Central America to Ecuador). – Eragrostidinae J. Presl in C. Presl, Reliq. Haenk. 1: 273. Jan-Jun 1830 [’Eragrostideae’]. Cladoraphis (2; C. cyperoides, C. spinosa; Namibia, Northern and Western Cape), Eragrostis (c 350; tropical to warm-temperate regions on both hemispheres; non-monophyletic), Richardsiella (1; R. eruciformis; Zambia), Steirachne (2; S. barbata, S. diandra; northeastern South America). – Tropical and subtropical (to warm-temperate) regions on both hemispheres.

[Zoysieae+Chlorideae]

Zoysieae Benth. in J. Linn. Soc. London, Bot. 19: 29. 24 Dec 1881

4–5/185–190. Zoysiinae Benth., Flora Austral. 7: 453, 505. 23-30 Mar 1878 [’Zoysieae’]. Zoysia (9; Mauritius to Polynesia), Urochondra (1; U. setulosa; Sudan and Somalia to Sind). – Sporobolinae Benth. in J. Linn. Soc. London, Bot. 19: 30. 24 Dec 1881 [’Sporoboleae’]. Psilolemma (1; P. jaegeri; East Africa), ’Sporobolus’ somalensis (East Africa); Sporobolus (c 175; warm-temperate to tropical regions on both hemispheres). – Subcosmopolitan.

Cynodonteae Dumort., Observ. Gramin. Belg.: 83. Jul-Sep 1824 [’Cynodoneae’]

95/785–800. Eleusininae Dumort., Anal. Fam. Plant.: 63. 1829 ['Eleusineae']. Dinebra (6; D. haareri, D. marquisensis, D. perrieri, D. polycarpha, D. retroflexa, D. somalensis; tropical East and northeastern Africa, Madagascar, southwestern Asia to India, southern United States), Eleusine (10; Africa, Madagascar, South America), Diplachne (5; D. cuspidata, D. fascicularis, D. festuciformis, D. fusca, D. gigantea; warm-temperate to tropical regions on both hemispheres), Leptochloa (27; eastern tropical and southern Africa, Australia, the Marquesas Islands, North and South America), Astrebla (4; A. elymoides, A. lappacea, A. pectinata, A. squarrosa; Australia), Disakisperma (4; D. dubium, D. eleusine, D. obtusiflorum, D. yemenicum; southern and tropical eastern Africa, southern United States), Enteropogon (17; tropical and subtropical regions on both hemispheres), Chloris (60–65; warm-temperate to tropical regions on both hemispheres), Tetrapogon (11; the Canary Islands, Africa, the Arabian Peninsula, the Middle East to India and Central Asia), Lepturus (c 15; coasts in East Africa, South Africa, Sri Lanka, Australia to Polynesia), Microchloa (6; M. altera, M. annua, M. caffra, M. ensifolia, M. indica, M. kunthii; tropical and southern Africa, tropical Asia to tropical Australia, southwestern United States to Argentina and Chile), Micrachne (5; M. fulva, M. obtusiflora, M. patentiflora, M. pilosa, M. simonii; Central and tropical East Africa), Eustachys (15; United States, Mexico, Central America, the West Indies, South America, one species, E. paspaloides, in eastern and southern Africa, Oman and Yemen, one species, E. tenera, in Southeast Asia to Taiwan and New Guinea), Chrysochloa (4; C. hindsii, C. hubbardiana, C. orientalis, C. subaequigluma; tropical Africa), Stapfochloa (6; S. berroi, S. canterae, S. ciliata, S. elata, S. lamproparia, S. parvispicula; tropical and subtropical Africa), Cynodon (10; tropical and subtropical regions on both hemispheres); unplaced Eleusininae: Afrotrichloris (2; A. hyaloptera, A. martinii; Somalia; in Schoenefeldia?), Schoenefeldia (2; S. gracilis, S. transiens; tropical and southern Africa, Madagascar, India; incl. Afrotrichloris?), Apochiton (1; A. burttii; Tanzania), Austrochloris (1; A. dichanthoides; Queensland), ’Coelachyrum’ (1–8; C. brevifolium, C. lagopoides, C. longiglume, C. piercei, C. poiflorum; tropical and southern Africa, the Arabian Peninsula, Pakistan; polyphyletic), Daknopholis (1; D. boivinii; East Africa, Madagascar, Aldabra), Oxychloris (1; O. scariosa; Australia), Harpochloa (1; H. falx; southern tropical and southern Africa), Neostapfiella (3; N. chloridiantha, N. humbertiana, N. perrieri; Madagascar), Pommereulla (1; P. cornucopiae; southern India, Sri Lanka), Rheochloa (1; R. scabiflora; central Brazil), Sclerodactylon (1; S. macrostachyum; coast of East Africa, islands in the Indian Ocean)? – Dactylocteniinae P. M. Peterson, Romasch. et Y. Herrera in Taxon 65(6): 1276. Dec 2016. Acrachne (3; A. henrardiana, A. perrieri, A. racemosa; tropical and southern Africa, Madagascar, Southeast Asia, Australia), Dactyloctenium (13; tropical and subtropical regions on both hemispheres), Brachychloa (2; B. fragilis, B. schiemanniana; KwaZulu-Natal, Mozambique), Neobouteloua (2; N. lophostachya, N. pauciracemosa; Chile, Argentina). – Triodiinae Benth. in J. Linn. Soc. London, Bot. 19: 30. 24 Dec 1881 ['Triodieae']. Triodia (c 70; Australia). – Aeluropodinae P. M. Peterson in Mol. Phylogen. Evol. 55: 591. May 2010. Aeluropus (6; A. badghyzii, A. laciniatus, A. lagopoides, A. littoralis, A. macrostachyus, A. pilosus; the Mediterranean, southwestern Asia to India and northern China), Odyssea (2; O. mucronata, O. paucinervis; coasts of the Red Sea, tropical and southern Africa). – Orininae P. M. Peterson, Romasch. et Y. Herrera in Taxon 65(69): 1277. Dec 2016. Cleistogenes (c 13; southern Europe, Turkey to temperate East Asia), Orinus (6; O. alticulmus, O. anomala, O. kokonorica, O. longiglumis, O. thoroldii, O. tibeticus; Central Asia, the Himalayas, Tibet, western China). – Orcuttiinae P. M Peterson et Columbus in Aliso 23: 592. Dec 2007. Neostapfia (1; N. colusana; California), Orcuttia (5; O. californica, O. inaequalis, O. pilosa, O. tenuis, O. viscida; California). – Zaqiqahinae P. M. Peterson, Romaschenko et Herrera Arrieta in Taxon 65(6): 1279. Dec 2016. Zaqiqah (1; Z. mucronata; Ethiopia, Somalia, Socotra, southwestern Arabian Peninsula). – Gouiniinae P. M. Peterson et Columbus in Aliso 23: 592. Dec 2007. Triplasis (2; T. americana, T. purpurea; central and eastern United States to Costa Rica), Triplasiella (1; T. eragrostoides; southern United States, Mexico, the West Indies, Venezuela), Schenckochloa (1; S. barbata; northern Brazil), Vaseyochloa (1; V. multinervosa; Texas), Tridentopsis (2; T. eragrostoides, T. mutica; southwestern United States, northern Mexico, the West Indies), Gouinia (12; Mexico, Central America, Cuba, Hispaniola, tropical South America). – Cteniinae P. M. Peterson, Romaschenko et Herrera Arrieta in Taxon 63(2): 283. Apr 2014. Ctenium (20; tropical and subtropical Africa, Madagascar, North America to Bolivia and Paraguay). – Trichoneurinae P. M. Peterson, Romaschenko et Herrera Arrieta in Taxon 63(2): 284. Apr 2014. Trichoneura (8; Africa, the Arabian Peninsula, Texas, Mexico, the Galápagos Islands, Peru, Chile). – Hubbardochloinae Auquier in Bull. Jard. Bot. Natl. Belg. 50: 246. 1980. Dignathia (5; D. aristata, D. ciliata, D. gracilis, D. hirtella, D. villosa; tropical East Africa, western India), Leptothrium (3; L. lineare, L. rigidum, L. senegalense; Senegal to Pakistan, the West Indies), Leptocarydion (1; L. vulpiastrum; northern Namibia, KwaZulu-Natal, Mpumalanga, Northern Province, Botswana, East Africa), Bewsia (1; B. biflora; northern Namibia, northern South Africa, Swaziland), Gymnopogon (14; southern and eastern United States, Mexico, Central America, the West Indies, tropical South America, one species, G. delicatulus, in Southeast Asia), Lophacme (2; L. digitata, L. parva; southern tropical and southern Africa), Hubbardochloa (1; H. gracilis; mountains in Central Africa), Decaryella (1; D. madagascariensis; Madagascar). – Perotidinae P. M. Peterson, Romaschenko et Herrera Arrieta in Taxon 63(2): 284. Apr 2014. Mosdenia (1; M. leptostachys; northern South Africa), Perotis (15; tropical and subtropical Africa, southern Madagascar, temperate to tropical Asia east to China, New Guinea and northern Australia), Trigonochloa (2; T. rupstris, T. uniflora; tropical, southern and eastern Africa, the Arabian Peninsula to India and Sri Lanka). – Farragininae P. M. Peterson, Romaschenko et Herrera Arrieta in Taxon 63(2): 283. Apr 2014. Farrago (1; F. racemosa; Tanzania), Craspedorhachis (9; southern tropical and southern Africa). – Tripogoninae Stapf in D. Oliver (ed. D. Prain), Flora Trop. Afr. 9: 22. 1 Jul 1917 ['Schizachyriastrae']. Desmostachya (1; D. bipinnata; North Africa, southwestern Asia to India and Southeast Asia), Melanocenchris (3; M. abyssinica, M. jacquemontii, M. monoica; Chad and tropical northeastern Africa to India and Sri Lanka), Halopyrum (1; H. mucronatum; coasts along the Indian Ocean), Eragrostiella (6; E. bifaria, E. brachyphylla, E. collettii, E. leioptera, E. lolioides, E. nardoides; East Africa, Sri Lanka to northern Australia), Tripogonella (3; T. minima: tropical and southern Africa, Madagascar; T. loliiformis: Australia; T. spicata: North America, Central America, the West Indies, South America), Oropetium (6; O. aristatum, O. capense, O. minimum, O. roxburghianum, O. thomaeum, O. villosulum; arid and semiarid regions in tropicl West Africa to India and Southeast Asia), Tripogon (c 50; tropical and southern Africa, Madagascar, India to Australia). – Pappophorinae Dumort., Anal. Fam. Plant.: 63. 1829 ['Pappophoreae']. Neesiochloa (1; N. barbata; northeastern Brazil), Pappophorum (7; P. bicolor, P. caespitosum, P. hassleri, P. krapovickasii, P. mucronulatum, P. pappiferum, P. philippianum; southern United States, Mexico, Central America, the West Indies, tropical South America), Tridens (15; southern Canada, eastern and southern United States, Mexico, Central America, Cuba, tropical South America). – Traginae P. M. Peterson et Columbus in Aliso 23: 592. Dec 2007. Polevansia (1; P. rigida; eastern Eastern Cape, Lesotho), Pogononeura (1; P. biflora; tropical East Africa), Monelytrum (1; M. luederitzianum; southern Angola, Namibia), Tragus (8; tropical to warm-temperate regions in the Old World east to China, Australia and New Caledonia, Argentina), Orthacanthus (1; O. pedunculatus; southern Africa), Willkommia (4; W. annua, W. newtonii, W. sarmentosa, W. texana; southern tropical and southern Africa, Texas). – Muhlenbergiinae Pilg. in H. G. A. Engler et K. A. E. Prantl, Nat. Pflanzenfam., ed. 2, 14d: 168. 16 Feb 1956. Muhlenbergia (155–160; Canada, United States, Mexico, Central America, tropical and subtropical South America, few species in Central to Northeast Asia, the Himalayas and Southeast Asia to New Guinea). – Sohnsiinae P. M. Peterson, Romasch. et Y. Herrera in Phytoneuron 2017-44: 6. 18 Jul 2017. Sohnsia (1; S. filifolia; Mexico). – Scleropogoninae Pilg. in H. G. A. Engler et K. A. E. Prantl, Nat. Pflanzenfam., ed. 2, 14d: 167. 16 Feb 1956. Scleropogon (1; S. brevifolius; southwestern United States, Mexico, Chile, Argentina), Swallenia (1; S. alexandrae; Eureka Valley Sand Dunes in Inyo County in California), Blepharidachne (4; B. benthamiana, B. bigelovii, B. hitchcockii, B. kingii; western United States, Argentina), Erioneuron (2; E. avenaceum, E. pilosum; southern United States, Mexico, Peru, Bolivia, Argentina), Munroa (5; M. andina, M. argentina, M. decumbens, M. mendocina, M. squarrosa; western and southwestern United States, northen Mexico, Chile, Argentina). – Jouveinae P. M. Peterson, Romasch. et Y. Herrera in Phytoneuron 2017-44: 5. 18 Jul 2017. Jouvea (2; J. pilosa, J. straminea; Baja California to Panamá). – Allolepiinae P. M. Peterson, Romasch. et Y. Herrera in Phytoneuron 2017-44: 4. 18 Jul 2017. Allolepis (1; A. texana; southern United States, Mexico). – Hilariinae P. M. Peterson et Columbus in Aliso 23: 592. Dec 2007. Hilaria (10; southern United States, Mexico, Guatemala). – Kaliniinae P. M. Peterson, Romasch. et Y. Herrera in Phytoneuron 2017-44: 5. 18 Jul 2017. Kalinia (1; K. obtusiflora; southwestern United States to central Mexico). – Monanthochloinae Pilg. ex Potztal in Willdenowia 5: 472. Nov 1969. Distichlis (11; Canada, United States, Mexico, the West Indies, South America, one species, D. distichophylla, in Australia). – Boutelouinae Stapf in D. Oliver (ed. D. Prain), Flora Trop. Afr. 9: 22. 1 Jul 1917. Bouteloua (c 40; southern Canada, United States, Mexico, Central America, the West Indies, tropical South America). – Cynodonteae incertae sedis: Kampochloa (1; K. brachyphylla; Zambia, Angola; possibly close to Ctenium?), Lepturidium (1; L. insulare; Isla de la Juventud West of Cuba), Vietnamochloa (1; V. aurea; Vietnam). – Tropical and subtropical (to warm-temperate) regions on both hemispheres.

