EMBRYOPSIDA Pirani & Prado

Gametophyte dominant, independent, multicellular, thalloid, with single-celled apical meristem, showing gravitropism; rhizoids +, unicellular; acquisition of phenylalanine lysase [PAL], phenylpropanoid metabolism [lignans +, flavonoids + (absorbtion of UV radiation)], xyloglucans +; plant [protoplasm dessication tolerant], ectohydrous [free water outside plant physiologically important]; cuticle +; cell wall also with (1->3),(1->4)-ß-D-MLGs [Mixed-Linkage Glucans]; chloroplasts per cell, lacking pyrenoids; glycolate metabolism in leaf peroxisomes [glyoxysomes]; centrioles in vegetative cells 0, metaphase spindle anastral, predictive preprophase band of microtubules, phragmoplast + [cell wall deposition spreading from around the spindle fibres], plasmodesmata +; antheridia and archegonia jacketed, stalked; spermatogenous cells monoplastidic; blepharoplast, bicentriole pair develops de novo in spermatogenous cell, associated with basal bodies of cilia [= flagellum], multilayered structure [4 layers: L1, L4, tubules; L2, L3, short vertical lamellae] + 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 dependent on gametophyte, embryo initially surrounded by haploid gametophytic tissue, plane of first division horizontal [with respect to long axis of archegonium/embryo sac], suspensor/foot +, cell walls with nacreous thickenings; sporophyte multicellular, with at least transient apical cell [?level], sporangium +, single, dehiscence longitudinal; meiosis sporic, monoplastidic, microtubule organizing centre associated with plastid, cytokinesis simultaneous, preceding nuclear division, sporocytes 4-lobed, with a quadripolar microtubule system; spores in tetrads, sporopollenin in the spore wall, initially laid down in association with several trilamellar layers [white-line centred layers, i.e. walls multilamellate]; nuclear genome size <1.4 pg, LEAFY and KNOX1 and KNOX2 genes present, ethylene involved in cell elongation; chloroplast genome with close association between trnLUAA and trnFGAA genes.

Many of the bolded characters in the characterization above are apomorphies of subsets of streptophytes along the lineage leading to the embryophytes, not apomorphies of crown-group embryophytes per se.

All groups below are crown groups, nearly all are extant. Characters mentioned are those of the immediate common ancestor of the group, [] contains explanatory material, () features common in clade, exact status unclear.

STOMATOPHYTES

Abscisic acid, ?D-methionine +; sporangium - tapetum +, columella + [developing from endothecial cells], seta developing from basal meristem [between epibasal and hypobasal cells]; stomata +, anomocytic, cell lineage that produces them with symmetric divisions [perigenous]; underlying similarities in the development of conducting tissue and in rhizoids/root hairs; spores trilete; polar transport of auxins and class 1 KNOX genes expressed in the sporangium alone; shoot meristem patterning gene families expressed; MIKC, MI*K*C* and class 1 and 2 KNOX genes, post-transcriptional editing of chloroplast genes; gain of three group II mitochondrial introns.

[Anthocerophyta + Polysporangiophyta]: archegonia embedded/sunken in the gametophyte; sporophyte long-lived, chlorophyllous; sporophyte-gametophyte junction interdigitate, sporophyte cells showing rhizoid-like behaviour.

POLYSPORANGIOPHYTA†

Sporophyte branched, branching apical, dichotomous; sporangia several, each opening independently; spore walls not multilamellate [?here].

EXTANT TRACHEOPHYTA / VASCULAR PLANTS

Photosynthetic red light response; plant homoiohydrous [water content of protoplasm relatively stable]; control of leaf hydration passive; (condensed or nonhydrolyzable tannins/proanthocyanidins +); sporophyte soon independent, dominant, with basipetal polar auxin transport; lignins +; vascular tissue +, G- and S-type tracheids, sieve cells + [nucleus degenerating], tracheids +, in both protoxylem and metaxylem, plant endohydrous [physiologically important free water inside plant]; endodermis +; leaves spirally arranged, blades with mean venation density 1.8 mm/mm2 [to 5 mm/mm2]; sporangia adaxial on the sporophyll, derived from periclinal divisions of several epidermal cells, wall multilayered [eusporangium]; columella 0; tapetum glandular; gametophytes exosporic, green, photosynthetic; basal body 350-550 nm long, stellate array in transition region initially joining microtubule triplets; placenta with single layer of transfer cells in both sporophytic and gametophytic generations, root lateral with respect to the longitudinal axis of the embryo [plant homorhizic].

