LIGNOPHYTA

True roots +; lateral meristems: cork cambium producing cork abaxially, vascular cambium producing phloem abaxially and xylem adaxially.

EXTANT SEED PLANTS/SPERMATOPHYTA

Plant woody, 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 derived from (some) sinapyl and particularly coniferyl alcohols, thus containing p-hydroxyphenyl and guaiacyl lignin units, (lignins derived from p-coumaryl alcohol, i.e. S [syringyl] lignin units); true roots present, apex multicellular, xylem exarch, and branching endogenous; arbuscular mycorrhizae +; shoot apical meristem multicellular, interface specific plasmodesmatal network; stem with ectophloic eustele, endodermis 0, xylem endarch, branching exogenous; vascular tissue in t.s. discontinuous by interfascicular regions; vascular cambium + [xylem ("wood") differentiating internally, phloem externally]; wood homoxylous, tracheids and rays alone, tracheid/tracheid pits circular, bordered; mature sieve tube/cell lacking functioning nucleus, plastids with starch grains; phloem fibres +; stem cork cambium superficial, root cork cambium deep seated; leaves with single trace from sympodium ["nodes 1:1"]; stomata ?; leaf vascular bundles collateral; leaves megaphyllous [determinancy evolved first, then ad/abaxial symmetry], spiral, simple, lamina with vein density up to 5 mm/mm² [mean for all non-angiosperms 1.8]; axillary buds associated with at most some leaves; prophylls [including bracteoles] two, lateral; plant heterosporous, sporangia eusporangiate, on sporophylls, sporophylls aggregated in indeterminate cones/strobili; true pollen [microspores, i.e. no distal pore for release of gametes] +, grains mono[ana]sulcate, exine and intine homogeneous; ovules unitegmic, crassinucellate, megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development first endo- then exosporic, tube developing from distal end of grain, to ca 2 mm from receptive surface to egg, gametes two, developing after pollination, with cell walls, with many flagellae; female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large", first cell wall of zygote transverse, embryo straight, endoscopic [suspensor +], short-minute, with morphological dormancy, white, cotyledons 2; plastid transmission maternal; 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 nad1 intron 2 and coxIIi3 intron and trans-spliced introns present.

MAGNOLIOPHYTA

Lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, [cyanogenesis in ANITA grade?], S [syringyl] lignin units common, positive Maüle reaction [syringyl:guaiacyl ratio more than 2-2.5:1], and 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; shoot apex with tunica-corpus construction, tunica 2-layered; reaction wood ?, with gelatinous fibres; 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 cells from same mother cell that gave rise to the sieve tube; sugar transport in phloem passive; nodes unilacunar [1:?]; stomata with ends of guard cells level with pore, paracytic, outer stomatal ledges producing vestibule; leaves petiolate, lamina [formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, veins (1.7-)4.1(-5.7) mm/mm², endings free; most/all leaves with axillary buds; flowers perfect, pedicellate, polysymmetric, parts spiral [esp. the A], free, numbers unstable, development in general centripetal; P not sharply differentiated, with a single trace, outer members not enclosing the rest of the bud, often smaller than inner members; A many, filament not sharply distinguished from anther, stout, broad, with a single trace, anther introrse, tetrasporangiate, sporangia in two groups of two [dithecal], ± embedded in the filament, with at least outer secondary parietal cells dividing, each theca dehiscing longitudinally by action of hypodermal endothecium, endothecial cells elongated at right angles to long axis of anther; tapetum glandular, binucleate; microspore mother cells in a block, microsporogenesis successive, walls developing by centripetal furrowing; pollen subspherical, tectum continuous or microperforate, ektexine columellar, endexine thin, compact, lamellate only in the apertural regions; nectary 0; G free, several, ascidiate, with postgenital occlusion by secretion, stylulus short, hollow, cavity not lined by distinct epidermal layer, stigma ± decurrent, dry [not secretory]; 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, megaspore tetrad linear, functional megaspore chalazal, lacking sporopollenin and cuticle; female gametophyte four-celled [one module, nucleus of egg cell sister to one of the polar nuclei]; P deciduous in fruit; seed exotestal; pollen binucleate at dispersal, trinucleate eventually, germinating in less than 3 hours, pollination siphonogamous, tube elongated, growing at 80-600 µm/hour, with pectic outer wall, callose inner wall and callose plugs, growing between cells, penetration of ovules via micropyle [porogamous] within ca 18 hours, distance to first ovule 1.1.-2.1 mm, tube moves between nucellar cells; double fertilisation +, endosperm diploid, cellular [micropylar and chalazal domains develop diffently, 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, embryo cellular ab initio, minute; germination hypogeal, seedlings/young plants sympodial; Arabidopsis-type telomeres [(TTTAGGG)_n]; whole genome duplication, 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]].

