EMBRYOPSIDA Pirani & Prado (crown group)

Gametophyte dominant, independent, multicellular, thalloid, with single-celled apical meristem, showing gravitropism; flavonoids + [absorbtion of UV radiation]; protoplasm dessication tolerant [plant poikilohydric]; cuticle +; cell walls with (1->4)-ß-D-glucans [xyloglucans], lignin +; rhizoids unicellular; several chloroplasts per cell; 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, centrioles develop de novo, associated with basal bodies of flagellae, multilayered structure +, proximal end of basal bodies lacking symmetry, stellate pattern associated with doublet tubules of transition zone; spermatozoids with a left-handed coil; male gametes with 2 lateral flagellae; oogamy; diploid 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, sporangium +, single, with polar transport of auxin, 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, wall with several trilamellar layers [white-line centred layers, i.e. walls multilamellate]; spores trilete; close association between the trnLUAA and trnFGAA genes on the chloroplast genome.

Note that many of the bolded characters in the characterization above are apomorphies in the streptophyte clade along the lineage leading to the embryophytes rather than being apomorphies of the crown embryophytes.

STOMATOPHYTES

Abscisic acid, ?D-methionine +; sporangium with seta, seta developing from basal meristem [between epibasal and hypobasal cells], sporangial columella + [developing from endothecial 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; polar transport of auxins and class 1 KNOX genes expressed in the sporangium alone; MIKC, MI*K*C* and class 1 and 2 KNOX genes, post-transcriptional editing of chloroplast genes; gain of three group II mitochondrial introns.

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

POLYSPORANGIOPHYTA

Sporophyte well developed, branched, free living, sporangia several; spore walls not multilamellate [?here]; apical meristem +.

EXTANT TRACHEOPHYTA / VASCULAR PLANTS

Photosynthetic red light response; water content of protoplasm relatively stable [plant homoiohydric]; control of leaf hydration passive; (condensed or nonhydrolyzable tannins/proanthocyanidins +); vascular tissue +, sieve cells + [nucleus degenerating], tracheids +, in both protoxylem and metaxylem; endodermis +; root xylem exarch [development centripetal]; stem with an apical cell; branching dichotomous; leaves spirally arranged, blades with mean venation density 1.8 mm/mm2 [to 5 mm/mm2]; sporangia adaxial on the sporophyll, sporangia derived from periclinal divisions of several epidermal cells, wall multilayered [eusporangium]; columella 0; tapetum glandular; stellate pattern split between doublet and triplet regions of transition zone; placenta with single layer of transfer cells in both sporophytic and gametophytic generations, embryonic axis not straight [root lateral with respect to the longitudinal axis; plant homorhizic].

[MONILOPHYTA + LIGNOPHYTA]

Branching ± monopodial; lateral roots +, endogenous, root apex multicellular, root cap +; 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; male gametes multiflagellate, basal bodies staggered, blepharoplasts paired; chloroplast long single copy ca 30kb inversion [from psbM to ycf2].

LIGNOPHYTA

Plant 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].

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 derived from (some) sinapyl and particularly coniferyl alcohols [hence with p-hydroxyphenyl and guaiacyl lignin units, so no Maüle reaction]; root with xylem and phloem originating on alternate radii, vascular tissue not medullated, cork cambium deep seated; arbuscular mycorrhizae +; shoot apical meristem interface specific plasmodesmatal 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 +; stem cork cambium superficial; branches exogenous; 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.; leaves with petiole and lamina, 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 linear, functional megaspore single, chalazal, lacking sporopollenin, megasporangium indehiscent; pollen grains landing on ovule; male gametophyte development first endo- then exosporic, tube developing from distal end of grain, gametes two, developing after pollination, with cell walls; 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, embryonic axis straight [shoot and root at opposite ends; plant allorhizic], white, cotyledons 2; plastid transmission maternal; ycf2 gene in inverted repeat, whole nuclear genome duplication [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 nad1 intron 2 and coxIIi3 intron and trans-spliced introns present.

ANGIOSPERMAE / MAGNOLIOPHYTA

Lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, 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 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, 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], ± 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 columellate, endexine thin, compact, lamellate only in the apertural regions; nectary 0; carpels present, superior, free, several, ascidiate, with postgenital occlusion by secretion, stylulus short, hollow, cavity not lined by distinct epidermal layer, stigma ± decurrent, carinal, 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, functional megaspore lacking cuticle; female gametophyte four-celled [one module, nucleus of egg cell sister to one of the polar nuclei]; supra-stylar extra-gynoecial compitum +; ovule not increasing in size between pollination and fertilization; pollen grains landing on stigma, bicellular at dispersal, mature male gametophyte tricellular, germinating in less than 3 hours, pollination siphonogamous, tube elongated, 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, 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, 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; embryogenesis cellular; dark reversal Pfr -> Pr; Arabidopsis-type telomeres [(TTTAGGG)n]; 2C genome size 1-8.2 pg [1 pg = 109 base pairs], whole nuclear genome duplication [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; 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).