[MONILOPHYTA + LIGNOPHYTA]

Sporophyte branching ± indeterminate; root apex multicellular, root cap +, lateral roots +, endogenous; endomycorrhizal associations + [with Glomeromycota]; G-type tracheids +, with scalariform-bordered pits; leaves with apical/marginal growth, venation development basipetal, growth determinate; sporangia borne in pairs and grouped in terminal trusses, dehiscence longitudinal, a single slit; cells polyplastidic, microtubule organizing centres not associated with plastids, diffuse, perinuclear; blepharoplasts +, paired, with electron-dense material, centrioles on periphery, male gametes multiciliate; chloroplast long single copy ca 30kb inversion [from psbM to ycf2]; LITTLE ZIPPER proteins.

LIGNOPHYTA†

Sporophyte woody; lateral root origin from the pericycle; branching lateral, meristems axillary; cork cambium + [producing cork abaxially], vascular cambium bifacial [producing phloem abaxially and xylem adaxially].

SEED PLANTS†

Plants heterosporous; megasporangium surrounded by cupule [i.e. = unitegmic ovule, cupule = integument]; pollen lands on ovule; megaspore germination endosporic [female gametophyte initially retained on the plant].

EXTANT SEED PLANTS / SPERMATOPHYTA

Plant evergreen; nicotinic acid metabolised to trigonelline, (cyanogenesis via tyrosine pathway); primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins particularly with guaiacyl and p-hydroxyphenyl [G + H] units [sinapyl units uncommon, no Maüle reaction]; root stele with xylem and phloem originating on alternate radii, cork cambium deep seated; mitochondrial density in whole SAM 1.6-6.2[mean]/μm2 [interface-specific mitochondrial network]; stem with vascular cylinder around central pith [eustele], phloem abaxial [ectophloic], endodermis 0, xylem endarch [development centrifugal]; wood homoxylous, tracheids and rays alone, tracheid/tracheid pits circular, bordered; mature sieve tube/cell lacking functioning nucleus, sieve tube plastids with starch grains; phloem fibres +; cork cambium superficial; leaves with single trace from vascular sympodium [nodes 1:1]; stomatal pore with active opening in response to leaf hydration, control by abscisic acid, metabolic regulation of water use efficiency, etc.; axillary buds exogenous, (none; not associated with all leaves); prophylls two, lateral; leaves with petiole and lamina, development basipetal, blade simple; plant heterosporous, sporangia borne on sporophylls, sporophylls spiral; microsporophylls aggregated in indeterminate cones/strobili; grains monosulcate, aperture in ana- position [distal], primexine + [involved in exine pattern formation with deposition of sporopollenin from tapetum there], exine and intine homogeneous; megasporangium indehiscent; ovules with parietal tissue 2+ cells across, megaspore tetrad linear, functional megaspore single, chalazal, sporopollenin 0; gametophyte development initially endosporic, dependent on sporophyte, apical cell 0, rhizoids 0, development continuing outside the spore; male gametophyte with tube developing from distal end of grain, male gametes two, developing after pollination, with cell walls; female gametophyte initially syncytial, walls then surrounding individual nuclei; embryo cellular ab initio, endoscopic, plane of first cleavage of zygote transverse, suspensor +, short-minute, embryonic axis straight [shoot and root at opposite ends; plant allorhizic], cotyledons 2; plastid transmission maternal; ycf2 gene in inverted repeat, whole nuclear genome duplication [ζ - zeta - duplication], two copies of LEAFY gene, PHY gene duplications [three - [BP [A/N + C/O]] - copies], nrDNA with 5.8S and 5S rDNA in separate clusters; mitochondrial trans- nad2i542g2 and coxIIi3 introns present.

