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.

VITALES + ROSIDS / ROSIDAE: anthers articulated [± dorsifixed, transition to filament narrow, connective thin]; (elaborate disc nectaries).   Back to Main Tree

Evolution. Divergence & Distribution. Using penalized likelihood, Wang et al. (2009: two estimates) suggested that the Vitaceae/rosid split occurred (115-)111(-109) or (96-)92(-88) million years ago; two Bayesian relaxed clock estimates were between 119 and 113 million years. Ages of (132-)125, 101(-97) million years are suggested by Bell et al. (2010 for details: Dilleniaceae in superasterids...). Wikström et al. (2001) suggested an age of (112-)108(-104)) million years for this split. Finally, Magallón and Castillo (2009) estimated ages of ca 90.6 and 90.8 million years (relaxed and constrained penalized likelihood datings respectively) for the crown group, while the stem group was 112.6 to 113.2 (relaxed and constrained repectively) million years old, while Argout et al. (2011) give a date for the crown clade of ca 123 million years.

Genes & Genomes. For the possible palaeohexaploidy of Vitales, see Jaillon, Aury et al. (2007), indeed, currently it is proposed that this whole genome triplication occured in the common ancestor of the [rosid + asterid] clade... There is further information on the Dilleniales page.

Chemistry, Morphology, etc. For disc nectaries, see Endress (2010c).

Phylogeny. Molecular data still do not link Vitaceae unambiguously with any other single core eudicot order, but they are definitely not nested within any group. They were placed sister to rosids, but with only moderate support (D. Soltis et al. 2000), and even this moderate support weakened in a subsequent four-gene analysis (D. Soltis et al. 2003a); however, Jansen et al. (2006a, b) using complete chloroplast genome sequences found quite strong support for this position (note that members of Berberidopsidales, Dilleniales, Santalales and Saxifragales were not included: see also Ruhlman et al. 2007; Jansen et al. 2007; Moore et al. 2007). Hilu et al. (2003: matK analysis [incomplete sequence] alone) suggest relationships between Vitales and Dilleniales (only moderate support in parsimony analysis, but 100% posterior probability in Bayesian analyses), the combined clade being just above Malpighiales and below Saxifragales in a pectinate tree of major clades within the core eudicots. Although this relationship was not recovered in the analysis of the matK gene by Worberg et al. (2007), the two do have a similar and rather distinctive testa anatomy (see also Kubitzki 2006a). See the Dilleniales page for further discussion on the relationships of Vitales. The placement adopted here, although somewhat tentative, seems quite likely.

VITALES Reveal  Main Tree, Synapomorphies.

Vessel elements with simple perforation plates; tension wood 0; nodes 3-7:3-7; sieve tube plastids with protein crystalloids and starch; raphide bundles +; food bodies [pearl glands] +; lamina with glandular teeth; C protective in bud; stamens = and opposite petals and from a common primordium; nectary gynoecial; style short; ovules 2/carpel, apotropous, micropyle?; fruit a berry, K deciduous; seeds perichalazal, vascular bundle ± surrounding seed, ± ruminate and with Y-shaped dorsal infold, testa multiplicative, exotesta fleshy, mesotesta 2-17 layers across, endotesta 2-5-layered, lignified, crystalliferous, exotegmen (crossed) tracheidal, endotegmen ± mucilaginous; embryo minute.

Evolution. Divergence & Distribution. Using penalized likelihood, Wang et al. (2009: two estimates) suggested that the stem Vitales age was (115-)111(-109) or (96-)92(-88) million years ago; two Bayesian relaxed clock estimates were between 119 and 113 million years. Wikström et al. (2001) suggested an age of (112-)108(-104)) million years for this clade, Magallón and Castillo (2009) estimated ages of ca 112.6 and 113.2 million years (relaxed and constrained penalized likelihood datings respectively).

Includes Vitaceae.

Synonymy: Leeales de Candolle

VITACEAE Jussieu, nom. cons.   Back to Vitales

Ellagic acid, myricetin +; (cork cambium deep-seated); cambium storied; wood with broad rays; nodes often swollen; petiole with ring of bundles; stomata variable; branching from previous flush; leaves palmately compound or -veined, lamina vernation conduplicate; inflorescences terminal, branched, paniculate-corymbose; flowers small, (3-)5(-7)-merous; K connate, C valvate; stigma capitate or 4-fid, dry; nucellar cap +, placental obturator + or 0; prominent raphe and chalazal knot, seed-coat ingrowths delimiting raphe, (endotesta palisade).