[CERATOPHYLLALES + EUDICOTS]: ethereal oils 0.

EUDICOTS: (Myricetin, delphinidin +), asarone 0 [unknown in some groups, + in some asterids]; root epidermis derived from root cap [?Buxaceae, etc.]; (vessels with simple perforation plates in primary xylem); nodes 3:3; stomata anomocytic; flowers (dimerous), cyclic; K/outer P members with three traces, ("C" +, with a single trace); A few, (polyandry widespread, initial primordia 5, 10, or ring, ± centrifugal), filaments fairly slender, anthers basifixed; microsporogenesis simultaneous, pollen tricolpate, apertures in pairs at six points of the young tetrad [Fischer's rule], cleavage centripetal, wall with endexine; G with complete postgenital fusion, stylulus/style solid [?here]; seed coat?

[PROTEALES [TROCHODENDRALES [BUXALES + CORE EUDICOTS]]]: (axial/receptacular nectary +).

[TROCHODENDRALES [BUXALES + CORE EUDICOTS]]: benzylisoquinoline alkaloids 0; euAP3 + TM6 genes [duplication of paleoAP3 gene: B class], mitochondrial rps2 gene lost.

[BUXALES + CORE EUDICOTS]: ?

CORE EUDICOTS / GUNNERIDAE: (ellagic and gallic acids +); leaf margins serrate; compitum + [one place]; micropyle?; whole nuclear genome duplication [palaeohexaploidy, gamma triplication], PI-dB motif +, small deletion in the 18S ribosomal DNA common.

[ROSIDS ET AL. + ASTERIDS ET AL.] / PENTAPETALAE: root apical meristem closed; (cyanogenesis also via [iso]leucine, valine and phenylalanine pathways); flowers rather stereotyped: 5-merous, parts whorled; P = calyx + corolla, the calyx enclosing the flower in bud, sepals with three or more traces, petals with a single trace; stamens = 2x K/C, in two whorls, internal/adaxial to the corolla whorl, alternating, (numerous, but then usually fasciculate and/or centrifugal); pollen tricolporate; G [5], G [3] also common, when [G 2], carpels superposed, compitum +, placentation axile, style +, stigma not decurrent; endosperm nuclear; fruit dry, dehiscent, loculicidal [when a capsule]; RNase-based gametophytic incompatibility system present; floral nectaries with CRABSCLAW expression.

[SANTALALES [BERBERIDOPSIDALES [CARYOPHYLLALES + ASTERIDS]]] / ASTERIDS ET AL. / SUPERASTERIDS : ?

[BERBERIDOPSIDALES [CARYOPHYLLALES + ASTERIDS]]: ?

[CARYOPHYLLALES + ASTERIDS]: seed exotestal; embryo long.

ASTERIDS / Sympetalae redux? / ASTERIDAE / ASTERANAE Takhtajan: nicotinic acid metabolised to its arabinosides; (iridoids +); tension wood decidedly uncommon; C enclosing A and G in bud, (connate [sometimes evident only early in development, petals then appearing to be free]); anthers dorsifixed?; (nectary gynoecial); G [2], style single, long; ovules unitegmic, integument thick, endothelium +, nucellar epidermis does not persist; exotestal [!: even when a single integument] cells lignified, esp. on anticlinal and/or inner periclinal walls; endosperm cellular.

[ERICALES [ASTERID I + ASTERID II]]: ovules lacking parietal tissue [tenuinucellate].

[ASTERID I + ASTERID II] / CORE ASTERIDS: ellagic acid 0, non-hydrolysable tannins not common; sugar transport in phloem active; inflorescence basically cymose; A = and opposite sepals or P, (numerous, usu. associated with increased numbers of C or G); style short[?]; duplication of the PI gene.

ASTERID II / CAMPANULIDAE Back to Main Tree

Myricetin 0; vessel elements with scalariform perforation plates; style shorter than the ovary; endosperm copious, embryo short/very short.

Age. Wikström et al. (2001) suggested an age of (112-)107, 99(-94) m.y., Bremer et al. (2004) an age of about 121 m.y. and Magallón and Castillo (2009) an age of ca 99.5 m.y. for the crown group, Janssens et al. (2009) an age of 113±9.8 m.y., Moore et al. (2010: 95% HPD) suggested ages of (81-)75(-71) m.y., Bell et al. (2010) ages of (109-)102, 100(-85) m.y., while Beaulieu et al. (2013a: 95% HPD) estimated ages of (115-)104(-95) m.y.; around 92.7 m.y. is an estimate in Naumann et al. (2013).