ANGIOSPERMAE / MAGNOLIOPHYTA

Lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, apigenin and/or luteolin scattered, [cyanogenesis in ANITA grade?], 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 apical meristem intermediate-open; root vascular tissue oligarch [di- to pentarch], lateral roots arise opposite or immediately to the side of [when diarch] 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; reaction wood ?, associated gelatinous fibres [g-fibres] with innermost layer of secondary cell wall rich in cellulose and poor in lignin; 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< µm across, cytoplasm with P-proteins, cytoplasm not occluding pores of sieve plate, companion cell and sieve tube from same mother cell; sugar transport in phloem passive; nodes 1:?; stomata brachyparacytic [ends of subsidiary cells level with ends of pore], outer stomatal ledges producing vestibule, reduction in stomatal conductance to increasing CO2 concentration; lamina formed from the primordial leaf apex, margins toothed, development of venation acropetal, overall growth ± diffuse, venation hierarchical-reticulate, secondary veins pinnate, veins (1.7-)4.1(-5.7) mm/mm2, endings free; most/all leaves with axillary buds; flowers perfect, pedicellate, ± haplomorphic; protogynous; parts spiral [esp. the A], free, numbers unstable, development in general centripetal; P +, members 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], sporangium pairs dehiscing longitudinally by a common slit, ± embedded in the filament, walls with at least outer secondary parietal cells dividing, endothecium +, endothecial cells elongated at right angles to long axis of anther; (tapetum glandular), 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; nectary 0; carpels present, superior, free, several, ascidiate, with postgenital occlusion by secretion, stylulus at most short [shorter than ovary], hollow, cavity not lined by distinct epidermal layer, stigma ± decurrent, carinal, stigma wet, extragynoecial compitum +; ovules few [?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 [crassinucellate], nucellar cap?; megasporocyte single, hypodermal, functional megaspore chalazal, lacking cuticle; female gametophyte lacking chlorophyll, not photsynthesising, four-celled [one module, nucleus of egg cell sister to one of the polar nuclei]; ovule not increasing in size between pollination and fertilization; pollen grains land on stigma, bicellular at dispersal, mature male gametophyte tricellular, germinating in less than 3 hours, pollen tube elongated, unbranched, growing between cells, growth rate (20-)80-20,000 µm/hour, apex of pectins, wall with callose, lumen with callose plugs, penetration of ovules via micropyle [porogamous], whole process takes ca 18 hours, distance to first ovule 1.1-2.1 mm; male gametes lacking cell walls, cilia 0, siphonogamy; double fertilization +, ovules aborting unless fertilized; P deciduous in fruit; mature seed much larger than ovule when fertilized, small [], dry [no sarcotesta], exotestal; endosperm diploid, cellular, heteropolar [micropylar and chalazal domains develop differently, first division oblique, micropylar end initially with a single large cell, divisions uniseriate, chalazal cell smaller, divisions in several planes], copious, oily and/or proteinaceous; dark reversal Pfr → Pr; Arabidopsis-type telomeres [(TTTAGGG)n]; nuclear genome size <1.4 pg [1 pg = 109 base pairs], whole nuclear genome duplication [ε - epsilon - duplication]; protoplasm dessication tolerant [plant poikilohydric]; ndhB gene 21 codons enlarged at the 5' end, single copy of LEAFY and RPB2 gene, knox genes extensively duplicated [A1-A4], AP1/FUL gene, paleo AP3 and PI genes [paralogous B-class genes] +, with "DEAER" motif, SEP3/LOFSEP and three copies of the PHY gene, [PHYB [PHYA + PHYC]].

[NYMPHAEALES [AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]]: wood fibres +; axial parenchyma diffuse or diffuse-in-aggregates; pollen monosulcate [anasulcate], tectum reticulate-perforate [here?]; ?genome duplication; "DEAER" motif in AP3 and PI genes lost, gaps in these genes.

[AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]: vessel elements with scalariform perforation plates in primary xylem; essential oils in specialized cells [lamina and P ± pellucid-punctate]; tension wood +; tectum reticulate; anther wall with outer secondary parietal cell layer dividing; carpels plicate; nucellar cap + [character lost where in eudicots?]; 12BP [4 amino acids] deletion in P1 gene.

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]] / MESANGIOSPERMAE: benzylisoquinoline alkaloids +; sesquiterpene synthase subfamily a [TPS-a] [?level], polyacetate derived anthraquinones + [?level]; outer epidermal walls of root elongation zone with cellulose fibrils oriented transverse to root axis; P more or less whorled, 3-merous [possible position]; pollen tube growth intra-gynoecial [extragynoecial compitum 0]; embryo sac bipolar, 8 nucleate, antipodal cells persisting; endosperm triploid.

[MONOCOTS [CERATOPHYLLALES + EUDICOTS]]: (extra-floral nectaries +); (veins in lamina often 7-17 mm/mm2 or more [mean for eudicots 8.0]); (stamens opposite [two whorls of] P); (pollen tube growth fast).

