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 Back to Main Tree

Myricetin 0; vessel elements with scalariform perforation plates; flowers rather small; style short; endosperm copious, embryo short/very short.

Evolution. Magallón and Castillo (2009) offer estimates of ca 99.3 and 99.7 million years for relaxed and constrained penalized likelihood datings for the crown group asterid II, while Moore et al. (2010: 95% highest posterior density) suggest ages of a mere (81-)75(-71) million years for diversification in this clade. Asterids II are represented in the fossil record by ca 83.5 million years, the Late Santonian-Early Campanian (Martinez-Millán 2010).

As mentioned on the asterid 1/Garryales page, taxa with relatively small flowers that are aggregated into flower-like inflorescences predominate here (see Leins & Erbar 2010: fig 157 for such flowers and inflorescences in members of the three large orders that are found in this clade).

Chemistry, Morphology, etc. Another synapomorphy for the asterid II clade may be a valvate corolla. The feature "small(!) flowers" may be assignable to the asterid node, since the first nodes of both the asterid I and II clades may have this feature, but I have tentatively assigned it to this node. The asterid II clade as a whole certainly has many taxa with small flowers (but cf. Campanulaceae, etc.) that are often aggregated into conspicuous inflorescences, and the fruits have few seeds; the apices of the petals tend to be pointed.

Phylogeny. Within the asterid II clade there is moderate (Olmstead et al. 2000; Soltis et al. 2000) to strong (B. Bremer et al. 2002; Janssens et al. 2009) support for Aquifoliales as being sister to the rest, and then Asterales may be sister to Apiales + Dipsacales (e.g. Olmstead et al. 2000; Lens et al. 2008a). Note, however, that studies on the duplication of the RPB2 gene suggested a relationship between the asterid I clade and Aquifoliaceae in particular (Oxelman et al. 2004b), and Aquifoliaceae also seem to lack the PI duplication of other members of the asterid I + II clade (Viaene et al. 2009), so the other Aquifoliales may need to be sampled see also asterid 1/Garryales.

Recent work is clarifying relationships within the whole clade (as the Campanulid clade, see Winkworth et al. 2008a). The basic relationships of the three major groups that make up this clade are [Asterales [Apiales + Dipsacales]], Aquifoliales being sister to them all, although with little support in analyses of non-coding chloroplast genes. Paracryphiaceae, Quintiniaceae and relatives (only Paracryphia and Quintinia were studied) are probably sister to Dipsacales, although not in analyses that included coding chloroplast genes (Winkworth et al. 2008a). A relationship between Apiales and Dipsacales has quite often been suggested, as by Nandi et al. (1998), Lundberg (2001c), Winkworth et al. (2008), etc. Bruniaceae are sister to [Columelliaceae + Desfontainiaceae], although not in analyses of non-coding chloroplast sequences, and in turn may be associated with Escalloniaceae and Asterales. Janssens et al. (2009: two genes) found weak support for [Dipsacales [Asterales + Apiales]] and Qiu et al. (2010) for [Apiales [Dipsacales + Asterales]] sets of relationships. Analyses by Tank and Donoghue (2010) suggest the relationships followed here, although the position of Escalloniales is still rather poorly supported, 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.

Classification. This group is also called campanulids (B. Bremer et al. 2002).

AQUIFOLIALES Senft  Main Tree, Synapomorphies.

Iridoids?; petiole bundles arcuate; C valvate, free, ?development; A free from P/C; nectary +; ovules 1-2/carpel, apical, apotropous; fruit a drupe, stones one-seeded. - 5 families, 21 genera, 536 species.

Evolution. Divergence & Distribution. Janssens et al. (2009) date stem group Aquifoliales to 113±9.8 million years ago, Magallón and Castillo (2009) offer estimates of ca 99.3 and 99.7 million years for relaxed and constrained penalized likelihood datings, also for the stem group of this clade.