[Oryzoideae+[Bambusoideae+Pooideae]] ("The BEP clade")

Phytoliths in Oryzoideae and Bambusoideae transverse-bilobate, in Brachyelytrum and Pooideae axial-bilobate or axial-polylobate. x = 12. Endosperm softness gene present. – Transverse-bilobates were possibly lost in the common ancestor of Brachyelytrum and Pooideae. Alternatively, this feature is a synapomorphy of [Oryzoideae+Bambusoideae] (Rudall & al. 2014).

Oryzoideae Kunth ex Beilschm., Flora 16(2): 52, 109. 1833

19/115. Widely distributed, especially on the Southern Hemisphere. Microhairs usually present. Fusoid cells absent. Longitudinal walls of epidermal cells sometimes straight. Spikelet developing basipetally. Glumes minute. Stamens (one to) six. Stylodia almost entirely separate. n = (10, 15).

Streptogyneae C. E. Hubb. ex Calderón et Soderstr. in Smithsonian Contr. Bot. 44: 18. 13 Feb 1980

1/2. Streptogyna (2; S. crinita: tropical West and Central Africa; S. americana: southern Mexico, Central America, tropical South America). – Epidermal papillae absent. Spikelet several-flowered. Lemma with awns. Lodiculae three. Multicellular microhairs present on lodiculae in S. crinita. Stamens two. Stigmas two or three. Hilum linear extending along entire fruit. Starch grains aggregated or simple. Embryo without mesocotyl internode. Epiblast prominent. First seedling leaf with strongly overlapping margins. n = 12. – Streptogyna may be sister to the clade [Panicoideae+Arundinoideae] (Peterson & al. 2011), although Soreng & al. (2012) assigned Streptogyna (Streptogyneae) to Ehrhartoideae.

Ehrharteae Nevski in Trudy Bot. Inst. Akad. Nauk S.S.S.R., ser. 1, Fl. Sist. Vyssh. Rast. 4: 227. 1937

1/c 35. Ehrharta (c 35; South Africa to Ethiopia, the Mascarenes, Malesia, the Philippines, Java to Australia and islands in the Pacific, mountains in New Zealand, Western and Eastern Cape). – Tropical and subtropical regions in the Old World. Arm cells absent. Epiblasts absent at least in Ehrharta. Roots in Ehrharta at scutellar node.

Oryzeae Dumort., Observ. Gramin. Belg.: 83. Jul-Sep 1824

13–14/72–73. Oryzinae Griseb. in C. F. von Ledebour, Flora Ross. 4: 465. Jun 1853 [‘Oryzeae‘]. Leersia (18; warm-temperate to tropical regions on both hemispheres), Maltebrunia (4; M. leersioides, M. letestui, M. maroana, M. schliebenii; tropical to southern Africa, Madagascar), Oryza (19; tropical and subtropical regions on both hemispheres; incl. Porteresia?), Prosphytochloa (1; P. prehensilis; Eastern Cape, KwaZulu-Natal, Mpumalanga, Northern Province). – Zizaniinae Benth. in J. Linn. Soc. London, Bot. 19: 54. 24 Dec 1881. Chikusichloa (3; C. aquatica, C. brachyathera, C. mutica; China, Japan, the Ryukyu Islands, Sumatra), Hygroryza (1; H. aristata; southern and Southeast Asia to China), Luziola (12; southern United States, Mexico, Central America, the West Indies, tropical South America), Potamophila (1; P. parviflora; northern New South Wales), Rhynchoryza (1; R. subulata; Paraguay, Argentina), Zizania (4; Z. aquatica, Z. latifolia, Z. palustris, Z. texana; eastern India to East Asia, North America), Zizaniopsis (5; Z. bonariensis, Z. killipii, Z. microstachya, Z. miliacea, Z. villanensis; southern United States, Mexico, tropical South America to Argentina). – Phyllorachideae C. E. Hubb.in Hooker’s Icon. Plant. 34: t. 3386, p. 5. Mar 19. Humbertochloa (2; H. bambusiuscula, H. greenwayi; tropical East Africa, Madagascar), Phyllorachis (1; P. sagittata; southern tropical Africa). – Subcosmopolitan. Arm cells present. First seedling leaf only sheath. – Suddia (1; S. sagittifolia; Sudan) may belong in Phyllorachideae.

[Bambusoideae+Pooideae]

Epiblast present. First seedling leaf with overlapping margins.

Bambusoideae Luerss., Grundz. Bot., ed. 5: 451. Jun 1893 [‘Bambusaceae’]

118/1.555–1.580. Tropical, subtropical and temperate regions, often forests. Woody, sometimes tree-like. Mesophyll differentiated into palisade and spongy tissues. Fusoid cells and strongly asymmetrically invaginated arm cells present. Microhairs elongate, with slender thin-walled cap cells (Panicoid type). Leaves with pseudopetiole, often with inner and outer distal ligules. Culm leaves often very different from other leaves (heterophylly). Lodiculae two or three. Stamens (two to) six (to c. 140). Filaments sometimes connate at base. Stigmas (one or) two or three. Fruit sometimes a berry. First seedling leaf only sheath. n = 7, 9–12. – Woody and some herbaceous species have a synchronized anthesis, and many are monocarpic (hapaxanthic); this may be a plesiomorphy in Bambusoideae. The woody temperate species perhaps form a sister-group to the remainder.

Arundinarieae Hackel in Engler et Prantl, Nat. Pflanzenfam. II, 2: 92, 93. Nov 1887

30/c 550. Ampelocalamus (13; the Himalayas to China), Bergbambos (1; B. tessellata; southern and eastern South Africa), Thamnocalamus (4; T. chigar, T. spathiflorus, T. tesellatus, T. unispiculatus; the Himalayas), ’Indocalamus’ (c 35; China, Japan; polyphyletic), Fargesia (c 80; eastern Himalayas, Tibet, China), Phyllostachys (c 50; the Himalayas, China to Japan), Gaoligongshania (1; G. megalothyrsa; southern Yunnan), Oldeania (1; O. alpina; tropical East Africa, Cameroon), Chimonocalamus (16; eastern Himalayas to Yunnan), Kuruna (7; K. debilis, K. densifolia, K. floribunda, K. scandens, K. serrulata, K. walkeriana, K. wightiana; southern India, Sri Lanka), Ferrocalamus (2; F. fibrillosus, F. strictus; southern Yunnan), Shibataea (7; S. chiangshanensis, S. chinensis, S. hispida, S. kumasasa, S. lancifolia, S. nanpingensis, S. strigosa; southeastern China, southwestern Japan), Acidosasa (11; southern China, Vietnam), Pleioblastus (23; China, the Korean Peninsula, Japan, Vietnam), Sasa (c 60; temperate East Asia, with their highest diversity in Japan), Arundinaria (3; A. appalachiana, A. gigantea, A. tecta; eastern United States), Bashania (4; B. abietina, B. fansipanensis, B. fargesii, B. qingchengshanensis; China; in Arundinaria?); unplaced Arundinarieae: Chimonobambusa (42; the Himalayas, China, Japan), Drepanostachyum (13; the Himalayas to southwestern China; incl. Himalayacalamus?), Himalayacalamus (9; the Himalayas to southwestern China; in Drepanostachyum?), Gelidocalamus (13; southeastern China), Indosasa (20; southern China, Vietnam), Oligostachyum (17; China), Pseudosasa (21; China, the Korean Peninsula, Japan), Sarocalamus (3; S. faberi, S. racemosus, S. spanostachyus; southern and eastern Himalayas to southwestern China), Sasaella (13; Japan), Semiarundinaria (7; S. densiflora, S. fastuosa, S. fortis, S. kagamiana, S. shapoensis, S. sinica, S. yashadake; China, Japan; hybrid origin), Sinobambusa (13; southwestern and southern China, northern Vietnam), Vietnamocalamus (1; V. catbaensis; Vietnam), ’Yushania’ (c 70; southern Himalayas to southern China and Peninsular Thailand, Taiwan, Indochina, the Philippines, northern Borneo; Madagascar; polyphyletic). – The Himalayas to East Asia, North America, with their largest diversity in China. – The Malagasy species of ’Yushania’ should probably be separated from the rest. Ampelocalamus may be sister to the remaining Arundinarieae. Arundinarieae aresister-group to[Bambuseae+Olyreae].