MONOCOTYLEDONS / MONOCOTYLEDONEAE / LILIANAE Takhtajan

Plant herbaceous, perennial, rhizomatous, growth sympodial; non-hydrolyzable tannins [(ent-)epicatechin-4] +, neolignans, benzylisoquinoline alkaloids 0, hemicelluloses as xylans; root apical meristem?; root epidermis developed from outer layer of cortex; trichoblasts in atrichoblast [larger cell]/trichoblast cell pairs, the former further from apical meristem, in vertical files; endodermal cells with U-shaped thickenings; cork cambium in root [uncommon] superficial; stele oligo- to polyarch, medullated [with prominent pith], lateral roots arise opposite phloem poles; primary thickening meristem +; vascular bundles in stem scattered, (amphivasal), vascular cambium 0 [bundles closed]; tension wood 0; vessel elements in root with scalariform and/or simple perforations; tracheids only in stems and leaves; sieve tube plastids with cuneate protein crystals alone; stomata parallel to the long axis of the leaf, in lines; prophyll single, adaxial; leaf blade linear, main venation parallel, the veins joining successively from the outside at the apex, transverse veinlets +, unbranched [leaf blade characters: ?level], vein/veinlet endings not free, margins entire, Vorläuferspitze +, base broad, ensheathing the stem, sheath open, petiole 0, colleters + ["intravaginal squamules"]; inflorescence terminal, racemose; flowers 3-merous [6-radiate to the pollinator], polysymmetric, pentacyclic; P = T, each with three traces, median T of outer whorl abaxial, aestivation open, members of whorls alternating, [pseudomonocyclic, each T member forming a sector of any tube]; stamens = and opposite each T member [primordia often associated, and/or A vascularized from tepal trace], anther and filament more or less sharply distinguished, anthers subbasifixed, endothecium from outer secondary parietal cell layer, inner secondary parietal cell layer dividing; pollen reticulations coarse in the middle, finer at ends of grain, infratectal layer granular; G [3], with congenital intercarpellary fusion, opposite outer tepals [thus median member abaxial], placentation axile; ovule with outer integument often largely dermal in origin, parietal tissue 1 cell across; antipodal cells persistent, proliferating; fruit a loculicidal capsule; seed small to medium sized [mean = 1.5 mg], testal; embryo long, cylindrical, cotyledon 1, apparently terminal, 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 +); cotyledon with a closed sheath, unifacial [hyperphyllar], both assimilating and haustorial; no dark reversion Pfr → Pr; duplication producing monocot LOFSEP and FUL3 genes [latter duplication of AP1/FUL gene], PHYE gene lost. (Some synapomorphies - almost whatever the immediate sister taxa to monocots might be - are in bold.)

[ALISMATALES [PETROSAVIALES [[DIOSCOREALES + PANDANALES] [LILIALES [ASPARAGALES + COMMELINIDS]]]]]: ethereal oils 0; 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]]]] / core monocots: cyanogenic glycosides uncommon; starch grains simple, amylophobic; leaf blade developing basipetally from hyperphyll/hypophyll junction; epidermis with bulliform cells [?level]; stomata anomocytic, (cuticular waxes as parallel platelets); colleters 0.

Age. The crown-group age of this group is ca 131 m.y. (Janssen & Bremer 2004). Magallón and Castillo (2009) suggest ca 150 m.y. for relaxed and 124 m.y. for constrained penalized likelihood datings of the crown group age, Bell et al. (2010) ages of (133-)124, 109(-99) m.y.; (127-)121, 109(-103) m.y. was estimated by Wikström et al. (2001: c.f. topology), and estimates were (132-)128(-121) m.y. in Merckx et al. (2008a), 133-108 m.y. in Mennes et al. (2013, see also 2015), ca 123.5 m.y.a. in Magallón et al. (2015) and similar estimates in Hertweck et al. (2015).

Evolution. Divergence & Distribution. Branching in this general part of the tree - i.e. the Petrosaviales, [Dioscoreales + Pandanales], and Liliales clades, and also in Petrosaviales itself, may be somewhere around 125-120 m.y.a. (ca 111 m.y. in Bremer 2000b), and the stem groups of all other orders, including those in the commelinid group, have diverged by ca 115 m.y.a. or soon afterwards (Janssen & Bremer 2004) or ca 106 m.y.a. (Magallón et al. 2015). Much of this divergence may have taken place in Southern Gondwana, i.e. Antarctica, Australasia, and southern South America (Bremer & Janssen 2006).

For a discussion on leaf development in monocots, see the Acorales page.

