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Lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, apigenin and/or luteolin scattered, [cyanogenesis in ANA grade? The three members of the old Xyridales that remain here may form a grade as follows: [Mayacaceae Xyridaceae Eriocaulaceae] [Thurniaceae [Juncaceae Cyperaceae (Givnish et al. Xyridaceae etc.] (the latter grouping was also recovered by Davis et al. There was weak support for an [Eriocaulaceae Xyridaceae] clade in the phylogenomic analysis of Mc Kain et al. Poaceae and their immediate relatives consistently form a clade, although details of relationships within it are still somewhat unclear (see below). ], lignin also with syringyl units common [G S lignin, positive Maüle reaction - syringyl:guaiacyl ratio more than 2-2.5:1], hemicelluloses as xyloglucans; root cap meristem closed (open); pith relatively inconspicuous, lateral roots initiated immediately to the side of [when diarch] or opposite xylem poles; origin of epidermis with no clear pattern [probably from inner layer of root cap], trichoblasts [differentiated root hair-forming cells] 0, hypodermis suberised and with Casparian strip [= exodermis]; shoot apex with tunica-corpus construction, tunica 2-layered; starch grains simple; primary cell wall mostly with pectic polysaccharides, poor in mannans; tracheid:tracheid [end wall] plates with scalariform pitting, wood parenchyma ; sieve tubes enucleate, sieve plate with pores (0.1-)0.5-10; protoplasm dessication tolerant [plant poikilohydric]; stomata randomly oriented, brachyparacytic [ends of subsidiary cells level with ends of pore], outer stomatal ledges producing vestibule, reduction in stomatal conductance with increasing CO concentration; lamina formed from the primordial leaf apex, margins toothed, development of venation acropetal, overall growth ± diffuse, secondary veins pinnate, fine venation hierarchical-reticulate, (1.7-)4.1(-5.7) mm/mm, vein endings free; flowers perfect, pedicellate, ± haplomorphic, protogynous; parts free, numbers variable, development centripetal; T , petal-like, each with a single trace, outer members not sharply differentiated from the others, not enclosing the floral bud; A many, filament not sharply distinguished from anther, stout, broad, with a single trace, anther introrse, tetrasporangiate, sporangia in two groups of two [dithecal], each theca dehiscing longitudinally by a common slit, ± embedded in the filament, walls with at least outer secondary parietal cells dividing, endothecium , cells elongated at right angles to long axis of anther; tapetal cells binucleate; microspore mother cells in a block, microsporogenesis successive, walls developing by centripetal furrowing; pollen subspherical, tectum continuous or microperforate, ektexine columellate, endexine lamellate only in the apertural regions, thin, compact, intine in apertural areas thick, pollenkitt ; nectary 0; carpels present, superior, free, several, spiral, ascidiate [postgenital occlusion by secretion], stylulus at most short [shorter than ovary], hollow, cavity not lined by distinct epidermal layer, stigma ± decurrent, carinal, dry; suprastylar extragynoecial compitum ; ovules few [? Note that in versions 6 [before November] and earlier of this site, Eriocaulaceae and their relatives were weakly associated with the Poaceae group.

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Lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, apigenin and/or luteolin scattered, [cyanogenesis in ANA grade? The three members of the old Xyridales that remain here may form a grade as follows: [Mayacaceae

bend in embryo axis], with a closed sheath, unifacial [hyperphyllar], both assimilating and haustorial, plumule apparently lateral; primary root unbranched, not very well developed, stem-borne roots numerous, hypocotyl short, (collar rhizoids ); no dark reversion P; duplication producing monocot LOFSEP and FUL3 genes [latter duplication of AP1/FUL gene], PHYE gene lost.