Evolution. Possible apomorphies for flowering plants are in bold. Note that the actual level to which many of these features, particularly the more cryptic ones, should be assigned is unclear. This is because some taxa basal to the [magnoliid + monocot + eudicot] group have been surprisingly little studied, there is considerable homoplasy as well as variation within and between families of the ANITA grade in particular for several of these characters, and also because details of relationships among gymnosperms will affect the level at which some of these characters are pegged. For example, if reticulate-perforate pollen is optimized to the next node on the tree (see Friis et al. 2009 for a discussion), it effectively makes the pollen morphology of the common ancestor of all angiosperms ambiguous... For other features such as details of sugar transport in the phloem, their placement on the tree is frankly speculative. Finally, for features such as parietal tissue/a nucellus only one (Nymphaeales) to three cells thick above the embryo sac and a stylar canal lacking an epidermal layer, although plesiomorphous for basal grade angiosperms (Williams 2009), I am unsure where on the tree a thicker nucellus and a stylar epidermal layer are acquired.

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

[ERICALES [ASTERID I + II]]: ovules tenuinucellate.

[ASTERID I + II] / CORE ASTERIDS: ellagic acid 0, non-hydrolysable tannins not common; sugar transport in phloem active; inflorescence basically cymose; C forming a distinct tube; A epipetalous, = and opposite sepals or P, polyandry associated with increased numbers of C or G, very uncommon; (pollen with orbicules); duplication of the PI gene.

ASTERID II / CAMPANULIDAE: myricetin 0; vessel elements with scalariform perforation plates; flowers rather small; style short; endosperm copious, embryo short/very short.

[ASTERALES [ESCALLONIALES [BRUNIALES [APIALES [PARACRYPHIALES + DIPSACALES]]]]] / APIIDAE: iridoids +; inflorescence?; C tube initiation early; G [2-3], inferior.

[ESCALLONIALES [BRUNIALES [APIALES [PARACRYPHIALES + DIPSACALES]]]]: ?

[BRUNIALES [APIALES [PARACRYPHIALES + DIPSACALES]]]: ?

[APIALES [PARACRYPHIALES + DIPSACALES]] / DIPSAPIIDAE: ?

[PARACRYPHIALES + DIPSACALES] / DIPSIDAE: nodes 3:3; true tracheids +; lamina serrate; inflorescence terminal.

Phylogeny. For the placement of Paracryphiaceae/Paracryphiales, see Winkworth et al. (2008) and especially Tank and Donoghue (2010); support for the position here was only slight in the full analysis of Soltis et al. (2011).

PARACRYPHIALES Reveal  Main Tree, Synapomorphies.

Inflorescence racemose; flowers 4-merous; anther thecae ± embedded in connective, filaments stout; G position?; capsule septicidal. - 1 family, 3 genera, 36 species.

Evolution. Divergence & Distribution. Magallón and Castillo (2009) offer estimates of ca 97.2 and 96.8 million years for relaxed and constrained penalized likelihood datings respectively for the stem group - but note topology, estimate probably too high.

Previous Relationships. Various families have been placed here or elsewhere in the asterid II clade, but with uncertain support; recent work is clarifying their relationships (Winkworth et al. 2008a; Tank & Donoghue 2010). For Polyosmaceae and Escalloniaceae, here as Escalloniaceae, see Escalloniales; Paracryphiaceae, Quintiniaceae, and Sphenostemonaceae are combined as Paracryphiaceae and are dealt with below; while Columelliaceae and Desfontainiaceae (as Columelliaceae) are close Bruniaceae, together making up Bruniales.

Includes Paracryphiaceae.

Synonymy: Sphenostemonineae I. Savin. - Quintiniales Doweld, Sphenostemonales Doweld

PARACRYPHIACEAE Airy-Shaw   Back to Paracryphiales

Leaves spiral.

3 [list]/36 - three genera below. S.W. Pacific, Philippines to Australia and New Caledonia.

1. Paracryphia Baker f.

Woody; chemistry?; petiole bundle flattened-annular, with medullary bundles; styloids +; hairs unicellular; plant andromonoecious; inflorescence branched-spicate; P (5), decussate-cochleate, caducous; A 8(-11); nectary 0; G [8-15], attached to central axial tissue, stigmas central, separate, conduplicate; ovules 4/carpel, crassinucellate; fruit with carpels pulling away acropetally and opening adaxially, columella persistent; seeds winged, exotesta? with sinuous anticlinal walls, inner walls lignified; embryo size?, radicle relatively long; n = ?

1/1: Paracryphia alticola. New Caledonia.

