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.

[[CANELLALES + PIPERALES]: flavonols, aporphine alkaloids +; nodes 3:3; G whorled.   Back to Main Tree

Chemistry, Morphology, etc. Doyle (2007) notes that the venation of members of this group is poorly differentiated and of low rank.

CANELLALES Cronquist  Main Tree, Synapomorphies.

Neolignans?, drimane-type sesquiterpenes +; indumentum 0; primary stem with continuous vascular cylinder; sieve tube plastids with starch and protein crystalloids and/or fibres; petiole bundle(s) arcuate; leaf cuticle waxes as tubules, nonacosan-10-ol the main wax; foliar sclereids +, branched; branching from previous innovation; flowers of moderate size; K and C distinct; fruit indehiscent, fleshy. - 2 families, 9-13 genera, 75-105 species.

Evolution. Divergence & Distribution. Magallón and Castillo (2009) offer estimates of ca 201 and 128 and 177 and 125 million years for relaxed and constrained penalized likelihood datings for stem and crown group Canellales respectively - but note topology.

Chemistry, Morphology, etc. Drimane-type sequiterpenoids are practically restricted to Canellales; they seem to be diverse in Canellaceae but are relatively uncommon in Winteraceae (Bastos et al. 1999).

For nodal anatomy, see Sugiyama (1979), for general vegetative anatomy, see Metcalfe (1987), and for a comparison of general embryology and seed coat anatomy, see Tobe and Sampson (2000).

Includes Canellaceae, Winteraceae.

Synonymy: Winterales Reveal - Winterineae Shipunov - Winteranae Doweld - Winteridae Doweld

CANELLACEAE Martius   Back to Canellales

Vessel elements with scalariform or reticulate perforations; true tracheids +; leaves spiral or two-ranked; inflorescence variable; receptacular cortical vascular system +; flowers red; K 3, C (4-)5-12, free (connate); A 6-12(-many), connate, (connective little produced), wall epidermis persistent; pollen (trichotomosulcate), ektexine microreticulate to psilate granular [Cinnamosma intectate]; G [2-6], also occluded by secretion, compitum +, placentation parietal, style short, stigma lobed; ovules 2-many/carpel, campylotropous, micropyle bistomal, zig-zag, outer integument 4-8 cells across, parietal tissue?, nucellar cap 0; fruit a berry, K persistent; seeds ruminate or not; exotesta sclerosed, rest undistinguished, (?pachychalazal - Cinnamosma); endosperm development?; n = 11, 13, 14.

5[list]/13. Tropics; U.S.A. (S. Florida), Antilles, South America, E. Africa, Madagascar (map: from Jackeline Salazar Lorenzo). [Photo - Flowers]

• Canellaceae are evergreen shrubs or trees often with rather pale bark that may be recognised by their rather thick, aromatic, exstipulate leaves with entire margins and flowers with three outer sepals, petals, connate stamens with extorse anthers, and ovary with parietal placentation. The fruit is a berry.

Evolution. Ecology & Physiology. For vascular flow in Cannelaceae, see Hudson et al. (2010) and Feild et al. (2011: Canella winterana a little odd.

Chemistry, Morphology, etc. Are the sepals "really" bracts - perhaps unlikely? - or bracteoles (for the latter, see Hiepko (1964). There has been discussion on the insertion of the petals, whether spiral or whorled, and if whorled, the number of whorls; Wilson (1966) also noted that the sepals have two traces and the petals three, although inner petals may have only a single trace. A vestigial aril has been reported from some genera (Igersheim & Endress 1997).

Some information is taken from Kubitzki (1993b); for chemistry, see Hegnauer 1964, 1989, 1990 - the last under Winteraceae), also Bastos et al. (1999: drimane-type sequiterpenoids).

Phylogeny. For a phylogeny of Canellaceae, see Salazar and Nixon (2008); Cinnamomodendron is polyphyletic.

WINTERACEAE Lindley   Back to Canellales

(Plant Al accumulator); vessels 0; rays 10+-seriate; (petiole bundles complex); leaves spiral, lamina vernation supervolute; inflorescence cymose (fasciculate); "K" 2, connate and forming calyptra, (calyptra splitting early), with 2+ traces, "C" with 1-3 traces, (outer members connate); A spiral, 3-many, (subintrorse), filaments stout, expanding during anthesis, (connective prolonged), fibrous outer layer of middle wall layer; pollen in tetrads, acalymmate, (monads), monoporate (trichotomocolpate), semitectate-coarsely reticulate; carpels postgenital occlusion by complete fusion, stigma bilobed, (not decurrent); ovules (1-)5-many/carpel, nucellar cap 2-3 cells across; endostome persistent, cells thick-walled, outermost enlarged; K deciduous in fruit; seed with palisade exotesta, (tegmen subfibrous).

5[list]/60-90 - two groups below. Montane tropics, not mainland Africa (map: from Vink 1993; Marquínez et al. 2009 [New World]: for fossil localities outside the range of extant members of the family, see Doyle 2000b - mauve = early Cretaceous, green = late Cretaceous to Tertiary [not in the sea]). [Photo - Flower]

1. Taktajanioideae Leroy

Lignification in stem diffuse; flowers red; "K" with four more members; anther thecae transverse; G [2], collateral, placentation parietal, placentae subapical, oblique to horizontal, compitum +; outer integument 4-5 cells across, (inner integument 4 cells across), parietal tissue 5-6 cells across; fruit ?"septifragally" dehiscent; n = 18.

1/1: Takhtajania perrieri. Madagascar.