Bambuseae Kunth ex Dumort., Anal. Fam. Plant.: 63. 1829 [‘Bambusaceae’]

67/890–915. Melocanninae Benth. in J. Linn. Soc. London, Bot. 19: 31. 24 Dec 1881 ['Melocanneae']. ’Cephalostachyum’ (14; Madagascar, eastern Himalayas, Tibet, Yunnan, Burma, Southeast Asia, the Philippines; polyphyletic), Melocanna (2; M. arundina, M. baccifera; India, Southeast Asia, Malesia), Schizostachyum (c 65; southeastern and northeastern India, Sri Lanka, the Himalayas, China, Southeast Asia), Annamocalamus (1; A. kontumensis; Vietnam). – Hickeliinae A. Camus in H. Lecomte, Compt. Rend. Hebd. Seances Acad. Sci. 179: 480. 1 Sep 1924. Hickelia (4; H. africana, H. alaotrensis, H. madagascariensis, H. perrieri; Tanzania, Madagascar), Nastus (c 15; Madagascar, Réunion, Southeast Asia, Malesia to New Guinea), Sirochloa (1; S. parvifolia; Madagascar, the Comoros), Cathariostachys (2; C. capitata, C. madagascariensis; Madagascar), Decaryochloa (1; D. diadelpha; Madagascar), Hitchcockella (1; H. baronii; Madagascar), Perrierbambus (2; P. madagascariensis, P. tsarasaotrensis; Madagascar), Sokinochloa (7; S. australis, S. bosseri, S. brachyclada, S. chapelieri, S. chiataniae, S. perrieri, S. viguieri; Madagascar), Valiha (2; V. diffusa, V. perrieri; Madagascar). – Bambusinae J. Presl in C. Presl, Reliq. Haenk. 1: 256. Jan-Jun 1830 ['Bambusaceae']. Bonia (5; B. amplexicaulis, B. laevigata, B. parvifloscula, B. saxatilis, B. tonkinensis; southern China, Vietnam), Neomicrocalamus (4; N. andropogonifolius, N. dongvanensis, N. prainii, N. yunnanensis; East Asia), Soejatmia (1; S. ridleyi; the Malay Peninsula), Pseudoxytenanthera (4; P. bourdillonii, P. monadelpha, P. ritcheyi, P. stocksii; southern India, Sri Lanka), Maclurochloa (1; M. montana; the Malay Peninsula), ’Melocalamus’ (13; India, southern China, Southeast Asia; polyphyletic), ’Dendrocalamus’ (55–60; India, Sri Lanka, Southeast Asia to China and the Philippines; polyphyletic), ‘Gigantochloa‘ (c 70; India, Sri Lanka, Southeast Asia, Malesia to New Guinea; non-monophyletic), ’Bambusa’ (145–150; tropical and subtropical regions in Asia, Australia and America; polyphyletic; incl. Fimbribambusa?), Fimbribambusa (2; F. horsfieldii, F. microcephala; Central and East Malesia; in Bambusa?), Oreobambos (1; O. buchwaldii; tropical East Africa), Oxytenanthera (1; O. abyssinica; tropical Africa), Vietnamosasa (3; V. ciliata, V. darlacensis, V. pusilla; southern Vietnam), Thyrsostachys (2; T. oliveri, T. siamensis; Burma and Thailand to China), Cochinchinochloa (1; C. braiana; southern Vietnam), Yersinochloa (1; Y. dalatensis; southern Vietnam), Temochloa (1; T. liliana; southern Thailand). – Racemobambosinae Stapleton in Edinburgh J. Bot. 51: 323. Oesterr. Bot. 1994. Racemobambos (19; India, East and Southeast Asia, Malesia to New Guinea), Chloothamnus (10; Southeast Asia, Malesia to New Guinea), Widjajachloa (2; New Guinea). – Dinochloinae K. M. Wong et W. L. Goh, Sandakania 22. 2016. Mullerochloa (1; M. moreheadiana; northern Queensland), Dinochloa (c 35; Burma to the Philippines), Neololeba (3; N. atra, N. glabra, N. hirsuta; Central Malesia to New Guinea and tropical Australia), Parabambusa (2; P. kaini, P. marginata; New Guinea), Sphaerobambos (3; S. hirsuta, S. philippinensis, S. subtilis; Malesia), Cyrtochloa (7; C. fenixii, C. hirsuta, C. luzonica, C. major, C. mindoroensis, C. puser, C. toppingii; the Philippines). – Greslaniinae K. M. Wong et W. L. Goh, Sandakania 22. 2016. Greslania (4; G. circinnata, G. montana, G. multiflora, G. rivularis; New Caledonia). – Holttumochloinae K. M. Wong et W. L. Goh, Sandakania 22. 2016. Kinabaluchloa (1; K. wrayi; Thailand, Vietnam, the Malay Peninsula), Holttumochloa (3; H. korbuensis, H. magica, H. pubescens; the Malay Peninsula, Borneo), Nianhochloa (1; N. bidoupensis; Mount Bidoup in southern Vietnam). – Temburongiinae K. M. Wong, Sandakania 22. 2016. Temburongia (1; T. simplex; Temburong River in Brunei). – Chusqueinae Soderstr. et R. P. Ellis in T. R. Soderstrom et al. (eds.), Grass Syst. Evol.: 238. Mar 1988. Chusquea (155–160; Mexico, Central America, tropical South America including the Andes, Chile, western Argentina). – Guaduinae Soderstr. et R. P. Ellis in T. R. Soderstrom et al. (eds.), Grass Syst. Evol.: 238. Mar 1988. Tibisia (3; T. angustifolia, T. farcta, T. pinifolia; the West Indies), Apoclada (1; A. simplex; Brazil), Eremocaulon (4; E. amazonicum, E. asymmetricum, E. aureofimbriatum, E. capitatum; Brazil), Guadua (26; Mexico, Central America, tropical South America), Olmeca (5; O. clarkiae, O. fulgor, O. recta, O. reflexa, O. zapotecorum; Mexico), Otatea (10; Mexico, Central America, northern Colombia). – Arthrostylidiinae Soderstr. et R. P. Ellis in T. R. Soderstrom et al. (eds.), Grass Syst. Evol.: 238. Mar 1988. Glaziophyton (1; G. mirabile; mountains in eastern Brazil), Cambajuva (1; C. ulei; southern Brazil), Actinocladum (1; A. verticillatum; Brazil), Athroostachys (2; A. capitata, A. shepherdiana; eastern Brazil), Merostachys (c 45; Central America, tropical South America), Filgueirasia (2; F. arenicola, F. cannavieira; Brazil), Alvimia (3; A. auriculata, A. gracilis, A. lancifolia; Brazil), Atractantha (5; A. amazonica, A. aureolanata, A. cardinalis, A. falcata, A. radiata, A. shepherdiana; eastern Brazil), Colanthelia (7; C. burchellii, C. cingulata, C. distans, C. intermedia, C. lanciflora, C. macrostachya, C. rhizantha; southern Brazil, northeastern Argentina), Aulonemia (c 45; Central America, tropical South America), ’Arthrostylidium’ (30–35; southern Mexico, Central America, the West Indies, tropical South America; non-monophyletic; incl. Rhipidocladum?), Rhipidocladum (16; southern Mexico, Central America, tropical South America; in Arthrostylidium?), Elytrostachys (2; E. clavigera, E. typica; Central America, Colombia, Venezuela), Didymogonyx (2; D. geminatum, D. longispiculatum; Colombia, Venezuela); unplaced Arthrostylidiinae: Myriocladus (12; the Roraimas in Venezuela). – Unplaced Bambuseae: Ruhooglandia (1; New Guinea). – Pantropical, the Himalayas, East Asia, New Caledonia, with their largest diversity in tropical Asia and tropical South America. – Bambuseae are sister-group to Olyreae. The American clade [Chusqueinae+[Guaduinae+Arthrostylidiinae]] is sister-group the the remaining Bambuseae (the Paleotropical clade; Zhou & al. 2017).

Olyreae Kunth ex Spenn. Fl. Friburg. 1: 172. 1825

21/c 115. Parianinae Hack. in H. G. A. Engler et K. A. E. Prantl, Nat. Pflanzenfam. II, 2: 88. Nov 1887 ['Parianeae']. ’Pariana’ (c 30; Costa Rica and Panamá to Amazonian and southeastern Brazil and Bolivia; paraphyletic; incl. Eremitis?), Eremitis (1; E. parviflora; Atlantic Brazil; in Pariana?), Parianella (2; P. carvalhoi, P. lanceolata; southern Bahia in Brazil). – Buergersiochloinae L. G. Clark et Judz. in Aliso 23: 311. Dec 2007. Buergersiochloa (1; B. bambusioides; New Guinea). – Olyrinae Kromb., Flore Luxembourg: 496. 1875. Agnesia (1; A. lancifolia; Amazonas), Cryptochloa (8; southern Mexico, Central America, tropical South America), ’Olyra’ (24; Mexico, Central America, tropical South America, one species, O. latifolia, also in tropical Africa, Madagascar, the Comoros and the West Indies; paraphyletic), Arberella (7; A. bahiensis, A. costaricensis, A. dressleri, A. flaccida, A. grayumii, A. lancifolia, A. venezuelae; Costa Rica to Brazil; in Olyra?), Lithachne (4; L. horizontalis, L. humilis, L. pauciflora, L. pinetii; southern Mexico, Central America, Cuba, tropical South America; in Olyra?), Reitzia (1; R. smithii; Venezuela and Trinidad to southern Brazil), Diandrolyra (3; D. bicolor, D. pygmaea, D. tatianae; southeastern Brazil), Ekmanochloa (2; E. aristata, E. subaphylla; eastern Cuba), Froesiochloa (1; F. boutelouoides; tropical South America), Maclurolyra (1; M. tecta; Panamá), Mniochloa (1; M. pulchella; Cuba), ’Parodiolyra’ (6; P. aratityopensis, P. colombiensis, P. lateralis, P. luetzelburgii, P. micrantha, P. ramosissima; Central America, tropical South America; paraphyletic), Piresiella (1; P. strephioides; Cuba), Raddiella (8; R. esenbeckii, R. kaieteurana, R. lunata, R. malmeana, R. minima, R. molliculma, R. potaroensis, R. vanessiae; Central America, tropical South America), ’Sucrea’ (3; S. maculata, S. monophylla, S. sampaiana; Brazil; paraphyletic), Raddia (9; Guyana, eastern Brazil), Rehia (1; R. nervata; tropical South America). – Tropical America, one species in New Guinea. – Olyreae are sister-group to Bambuseae. Species of Olyra and Pariana are said to be pollinated by insects (Soderstrom & Calderón 1971).

Pooideae Benth., Fl. Hongk.: 407. Feb 1861 [’Poaeaceae’]

186–188/3.630–3.680. Mainly temperate regions in the Northern Hemisphere. Epichloe endophytes pervasive. C₃ photosynthesis present. C₄ photosynthesis probably absent. Asymmetric division of root epidermal cells forming trichoblast-atrichoblast pair. Longitudinal walls of epidermal cells straight. Microhairs absent. Stomatal subsidiary cells with parallel margins. Primary inflorescence branches distichous. Lemma usually with five veins. Lodiculae slightly or not vascularized. Styles almost entirely separate (connate only at base). Hilum often short. Embryo small. Epiblast present at least in most Pooideae (absent in Bromus and in some Brachypodium and Hordeeae). Scutellum not peltate. Scutellum tail absent (also from some Brachyelytrum). n = (2, 4–)7(–13). Duplication of β-amylase gene. Fructose oligosaccharides present in stem. – Epichloe (Clavicipitaceae), an endophytic group of ascomycetes, is only known in Pooideae. A similar group is Neotyphodium. At least 30% of the species in Pooideae are associated with this kind of symbioses and vertical as well as horizontal transfers of these fungi take place. Numerous alkaloids (e.g. ergotamins) are produced as a result of these symbioses.

Brachyelytreae Ohwi in Bot. Mag. (Tokyo) 55: 361. 1941

1/3. Brachyelytrum (3; B. aristosum, B. erectum, B. japonicum; Japan, southeastern China, eastern United States). – Stomatal subsidiary cells with parallel sides. n = 11. – Brachyelytrum is probably sister to the remaining Pooideae. On the other hand, it is sometimes placed as part of a polytomy including also Bambusoideae and Ehrhartoideae (see, e.g., Blaner & al. 2014).

Nardeae W. D. J. Koch, Syn. Fl. Germ. Helv.: 830. Jan-Oct 1837

1/1. Nardus (1; N. stricta; Europe except the Mediterranean, western Asia). – Europe, western Asia, the Mediterranean. Microhairs bicellular. Lodiculae absent. Style and stigma single. n = 10, 13. – Nardeae are successive sister-group to the remaining Pooideae except Brachyelytrum, but see Soreng & al. (2017).

Lygeeae J. Presl, Wsobecny Rostl. 2: 1708, 1753. 1846

1/1. Lygeum (1; L. spartum; the Mediterranean, North Africa). Microhairs bicellular. Lodiculae absent. Inflorescence spatheate. Spikelet very modified. Style and stigma single. – Nardeae, Lygeeae, Duthieeae and Phaenospermateae are joined in the supertribe Nardodae by Soreng & al. (2017).