2. Quintinia A. de Candolle

Trees; plants Al accumulators, group 1 secoiridoids, ellagic acid +; vessels vestured; petiole bundle?; glands peltate; (lamina margins entire); flowers also 5-merous; C free; filaments less stout, anthers ± basifixed, placentoid ?; pollen 4-6 colporate; G [3-5], inferior, placentation parietal or axile, nectary on top of ovary, style long, stigma lobed, wet; ovules many/carpel, bitegmic, micropyle endostomal; K persistent; micropylar haustoria ?; n = ?22.

1/25. Philippines and New Guinea to New Zealand and New Caledonia (map: from Heywood 2007, in part).

Synonymy: Quintiniaceae Doweld

3. Sphenostemon Baillon

Evergreen shrubs or trees; iridoids?; phloem stratified; petiole bundles three, arcuate, or annular with wing bundles; styloids + or 0; hairs unicellular; leaves (subopposite), (lamina margins entire), stipules cauline, minute; K and C decussate, free, (C 0); stamens = and opposite sepals-12; pollen por(or)ate; ?nectary; G [2], placentation apical, style 0, stigma large, capitate; ovules 1(2)/carpel, funicular obturator +, endothelium?; fruit a berry; seeds ruminate or not, exo- or exoendotestal, endotestal cells with dark contents; embryo short?; n = ?

1/10. New Guinea, Australia (Queensland) and New Caledonia (map: from van Balgooy 1984; Mark Newman, pers. comm.).

Synonymy: Sphenostemonaceae P. van Royen

• Quintinia is an evergreen tree or shrub with spiral, serrate, estipulate leaves; the plant is covered with peltate scales. The inflorescence is terminal and the flowers have free sepals and petals and an inferior ovary. The fruit is a septicidal capsule.

• Sphenostemon is usually a small, evergreen tree with spiral, strongly serrate leaves that lack stiples, and terminal inflorescences with rather small flowers. The sepals and petals are free, the anther thecae are borne on stout filaments, and there is no style; the fruit is fleshy and two-seeded.

Evolution. Divergence & Distribution. The Late Cretaceous Silvianthemum suecicum, from Sweden, has many features suggesting a relationship with Quintinia (Friis 1990; see also Martínez-Millán 2010; Friis et al. 2011). However, it has tricolp(or)ate pollen, the anthers appear to be dorsifixed, and there are three short, adaxially grooved styles; Quntinia has stamens.

Other. Sphenostemon and Paracryphia are two of the few asterid I + II taxa which sometimes have more than twice as many stamens as petals; the perianth is either uniseriate and/or decussate. The apomorphies of Paracryphia in particular represent a very odd combination for a member of the asterid I + II clade.

Chemistry, Morphology, etc. Quintinia is almost unknown embryologically. Its placentation is perhaps fundamentally parietal (Bensel & Palser 1975b); the ovules may be bitegmic and crassinucellate (Mauritzon 1933, parietal tissue ca 2 cells across?; Philipson 1974), but this needs confirming. Spenostemon, too, is poorly known. Styloids are visible on the abaxial surface of the lamina of Papuasian species; they look rather like cystoliths. For some details of the flower, see Endress (2008c). The fruit is often described as being a drupe, but Lundberg (2001c) characterized it as being a pseudo-drupe (and the seeds as being pachychalazal), while Savinov (2003) described it as being a berry.

For Paracryphia, see Dickison and Baas (1977: anatomy) and Lundberg (2001e: general); for Quintinia, see Lundberg (2000d: general); and for Sphenostemon, see Jéremie (1997) for general information, Savinov (2003) for fruit and seed anatomy.

Phylogeny. Paracryphiaceae form a clade in Lundberg's three-gene Bayesian analysis (Lundberg 2001e); Cameron (2001, see also 2003) also suggested an association between Paracryphia and Sphenostemon. The vessel elements of the three are very long and the perforation plates have many bars (aspects of the wood anatomy of Paracryphia have previously been considered to be among the most primitive in angiosperms...).

In a 17-gene analysis by Soltis et al. (2011), Quintinia linked with Polyosma (Escalloniales here), but support for this position came from the mitochondrial component of the analysis. Soltis et al. (2011) were inclined to think that horizontal gene transfer of the mitochondrial genes might be involved.

Classification. Since the three genera are morphologically rather different they are characterised separately above. They have all been placed in monogeneric families, but only relatively recently; they are combined here (see also APG 2009).

Previous Relationships. Paracryphia was included in Theales by Cronquist (1981) and in Theanae by Takhtajan (1997) as a monotypic family. It was linked quite strongly with Rutaceae + Meliaceae + Simaroubaceae by Källersjö et al. (1998), but cf. Savolainen et al. (2000a). Quintinia has long been included in woody Saxifragaceae/Hydrangeaceae. Baas (1975) thought that Sphenostomonaceae (and Phellinaceae) were members of Celastrales, close to Icacinaceae, while Takhtajan (1997) included them in Icacinales.