Pirani & Prado Gametophyte dominant, independent, multicellular, initially ±globular, not motile, branched; showing gravitropism; acquisition of phenylalanine lysase* [PAL], flavonoid synthesis*, microbial terpene synthase-like genes , triterpenoids produced by CYP716 enzymes, phenylpropanoid metabolism [development of phenolic network], xyloglucans in primary cell wall, side chains charged; plant poikilohydrous [protoplasm dessication tolerant], ectohydrous [free water outside plant physiologically important]; thalloid, leafy, with single-celled apical meristem, tissues little differentiated, rhizoids , unicellular; chloroplasts several per cell, pyrenoids 0; glycolate metabolism in leaf peroxisomes [glyoxysomes]; centrioles/centrosomes in vegetative cells 0, microtubules with γ-tubulin along their lengths [? (2013, 2015 respectively), while ages in Hertweck et al. This is partly because homoplasy is very common, in addition, basic information for all too many characters is very incomplete, frequently coming from taxa well embedded in the clade of interest and so making the position of any putative apomorphy uncertain. 1965; Hultén & Fries 1986; Flora Base 2005 - somewhat notional: the map in Knobloch & Mai 1986 differs very considerably from its source, Meusel et al.

here], interphase microtubules form hoop-like system; metaphase spindle anastral, predictive preprophase band [with microtubules and F-actin; where new cell wall will form], phragmoplast [cell wall deposition centrifugal, from around the anaphase spindle], plasmodesmata ; antheridia and archegonia , jacketed*, surficial; blepharoplast , centrioles develop de novo, bicentriole pair coaxial, separate at midpoint, centrioles rotate, associated with basal bodies of cilia, multilayered structure [4 layers: L1, L4, tubules; L2, L3, short vertical lamellae] (0), spline [tubules from L1 encircling spermatid], basal body 200-250 nm long, associated with amorphous electron-dense material, microtubules in basal end lacking symmetry, stellate array of filaments in transition zone extended, axonemal cap 0 [microtubules disorganized at apex of cilium]; male gametes [spermatozoids] with a left-handed coil, cilia 2, lateral; oogamy; sporophyte *, multicellular, growth 3-dimensional*, cuticle *, plane of first cell division transverse [with respect to long axis of archegonium/embryo sac], sporangium and upper part of seta developing from epibasal cell [towards the archegonial neck, exoscopic], with at least transient apical cell [? Then there are the not-so-trivial issues of how character states are delimited and ancestral states are reconstructed (see above).

[ALISMATALES [PETROSAVIALES DIOSCOREALES PANDANALES] [LILIALES [ASPARAGALES COMMELINIDS (Givnish et al. Xyridaceae etc.] (the latter grouping was also recovered by Davis et al. There was weak support for an [Eriocaulaceae Xyridaceae] clade in the phylogenomic analysis of Mc Kain et al. Poaceae and their immediate relatives consistently form a clade, although details of relationships within it are still somewhat unclear (see below).

], lignin also with syringyl units common [G S lignin, positive Maüle reaction - syringyl:guaiacyl ratio more than 2-2.5:1], hemicelluloses as xyloglucans; root cap meristem closed (open); pith relatively inconspicuous, lateral roots initiated immediately to the side of [when diarch] or opposite xylem poles; origin of epidermis with no clear pattern [probably from inner layer of root cap], trichoblasts [differentiated root hair-forming cells] 0, hypodermis suberised and with Casparian strip [= exodermis]; shoot apex with tunica-corpus construction, tunica 2-layered; starch grains simple; primary cell wall mostly with pectic polysaccharides, poor in mannans; tracheid:tracheid [end wall] plates with scalariform pitting, wood parenchyma ; sieve tubes enucleate, sieve plate with pores (0.1-)0.5-10; protoplasm dessication tolerant [plant poikilohydric]; stomata randomly oriented, brachyparacytic [ends of subsidiary cells level with ends of pore], outer stomatal ledges producing vestibule, reduction in stomatal conductance with increasing CO concentration; lamina formed from the primordial leaf apex, margins toothed, development of venation acropetal, overall growth ± diffuse, secondary veins pinnate, fine venation hierarchical-reticulate, (1.7-)4.1(-5.7) mm/mm, vein endings free; flowers perfect, pedicellate, ± haplomorphic, protogynous; parts free, numbers variable, development centripetal; T , petal-like, each with a single trace, outer members not sharply differentiated from the others, not enclosing the floral bud; A many, filament not sharply distinguished from anther, stout, broad, with a single trace, anther introrse, tetrasporangiate, sporangia in two groups of two [dithecal], each theca dehiscing longitudinally by a common slit, ± embedded in the filament, walls with at least outer secondary parietal cells dividing, endothecium , cells elongated at right angles to long axis of anther; tapetal cells binucleate; microspore mother cells in a block, microsporogenesis successive, walls developing by centripetal furrowing; pollen subspherical, tectum continuous or microperforate, ektexine columellate, endexine lamellate only in the apertural regions, thin, compact, intine in apertural areas thick, pollenkitt ; nectary 0; carpels present, superior, free, several, spiral, ascidiate [postgenital occlusion by secretion], stylulus at most short [shorter than ovary], hollow, cavity not lined by distinct epidermal layer, stigma ± decurrent, carinal, dry; suprastylar extragynoecial compitum ; ovules few [? Note that in versions 6 [before November] and earlier of this site, Eriocaulaceae and their relatives were weakly associated with the Poaceae group.