Chemistry, Morphology, etc. With the partial exception of Aquifoliaceae, embryology, testa anatomy, and chemistry of this order are little known, and the polarity of features for and within the order is very unclear.

Phylogeny Aquifoliaceae were included in Asteridae II by Gustafsson et al. (1996) and B. Bremer (1996). rbcL and other data suggested the following relationships: Phyllonoma (Hellwingia + Ilex) (Morgan & Soltis 1993; see also Soltis & Soltis 1997; Olmstead et al. 2000; Kårehed 2002b; Whitworth et al. 2008; Manen et al. 2010). It seemed a little odd that there was little evidence that the two taxa with epiphyllous inflorescences form a monophyletic clade - they also have fimbriate stipules, brochidodromous venation, absence of hairs, etc., in common - but recent comprehensive analyses of the asterid II clade (Tank et al. 2007) recovered a sister group relationship between them (1.0 p.p.), as did Soltis et al. (2011: 99% ML bootstrap support). Irvingbaileya + Gomphandra were placed with strong support in Aquifoliales (D. Soltis et al. 2000), and the group expanded by Kårehed (2001). Cardiopteridaceae are sister to Stemonuraceae, in turn sister to Ilex (one species!), Phyllonoma and Helwingia, with very strong support for this basic structure (Kårehed 2001 - see tree). 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: Vessel elements with simple and scalariform perforation plates; pits usually not bordered; apotracheal parenchyma and variants common; (crystal sand in wood rays); nodes 3:3; stomata cyclocytic to anisocytic; hairs unicellular (adpressed); leaves two-ranked or spiral, lamina margins entire; (plant dioecious); petals inflexed at apex, (ridged adaxially); A basifixed; G [2], unilocular, adaxial carpel alone fertile; ovules 2/carpel, apical, thinly crassinucellate, integument vascularized, funicular obturator +; fruit a 1-seeded drupe; testa vascularized; embryo very short.

Chemistry, Morphology, etc.In general, the gynoecial morphology, embryology, etc., of this clade need study. The two sides of the gynoecium/fruit are sometimes dramatically different in this clade, and this needs to be related to gynoecial development.

For information, see Bailey and Howard (1941, their group II: anatomy), Mauritzon (1936c) and Fagerlind (1945a), both embryology, Heintzelmann and Howard (1948: indumentum and crystals), Padmanabhan (1961: embryology), Sleumer (1971a: general), van Staveren and Baas (1973: epidermis), Baas (1973: epidermis, 1974: stomata), Lobreau-Callen (1977, 1980: pollen), Kaplan et al. (1991: chemistry), Teo and Haron (1999: anatomy). Kårehed (2001, 2002b) discusses the taxa in their current circumscriptions, while Lens et al. (2008a) provide a detailed anatomical survey in a phylogenetic context.

Classification. These two families have quite a lot in common, however, Kårehed (2002c) recognised them as separate.

Previous Relationships. For the other genera that were until recently included in Icacinaceae s.l., see Icacinaceae themselves and relatives (near Garryales) and Pennatiaceae (Apiales).

CARDIOPTERIDACEAE Blume, nom. cons.   Back to Aquifoliales

(Twining vine); plants Al accumulators [?all], iridoids?; (vessel elements with scalariform perforation plates only); petiole bundles annular (+ medullary); (articulated laticifers + - Cardiopteris); stomata also anomocytic and paracytic; (lamina margins toothed), (2ndary veins palmate); (plant dioecious, andromonoecious); (inflorescence branched, ultimate units clearly cymose or not), (bracts 0); K basally connate, quincuncial, C usu. ± connate, (C imbricate); A often adnate to C, also dorsifixed; (nectary 0); (pseudoloculus - Pseudobotrys, Citronella), style usu. slender, stigma truncate or capitate, (styles 2, heteromorphic, one stout, lobed and grooved, the other slender, with capitate stigma – Cardiopteris); (ovule 1); (fruit 2-winged samara, wings horizontally striate, stout style accrescent – Cardiopteris); endosperm ?development, (ruminate), (embryo long, with foliaceous cotyledons); n = 14 [Leptaulus].