14/850 - 2 groups below. Pantropical and (warm) temperate.

1. Leeoideae Burmeister

Herbs to trees; raphides barbed; leaves spiral, to twice compound, teeth with small glandular apex, one lateral vein continues its course above the tooth, stipules borne along petiole margin, sheathing; C basally connate, stamens adnate to corolla, connate, with a lobed tube; nectary disc 0; G [3 (4)], (semi-inferior), odd member abaxial, loculi divided; micropyle exostomal; rarely raphides in the seed coat; n = (10-)12.

1[list]/34. Most Indo-Malesian, few Africa and Madagascar (map: from Ridsdale 1976). [Photo - Flower]

Synonymy: Leeaceae Dumortier, nom. cons.

2. Vitoideae Eaton

Vines or lianes (stout trees, herbs; rootstock swollen) climbing by leaf-opposed tendrils; raphides smooth; (cuticle waxes as tubular rodlets); (more than one bud/node); leaves opposite, spiral or two-ranked, simple or palmate, lamina teeth with gland broadening distally and with foramen, veins from above and below, (stipules adaxially connate); inflorescences leaf-opposed (terminal); (flowers 4-merous); (C connate by papillae and calyptrate); (tapetum plasmodial); disc annular or as five glands, (investing G; 0); G [2], transverse or vertical, (style long); micropyle endo(bi)stomal, outer integument 4-7 cells across, inner integument (1 - Vitis)2-3(-4) cells across, parietal tissue to 17 cells across, nucellar cap to 8 cells across; raphides in the seed coat, (outer integument multiplicative); n = 10-16, 19, 20.

14[list]/825: Cissus (350), Cyphostemma (150: ?= Vitis), Ampelocissus (100), Tetrastigma (95), Vitis (65), Cayratia (65). Pantropical and (warm) temperate (map: from Wickens 1971; Morley and Toelken 1983; Lombardi 2000; FloraBase 2006, still a lot to do). [Photo - Flower]

Synonymy: Ampelidopsaceae Kosteletzky, Cissaceae Drejer, Pterisanthaceae J. Agardh

• Vitaceae are often vines, and leaf-opposed tendrils or inflorescences are common in the family and the nodes are often swollen. Vitaceae may also be recognised by their caducously stipulate leaves with palmately compound, -lobed or -veined blades and often rather coarse teeth; the inflorescences are cymose and the smallish flowers have valvate petals. The stamens are equal in number to and opposite the petals, the usually 4-ovulate, bicarpellate gynoecium has more or less sessile stigmas, and the berry often has ruminate seeds. The stems are often strongly lenticellate, with the bark ultimately flaking, and the leaf blades are very brittle when dry, the fine details of the venation being obscure. Even individual seeds are distinctive, the endosperm being T-shaped in transverse section, the arms often being incurved.

Evolution. Divergence & Distribution. Wikström et al. (2001) suggested that crown group Vitaceae started diverging 92-78 million years before present and Magallón and Castillo (2009) estimated ages of ca 90.7 and 90.8 million years (relaxed and constrained penalized likelihood datings respectively). Nevertheless, fossils of the family are not yet known from the Cretaceous, although the distinctive seeds are found quite commonly in both North America and Europe in deposits of Tertiary age; there may be more than twelve seeds per berry, which suggests a gynoecium unlike that of any extant member of the family (Manchester & Chen 2009).

Ecology & Physiology. Vitaceae are a major clade of vines; they are all tendrillate and have adhesive pade. For how the pads may function and the possible role of small hooks on the shoots as mechanoreceptors - in Parthenocissus at least - see Steinbrecher et al. (2011). The tendrils are clearly stem structures, and some are replaced by inflorescences in fertile shoots (and part tendril-part inflorescences are not uncommon - Calonje et al. 2002 for development). In some species not all leaves have axillary buds, and there has been discussion as to whether the inflorescence/tendril is an evicted terminal shoot, or not (Wilson et al. 2002, and references).

In drier parts of Africa there are a number of non-climbing species of Cissus s.l. with swollen stems, and these species may have crassulacean acid metabolism.