Phylogeny. Relationships between commelinids, Asparagales, Dioscoreales, Liliales, and Pandanales were unclear for some time. In a smallish early study, Liliales were sometimes embedded in Asparagales (Eguiarte et al. 1994). A three-gene (rbcL, atpB, 18S RNA) study (Chase et al. 2000a) showed a polytomy of Petrosaviaceae, Dioscoreales, Pandanales, Liliales, Asparagales and commelinids, although a single shortest tree showed a pectinate structure with the taxa in the sequence followed here; another analysis with placeholders for taxa missing some sequences gave a similar structure, except that Pandanales and Liliales were sister taxa. (A combined morphological plus molecular tree in the same volume [Stevenson et al. 2000] suggested a substantially different set of relationships; bootstraps were not given.) Fay et al. (2000a) also suggested a sister relationship between Asparagales and commelinids, although sampling outside Asparagales was sketchy since it was outside their immediate interest. Hilu et al. (2003: matK) i.a. suggested that Orchidaceae might be separate from other Asparagales (the latter being sister to commelinids) and that Dioscoreales and Pandanales formed a clade.

However, a two-gene (matK, rbcL) study (Tamura et al. 2004a) began to clarify the situation. Petrosaviaceae (both genera - but see below for a possible third - were studied) were sister to a clade [[Dioscoreaceae + Pandanaceae] [Liliales [Asparagales + commelinids]]] (see Chase et al. 2000a above). Support was quite high (³85% bootstrap) for all order and family branches, although rather lower for [Asparagales + commelinids] (68%) (see also Tamura et al. 2004b, a smaller study). Davis et al. (2004) also found Petrosaviales to be sister to the same monocots, but with moderate to weak (>72%) support. Givnish et al. (2005: ndhF gene alone) found very much the set of relationships in the tree here, although Pandanales grouped with Liliales (low support) and Dasypogonaceae were sister to [Commelinales + Zingiberales]; a grouping [Liliales [Pandanales + Dioscoreales]] also appeared - and had moderate support - in MP, but not in ML analyses of plastid genomes in Barrett et al. (2013: sampling). Graham et al. (2006) in a study analysing considerable amounts of data also recovered relationships similar to those suggested by Tamura et al. (2004a), all sister taxon relationships in this area having 94% or more support, although that for [Liliales [commelinids + Asparagales]] was only 70% (see also Givnish et al. 2006b; Chase et al. 2006; Magallón et al. 2015). Dioscoreales and Pandanales are sister taxa in most studies (e.g. Hilu et al. 2003; Chase et al. 2006; Qiu et al. 2010: support strong; Magallón et al. 2015; Hertweck et al. 2015; Lam et al. 2015).

There is perhaps still some uncertainty. In some reconstructions Dioscoreales and Pandanales are adjacent along the spine (e.g. Janssen & Bremer 2004; Bremer & Janssen 2006; Givnish et al. 2006b: not strongly supported), or Nartheciaceae link with Pandanales, the combined group in turn joining with Liliales (Davis et al. 2004: summary of earlier literature on relationships of the two). A four-gene mitochondrial tree suggested the relationships [Asparagales [[Dioscoreales + Pandanales] [Liliales + Commelinids]]], but support was not strong (Qiu et al. 2010), while Davis et al. (2013) recovered a very weakly supported topology [Asparagales s. str. [Orchidaceae + Liliaceae]] in parsimony but not in maximum likelihood analyses (see also Barrett & Davis 2011). Ruhfel et al. (2014) found a [Liliales [Dioscoreales + Pandanales]] clade while in analyses of the nuclear PHYC gene alone Orchidaceae was not sister to other Asparagales - relationships were [Asparagales [Liliales [Orchidaceae + commelinids]]], but support was weak (Hertweck et al. 2015).

Some myco-heterotrophic taxa cause problems. Thus Neyland (2002) found that Thismia was sister to a well supported Burmannioideae, but with less support, but Burmanniaceae s.l. did not link with other Dioscoreales. Analysis of 26S rDNA sequences suggested that Corsiaceae were polyphyletic; Arachnitis perhaps being sister to Thismia and/or Burmannia (Neyland & Hennigan 2003; G. Petersen et al. 2006b: combined analysis). A recent analysis of plastid loci also failed to include Arachnitis in Liliales, and perhaps it is to be included in the commelinids (Kim et al. 2012). However, these relationships need to be confirmed.

Classification. Since no firm association of Petrosaviaceae with any other order has been supported, and its phylogenetic position, as in the tree here, seems well supported, a monofamilial Petrosaviales is appropriate.

PETROSAVIALES Takhtajan  Main Tree.