Synonymy: Takhtajaniaceae J.-F. Leroy

2. Winteroideae Arnott

Sesquiterpene dialdehyde cinnamates +; (tracheid walls vestured); sieve elements with non-dispersive protein bodies; (petiole bundles numwerous, arrangement complex); stomatal apertures often occluded by wax-cutin plugs; flowers ± white; "K" with 0-4 more members, "C" 2-13; (anther thecae transverse); G (1-)5-many; outer integument 3-4 cells across, inner integument 2(-3) cells across, parietal tissue 2-5 cells across, hypostase +; fruit berrylet (follicle); cotyledons convolute [Drimys]; n = 13, (19 43).

3-6/60-90. New Guinea to New Zealand and New Caledonia, few Borneo and the Philippines and South America, usu. not lowlands.

Synonymy: Drymidaceae Baillon

• Winteraceae are often rather small evergreen trees or shrubs that can be recognised by their aromatic and/or peppery taste and their spiral, entire leaves that are often glaucous below. There is a more or less obviously calyptrate calyx more or less covering the flower although the other flower parts are nearly always free. The fruit is often a berrylet.

Evolution. Divergence & Distribution. Fossils assigned to Winteraceae are have a much wider distribution than that of the family today (Doyle 2000b, see map above). Pollen tetrads about 122.5 million years old from Gabon have been assigned to Winteraceae, and other slightly younger, common, and more widespread pollen types are less certainly associated with the family (e.g. Doyle et al. 1990a; Doyle 1999). The earlier Cretaceous records are of calymmate tetrads with fine ornamentation, and grains of this type (Walkeripollis have been placed sister to Winteraceae in a constrained morphological analysis (Doyle & Endress 2010). Fossil wood has been reported from the Maastrichtian of California (see Vink 1993, for literature). However, although estimates of the divergence of Takhtajania from other Winteraceae are a mere 46-41 million years ago (Wikström et al. 2001), Marquínez et al. (2009) suggested that Tasmannia diverged from the remainder of Winteroideae 92.4-77 million years ago....

Ecology & Physiology. The occlusion of the stomatal apertures that is common in Winteroideae seems to prevent these apertures from being wetted (Feild et al. 1998, 2000); Winteraceae in general grow in moist environments. For vascular flow in Winteraceae, see

Floral Biology & Seed Dispersal. The flowers of some Winteraceae show thermogenesis (Seymour 2001), and pollination is by a diversity of small insects, including thrips. In New Caledonia the primitive jawed moths, Sabatinca (Micropterigidae, sister to all other Lepidoptera), eat the pollen of Zygogynum; the tetrads are covered by much oily pollen-kitt which makes them stick to the moths. Sabatinca also uses the flowers as a place of assembly prior to mating (Thien et al. 1985: cf. Nothofagaceae).

Genes & Genomes. The substitution rate of ITS rDNA seems to have been notably slow (Suh et al. 1993).

Chemistry, Morphology, etc. Takhtajan (1997) suggests there are no alkaloids; Cronquist (1981) claims that there are at least some... Sesquiterpene dialdehyde cinnamates are known only from Winteroideae, and prenylated flavanones have recently been found in Pseudowintera (Larsen et al. 2007).

Compression wood has been reported from Drimys (Westing 1965). Keating (2000a) found that the leaf traces may sometimes have paired vascular bundles. The pericyclic sclerenchyma is of various origins (Metcalfe 1987). The petiole vasculature is often quite complex (Keating 2000a and references). For the stomatal morphology of Takhtajania, see Baranova (2004a), but cf. Keating (2000a: brachyparacytic). Deroin (2000) notes that there are no cortical bundles in the flower. Tasmannia is usually dioecious. Petals of Drimys s.l. and some other taxa have either one or three vascular bundles at the base (see also Canellaceae!), as have the sepals, but only a single trace leaves the stele (Endress et al. 2000). Doust (2000) noted the complexity of phyllotactic patterns in th perianth; he also described the sepals as being lateral in position, and there is no mention of bracteoles... The carpels of Tasmannia may be plicate. For the fruit anatomy of Takhtajania, see Deroin (2000).

See also Nast (1944) and Tucker (1959: both floral vasculature - note variation in vascularization of the corolla), Bailey and Nast (1945 and references: comparative studies of family), Bhandari (1963) and Bhandari and Venkataraman (1968), both embryology, Hegnauer (1973, 1990: chemistry), Vink (1985, 1993: general), Svoma (1998b: ovules), papers not otherwise cited in Ann. Missouri Botanical Gard. 87(3). 2000, all focusing on Takhtajania, Doust (2000: floral morphology), Floyd and Friedman (2000: endosperm development), Ehrendorfer and Lambrou (2000: chromosome numbers), and van der Ham and van Heuven (2002) and Sampson (2007), both pollen, for further information.

Phylogeny. For the phylogeny of Winteraceae, with Drimys s.l. being paraphyletic and the limits of Zygogynum being extended, see Suh et al. (1993), Karol et al. (2000), Doust (2003), Doust and Drinnan (2004) and Marquínez et al. (2009). Morphological analyses by Endress et al. (2000) suggested that Takhtajania, which they thought might be polyploid, was more or less associated with Psuedowintera within Winteraceae.

Classification. Is Drimys piperita the only species in Malesia, with Drimys also in S. America, or should there be two genera - Drimys (flowers perfect: South America), and Tasmannia (plant dioecious: 30+ species in New Guinea alone)? A generic distinction between the Old and New World taxa is strongly supported by Suh et al. (1993), Karol et al. (2000), Doust (2003) and Doust and Drinnan (2004), and there are clearly numerous (30+) species of Tasmannia in New Guinea and a few more at least in Borneo and Australia. In the field in Papua New Guinea at least there are morphologically and ecologically very distinctive species that grow together yet show at most uncommon hybridisation.