(2015) described a spikelet assigned to Poaceae (parts spirally arranged) that was infected with a Claviceps-like fungus, and they noted "With the discovery of a Claviceps-like fossil infecting a floret of an Earlymid Cretaceous Asian grass, we propose that the progenitor of Claviceps evolved in Asia among early grasses sometime in the mid- to Late Jurassic. old and from Brazil (Leme & Brown 2011), is confirmed, older dates for the clade are again suggested.

Similarly, if the identity of the putative stem Bromeliaceae Protoananas lucenae, 114-112 m.y.

position of transfer cells; MTOCs not associated with plastids, basal body 350-550 nm long, stellate array in transition region initially joining microtubule triplets; suspensor , shoot apex developing away from micropyle/archegonial neck [from hypobasal cell, endoscopic], root lateral with respect to the longitudinal axis of the embryo [plant homorhizic]. For primary cell wall composition, see literature in Harris (2005); Arecaceae sampled are somewhat intermediate between this clade and other monocots. (2005) suggested an age of 109-106 m.y., Magallón and Castillo (2009) ages of ca 109 and 99.2 m.y., Bell et al. There is variation in the way pollen is arranged in the pollen loculi. (2004) for Si O bodies (phytoliths) and Tillich (2007) for seedling morphology and evolution. Poales do not always have very strong support, but c.f. A three-nucleotide deletion in the atp A gene was found to characterise Typhaceae and Bromeliaceae (Davis et al. Other work also suggests that Typhaceae and Bromeliaceae form a clade sister to other Poales, and Rapateaceae are in turn consistently sister to the remainder (Givnish et al. 2006 also found the position of Hydatellaceae to be problematic; for the association of Mayacaceae with Eriocaulaceae and Xyridaceae, see also Campbell et al. Members of this group of families were in adjacent branches along the spine of the tree, with one including Flagellariaceae, the Juncaceae group, some Xyridaceae, Mayacaceae, and perhaps Hydatellaceae.

]] DIOSCOREALES PANDANALES] [LILIALES [ASPARAGALES COMMELINIDS]]]: ethereal oils 0; (trichoblasts in vertical files, proximal cell smaller); raphides (druses 0); leaf blade vernation supervolute-curved or variants, (margins with teeth, teeth spiny); endothecium develops directly from undivided outer secondary parietal cells; tectum reticulate with finer sculpture at the ends of the grain, endexine 0; (septal nectaries ) [intercarpellary fusion postgenital].

[PETROSAVIALES DIOSCOREALES PANDANALES] [LILIALES [ASPARAGALES COMMELINIDS]]: cyanogenic glycosides uncommon; starch grains simple, amylophobic; leaf blade developing basipetally from hyperphyll/hypophyll junction; epidermis with bulliform cells [?