Duthieeae Röser et Jul. Schneider in Syst. Biodivers. 9: 41. Mar 2011

7/11. Stephanachne (2; S. nigrescens, S. pappophorea; Central Asia, western China), Sinochasea (1; S. trigyna; western China), Danthoniastrum (1; D. compactum; the Balkan Peninsula, the Caucasus), Pseudodanthonia (1; P. himalaica; mountains in northern India), Metcalfia (1; M. mexicana; Mexico), Anisopogon (1; A. avenaceus; eastern New South Wales, eastern Victoria), Duthiea (4; D. brachpodium, D. bromoides, D. oligostachya, D. nepalensis; Afghanistan to China). – Temperate and subtropical regions in the Old World, Mexico.

Phaenospermateae Renvoize et Clayton in Kew Bull. 40: 478. 4 Jul 1985

1/1. Phaenosperma (1; P. globosa; Assam, southeastern China, the Korean Peninsula, Japan, Taiwan). – Northeastern India to East Asia including Taiwan.

Meliceae Link ex Endl., Fl. Poson.: 116. Mai 1830 [’Melicaceae’]

8/133. Brylkiniinae Ohwi in Acta Phytotax. Geobot. 10: 107. 1 Jun 1941. Brylkinia (1; B. caudata; eastern Russia, China, Japan). – Melicinae Fr., Flora Scan.: 204, 208. 1835 [’Meliceae’]. Glyceria (38; temperate regions on the Northern Hemisphere, with their largest diversity in North America), Lycochloa (1; L. avenacea; Lebanon), Schizachne (1; S. purpurascens; arctic and temperate Russia, Siberia, Central Asia, Mongolia, China, the Korean Peninsula, temperate and arctic North America, mountains in southwestern United States), Melica (78; temperate regions on both hemispheres except Australia), Pleuropogon (6; P. californicus, P. davyi, P. hooverianus, P. oregonus, P. refractus, P. sabinei; western United States, one species, P. sabinei, arctic circumpolar), Triniochloa (6; T. andina, T. gracilis, T. laxa, T. micrantha, T. stipoides, T. talpensis; Mexico, Central America, northwestern South America), Koordersiochloa (2; K. longiarista, K. sanjappae; tropical and southern Africa, Réunion, southern India, mountains in Indonesia and the Philippines). – Temperate regions on the Northern Hemisphere, few species in tropical regions and Chile. – Meliceae may be sister-group to the remaining Pooideae. Brylkinia and Koordersiochloa have a cylindrical to lanceoloid style and a punctate hilum. Some species of Melica have seeds possessing elaiosome formed by clavate lemmas or sterile florets.

Ampelodesmeae (Conert) Tutin in Bot. J. Linn. Soc. 76(4): 369. Jun 1978

1/1. Ampelodesmos (1; A. mauritanicus; the Mediterranean). – Ampelodesmos probably originated from ancient hybridization between members of Duthieeae and Stipeae (Soreng & al. 2017). It is often recovered as sister to Stipeae (e.g. Hochbach & al. 2015).

Stipeae Dumort., Observ. Gramin. Belg.: 83. Jul-Sep 1824 [’Stipaceae’]

23–25?/545–550. Macrochloa (1; M. tenacissima; the Mediterranean), ’Stipa’ (c 150; Europe, North Africa, temperate and drier subtropical regions in Asia, with their highest diversity in southwestern and southern Asia; paraphyletic), Ortachne (3; O. breviseta, O. erectifolia, O. rariflora; Costa Rica to Peru, Chile and Argentina), Psammochloa (1; P. villosa; the Gobi Desert), Trikeraia (4; T. hookeri, T. oreophila, T. pappiformis, T. tianshanica; Pakistan to China), ’Ptilagrostis’ (13; Russia to China; polyphyletic), Piptatheropsis (5; P. canadensis, P. exigua, P. micrantha, P. pungens, P. shoshoneana; Canada, United States), Hesperostipa (5; H. comata, H. curtiseta, H. neomexicana, H. saxicola, H. spartea; Canada, United States, northern Mexico), Piptochaetium (35; semi-arid grasslands in the United States and Mexico to Argentina), Aciachne (3; A. acicularis, A. flagellifera, A. pulvinata; the High Andes in Costa Rica to northern Argentina), Anatherostipa (10; the Andes in Costa Rica till Chile), Patis (3; P. coreana, P. obtusa, P. racemosa; temperate regions on the Northern Hemisphere), Oryzopsis (1; O. asperifolia; North America), ’Achnatherum’ (c 40; Europe, North Africa, temperate regions in Asia east to Japan), Eriocoma (>3?; North America)?, Celtica (1; C. gigantea; western Mediterranean), Anemanthele (1; A. lessoniana; New Zealand), Austrostipa (c 60; Australia, one species also in New Zealand), Nassella (115–120; western and southwestern United States, Mexico, Central America, the West Indies, tropical and subtropical South America, with their largest diversity in the Andes), Jarava (c 55; South America), Oloptum (1; O. miliaceum; the Mediterranean, southwestern Asia), Orthoraphium (1; O. roylei; the Himalayas), Pappostipa (23; the United States and Mexico to Chile and Argentina), Stipellula (5; S. capensis, S. nitens, S. parviflora, S. staintonii, S. tigrensis; warm-temperate and subtropical regions in the Old World), Timouria (4?; Pakistan and Central Asia to Mongolia and China; in Psammochloa?)? – Subcosmopolitan, with their highest diversity in temperate and alpine regions. – The clade [Stipeae+Ampelodesmos] may be sister-group to all the Pooideae below but see Soreng & al. (2017). The taxonomy within Stipeae is still unclear. However, Macrochloa is a potential sister to the remaining Stipeae.

Diarrheneae (Ohwi) Tateoka ex C. S. Campb. in J. Arnold Arbor. 66: 188. 8 Apr 1985

1/5. Diarrhena (5; D. americana, D. fauriei, D. japonica, D. mandshurica, D. obovata; East Asia, North America). – Embryo of Bambusoid type.

Brachypodieae Harz in Linnaea 43: 15. 1880

1/22. Brachypodium (22; Europe, the Mediterranean, temperate regions in Asia, southern Africa, tropical mountains in Mexico to Bolivia). – Brachypodium may be sister to the rest of Pooideae, although Soreng & al. (2017) joined Ampelodesmeae, Stipeae and Diarrheneae together with Brachypodieae in the supertribe Stipodae L. Liu.

Triticeae Dumort., Observ. Gramin. Belg.: 82. Jul-Sep 1824

20/510–520. Littledaleinae Röser in J. Schneider et al., Taxon 58: 420. 28 Mai 2009. Littledalea (4; L. alaica, L. przevalskyi, L. racemosa, L. tibetica; Central Asia and Tibet to western China). – Brominae Dumort., Anal. Fam. Plant.: 63. 1829 ['Bromeae']. Bromus (c 165; temperate regions on the Northern Hemisphere, the Mediterranean, southern Africa, tropical mountains in South America). – Triticinae Fr., Flora Scan.: 210. 1835 ['Triticeae']. Aegilops (c 25; the Canary Islands, the Mediterranean to Pakistan and Central Asia; in Triticum?), Triticum (17; the Mediterranean and southwestern Asia to China; incl. Aegilops?), Dasypyrum (2; D. hordeaceum, D. villosum; the Mediterranean), Thinopyrum (11; coastal regions in Europe, the Mediterranean, southwestern Asia to Iran). – Hordeinae Dumort., Anal. Fam. Plant.: 63. 1829 ['Hordeaceae']. Agropyron (13; temperate regions in the Old World), Australopyrum (5; A. calcis, A. pectinatum, A. retrofractum, A. uncinatum, A. velutinum; New Guinea, eastern Australia, Tasmania, New Zealand), Crithopsis (1; C. delileana; Crete and Libya to Iran), Elymus (150–155; temperate regions on the Northern Hemisphere, one species, Connorochloa tenuis, in New Zealand), Eremopyrum (4; E. bonaepartis, E. distans, E. orientale, E. triticeum; southern Europe, the Mediterranean, Morocco to western China), Henrardia (2; H. persica, H. pubescens; Turkey and Iran to Central Asia), Heteranthelium (1; H. piliferum; Turkey to Pakistan), Hordeum (c 35; temperate regions on both hemispheres), Hordelymus (1; H. europaeus; Europe and North Africa to the Caucasus; in Leymus?), Leymus (c 55; temperate regions on the Northern Hemisphere, one species, L. erianthus, also in Chile and Argentina), Peridictyon (1; P. sanctum; the Balkan Peninsula), Psathyrostachys (10; eastern Mediterranean to Central Asia), Secale (9; eastern Europe and eastwards to Central Asia, the Mediterranean, the Middle East, Roggeveld in Western Cape), Taeniatherum (1; T. caput-medusae; the Iberian Peninsula, the Mediterranean to Pakistan and Central Asia). – Temperate and subtropical regions on both hemispheres, with their highest diversity in temperate Eurasia. – Triticeae are sister-group to Poeae. Littledalea is sister to the remaining Triticeae. Bromus is successive sister to the rest.

Poeae R. Br. in M. Flinders, Voy. Terra Austral. 2: 582. 19 Jul 1814 [’Poaceae’]