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

·        Cardiopteridaceae are a rather heterogeneous group of genera, although one or sometimes two rather slender styles are usually evident. The fruits are more or less flattened and/or ridged drupes.

Chemistry, Morphology, etc. Cardiopteris itself looks rather like Dioscorea, but the latter has three-merous flowers and an inferior ovary. Leptaulus has a violet-colored flavonoid; in L. daphnoides the shoot apex aborts.

Tobe (2011b) clarified the gynoeical morphology of Cardiopteris and noted that there was a nectary on the bottom part of the gynoecium. Kong et al. (2002) described the ovules of Cardiopteris as being ategmic, straight, tenuinucellate and with the egg at the chalazal end. This is rather remarkable and needs confirmation; the ovules may in fact be anatropous.

For additional literature, see above, also Lobreau-Callen (1982 - pollen, Peripterygium), and Vera-Caletti and Wendt (2001).

Phylogeny. Dendrobangia does not (see Apodytes group - Garryales) and Pseudobotrys may not belong here (M. Schori, pers. comm. vi.2010).

Previous Relationships. The relationships of Cardiopteris, a vine with distinctive morphology, were previously obscure. It was included in Celastrales by Cronquist (1981), near there by Takhtajan (1997), and placed as sister to Pentaphylacaceae (Ericales), but with weak support, by Savolainen et al. (2000b).

Synonymy: Leptaulaceae van Tieghem, Peripterygiaceae G. King

STEMONURACEAE Kårehed   Back to Aquifoliales

Iridoids +; petiole bundles arcuate + wing or annular + medullary; sclereids +/0; plant dioecious 0r flowers perfect, pedicels articulated; flowers 4-5(-7) merous; K ± connate, (C connate; keeled); filaments often stout and with club-shaped hairs, connective well developed, tapetum multinucleate, (staminodes +/0 - female flowers); pollen usu. colpate or 2-6 porate; nectary (semi)annular (0); (pistillode + - male flowers); style 0, stigma broad; ovule with integument >10 cells across; drupe compressed dorsi-ventrally (rounded), often sulcate, the two sides sometimes very different and one with a fleshy "appendage" developing over the sulcus, (pseudoloculus – Cantleya), endocarp with ridges and grooves on the side of the appendage, when present; testa thick, outer cells thick-walled, elongate, inner cells not thickened, post-chalazal bundle +; endosperm free-nuclear; n = 22. ILLUSTRATION.

12[list]/95: Gomphandra (55), Stemonurus (15). Tropics, esp. Indo-Malesia to Australia (Queensland) (map: from Sleumer 1971a; Utteridge & Brummitt 2007).

• Stemonuraceae are woody plants whose entire, exstipulate leaves can lack much in the way of venation when dry. The flowers are rather small and are borne in cymose inflorescences; the usually free petals are inflexed at the apex, stout filaments are often conspicuously hairy, the broad stigma is sessile, and the gynoecium appears to have a single carpel. The fruit is a more or less flattened and ridged drupe with one side sometimes very different in appearance, both in colour and texture, from the other, as in Medusanthera.

Chemistry, Morphology, etc. The carpellate flowers of Gomphandra are monosymmetric, having a single reflexed staminode (M. Schori, pers. comm.), Schori et al. (2009) discuss fruit variation in the family; for additional literature, see above. I am grateful to Melanie Schori for comments.

Phylogeny. Note that Lasianthera, with colporate pollen, is sister to the other Stemonuraceae, but only five genera were sampled (Kårehed 2001). Still…

[Aquifoliaceae [Phyllonomaceae + Helwingiaceae]] : nodes 1:1; petiole bundle arcuate; leaves spiral, lamina margins toothed, stipules +, small, cauline; drupe with separate pyrenes.

Evolution. Janssens et al. (2009) dated the crown group in this part of Aquifoliales to 62±11.9 million years.