Plant-Animal Interactions. Caterpillars of some lepidoptera are found on Vitaceae and Onagraceae alone (Forbes 1956) - and both contain raphides. The raphides of Vitis are bipartite, square in transverse section, and like an arrow-head in longitudinal section (Horner & Wagner 1995).

Tetrastigma in West Malesia is the only host of the giant parasite Rafflesia (Rafflesiaceae, Malpighiales).

Food bodies, often called pearl glands, are common on the surface of the plant. They are multicellular, with a multiseriate stalk, sometimes with a stoma on the swollen head, and the central parenchymatic cells accumulate oils and sugars (Pavia et al. 2009).

Floral Biology. The Malesian Pterisanthes has remarkable flattened red to purplish inflorescence axes on which the flowers are scattered.

Genes & Genomes. Vitis, at least, has a massive mitochondrial genome largely as the result of the the expansion of intergenic spacers; some DNA from the mitochondrion may have migrated to the nucleus - very unusual (Goremykin et al. 2009a).

Economic Importance. See Terral et al. (2010) for the early history of domestication of the grape.

Chemistry, Morphology, etc. In temperate Vitaceae there is pronounced vessel dimorphism while in tropical members of the family there is often distinctive cambial structure and hence secondary thickening patterns. The raphides of Vitis are bipartite, square in transverse section and like an arrow-head in longitudinal section (Horner & Wagner 1995). Flowers have a common stamen-petal primordium. There is considerable variation in nectary morphology, from enveloping the ovary and forming little projections on top to being absent. Although Leea lacks an obvious nectary like that of Vitis, etc., developmental data show that the lobes on the staminal tube are comparable to a nectary (Gerrath et al. 1990). The ovary has been described as being "anatomically parietal" (Brizicky 1965, for references). The egg apparatus of Cissus is reported to lie outside the ovule (Nair 1970 for references). The raphe in the seed is flanked by a deep groove.

Some general information is taken from Lombardi (2000) and Timmons (2006); for vascular anatomy, see Wheeler and LaPasha (1994), for nodal anatomy and stipules, see Shah (1959), for phyllotaxis, Gerrath et al. (1998), for leaf teeth, Hickey and Wolfe (1975), for floral development, see Gerrath and Posluszny (1989 and references - Viticoideae) and Timmons et al. (2007: some Viticoideae, useful table), and Gerrath et al. (1990 - Leeoideae), for embryology, etc., see Nair (1970), for seed anatomy in extant and fossil taxa, see Chen and Manchester (2007, 2011), and for a recent general summary, see Wen (2006, as Leeaceae and Vitaceae).

Phylogeny. Ingrouille et al. (2002) in a study of rbcL phylogeny considered in the context of morphological variation found little strong support for clades within Vitaceae. Subsequent studies using more genes (e.g. Soejima & Wen 2006) find somewhat more resolution, although support values and relationships of the clades other than the Cyphostemma-Cayratia-Tetrastigma clade are still rather uncertain (Rossetto et al. 2002; Wen et al. 2007; Wen 2008). Ren et al. (2011) found two main clades in Viticoideae, one included species with 4-merous flowers, the other, species with 5-merous flowers, and within which - especially the former - there is a fair bit of well-supported phylogenetic structure that is correlated with names in current use.

For the phylogeny of Tetrastigma, host of Rafflesia, see P. Chen et al. (2011). Vitis seems to be monophyletic, and the species hybridize (Tröndle et al. 2010).

Classification. Although Vitis and relatives and Leea are morphologically distinguishable, there are numerous features that unite the two and they are sister taxa in all phylogenetic studies; inclusion in a single family seems reasonable. The two agree in numerous features (see above), and the wood anatomy of Leea and Rhoicissus in particular is very similar.

Generic limits in Viticoideae need attention, for example, species of Cissus occur all over the tree, while Tetrastigma, although monophyletic, is embedded in Cayratia (e.g. Chen et al. 2011).

Previous Relationships. The affinities of Vitaceae have long been uncertain. They were often associated with Rhamnaceae, since both have stamens opposite the petals, and Takhtajan (1997) placed them near Proteanae, in his Rosidae; Proteanae also have stamens opposite their tepals, but are otherwise very different.