121/2.395–2.430. Torreyochloinae Soreng et J. I. Davis in Contr. U. S. Natl. Herb. 48: 721. Oct 2003. Amphibromus (12; Australia, New Zealand, temperate South America), Torreyochloa (5; T. erecta, T. fernaldii, T. natans, T. pallida, T. pauciflora; northeast Asia, Canada, United States). – Phalaridinae Fr., Flora Scan.: 195. 1835 ['Phalarideae']. Phalaris (17; temperate regions on the Northern Hemisphere, the Mediterranean, Andean and southern South America). – Aveninae J. Presl in C. Presl, Reliq. Haenk. 1: 246. Jan-Jun 1830 ['Avenaceae']. Arrhenatherum (7; A. album, A. calderae, A. elatius, A. kotschyii, A. longifolium, A. palaestinum, A. pallens; Europe, the Mediterranean, northern and western Asia), Avena (22; Europe, the Mediterranean, North Africa to Ethiopia, southwestern Asia), Helictotrichon (60; western United States, Eurasia, the Mediterranean, East and Southeast Asia, tropical mountains). – Koeleriinae Asch. et Graebn., Syn. Mitteleur. Fl. 2(1): 342. 22 Mai 1900. Sphenopholis (7; S. filiformis, S. intermedia, S. interrupta, S. longiflora, S. nitida, S. obtusata, S. pensylvanica; Canada, United States, Mexico, Hispaniola, the Hawaiian Islands), Cinnagrostis (80–90; Central America, South America), Peyritschia (7; P. conferta, P. deyeuxioides, P. erythraea, P. humilis, P. koelerioides, P. pringlei, P. roederi; Mexico, Central America, northwestern tropical South America), Trisetopsis (20–25; tropical to southern Africa, Madagascar, Yemen, the Himalayas, central China, southern India and Sri Lanka, Sumatra, Java), Trisetaria (16; the Mediterranean to western Himalayas), Koeleria (c 45; temperate regions on both hemispheres, tropical and subtropical African mountains), Rostraria (12; Europe, the Mediterranean, southern Africa), Gaudinia (4; G. coarctata, G. fragilis, G. hispanica, G. maroccana; the Azores, the Mediterranean), Graphephorum (2; G. melicoides, G. wolfii; North America, Mexico, Central America), Trisetum (c 75; temperate regions on the Northern Hemisphere), Tzveleviochloa (3; T. burmanica, T. parviflora, T. potaninii; Assam, northern Burma, central China). – Lagurus (1; L. ovatus; the Mediterranean), Tricholemma (2; T. breviaristatum, T. jahandiezii; Morocco, Algeria). – Anthoxanthinae A. Gray, Man. Bot., ed. 2: 538. 1 Sep 1856 ['Anthoxantheae']. Anthoxanthum (c 50; temperate and alpine regions on the Northern Hemisphere, tropical mountains in Africa and Asia). – Brizinae Tzvelev in Bot. Zhurn. (Moscow et Leningrad) 53: 310. 5 Mar 1968. Airopsis (1; A. tenella; southern Europe, northwestern Africa), Briza (5; B. humilis, B. marcowiczii, B. maxima, B. media, B. minor; Europe, temperate Asia, the Mediterranean, South America). – Echinopogoninae Soreng in J. Syst. Evol. 55(4): 262. 2017. Ancistragrostis (1; A. uncinioides; New Guinea, tropical Australia)?, Bromidium (2; B. ramboi, B. trisetoides; South America)?, Dichelachne (9; East Malesia to New Guinea and Australia, New Zealand), Echinopogon (7; E. caespitosus, E. cheelii, E. intermedius, E. mckiei, E. nutans, E. ovatus, E. phleoides; New Guinea, Australia, New Zealand), Pentapogon (1; P. quadrifidus; southeastern South Australia, southeastern New South Wales, Victoria, Tasmania), Relchela (1; R. panicoides; Chile, Argentina). – Calothecinae Soreng 2015. Chascolytrum (23; Mexico, Central America, South America). – Agrostidinae Fr., Flora Scan.: 196. 1835 ['Agrostideae']. Agrostis (190–200; temperate regions on both hemispheres, tropical mountains, Macquarie Island, subAntarctic South America, Falkland Islands, Crozet Islands, Kerguélen, Prince Edward Islands), ’Calamagrostis’ sensu stricto (c 260; temperate regions on both hemispheres; non-monophyletic), Gastridium (2; G. phleoides, G. ventricosum; western Europe, Macaronesia, the Mediterranean to the Caucasus and Iran), Hypseochloa (2; H. cameroonensis, H. matengoensis; Mount Cameroon, mountains in Tanzania), Limnodea (1; L. arkansana; southern United States), Polypogon (21; warm-temperate regions on both hemispheres, tropical mountains), Triplachne (1; T. nitens; the Mediterranean). – Scolochloinae Tzvelev in Komarov. Chten. 37: 33. 1987. Dryopoa (1; D. dives; southeastern New South Wales, Victoria, Tasmania), Scolochloa (1; S. festucacea; temperate regions on the Northern Hemisphere). – Sesleriinae Parl., Flora Palerm.: 127. Mar-Dec 1845 ['Seslerieae']. Sesleriella (1; S. sphaerocephala; central and eastern Alps), Mibora (2; M. maroccana, M. minima; western Europe, northwestern Africa), Oreochloa (5; O. blanka, O. confusa, O. disticha, O. elegans, O. seslerioides; the Pyrenees, the Alps, the Carpathians), Echinaria (1; E. capitata; southern and southeastern Europe, the Mediterranean, southwestern Asia to the Caucasus), Psilathera (1; P. ovata; central and eastern Alps), Sesleria (c 30; Europe, Morocco, Libya, western Asia to the Caucasus and Iran). – Airinae Fr., Flora Scan.: 196, 198. 1835 ['Aireae']. Avenella (1; A. flexuosa; temperate regions on the Northern Hemisphere), Aira (8; Europe, the Mediterranean to Iran), Antinoria (2; A. agrostidea, A. insularis; the Mediterranean), Corynephorus (5; C. canescens, C. deschampsioides, C. divaricatus, C. fasciculatus, C. macrantherus; Europe, the Mediterranean to Iran), Helictochloa (22; central and southern Europe, the Mediterranean, the Canary Islands, North Africa, the Balkan Peninsula, eastern Europe to Crimea and the Caucasus, the Middle East, one species, H. hookeri, in Russia, Ukraine, Central Asia, Mongolia, China and North America), Periballia (3; P. involucrata, P. laevis, P. minuta; the Mediterranean). – Holcinae Dumort. in Bull. Soc. Roy. Bot. Belgique 7: 68. 1868 ['Holceae']. Holcus (10; Europe, the Mediterranean, North and southern Africa, southwestern Asia), Vahlodea (1; V. atropurpurea; cold-temperate and arctic-alpine regions on the Northern Hemisphere, subAntarctic South America). – Aristaveninae F. Albers et Butzin in Willdenowia 8: 82. 25 Aug 1977. Deschampsia (c 40; temperate and polar regions on both hemispheres, the Andes from 34°10’ south latitude to Tierra del Fuego, Falkland Islands, South Georgia Island, South Shetland Islands, the Antarctic Peninsula and adjacent islands, South Orkney Islands, South Sandwich Islands, Prince Edward Islands, Crozet islands, Kerguelen Islands, Heard Island). – Loliinae Dumort., Anal. Fam. Plant.: 63. 1829 ['Loliaceae']. Castellia (1; C. tuberculosa; Macaronesia, the Mediterranean, North Africa to Pakistan), Festuca (470–480?; temperate, polar and alpine regions on both hemispheres, Tierra del Fuego, Falkland Islands, South Georgia Islands, Kerguelen Islands, Macquarie Island, tropical mountains), Leucopoa (10?; temperate Asia, Canada, United States), Drymochloa (5; D. donax, D. drymeja, D. grandis, D. lasto, D. sylvatica; Europe, western Mediterranean, North Africa; in Leucopoa?), Lolium (12; temperate regions on the Northern Hemisphere, the Canary Islands, the Mediterranean), Megalachne (3; M. berteroniana, M. robinsoniana, M. masafuerana; Juan Fernandez Islands), Patzkea (1–5; P. paniculata; Europe, the Mediterranean), Podophorus (1; P. bromoides; Juan Fernandez Islands). – Pseudobromus (6; P. africanus, P. ambilobensis, P. breviligulatus, P. engleri, P. humbertianus, P. tenuifolius; tropical and subtropical Africa)? – Dactylidinae Stapf in W. H. Harvey et O. W. Sonder (ed. W. T. Thiselton-Dyer), Flora Cap. 7: 317. Jul 1898 ['Dactylideae']. Dactylis (2; D. glomerata: Europe, temperate Asia; D. smithii: Macaronesia), Lamarckia (1; L. aurea; the Mediterranean, the Middle East to Pakistan). – Cynosurinae Fr., Flora Scan.: 204. 1835 ['Cynosureae']. Cynosurus (9; Europe, the Mediterranean, southwestern Asia). – Ammochloinae Tzvelev, Zlaki SSSR: 535. 1976. Ammochloa (3; A. involucrata, A. palaestina, A. pungens; the Mediterranean, southwestern Asia). – Parapholiinae Caro in Dominguezia 4: 41. 1982. Agropyropsis (1; A. lolium; Algeria), Cutandia (6; C. dichotoma, C. divaricata, C. maritima, C. memphitica, C. rigescens, C. stenostachya; the Mediterranean, Macaronesia, southwestern Asia), Desmazeria (4; D. lorentii, D. philistaea, D. pignattii, D. sicula; Europe, the Mediterranean, North Africa, southwestern Asia to Iran), Hainardia (1; H. cylindrica; the Mediterranean), Parapholis (6; P. filiformis, P. gracilis, P. incurva, P. marginata, P. pycnantha, P. strigosa; Europe, the Mediterranean, southern Asia), Sphenopus (2; S. divaricatus, S. ehrenbergii; the Mediterranean, southwestern Asia to Iran), Vulpiella (1; V. tenuis; western Mediterranean). – Avenula (1; A. pubescens; temperate Eurasia). – Coleanthinae Rouy, Flore France 14: 28, 55. Apr 1913. Catabrosa (3; C. aquatica: temperate regions on the Northern Hemisphere, southern South America; C. drakensbergensis: Lesotho, KwaZulu-Natal; C. werdermannii: the Andes in Bolivia, Chile and Argentina), Coleanthus (1; C. subtilis; temperate regions on the Northern Hemisphere), Phippsia (3; P. algida, P. concinna, P. wilczekii; arctic regions on the Northern Hemisphere), Puccinellia (c 125; temperate and arctic regions on the Northern Hemisphere, southern Africa, one species, P. stricta, in Australia), Sclerochloa (2; S. dura, S. woronowii; the Mediterranean, North Africa and the Middle East to Central Asia and China). – Poinae Dumort., Anal. Fam. Plant.: 63. 1829 ['Poeae']. Poa (>500; temperate, alpine and polar regions on both hemispheres, tropical mountains). – Miliinae Dumort., Anal. Fam. Plant.: 64. 1829 [’Miliaceae’]. Milium (6; M. atropatanum, M. effusum, M. pedicellare, M. schmidtianum, M. transcaucasicum, M. vernale; Europe, temperate Asia, eastern North America). – Phleinae Dumort. in Bull. Soc. Roy. Bot. Belgique 7: 69. 1868 [‘Phleae‘]. Phleum (16; temperate regions on the Northern Hemisphere, temperate South America). – Beckmanniinae Nevski in Trudy Bot. Inst. Akad. Nauk S.S.S.R., ser. 1, Fl. Sist. Vyssh. Rast. 4: 228. 1937. Beckmannia (2; B. eruciformis, B. syzigachne; temperate regions on the Northern Hemisphere), Pholiurus (1; P. pannonicus; southeastern Europe to Central Asia), Pseudophleum (2; P. anatolicum, P. gibbum; Turkey to Iran), Rhizocephalus (1; R. orientalis; the Mediterranean to Iran). – Cinninae Caruel, Epit. Flor. Europ. 1: 69. Jan 1892 [’Cinneae’]. Cinna (4; C. arundinacea, C. bolanderi, C. latifolia, C. poiformis; temperate regions on the Northern Hemisphere), Aniselytron (2; A. agrostoides, A. treutleri; northern India to Japan), Cyathopus (1; C. sikkimensis; the Himalayas), Simplicia (2; S. buchananii, S. laxa; New Zealand), Agrostopoa (3; A. barclayae, A. wallisii, A. woodii; Colombia)? – Alopecurinae Dumort., Anal. Fam. Plant.: 64. 1829 [’Alopecureae’]. Alopecurus (c 40; temperate regions on the Northern Hemisphere, temperate South America), Cornucopiae (2; C. alopecuroides, C. cucullatum; eastern Mediterranean to Iraq), Limnas (3; L. malyschevii, L. stelleri, L. veresczagini; Central Asia to northeastern Siberia). – Ventenatinae Holub ex L. J. Gillespie, Cabi et Soreng in J. Syst. Evol. 55(4): 264. 2017. Ventenata (8; southern Europe, the Mediterranean to the Caspian Sea), Bellardiochloa (5; B. argaea, B. carica, B. polychroa, B. variegata, B. violacea; central and southern Europe, the Mediterranean, southwestern Asia), Nephelochloa (1; N. orientalis; Turkey), Apera (5; A. baytopiana, A. intermedia, A. interrupta, A. spica-venti, A. triaristata; Europe, southwestern Asia to Afghanistan), Parvotrisetum (1; P. myrianthum; northern Italy, southern and western Balkan Peninsula). – Cosmopolitan, with their largest diversity in temperate regions on the Northern Hemisphere. – Poeae are sister-group to Triticeae. – Poeae incertae sedis: Brizochloa (1; B. humilis; eastern Europe to the Caucasus and western Asia), Dupontia (2; D. fisheri, D. fulva; arctic regions), Dupontiopsis (1; D. hayachinensis; mountains in northern Japan), Arctagrostis (1; A. latifolia; arctic regions), Hookerochloa (1; H. hookeriana; eastern New South Wales, Victoria, Tasmania), Nicoraepoa (6; N. andina, N. chonotica, N. erinacea, N. pugionifolia, N. robusta, N. subenervis; the Andes in South America), Saxipoa (1; S. saxicola; southeastern New South Wales, eastern Victoria, Tasmania), Sylvipoa (1; S. queenslandica; southeastern Queensland, eastern New South Wales).