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bend in embryo axis], with a closed sheath, unifacial [hyperphyllar], both assimilating and haustorial, plumule apparently lateral; primary root unbranched, not very well developed, stem-borne roots numerous, hypocotyl short, (collar rhizoids ); no dark reversion P; duplication producing monocot LOFSEP and FUL3 genes [latter duplication of AP1/FUL gene], PHYE gene lost. Pirani & Prado Gametophyte dominant, independent, multicellular, initially ±globular, not motile, branched; showing gravitropism; acquisition of phenylalanine lysase* [PAL], flavonoid synthesis*, microbial terpene synthase-like genes , triterpenoids produced by CYP716 enzymes, phenylpropanoid metabolism [development of phenolic network], xyloglucans in primary cell wall, side chains charged; plant poikilohydrous [protoplasm dessication tolerant], ectohydrous [free water outside plant physiologically important]; thalloid, leafy, with single-celled apical meristem, tissues little differentiated, rhizoids , unicellular; chloroplasts several per cell, pyrenoids 0; glycolate metabolism in leaf peroxisomes [glyoxysomes]; centrioles/centrosomes in vegetative cells 0, microtubules with γ-tubulin along their lengths [? (2013, 2015 respectively), while ages in Hertweck et al. This is partly because homoplasy is very common, in addition, basic information for all too many characters is very incomplete, frequently coming from taxa well embedded in the clade of interest and so making the position of any putative apomorphy uncertain. 1965; Hultén & Fries 1986; Flora Base 2005 - somewhat notional: the map in Knobloch & Mai 1986 differs very considerably from its source, Meusel et al. here], interphase microtubules form hoop-like system; metaphase spindle anastral, predictive preprophase band [with microtubules and F-actin; where new cell wall will form], phragmoplast [cell wall deposition centrifugal, from around the anaphase spindle], plasmodesmata ; antheridia and archegonia , jacketed*, surficial; blepharoplast , centrioles develop de novo, bicentriole pair coaxial, separate at midpoint, centrioles rotate, associated with basal bodies of cilia, multilayered structure [4 layers: L1, L4, tubules; L2, L3, short vertical lamellae] (0), spline [tubules from L1 encircling spermatid], basal body 200-250 nm long, associated with amorphous electron-dense material, microtubules in basal end lacking symmetry, stellate array of filaments in transition zone extended, axonemal cap 0 [microtubules disorganized at apex of cilium]; male gametes [spermatozoids] with a left-handed coil, cilia 2, lateral; oogamy; sporophyte *, multicellular, growth 3-dimensional*, cuticle *, plane of first cell division transverse [with respect to long axis of archegonium/embryo sac], sporangium and upper part of seta developing from epibasal cell [towards the archegonial neck, exoscopic], with at least transient apical cell [? Then there are the not-so-trivial issues of how character states are delimited and ancestral states are reconstructed (see above). [ALISMATALES [PETROSAVIALES DIOSCOREALES PANDANALES] [LILIALES [ASPARAGALES COMMELINIDS]]]: ethereal oils 0; (trichoblasts in vertical files, proximal cell smaller); raphides (druses 0); leaf blade vernation supervolute-curved or variants, (margins with teeth, teeth spiny); endothecium develops directly from undivided outer secondary parietal cells; tectum reticulate with finer sculpture at the ends of the grain, endexine 0; (septal nectaries ) [intercarpellary fusion postgenital]. [PETROSAVIALES DIOSCOREALES PANDANALES] [LILIALES [ASPARAGALES COMMELINIDS]]: cyanogenic glycosides uncommon; starch grains simple, amylophobic; leaf blade developing basipetally from hyperphyll/hypophyll junction; epidermis with bulliform cells [?

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1]/carpel, marginal, anatropous, bitegmic, micropyle endostomal, outer integument 2-3 cells across, often largely subdermal in origin, inner integument 2-3 cells across, often dermal in origin, parietal tissue 1-3 cells across, nucellar cap? Includes Bromeliaceae, Cyperaceae, Ecdeiocoleaceae, Eriocaulaceae, Flagellariaceae, Joinvilleaceae, Juncaceae, Mayacaceae, Poaceae, Rapateaceae, Restionaceae, Thurniaceae, Typhaceae, Xyridaceae. 2015: Table S2) - or, if they are not sister taxa, the crown-group age for Poales as a whole - is estimated at around (105-)100(-95) m.y.a.

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