The oldest fossils of this clade are ca 83.5 m.y., from the Late Santonian-Early Campanian, and have been assigned to Paracryphiales (q.v., although their identity is suspect (see also Cornales). Fossils assigned to Aquifoliaceae are about the same age (Loizeau et al. 2005; Martinez-Millán 2010; Friis & Pedersen 2012, c.f. Beaulieu et al. 2013a).

Evolution. Divergence & Distribution. Diversification at this node probably occurred in the southern hemisphere (Beaulieu et al. 2013a). For the evolution of plant habit in the asterid II clade, see Beaulieu et al. (2013b). Aquifoliales (and the three basal lamiid clades), Bruniales, Escalloniales, and Paracryphiales are all (largely) woody and species poor, as are basal Apiales and woody clades in Asterales.

Pollination Biology & Seed Dispersal. In the campanulids the fruits often have few (often 1-2) seeds - but c.f. families like Campanulaceae and Goodeniaceae. Even when each flower has only one or two seeds, these are generally small, indeed, euasterids as a whole have rather small seeds (Linkies et al. 2010). Beaulieu and Donoghue (2013) examined fruit evolution in campanulids from an ecological point of view and conclude that the plesiomorphic fruit type for the whole group was likely to be dry, dehiscent, and with two or more seeds, although this may need to be revised - it is more likely that the plesiomorphic fruit morphology was quite large, fleshy, indehiscent and single-seeded (see . Achenes, dry, indehiscent, and single seeded fruits (they include Apiaceae, although such fruits there are the individual mericarps that result from the separation/dehiscence of the two-seeded fruits) and were often associated with increased diversification rates. However, they were unclear as to any causal connections. These results may have to be modified somewhat when outgroups are included (e.g., what are the likely states for the euasterids as a whole?) and if fruit types are redefined.

Plant-Animal Interactions. Clades of the dipteran agromyzid leaf miner Phytomyza diversified considerably in the campanulids; they moved from Ranunculaceae hosts (Winkler et al. 2009: >700 species in the genus).

Phylogeny. For discussion about the position of Aquifoliaceae and the circumscription of Aquifoliales and the asterid I and II clades, see the euasterids.

There is now moderate (Olmstead et al. 2000; Soltis et al. 2000) to strong (B. Bremer et al. 2002; Janssens et al. 2009) support for Aquifoliales as sister to the rest of the asterid II clade. The position of Dipsacales within the asterid II clade in early phylogenetic analyses was unclear. Downie and Palmer (1992) associated Adoxaceae with Asterales, while they were sister to Apiales in some studies (Backlund & Bremer 1997). The clade [Asterales [Apiales + Dipsacales]] (e.g. Nandi et al. 1998; Olmstead et al. 2000; Lundberg 2001c; Lens et al. 2008a; see also B. Bremer et al. 2002; Winkworth et al. 2008a; Beaulieu et al. 2013b) is generally supported. On the other hand, Janssens et al. (2009: two genes) found weak support for the clade [Dipsacales [Asterales + Apiales]], Qiu et al. (2010) for [Apiales [Dipsacales + Asterales]], while Soltis et al. (2011) in their 17-gene study found little support for any relationships other than strong support for Aquifoliales as sister to the rest of the clade.

For the positions of the smaller clades along the asterid II spine, which are now mostly settling down, see Escalloniales.

AQUIFOLIALES Senft  Main Tree.

Shrubs or trees; iridoids?; petiole bundles arcuate; inflorescence axillary; nectary +. - 5 families, 21 genera, 536 species.

Age. K. Bremer et al. (2004) suggested an age of about 113 m.y.a. for this node, Bell et al. (2010) ages of (101-)88, 87(-85) m.y. ago.

Note: 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 is the not-so-trivial issue of how ancestral states are reconstructed...

Evolution. Divergence & Distribution. In their study of the evolution of plant habit in the asterid II clade, Beaulieu et al. (2013b) noted that growth form was notably constrained in Aquifoliales; all members of which are woody; see also the node.

Chemistry, Morphology, etc. With the partial exception of Aquifoliaceae, gynoecial morphology, embryology, testa anatomy, and chemistry of this order are little known. For a summary of pollen variation, see Schori and Furness (2011, esp. 2014).

Phylogeny rbcL and other data suggested the relationships [Phyllonoma [Hellwingia + Ilex]] (Morgan & Soltis 1993; see also Soltis & Soltis 1997; Olmstead et al. 2000; Kårehed 2002b; Winkworth et al. 2008; Lens et al. 2008b; Manen et al. 2010; Bell et al. 2010). The absence of evidence that the two taxa with epiphyllous inflorescences formed a monophyletic clade - they also have several other features in common (see below) - seemed a little odd, but a comprehensive analysis of the campanulids (Tank et al. 2007) recovered a sister group relationship between them (1.0 p.p.), as did Soltis et al. (2011: 99% ML bootstrap).