AQUIFOLIACEAE Berchtold & J. Presl, nom. cons.   Back to Aquifoliales

Evergreen (deciduous) trees or shrubs (climbers); tanniniferous, iridoids 0; resiniferous, laticiferous idioblasts +; nodes also 3:3, etc.; petiole bundles arcuate to annular and complex; branching from previous flush; (leaves opposite; two-ranked), lamiona vernation supervolute (conduplicate), teeth with single vein and opaque, glandular deciduous apex, (margins entire), (stipules 0); plants often dioecious; flowers 4-9-merous, C imbricate, often connate basally; A adnate to base of C (free); pollen surface conspicuously gemmate/clavate; G [(2-)4-6(-many)], opposite petals, placentation axile basally, becoming free-central, stigma broad, wet; ovules 1 (2)/carpel, integument 12-15 cells across, parietal tissue ca 1 cell across, hypostase +, funicular obturator papillate (0); stones several, stigma prominent, K deciduous (semipersistent); exotestal cells cuboid, tangentially elongated, inner walls lignified, rest crushed, endotesta tanniniferous; endosperm hemicellulosic; n = 9, 10; mitochondrial coxII.i3 intron 0.

1[list]/405: Ilex. ± World-wide, esp. America and South East Asia-Malesia, one species in Africa (map: see Meusel et al. 1978; Loiseau et al. 2005). [Photo - Staminate Flower, Carpellate Flower.]

• The family may be recognised by its usually spiral, toothed leaves which may have small cork-warts on the lower surface of the lamina; the stipules are minute and blackish. The inflorescence is cymose and frequently subumbellate. The fruits often dry broadly bottle-shaped, being capped by the distinctive, sessile, broad, black stigma; there are several stones.

Evolution. Divergence & Distribution. Pollen of Ilex is known from Cretaceous Turonian deposits ca 80 million years before present in S.E. Australia, and there are perhaps older records, although without photographs of the distinctive pollen (Martin 1977; Loizeau et al. 2005); Manen et al. (2010) use a date of ca 69 million years for the oldest fossils. In any event, crown group Ilex, or at least its plastome, may be a mere 15 million years old (Miocene), which suggests that there has been much extinction in the clade, and there is also evidence of extensive hybridization within the crown group (Manen et al. 2010). Ilex has become extinct in New Zealand in the Tertiary (Lee et al. 2001).

Chemistry, Morphology, etc. Pallisade glandular tissues with protein rich secretions are found on the leaf teeth and stipules; the latter have been called colleters as a result (Gonzalez & Tarragó 2009). Spiral strands may join the two halves of a tranversely-torn leaf.

For the gynoecial nectary, see Erbar and Leins (2010); the nectary may seem to be coming from the petals. The integument is about 15 cells across (e.g. van Tieghem 1898). The embryo is often minute and barely developed when the fruit is dispersed, only slowly maturing afterwards (Herr 1961 for references).

See Copeland (1964) for general information, Baas (1975) for vegetative anatomy, Lobreau-Callen (1977), and Martin (1977) for pollen; Galle (1997) provides an account of the cultivated members of the family.

Phylogeny. The erstwhile genus Nemopanthus is deeply embedded in Ilex (Powell et al. 2000), the two having the same distinctive pollen, etc. Cuénoud et al. (2000) found several clades correlating with geography/morphology in their study of Ilex s. str., however, support for some was weak; Ilex canariensis was not associated with any of these clades. However, a South American group may be sister to the rest of the genus, including Ilex canariensis itself (Selbach-Schnadelbach et al. 2009). Manen et al. (2010) also found strong geographic structure within the genus, although the position of Ilex canariensis was still unclear; in particular, Hawaiian and New Caledonian species were embedded in an American clade (MCC value of 0.84).

Previous Relationships. Phelline and Sphenostemon have sometimes been included in Aquifoliaceae (e.g. Mabberley 1997), but Phelline is here recognised as a separate family in Asterales (Phellinaceae) while Sphenostemon is in Paracryphiaceae (Paracryphiales).