Cladogram (simplified and slightly modified) of Poaceae based on DNA sequence data (Bouchenak-Khelladi & al. 2008; Schneider & al. 2009). Anomochloa and Streptochaeta are sister-groups in some analyses. Isachne is sister toEriachne inMicrairoideae (=Micraireae)in, e.g., Peterson & al. (2011).

Habit Bisexual, perennial herbs or, in older individuals, with lignified up to one metre tall (sometimes taller) aerial stem. Graminids. Helophyte. Culm angular in cross-section, erect.

Vegetative anatomy Mycorrhiza absent? Aerial stem with thin-walled epidermis, cortex with loosely packed parenchyma cells and scattered vascular bundles, and central cylinder with scattered amphivasal bundles. Phellogen absent. Secondary lateral growth absent. Vessels present in roots, stem and leaves. Vessel elements usually with scalariform (in stems sometimes simple) perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Silica bodies spherical, in epidermal cells; silica bodies absent from parenchyma cells. Tanniniferous cells scattered. Calciumoxalate crystals absent.

Leaves Alternate (tristichous), simple, entire, linear, flat or V-shaped in cross-section (with spinose-serrate keel), unifacial, with convolute (supervolute) ptyxis. Stipules absent; leaf sheath closed, without ligule? Venation parallelodromous. Stomata paracytic. Cuticular wax crystalloids as longitudinally aggregated rodlets, Strelitziatype. Air canals with chlorenchyma. Epidermis with silica bodies? Mesophyll without mucilaginous idioblasts or calciumoxalate crystals. Leaf margin spinose-serrate.

Inflorescence Terminal, richly branched, panicle. Floral prophyll (bracteole) absent.

Flowers Actinomorphic, small. Hypogyny. Tepals 3+3, sepaloid, membranous, persistent, free. Nectary absent. Disc absent.

Androecium Stamens 3+3. Filaments filiform, free from each other and from tepals. Anthers basifixed, non-versatile?, tetrasporangiate, latrorse?, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with uninucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains with indistinct aperture (ulcerate), shed as tetrahedral and tetragonal tetrads, tricellular at dispersal. Exine tectate, with granular? infratectum, scabrate or ulcerate.

Gynoecium Pistil composed of three carpels, connate only at base. Ovary superior, trilocular, narrowing towards apex. Stylodia three, free or slightly connate at base. Stigmas three, papillate, Dry? type, Pistillodium absent.

Ovules Placentation axile. Ovules one to seven per carpel, anatropous, ascending, apotropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument one or two cell layers thick. Inner integument two cell layers thick. Parietal cell formed from archesporial cell. Megagametophyte monosporous, Polygonumtype. Synergids with a filiform apparatus. Antipodal cells persistent. Endosperm development ab initio helobial. Endosperm haustoria? Embryogenesis onagrad (Juncusvariation).

Seeds Aril absent. Seed coat testal-tegmic. Exotesta consisting of one to three layers of sclerenchymatous fibres. Endotesta consisting of one or two parenchymatous layers. Tegmen consisting of elongate tightly packed tanniniferous cells. Perisperm not developed. Endosperm copious, starchy. Embryo small, straight, well differentiated, Xyris-Scirpustype, with chlorophyll. Cotyledon one, photosynthesizing. Cotyledon hyperphyll? Hypocotyl absent. Mesocotyl? Coleoptile absent. Plumule terminal. Collar very small, with rhizoids. Germination phanerocotylar.

Phytochemistry Very insufficiently known. Flavone-C-glycosides present. Luteolin-5-methylether? Cyanogenic compounds not found.

Rapateales (Meisn.) Colella ex Reveal et Doweld in Novon 9: 551. 30 Dec 1999; Rapateanae Doweld, Tent. Syst. Plant. Vasc.: lxii. 23 Dec 2001

Distribution Tropical Central and South America, tropical West Africa, with their highest diversity in the Guayana and Venezuelan Highlands.

Habit Bisexual, perennial herbs. Usually hygrophytic (a few species are epiphytic).

Vegetative anatomy Phellogen absent. Stems with wide outer cortex consisting of parenchyma cells with starch; inner cortex consisting of several sclereid layers. Central stem cylinder consisting of sclerified parenchyma and collateral and amphivasal vascular bundles. Secondary lateral growth absent. Vessels present in roots and stem (rarely in leaves). Vessel elements with simple and/or scalariform perforation plates; lateral pits scalariform. Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Mucilage canals present in some species. Tanniniferous cells often frequent (at least in hypodermis). Silica bodies (druses) and/or silica sand usually present. Calciumoxalate probably absent.

Trichomes Axillary, multicellular, uniseriate, mucilage secreting eglandular hairs often abundant, especially on young shoots.

Leaves Alternate (usually distichous, sometimes spiral), simple, entire, often linear, unifacial to bifacial, usually V-shaped, often twisted 90°, with conduplicate ptyxis, sometimes with spines. Stipules absent; leaf sheath open (rarely with distal ligule), symmetrical or asymmetrical. Colleters uniseriate, secreting mucilage. Venation parallelodromous. Stomata paracytic, with dumbbell-shaped Poaceae type guard cells; subsidiary cells oily. Cuticular wax crystalloids usually absent (sometimes amorphous and spherical, non-orientated). Epidermis with silica druses. Mesophyll without mucilaginous cells and calciumoxalate crystals. Leaf margin entire.

Inflorescence Terminal or axillary, simple or compound spherical or flattened thyrses or thyrsoid inflorescences with one to more than 70 many-flowered cymose partial inflorescences as spike-like bostryces with terminal flower and coriaceous bracts, often with two or more, sometimes more or less connate, spathe-like bracts.

Flowers Actinomorphic or somewhat zygomorphic, large. Hypogyny. Tepals 3+3; outer tepals with imbricate aestivation, sepaloid, free or connate at base; inner tepals usually with imbricate aestivation, petaloid, caducous, connate at base. Septal nectaries present at least in Monotrematoideae. Disc absent. Colleters present on floral parts.

Androecium Stamens 3+3. Filament bases usually connate and/or adnate to inner tepals (in one species of Rapatea hairy). Anthers basifixed to subbasifixed, non-versatile, usually tetrasporangiate (sometimes disporangiate), extrorse to introrse, poricidal (dehiscing by usually one or two, in Schoenocephalium four, apical to adaxial pores or short slits); connective sometimes with apical appendages. Tapetum secretory, with mostly binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains monosulcate or trichotomosulcate (sometimes zonisulcate or zonisulculate, rarely disulculate), shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, reticulate, foveolate or scrobiculate.

Gynoecium Pistil composed of three connate carpels. Ovary superior, trilocular or (due to reduction of two carpels) unilocular; septa often incomplete. Style single, simple. Stigma capitate to punctate, often finely hairy, type? Pistillodium absent.

Ovules Placentation usually axile (sometimes basal). Ovules usually one to eight (sometimes up to c. 50) per carpel, anatropous, ascending, apotropous, bitegmic, crassinucellar. Micropyle usually bistomal (sometimes endostomal), directed downwards. Outer integument three to ten cell layers thick. Inner integument ? cell layers thick. Funicular obturator present in some representatives. Parietal cell formed from archesporial cell. Epidermal megasporangium cells often radially prolonged. Megagametophyte monosporous, Polygonum type. Antipodal cells sometimes more than three. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis asterad.

Seeds Aril absent. Sometimes with a conical or cap-like spongy caruncle or elaiosome at chalazal end. Seed coat testal-tegmic, sometimes winged. Exotesta thin, with silica bodies. Endotesta sometimes with silica bodies; endotestal cells thick-walled (in Spathanthus with U-shaped wall thickenings). Cuticular layer present between testa and tegmen. Tegmic cells tanniniferous. Perisperm not developed. Endosperm copious, starchy, oily and proteinaceous; starch grains simple. Embryo small, little differentiated, micropylar, Xyris-Scirpus type. Cotyledon one. Coleoptile absent. Collar absent. Rhizoids absent. Radicula little developed. Germination?

Cytology n = 11 (Maschalocephalus); n = 26 (Cephalostemon, Spathanthus)

Phytochemistry Virtually unknown. Aluminium accumulation occurring in many species.

Restionales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 266. Jan-Apr 1820 [’Restiaceae’]; Restionineae Link, Handbuch 1: 134. 4-11 Jul 1829 [‘Restiaceae’]; Devauxiaceae Dumort., Anal. Fam. Plant.: 62, 63. 1829, nom. illeg.; Centrolepidaceae Endlicher, Gen. Plant.: 119. Dec 1836 [’Centrolepideae’], nom. cons.; Elegiaceae Raf., Fl. Tellur. 4: 32, 33. med 1838 [’Elegides’]; Centrolepidales R. Dahlgren ex Takht., Divers. Classif. Fl. Pl.: 553. 24 Apr 1997

Distribution Tropical and southern Africa, Madagascar, southern China, Southeast Asia and Malesia to New Guinea, Australia, Tasmania, New Zealand, Chile, with their largest diversity in southwestern Australia and the Western Cape Province in South Africa.

Fossils Restiocarpus comprises fossil fruits from the Eocene/Oligocene border of Queensland and may be remnants of Restionaceae or some allied group. Pollen grains with the formal name of Milfordia are known from Eocene strata in Queensland, Europe, Asia and North America. They have often been assigned to Restionaceae, although they may emanate from other graminoid groups. Plausible Restionaceae pollen have been described from the Neogene at several places in the Southern Hemisphere, from the earliest Paleocene onwards.

Habit Usually dioecious (in Coleocarya and Lepyrodia monoecious; in, e.g., some species of Lepyrodia sometimes bisexual; in Centrolepidoideae monoecious, andromonoecious or polygamomonoecious), perennial (sometimes annual) herbs. Graminids. Often xeromorphic. Culm photosynthesizing, simple or branched (branches sometimes verticillate), usually terete (rarely quadrangular or flattened in cross-section), smooth, verrucose, striate, furrowed or pitted, medullated or with a hollow centre, often with compact swollen nodes.

Vegetative anatomy Mycorrhiza absent. Root hairs developing from any epidermal cell, usually persistent, lignified, very long and densely spaced. Root in Centrolepidoideae without pericycle. Lateral roots developing from a zone usually opposite protoxylem poles (in Centrolepidoideae from a zone opposite protophloem poles). Rhizome with endodermis-like envelope. Phellogen absent. Rhizome with endodermoid sheath. One or several subepidermal palisade-like chlorenchyma layers with peg cells (in Centrolepidoideae usually without peg cells); inside these layers a parenchymatic cylinder and inside this a sclerenchymatous cylinder; medulla with scattered collateral vascular bundles. Culm with protective lignified chlorenchymatous cells lining substomatal cavities. Secondary lateral growth absent. Vessels present in roots and stem (rarely in leaves). Vessel elements with scalariform or simple (sometimes reticulate) perforation plates; lateral pits scalariform or alternating. Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c_c type (with cuneate protein crystals), P2c_cl type (with cuneate and several additional loosely packed protein crystals) or P2cf (with cuneate protein crystals and peripheral protein filaments). Nodes? Silica as spheroidal-nodular bodies or granular crystal sand (in Centrolepidoideae silica bodies and crystals usually absent?); parenchyma and sclerenchyma cells often filled with silica bodies, usually in association with vascular strands (especially in parenchyma sheath between chlorenchyma and sclerenchyma cylinders, or in outer sclerenchymatous layer). Calciumoxalate crystals absent.

Trichomes Hairs unicellular or multicellular, usually uniseriate (rarely T-shaped, in Gaimardia often branched), multiseriate-stellate, stalked (sometimes peltate) or absent; multicellular microhairs rarely? present.

