Plant a shrub or tree; true roots +, origin endogeneous, root cap +, apex multicellular; endodermis +; shoot apical meristem multicellular; lateral meristems +, cork cambium producing cork abaxially, vascular cambium producing phloem abaxially and xylem adaxially; lamina with mean venation density 1.8 mm/mm2 (to 5 mm/mm2).
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, so no Maüle reaction; root xylem exarch, cork cambium deep seated; arbuscular mycorrhizae +; shoot apical meristem interface specific plasmodesmatal network; stem with vascular tissue around central pith [eustele], vascular bundles with interfascicular tissue, ectophloic, endodermis 0, xylem endarch; 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 +; stem cork cambium superficial; branches exogenous; leaves with single trace from vascular sympodium ["nodes 1:1"]; vascular bundles collateral [stem: phloem external; leaf: phloem abaxial]; stomata morphology?, pore opening controlled by abscisic acid; leaves with petiole and lamina, spiral, development basipetal, blade simple; axillary buds +, not associated with all leaves; prophylls two, lateral; plant heterosporous, sporangia borne on sporophylls; microsporophylls aggregated in indeterminate cones/strobili; true pollen +, grains mono[ana]sulcate, exine and intine homogeneous; ovules unitegmic, parietal tissue 2+ cells across, 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, flagellae numerous; ovules increasing considerably in size between pollination and fertilization, female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large" [ca 8 mm3], but not much bigger than ovule, with morphological dormancy; embryo cellular ab initio, endoscopic, plane of first cleavage of zygote transverse, suspensor +, short-minute, embryo straight, shoot and root at opposite ends [allorrhizic], white, cotyledons 2; plastid transmission maternal; ycf2 gene in inverted repeat, 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.
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, 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 unilacunar [1:?]; stomata brachyparacytic [ends of subsidiary cells level with ends of pore], outer stomatal ledges producing vestibule; lamina formed from the primordial leaf apex, margins toothed, development of venation acropetal, secondary veins pinnate, overall growth ± diffuse, venation hierarchical, fine venation reticulate, veins (1.7-)4.1(-5.7) mm/mm2, endings free; most/all leaves with axillary buds; flowers perfect, pedicellate, ± haplomorphic, 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, 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 columellar, endexine thin, compact, lamellate only in the apertural regions; nectary 0; G superior, 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]; ovule not increasing in size between pollination and fertilization; pollen binucleate at dispersal, male gametophyte trinucleate, germinating in less than 3 hours, pollination siphonogamous, tube elongated, growing between cells, growth rate 20-20,000 µm/hour, outer wall pectic, inner wall callose, 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, flagellae 0, double fertilization +, ovules aborting unless fertilized; P deciduous in fruit; seed exotestal, becoming much larger than ovule at time of fertilization; endosperm diploid, cellular [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; embryogenesis cellular; 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]].
[NYMPHAEALES [AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]]: vessels +, elements with elongated scalariform perforation plates; 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]]]]: essential oils in specialized cells [lamina and P ± pellucid-punctate]; tension wood 0; 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 +; 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 positiion]; embryo sac bipolar, 8 nucleate, antipodal cells persisting; endosperm triploid; ?germination.
[CHLORANTHALES [[MAGNOLIALES + LAURALES] [CANELLALES + PIPERALES]]]: sesquiterpenes +; seed endotestal.
[[MAGNOLIALES + LAURALES] [CANELLALES + PIPERALES]] / MAGNOLIIDS / MAGNOLIANAE Takhtajan: (neolignans +); vessels solitary and in radial multiples; (sieve tube plastids with polygonal protein crystals); lamina margins entire; A many, spiral [possible position here], extrorse; ovules with hypostase, nucellar cap +, raphal bundle branches at the chalaza; antipodal cells soon die.
[[CANELLALES + PIPERALES]: flavonols, aporphine alkaloids +; nodes 3:3; G whorled.
Evolution. Divergence & Distribution. Magallón and Castillo (2009: note topology) offer estimates of ca 201 and 128 m.y. (relaxed and constrained penalized likelihood datings) for the beginning of divergence in this clade; an age suggested by Magallón et al. (2013) is around 137.9 m.y..
Chemistry, Morphology, etc. Doyle (2007) noted that the venation of members of this group was 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.
Note: Possible apomorphies are now being added throughout the site; they 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 because there is very considerable homoplasy, with variation within and between clades, for most characters. Furthermore, the basic information for all too many characters is very incomplete, often coming from taxa well embedded in the clade of interest and so making the position of any putative apomorphy uncertain. Then there is the not-so-trivial issue of how ancestral states are reconstructed...
Evolution. Divergence & Distribution. Magallón and Castillo (2009: relaxed and constrained penalized likelihood datings) offer estimates of 128 and 125 m.y. for crown group Canellales; Magallón et al. (2013) an age of around 122.8 m.y..
Chemistry, Morphology, etc. Drimane-type sequiterpenoids are practically restricted to Canellales; they are quite 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 +; sieve tube plastids with peripheral protein fibrils; 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]
Evolution. Ecology & Physiology. For vascular flow in Canellaceae, see Hudson et al. (2010) and Feild et al. (2011: Canella winterana is 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), both secondary parietal cell layers dividing, fibrous outer layer of middle wall layer; pollen in tetrads, acalymmate [each grain has separate exine], (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]).
1. Taktajanioideae Leroy
Lignification in stem diffuse; reaction wood 0; flowers red; "K" with four more members; anther thecae transverse; G , 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 in the tropics. [Photo - Flower.]
Synonymy: Drymidaceae Baillon
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 m.y. 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 (the exine forms a continuous covering over the tetrad) 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 end-Cretaceous Maastrichtian of California (see Vink 1993, for literature).
Although estimates of the divergence of Takhtajania from other Winteraceae are only 46-41 m.y. (Wikström et al. 2001), Marquínez et al. (2009) suggested that Takhtajania diverged from the rest of the family ca 120 m.y.a. and that Tasmannia diverged from all other Winteroideae 92.4-77 m.y.a.; Feild et al. (2012: suppl.) estimated divergence within Tasmannia (T. lanceolata splitting from other Australian and New Guinean species) at 59.1-49.2 m.y.a..
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, 2000a); most Winteraceae grow in moist, cool environments. For vascular flow in Winteraceae in conditions where freezing may occur, see Feild et al. (2002); their tracheids are relatively immune to freezing-induced xylem embolism, particularly compared to the vessel-containing wood of Canella. Drimys (= Tasmannia) cordata is a rather odd vessel-less liana (Feild et al. 2012).
Pollination 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: see also Nothofagaceae). Tasmannia is usually dioecious.
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 c.f. Keating (2000a: brachyparacytic).
Deroin (2000) noted that there are no cortical bundles in the flower. 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. 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). 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, variation in vascularization of the corolla), Bailey and Nast (1945: comparative studies), 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 - Drimys s.l. is paraphyletic and the limits of Zygogynum are to be 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 Pseudowintera 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: Australia-Malesia)? A generic distinction between the Old and New World taxa is strongly supported (see above). Furthermore, there are clearly numerous (30+) species of Tasmannia in New Guinea where morphologically and ecologically very distinctive species grow together yet show at most uncommon hybridisation, and there are more species in in Borneo and Australia at least.