Irvingbaileya and Gomphandra, ex-Icacinaceae, were placed with strong support in Aquifoliales (D. Soltis et al. 2000), and the group was expanded by Kårehed (2001). There is very strong support for the basic structure [[Cardiopteridaceae + Stemonuraceae] [Ilex, Phyllonoma, Helwingia]] (Kårehed 2001; only 1 sp. of Ilex included; Lens et al. 2008b). The grouping [Cardiopteridaceae + Pentaphylacaceae] had weak support in an earlier single gene analysis (Savolainen et al. 2000b); for the latter family, see Ericales.



Includes Aquifoliaceae, Cardiopteridaceae, Helwingiaceae, Phyllonomaceae, Stemonuraceae.

Synonymy: Aquifoliineae Shipunov - Cardiopteridales Takhtajan, Helwingiales Takhtajan, Ilicales Martius

[Cardiopteridaceae + Stemonuraceae]: iridoids +; vessel elements also with simple perforation plates; pits usually not bordered; apotracheal parenchyma and variants common; (styloids +); stomata cyclocytic to anisocytic; hairs unicellular (adpressed); leaves two-ranked or spiral, lamina margins entire; (plant dioecious); A basifixed; (pollen grains ± asymmetric); G unilocular, adaxial carpel alone fertile; ovules epitropous, with parietal tissue, integument vascularized, funicular obturator +; (seed ruminate).

Age. An age for this clade of (90-)83, 65(-58) m.y. is suggested by Wikström et al. (2001) and of (90-)73, 66(-43) m.y. by (Bell et al. 2010).

Evolution. Divergence & Distribution. Kårehed (2001, 2002b) discussed the taxa in their current familial circumscriptions, while Lens et al. (2008a) provided a detailed anatomical survey in a phylogenetic context.

Chemistry, Morphology, etc. The two sides of the gynoecium/fruit are sometimes dramatically different in appearance, as in Medusanthera; this needs to be related to gynoecial development (see also the node).

For additional information, see Sleumer (1942a, b, 1971a), Howard (1942b), and Utteridge et al. (2005), all general, also Kaplan et al. (1991: chemistry), Lens et al. (2008a: measurements = range of means and upper end of variation, Cardiopteris [immature] excluded) and Bailey and Howard (1941a-d), all vascular anatomy, Heintzelmann and Howard (1948: crystals and indumentum), van Staveren and Baas (1973) and Baas (1973, 1974:), all epidermis and stomata, Teo and Haron (1999: anatomy), Lobreau-Callen (1972, 1973, 1977, 1980: pollen), and Mauritzon (1936c), Fagerlind (1945a) and Padmanabhan (1961: Gomphandra), all embryology.

Classification. These two families have quite a lot in common morphologically, however, Kårehed (2001) recognised them as separate.

Previous Relationships. For the other genera that were until recently included in Icacinaceae s.l., see Icacinaceae themselves and relatives, i.e. Metteniusaceae, in Metteniusales, Icacinaceae, in Icacinales, and Pennantiaceae, in Apiales.

CARDIOPTERIDACEAE Blume, nom. cons.   Back to Aquifoliales

Cardiopteridaceae

(Lianes); plants Al accumulators [?all], secoiridoids +; (vessel elements with scalariform perforation plates only - Citronella), 500-1,390(-1,950) µm long, fibres 2,190-2,970(-3,450) µm long; petiole bundles annular (+ medullary); (articulated laticifers + - Cardiopteris); stomata also anomocytic and paracytic; (lamina margins toothed), (secondary veins palmate); (plant dioecious, andromonoecious); (inflorescence branched, ultimate units clearly cymose or not), (bracts 0); K basally connate, quincuncial, C (imbricate - Cardiopteris); A adnate to C or not, also dorsifixed; (pollen porate), (endoapertures enlarged - not Citronella); (nectary 0); (G [3], odd carpel adaxial), carpels superposed, (pseudoloculus - Pseudobotrys, Citronella), style usu. slender, (3), about as long as ovary, (± laterally positioned), stigma truncate or capitate, (branched to the base, branches heteromorphic, two stout, subconnate, lobed and grooved, the other slender, with capitate stigma Cardiopteris); ovule (1), (integument vascularized), (ategmic - Cardiopteris), parietal tissue ca 2 cells across; (fruit 2-winged samara, wings horizontally striate, stout styles accrescent Cardiopteris); ?testa; endosperm ?development, (ruminate), (embryo long, with foliaceous cotyledons - Gonocaryum); n = 14 [Leptaulus].

5[list]/43: Citronella (21). Tropics, inc. the Pacific, to Taiwan (map: from Sleumer 1971a, c; Utteridge & Brummitt 2007; Trop. Afr. Fl. Pl. Ecol. Distr. 5. 2010).