Leaves Alternate (spiral or distichous to spirodistichous), simple, entire, linear (rarely subulate), unifacial, often with convolute (supervolute) ptyxis, usually reduced to mere sheaths or bristle-like. Stipules absent; leaf sheath well developed, usually open (rarely closed), usually persistent (sometimes caducous), often with one pair of membranous lobes or a membranous distal edge (ligule?), usually without ligule (in Gaimardia with distal ligule). Venation parallelodromous. Stomata (brachy)paracytic; often Poaceae type guard cells, sometimes sunken. Cuticular wax crystalloids as non-orientated platelets, only associated with stomata. Epidermis with or without rows of alternately long and short cells; epidermal cells with or without silica bodies. Mesophyll without mucilaginous idioblasts and usually without calciumoxalate crystals (raphides absent). Leaf margin entire. Epidermis with or without hairs and papillae.

Inflorescence Terminal, compound, simple or branched panicle, corymb, spike- or head-like with one- or many-flowered partial inflorescences as spikelets (inflorescence sometimes consisting of a single spikelet); inflorescence branches in axils of often spatha-like bracts; lower spikelet bracts often without flowers; flowers usually with a single bract (rarely two bracts). Inflorescences often with large sexual dimorphism. Floral prophylls (bracteoles) often absent (sometimes two). Inflorescence in Centrolepidoideae terminal, few- to many-flowered usually bisexual pseudanthia, each pseudanthial unit being a spikelet with usually two (sometimes one, three or no) hyaline membranous secondary bracts (floral prophylls?); pseudanthia concentrated in capitate or distichous spike-like inflorescences, each subtended by two (or more) scale-like or green primary bracts.

Flowers Actinomorphic to somewhat zygomorphic, small. Hypogyny. Tepals usually 3+3 (rarely two, three, 2+2, 3+2, or absent), dry, membranous to hard (in male flowers rarely reduced), often persistent, usually free (sometimes connate at base); outer lateral tepal often with a hairy (sometimes winged) keel. Nectary absent. Disc absent.

Androecium Stamens usually three (rarely – such as in Centrolepidoideae – one or two), antepetalous (outer staminal whorl absent). Filaments filiform, free from each other and from tepals. Anthers dorsifixed, sometimes versatile, usually disporangiate and monothecal (rarely tetrasporangiate and dithecal), usually introrse (rarely latrorse), longicidal (dehiscing by longitudinal slits); connective sometimes slightly prolonged. Tapetum secretory, with uninucleate to quadrinucleate cells; anther walls highly tanniniferous. Female flowers often with staminodia.

Pollen grains Microsporogenesis successive. Pollen grains graminoid, monoporate or monoulcerate (aperture with various shape; most African representatives with a very wide pore), with or without operculum or annulus, shed as monads, usually tricellular (sometimes bicellular) at dispersal. Exine tectate, with columellate (sometimes granular) infratectum, perforate, rugulate or scrobiculate, usually microverrucate or smooth (rarely verrucate or spinulate).

Gynoecium Pistil composed of one to three ascidiate connate antesepalous carpels (sometimes only one out of three carpels fertile leading to pseudomonomery); pistil in Centrolepidoideae composed of a single carpel or, alternatively, one to 30 (rarely up to 45) free or (seemingly) connate carpels. Ovary superior, unilocular to trilocular, sometimes shortly stipitate; ovary walls richly tanniniferous. Stylodia usually three (rarely one or two), usually free (rarely connate at base), with adaxial stigmatic surfaces; style in Centrolepidoideae single, simple, filiform, persistent, with adaxial pollen receptive surface. Stigmas usually three (rarely one or two, in Centrolepidoideae one), much branched, with papillate branches, Dry type. Male flowers often with pistillodium (at least rudimentary).

Ovules Placentation usually axile to apical (apical or marginal, when ovary monomerous then unilocular). Ovule one per carpel, orthotropous, pendulous, bitegmic, usually tenuinucellar (in at least Alexgeorgea crassinucellar). Micropyle usually bistomal (in Willdenowia and some species of Leptocarpus endostomal). Outer integument two? cell layers thick. Inner integument two? cell layers thick; inner epidermis of inner integument strongly tanniniferous. Parietal cell usually not formed (parietal tissue one cell layer thick present in at least, e.g., Alexgeorgea). Hypostase usually present (no widening of micropylar part of integument; in Centrolepidoideae absent). Megasporangial epidermis with anticlinally elongate cells (radial elongation of megasporangial epidermis). Megagametophyte monosporous, Polygonum type (sometimes Poaceae variation), with large compound starch grains (polar nuclei surrounded by large starch bodies). Synergids sometimes with a filiform apparatus. Antipodal cells usually binucleate?, in Restioneae at least sometimes proliferating. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad.

Fruit A loculicidal usually three-seeded capsule or a one-seeded unilocular nutlet; fruit stalk sometimes fleshy and elaiosome-like (spikelet axis in Hypolaena fleshy); nutlet sometimes with persistent tepals with wing-shaped keel; fruit in Centrolepidoideae a one-seeded, abaxially dehiscing follicle with membranous pericarp (in Aphelia indehiscent), collateral ovaries often fused into a capsule-like coenocarp. Fruit in Alexgeorgea hypogeal.

Seeds Aril absent. Elaiosome present in some African genera. Seed coat endotegmic. Exotesta persistent (testa and exotegmen in at least Centrolepidoideae degenerating). Endotesta? Tegmen? (endotegmen in at least Centrolepidoideae membranous, tanniniferous, persistent) Cuticle between integument well developed. Perisperm not or only slightly developed. Endosperm copious, starchy; starch grains in at least Centrolepidoideae compound. Embryo small, often lens-shaped or conical, little differentiated, rudimentary, Xyris-Scirpus type, chlorophyll? Cotyledon one, usually photosynthesizing. Cotyledon hyperphyll prolonged, assimilating. Hypocotyl and collar rudimentary. Hypocotyl internode absent. Coleoptile absent. Collar rhizoids present. Germination phanerocotylar. First leaf of seedling unifacial, with sheath-like or (in Centrolepidoideae) disc-shaped part; with isodiametric or palisade chlorenchyma cells. Photosynthesizing unifacial cotyledon hyperphyll (phanomer) usually present in at least Centrolepidoideae.

Cytology n = 6, 7, 9, 10 (Centrolepidoideae), 11, 12 (Australian genera); n = 16, 20 (African genera) – Agamospermy probably occurs in some genera. Some species are parthenocarpic and known as female plants only.

DNA The plastid genome has an inversion of 28 kb in some species (present at least in Askidiosperma, Baloskion, Leptocarpus, Rhodocoma, and Elegia fenestrata; absent from Desmocladus, Elegia cuspidata, and Centrolepis).

Phytochemistry Flavonols (kaempferin, quercetin), flavones, flavone-C-glycosides and flavone sulphates sometimes present (flavones and glycoflavones present in some African clades). Proanthocyanidins present in almost all African species, whereas remaining species (including a few African species) possess 8-hydroxyflavonoids gossypetin and hypolaetin (8-hydroxyluteolin). Glycosides of myricetin, laricitrin and syringetin present in ’Elegia’ and Chondropetalum. S-methylcysteine (an amino acid) present in Lepyrodia. Tricin, ellagic acid, alkaloids, saponins, and cyanogenic compounds not found. Aluminium accumulation occurring in Centrolepis.

Use Ornamental plants (Chondropetalum tectorum, Elegia capensis etc.), thatching (Thamnochortus insignis etc.), forage plants.

Habit Bisexual, perennial herbs. Graminids. Often helophytic or aquatic. Culm angular in cross-section, erect.

Vegetative anatomy Mycorrhiza absent? Phellogen absent. Secondary lateral growth absent. Vessels present in roots, stem and leaves. Vessel elements usually with scalariform (in stems sometimes simple) perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Silica bodies spherical, in epidermal cells, and silica sand in parenchyma cells. Tanniniferous cells scattered. Calciumoxalate crystals absent.

Leaves Alternate (tristichous or tetrastichous), simple, entire, linear, flat or V-shaped in transverse section, unifacial, with convolute (supervolute) ptyxis. Stipules absent; leaf sheath closed?, with ligule? Venation parallelodromous; vascular bundles in vertical pairs (superposed) with a larger adaxial non-inverted (with phloem on lower side) and a smaller abaxial inverted bundle (with phloem on upper side), the two phloems facing each other. Stomata usually paracytic (sometimes tetracytic). Cuticular wax crystalloids as aggregated rodlets. Epidermis with silica bodies. Mesophyll without mucilaginous idioblasts or calciumoxalate crystals. Leaf margin entire to hispid.

Inflorescence Flowers in one or several terminal dense racemose heads subtended by narrowly elongate bracts. Peduncle (culm) triangular or quadrangular in transverse section.

Flowers Actinomorphic, small. Hypogyny. Tepals 3+3, sepaloid, membranous, persistent, free. Nectary absent. Disc absent.

Androecium Stamens 3+3. Filaments filiform, free from each other, somewhat adnate at base to tepals. Anthers basifixed, non-versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with uninucleate? cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains with indistinct aperture (ulcerate), shed as tetrahedral tetrads, tricellular at dispersal. Exine tectate, with granular? infratectum, scabrate or ulcerate.

Gynoecium Pistil composed of three connate carpels. Ovary superior, trilocular, narrowing towards apex. Style single, trilobate. Stigmas three, papillate, Dry type. Pistillodium absent.

Ovules Placentation axile to axile-basal or lateral. Ovules one to seven per carpel, anatropous, ascending, apotropous, bitegmic, crassinucellar. Micropyle bistomal, Z-shaped (zig-zag). Outer integument one or two cell layers thick. Inner integument two cell layers thick. Parietal cell formed from archesporial cell. Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Seeds Aril present. Seed coat testal-tegmic, with one subulate outgrowth at each end. Testa puberoulous. Exotesta consisting of one to three layers of sclerenchymatous fibres. Endotesta consisting of one or two parenchymatous layers. Tegmen consisting of elongate tightly packed tanniniferous cells. Perisperm not developed. Endosperm copious, starchy. Embryo small, straight, well differentiated, Xyris-Scirpus type, chlorophyll? Cotyledon one, photosynthesizing. Cotyledon hyperphyll? Hypocotyl absent. Mesocotyl? Coleoptile absent. Plumule terminal. Collar very small, with rhizoids. Germination phanerocotylar.

Phytochemistry Virtually unknown. Flavone-C-glycosides? Luteolin-5-methylether? Cyanogenic compounds not found.

Sparganiaceae Hanin, Cours Bot.: 400. 16-23 Apr 1811 [’Spargania’], nom. cons.; Typhales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 263. Jan-Apr 1820 [‘Typhaceae’]; Typhanae Thorne ex Reveal in Novon 2: 237. 13 Oct 1992

Fossils Fruits assigned to Typha have been found in the Maastrichtian of Central Europe and Sparganium fruits were reported from Paleocene and younger layers. The oldest fruits are quinquelocular. Sparganium and Typha have been recorded more frequently from the Eocene onwards. Typhaspermum, representing seeds similar to Typha, is known from Late Eocene to Oligocene layers in Queensland.

Habit Monoecious, perennial herbs. Aquatic or helophytic. Rhizome rich in starch.

Vegetative anatomy Mycorrhiza probably absent. Lateral roots developing from a zone opposite protoxylem poles. Phellogen absent. Secondary lateral growth absent. Vessels present in roots, stem and leaves. Vessel elements with scalariform perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2cs type, with cuneate protein crystals and starch grains. Nodes? Silica bodies present or absent. Calciumoxalate as raphides, druses or single prismatic or rod-shaped crystals. Calciumoxalate styloids present in rows of cells above sclerenchyma bundles in Typha. Starch grains simple.

Leaves Alternate (distichous), simple, entire, aerial leaves linear, in Sparganium sometimes keeled and laterally flattened, with ? ptyxis. Stipules absent; leaf sheath well developed, in Typha with adaxial mucilaginous cells. Venation parallelodromous. Stomata paracytic (subsidiary cells obliquely divided). Cuticular wax crystalloids as longitudinally aggregated rodlets (Strelitzia type), chemically dominated by wax esters. Epidermis with or without silica bodies; epidermis with tanniniferous myriophyllin cells. Mesophyll with or without mucilaginous idioblasts and sacs containing calciumoxalate raphides. Leaf margin entire.