Synonymy: Ilicaceae Dumortier

[Phyllonomaceae + Helwingiaceae]: hairs 0; lamina with brochidodromous venation, stipules fimbriate; inflorescence epiphyllous, on adaxial side of lamina.

PHYLLONOMACEAE Small   Back to Aquifoliales

Trees or shrubs; plants Al accumulators; cork ?; young stem with separate bundles; petiole bundle annular; plant glabrous, leaves ?two-ranked; inflorescences usu. branched; flowers perfect; K 4-5, open, C 3-5; G [2], inferior, placentation parietal, stigmas recurved; ovules many /carpel, ?crassinucellate, parietal tissue ca 1 cell across, nucellus base broad; fruit a few-seeded berry; testa multilayered, exotestal cells large, thick-walled, mucilaginous, 2-3 layers of flattened cells; endosperm hemicellulosic; n = ?

1/4. Mexico to Peru (map: see Mori & Kallunki 1977). [Photo - Leaves, Flowers]

• Phyllonomaceae are shrubs or small trees that may be recognised by their leaves which have epiphyllous inflorescences borne on the upper surface of the lamina towards the apex; the stipules are minute. The ovary is inferior and has a prominent nectariferous disc on top; the baccate fruit contains only a few seeds.

Chemistry, Morphology, etc. Although the inflorescence of Phyllonoma has been described as being "truly phyllogenous", it appears to be produced from a displaced axillary shoot, as in Helwingia (Weber 2004c, and references; see also Dickinson and Sattler 1974).

See Thouvenin (1890) for general information, Mauritzon (1933) for some embryology, Krach (1976) and Takhtajan (2000) for seed anatomy, Mori and Kallunki (1977) for a revision.

Previous Relationships. Krach (1977) suggested that the seeds of Phyllonoma and those of Grossulariaceae were similar. Phyllonoma was included in Grossulariaceae by Cronquist (1981), and as Phyllonomaceae, in Hydrangeales, by Takhtajan (1997).

Synonymy: Dulongiaceae J. G. Agardh, nom. illeg.

HELWINGIACEAE Decaisne   Back to Aquifoliales

Trees or shrubs; flavones, chlorogenic acid, unidentified iridoids +; pericyclic fibres 0; petiole with arcuate and two small adaxial inverted bundles; cuticle wax crystalloids 0; lamina vernation supervolute-curved; plant dioecious, inflorescence fasciculate; P 3-5, valvate, ?nectary; staminate flowers: stamens = and alternating with P, pollen with diffuse endoapertures; carpellate flowers: staminodes 0, G [2-4], alternating with P, stigma dry; ovule one/carpel, pachychalazal, integument thick, incompletely tenuinucellate; stones few; testa thin; endosperm weakly ruminate, ?development; n = 19.

1/3. Himalayas to Japan (map: from Hara 1972). [Photo - Fruit]

• Helwingiaceae are evergreen shrubs or small trees that may be recognised by their leaves which have epiphyllous fasciculate inflorescences on the upper surface of the blade; the stipules are small and fimbriate. The flowers have only a single perianth whorl, the ovary is superior, and the fruit is a drupe.

Chemistry, Morphology, etc. The fasciculate inflorescence is clearly cymose (Weber 2004c). Helwingiaceae may be interpreted as lacking a calyx, and so anthers and corolla alternate (Takhtajan 1997); however, external morphology shows no trace of a missing calyx, so is the corolla absent (the interpretation prefered here)? The carpel ventral bundles are central.

For chemistry, see Iwashina et al. (1997), for embryo and seed, see Korobova (1980), while some other information is taken from Wangerin (1906), Horne (1914), Dickinson and Sattler (1975) and Hara and Kurosawa (1975: revision).

Previous Relationships. Helwingiales were included in Aralianae by Takhtajan (1997); Helwingia was included in Cornaceae by Cronquist (1981) and Mabberley (1997 - but with hesitation).