Stem with a ring of bundles; sieve tube plastids also with polygonal protein crystalloids; inflorescence bracts +; microsporogenesis simultaneous; pollen surface gemmate; septal nectaries +; styluli +; fruit a follicle; seed endotestal. - 1 family, 2 genera, 4 species.

Note: (....) denotes a feature common in the clade, exact status uncertain, [....] includes explanatory material. Possible apomorphies are in bold. However, the actual level at which many of these features, particularly the more cryptic ones, should be assigned is unclear. This is partly because many characters show considerable homoplasy, in addition, basic information for all too many is very incomplete, frequently coming from taxa well embedded in the clade of interest and so making the position of any putative apomorphy uncertain. Then there are the not-so-trivial issues of how character states are delimited and ancestral states are reconstructed (see above).

Evolution. Divergence & Distribution. This is not exactly a clade in which there has been much divergence Hertweck et al. 2015).

Includes Petrosaviaceae.

Synonymy: Miyoshiales Nakai - Petrosaviineae Shipunov

PETROSAVIACEAE Hutchinson

Petrosaviaceae

(Plant echlorophyllous, myco-heterotrophic, vascular bundles in stem forming a cylinder, vessels 0 - Petrosavia); plant glabrous; leaves spiral, scaly on rhizome, base?; bracteoles sublateral (0); T members with a single trace, whorls slightly differentiated [outer somewhat smaller], tube at most short; A inserted at base of T or free; (pollen gemmate - Japonolirion); ovary superior to semi-inferior, partly connate, plicate, fusion (congenital and) postgenital, stigma subcapitate or decurrent; ovules 4-many/carpel, ana-campylotropous, integumentary obturator +; fruit also septicidal [Japonolirion], T persistent [?Petrosavia]; seeds obliquely arranged, winged or not, endotegmen crushed, contents persist; embryo small; n = 12, 13, 15; seedling?

2[list]/4. Japan and China, W. Malesia (map: from Jessop 1979).

Age. Crown group Petrosaviaceae date to ca 123 m.y. (Janssen & Bremer 2004) or (134-)99, 89(-62) m.y. (Bell et al. 2010); estimates were (102-)108(-87) m.y. in Merckx et al. (2008a), 96-8 m.y. and 91-9 m.y. in Mennes et al. (2013, 2015 respectively) and (115-)108(-99) or (64-)60(-57) m.y. in Hertweck et al. (2015).

Evolution. Bacterial/Fungal Associations. Japonolirion osense has about 22 phylotypes of Glomales associated with it (Paris-type mycorrhizae here); the mycoheterotrophic Petrosavia sakuraii has but a single one of these (Yamato et al. 2014).

Genes & Genomes. For evolution (loss of genes, etc.) in the chloroplast genome of the myco-heterotroph Petrosavia, see Logacheva et al. (2014); rather unusually for such systems, there has been extensive rearrangment of the genome.

Chemistry, Morphology, etc. The roots of Petrosavia have an unmedullated, four-radiate stele. Remizowa (2011) suggested that the position of the septal nectaries in both the ascidiate and plicate zone of the gynoecium might be unique and so a synapomorphy for this tiny but quite heterogeneous clade.

For general information, see Tamura (1998: Nartheciaceae) and especially Cameron et al. (2003), for anatomy, see Stant (1970), for pollen, see Caddick et al. (1998), for sieve tube plastids, see Behnke (2003), for floral and inflorescence morphology, see Remizowa et al. (2006a, b) and Tobe (2008), and for the embryology of Petrosavia, see Tobe and Takahashi (2009: nice comparative table).

Phylogeny. Isidrogalvia (as I. schomburgkiana, Tofieldiaceae!) was placed sister to Petrosavia (L.-Y. Chen et al. 2013) in a study focusing on the aquatic Alismatales - a rather remarkable position that needs confirmation.

Previous Relationships. Petrosaviaceae have often been included in other families. Dahgren et al. (1985) placed them - along with genera here placed in Nartheciaceae and Tofieldiaceae - in Melianthaceae, and while Tamura (1998) recognised a Petrosaviaceae, this also included members of Tofieldiaceae and Nartheciaceae. Petrosaviaceae s. str. (i.e. Petrosavia alone) were placed in Triuridales by Cronquist (1981) and in Triurididae by Takhtajan (1997); the latter included a monogeneric Japonoliriaceae in his Melanthiales-Liliidae.

Synonymy: Japonoliriaceae Takhtajan, Miyoshiaceae Nakai