Inflorescence Terminal (in Sparganium often axillary), complex multiple compound racemose, with male inflorescence(s) in upper part and female inflorescence(s) in lower part; in Typha spike-like (stachyoid) or spadix-like (spadicioid); in Sparganium spherical and capitate (cephalioid); inflorescences compact and consisting of reduced one- or few-flowered branched partial inflorescences. Male and female inflorescences in Typha at least initially separated by a bract.

Flowers Actinomorphic, small to minute. Hypogyny. Tepals one to c. 20, in male flowers usually three, in female flowers one to four, filiform (Typha), or three or 3+3 sepaloid and scale-like and with quincuncial aestivation (Sparganium), persistent, free or connate at base. Nectary absent. Disc absent.

Androecium Stamens in Typha usually three (sometimes one, two or up to eight); in Sparganium (one to) three or 3+3, on androphore. Filaments linear to filiform, free from each other or connate at base, free from tepals. Anthers basifixed, non-versatile, tetrasporangiate, extrorse to latrorse, longicidal (dehiscing by longitudinal slits); connective in Typha somewhat prolonged. Tapetum intermediate between secretory and amoeboid-periplasmodial, in Sparganium with 8-nucleate cells (Typha type: tapetal nuclei dividing, each cell becoming ab initio binucleate, subsequently dividing twice). Staminodia absent.

Pollen grains Microsporogenesis successive. Pollen grains monoulcerate with indistinct pore-like aperture, usually shed as monads (in Typha occasionally as tetrads), usually bicellular at dispersal. Exine tectate, with columellate infratectum, finely reticulate, echinulate or smooth.

Gynoecium Pistil in Typha composed of one stipitate carpel; pistil in Sparganium composed of two or three connate carpels (usually only one carpel fertile, pseudomonomerous). Ovary superior, unilocular, sometimes on gynophore. Style (stylodium) single, simple or bifid or trifid in upper part. Stigma two or three, decurrent, fairly elongate, Dry type, in Sparganium papillate. Pistillodium absent.

Ovules Placentation apical or subapical. Ovule one per carpel, anatropous, pendulous, apotropous, bitegmic, crassinucellar. Micropyle bistomal (in Typha endostomal?). Outer integument ? cell layers thick. Inner integument ? cell layers thick. Obturator present. Parietal cell formed from archesporial cell and further developing into parietal tissue. Nucellar cap approx. two cell layers thick. Megagametophyte monosporous, Polygonum type. Antipodal cells in Sparganium often proliferating (to c. 150 cells). Endosperm development ab initio helobial of aberrant type: small chalazal endosperm chamber with rapid cell divisions (free nuclear divisions up to 16-nucleate stage or cell wall formation earlier), large micropylar chamber developing into nutrient tissue. Endosperm haustoria absent. Embryogenesis asterad (Typha) or onagrad (Sparganium).

Fruit In Typha a hairy follicular nutlet with strongly reduced endocarp and mesocarp; in Sparganium a drupe with usually spongy exocarp, decaying during maturation, and hard endocarp.

Seeds Aril absent. Operculum present. Seed coat in Sparganium testal-tegmic, with testa and tegmen membranous except at micropylar end, thick-walled cells here forming a conical double operculate structure; testa in Typha compressed, with thin-walled cells, tegmen with thick-walled cells. Perisperm thin, weakly developed. Endosperm copious, with micropylar chamber containing aleurone, starch (starch grains pteridophyte type, amylophilic), proteins (in Sparganium), and oils (in Typha); chalazal chamber with very small amounts of nutrients. Embryo long, straight, slender, without chlorophyll, Trillium type. Cotyledon one, terminal. Cotyledon hyperphyll elongate, assimilating. Hypocotyl present. Hypocotyl internode absent (Typha) or short (Sparganium). Mesocotyl absent. Coleoptile absent (Typha) or present (Sparganium). Plumule lateral. Collar rhizoids present. Germination?

Phytochemistry Flavonols (in Typha kaempferol, quercetin, cinnamic acids; in Sparganium quercetin, myricetin), catechin, and cyanidin (in Sparganium also delphinidin?) present. Alkaloids and cyanogenic compounds rare. Ellagic acid, chelidonic acid, and saponins not found. Ferulic acid (ferulate, esterified) component of non-lignified cell walls.

Use Ornamental plants, Typha also for matting, basketry, vegetables (starchy rhizome), etc.

Xyridales Lindl., Veg. Kingd.: lvii, 185. 14-28 Mar 1846; Abolbodaceae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 221. 20 Jul 1943; Xyridineae thorne et Reveal in Bot. Rev. (Lancaster) 73: 85. 29 Jun 2007

Distribution Tropical and subtropical regions on the Southern and Northern Hemispheres, with their largest diversity in northern South America; few species in temperate areas.

Fossils Fossilized seeds attributed to Xyris have been found in the mid- and Late Miocene of Germany (Mai 2000).

Habit Bisexual, usually perennial (sometimes annual) herbs. Often helophytic (rarely aquatic); some species are xerophytic.

Vegetative anatomy Mycorrhiza absent. Roots fibrous. Root endodermis uni- or multilayered, consisting of large evenly thick cells with specific orientation. Root stele relatively narrow. Root phloem and xylem elements usually irregularly scattered in parenchyma. Lateral roots developing from a zone opposite protophloem poles. Phellogen absent. Stem vascular bundles amphivasal. Secondary lateral growth absent. Vessels present in roots and, usually, in stem and leaves. Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits scalariform. Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c_cof type, with cuneate and orthogonal protein crystals and peripheral protein filaments. Nodes? Silica bodies absent. Calciumoxalate as single crystals; raphides absent.

Trichomes Hairs unicellular or multicellular (few-celled), uniseriate, usually simple; in Xyris uniseriate hairs with bulb-like terminal cell producing mucilage present in leaf axils.

Leaves Alternate (usually distichous, sometimes spiral), simple, entire, linear, usually equitant, laterally flattened, sometimes subulate, with convolute (supervolute) ptyxis? Stipules absent; leaf sheath distinct, open, some species of Xyris with distal ligule (or auriculae). Venation parallelodromous. Stomata paracytic or anomocytic (in Abolboda sometimes tetracytic). Cuticular waxes absent. Cuticle with insoluble secretions. Outer epidermal cell walls often with papillae, tubercles or other types of outgrowths, or thickened. Mesophyll without mucilaginous cells, often with calciumoxalate as druses or solitary prismatic crystals (raphides absent). Leaf margin entire.

Inflorescence Terminal or lateral, dense, spike- or head-like condensed panicle (usually stachyoid; in Achlyphila open) consisting of one- or sometimes two- or three-flowered cymose partial inflorescences (in some species spikes in panicles; in Aratitiyopea racemose; in Achlyphila solitary flowers, each of which enclosed by an involucral bract), on long peduncle bracteate in lower part. Each flower usually subtended by one or several pairs of bracts. Floral prophyll (bracteole) absent.

Flowers Actinomorphic or zygomorphic (in Orectanthe bilabiate). Hypogyny. Tepals usually 3+3 (in Abolboda and Orectanthe sometimes 2+3); outer tepals usually three (sometimes two, carinate), sepaloid, free or two lateral connate at base; median tepal in Xyris membranous, caducous, or all outer tepals persistent (median outer tepal in Abolboda strongly reduced or absent); inner tepals with imbricate aestivation, petaloid, clawed, early caducous, usually free (sometimes large and often connate at base into a tube). Nectary absent. Disc absent.

Androecium Stamens usually three (outer staminal whorl absent), antepetalous (sometimes also one to three staminodia; one species of Xyris with six fertile stamens). Filaments often filiform, with flattened base, usually connate, usually adnate to inner tepals (epipetalous; in Achlyphila free). Anthers basifixed, non-versatile, tetrasporangiate (microsporangia sometimes connate), usually latrorse or extrorse (rarely introrse), longicidal (dehiscing by longitudinal slits). Tapetum usually secretory (in Abolbodoideae amoeboid-periplasmodial), with binucleate cells. Exothecium present. Staminodia (in Xyris and sometimes in Abolboda) usually three (sometimes one or two), alternipetalous, extrastaminal, free from tepals, simple, bifid or quadrifid, usually provided with long (often plumose) hairs (secondary pollen presentation).

Pollen grains Microsporogenesis successive. Pollen grains monosulcate (in Xyris; sometimes disulcate) or inaperturate (in Abolbodoideae), shed as monads, usually bicellular (sometimes tricellular) at dispersal. Exine tectate, with columellate? infratectum, spinulate or smooth.

Gynoecium Pistil composed of three connate carpels. Ovary superior, unilocular to entirely or partially (at least basally) trilocular. Style single, filiform, terete or triangular in cross-section in lower part, usually trilobate (in Achlyphila simple), in Abolboda, Aratitiyopea and Orectanthe trifid in upper part and with three lateral appendages. Stigmas three, lobate or U-shaped, often infundibuliform to subcapitate, usually fimbriate or papillate, type? Pistillodium absent.

Ovules Placentation in Xyris marginal, parietal (sometimes intrusively so), basal, basal-axile, free central or axile, in Abolbodoideae parietal. Ovules usually numerous (sometimes one) per carpel, anatropous (Achlyphila, Abolboda; in Orectanthe anatropous to slightly campylotropous)ororthotropous(Xyris), bitegmic, tenuinucellar (Xyris, Abolboda) to more or less crassinucellar (Achlyphila). Micropyle bistomal. Outer integument two cell layers thick. Inner integument two cell layers thick. Hypostase absent (in Xyris) or present (in Abolbodoideae). Obturator absent. Parietal cell not formed (parietal tissue absent). Megasporangial epidermis in Xyris sometimes with periclinal cell divisions. Megagametophyte monosporous, Polygonum type, or disporous, Allium type. Endosperm development ab initio helobial (or possibly sometimes nuclear?). Endosperm haustoria absent. Embryogenesis asterad.

Fruit Usually a loculicidal, many-seeded (rarely one-seeded) capsule (in Xyris rarely a pyxidium) with persistent tepals.

Seeds Aril absent. Seed coat testal-endotegmic, sometimes winged. Exotestal cells thin-walled. Endotestal cells thick-walled. Seeds sometimes operculate; operculum micropylar, formed from exotegmen or (in Xyris) both integuments. Tegmen mechanical, rich in resins and tannins. Perisperm not developed. Endosperm copious, starchy and proteinaceous (and sometimes oily); starch grains compound (Xyris) or simple (Abolboda). Embryo small (Xyris) or large, straight or curved (in Abolbodoideae large and curved), little differentiated or rudimentary, with chlorophyll, Xyris-Scirpus type. Cotyledon one. Cotyledon hyperphyll prolonged, assimilating. Hypocotyl internode short, with rhizoids. Coleoptile absent. Collar rhizoids usually present. Germination?

Phytochemistry Flavonols (kaempferol, quercetin), flavone glycosides, cyanidin, saponins, and chrysazine (an anthraquinone) present. Esterified ferulic acid (diferulic acid?, p-coumaric acid?) component of non-lignified cell walls. Phenolic compounds present or absent. Ellagic acid and alkaloids not found. Aluminium accumulated.

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CYPERALES Juss. ex Bercht. et J. Presl

ANARTHRIACEAE D. F. Cutler et Airy Shaw

BROMELIACEAE Juss.

CYPERACEAE Juss.

ECDEIOCOLEACEAE D. F. Cutler et Airy Shaw

ERIOCAULACEAE Martinov

FLAGELLARIACEAE Dumort.

JOINVILLEACEAE Toml. et A. C. Sm.

JUNCACEAE Juss.

MAYACACEAE Kunth

POACEAE (R. Br.) Barnhart

PRIONIACEAE S. L. Munro et H. P. Linder

RAPATEACEAE Dumort.

RESTIONACEAE R. Br.

THURNIACEAE Engl.

TYPHACEAE Juss.

XYRIDACEAE C. Agardh