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.

Taxa that have flowers with many stamens are scattered throughout the core eudicots. Such flowers may initially have only five or ten primordia, in the former case, the primordia usually arise opposite the petals, rather than alternating with them. Numerous individual stamens then develop from these few initial primordia, and development is often centrifugal (cf. esp. magnoliids and the ANITA grade). At maturity, the stamens themselves may be more or less connate or in fascicles (especially Corner 1946b; Weberling 1989; Leins 2000 and references; Prenner et al. 2008). Note, however, that there may be considerable variation in staminal development between closely related multistaminate taxa (e.g. Hufford 1990; Ge et al. 2007). Polyandry is much less common in the asterid I + II clade (q.v. for discussion) and it often appears to be of a rather different nature there. Tentatively, then, and based entirely on gross morphology, there seem to have been major changes in basic floral organisation at the monocot node, the [rosids et al. + asterids et al.] node ("Pentapetalae"), the [asterid I + asterid II] node, and perhaps the [monocot [Ceratophyllales + eudicot]] node. Note that although Gunnerales are now included in core eudicots, the floral morphology of extant Gunnerales is very different from that of other core eudicots and is more similar to that of the eudicots immediately basal to them.

For the floral development of Berberidopsis corallina and Aextoxicon (Berberidopsidales), possible "links" in the evolution of the flower of core eudicots, see Ronse De Craene (2004, 2007, 2010). The link is at best usable as an analogy, since several elements of this floral morphology are probably parallelisms within core eudicots and others even reversals; there is considerable variation in floral morphology in this small clade. Berberidospidales are part of the pectination immediately basal to asterids, relationships being [Santalales [Berberidopsidales [Caryophyllales + asterids]]] (e.g. Moore et al. 2008, esp. 2010 - see below; Wang et al. 2009). Chase (2005) noted that in Santalales some floral parts, particularly stamens, might have several whorls, and this perhaps suggested that canalisation of floral development was less than in some other core eudicots; whether Santalales really are different in this respect from other core eudicot group remains to be established.

For the possible palaeohexaploidy of Vitales, see Jaillon, Aury et al. (2007). If this is a feature of rosids as a whole, then by the time one gets to genera like Brassica and Arabidopsis, the genome will have duplicated many, many times... It has more recently been suggested that there has been gene duplication, possibly because of hybridization, within the Vitis lineage itself, apparently bringing the Vitis genome more into line with that of other rosids (Velasco et al. 2007). Nevertheless, Freeling et al. (2008: the Carica papaya genome was included in their study) suggest that most rosids, i.e. the node [Vitales + rosids s. str], are indeed palaeohexaploids in which two of the three genomes involved have lost notably more ancestral genes than has the other, the Atg event. Furthermore, collinearity between genomes suggests that there is also evidence for this palaeohexaploidy event in Coffea (Cenci et al. 2010); see also Tang et al. (2008a, b), Diaz-Riquelme et (al. 2009), Barker et al. (2009), Abrouk et al. (2010) and Severin et al. (2011) for this genome triplication, whether or not also involving asterids. In any event, evidence of large genome duplications may be lost, as in Fragaria vasca (Rosaceae) (Shulaev et al. 2010).

There are quite a number of gene duplications known from this general area, perhaps a whole genome duplication is involved (e.g. Litt & Irish 2003; Kramer et al. 2004; Kim et al. 2004; Zahn et al. 2005b; Howarth & Donoghue 2006; especially Kramer & Zimmer 2006; Shan et al. 2007 - see immediately above!). Duplications include: euAP1 + euFUL + AGL79 genes [duplication of AP1/FUL or FUL-like gene], PLE + euAG [duplication of AG-like gene: C class], SEP1 + FBP6 genes [duplication of AGL2/3/4 gene]. Looking more specifically at floral genes, note that not all major core eudicot groups have been sampled for the euAP1 gene, the situation in Santalales, for example, being unknown; there has been another duplication of this gene (and also of the AGL1/2/3 gene) perhaps immediately below this node, but above the Ranunculales node; and the roles these genes may have in many eudicot groups is unknown. However, the eu AP1clade includes key regulators that have been implicated in the specification of perianth identity (Litt & Irish 2003). Duplications of the CYC2 gene clade are widespread here, and they are often associated with the evolution of monosymmetric flowers (Howarth et al. 2011 and references).

Chemistry, Morphology, etc. The occurrence of ellagic acid has a distribution similar to that of polyandry in the eudicots. Sampling of variation in the root apical meristem is poor, and possible reversals (for which, see Groot et al. 2004) have not been placed on the tree. Lee et al. (2004) suggest that the CRABS CLAW gene is expressed in core eudicot nectaries (including extrafloral nectaries), or at least in the rosids and asterids that they sampled; it was not expressed in nectaries of Ranunculaceae, and what happens in Proteaceae (with axial nectaries, like rosids - see Smets 1988) and Sabiaceae is unfortunately unknown. This gene is not expressed in the extrafloral nectaries of Passifloraceae (Krosnick et al. 2008a). Seed coats with a mechanical layer more than a single cell thick occur throughout BLAs, but again, seed coats of the asterid I + II clade are rather different, usually being only one or two cells across.

Phylogeny. This clade is strongly supported, e.g. Chase et al. (1993), D. Soltis et al. (1997, 1999, 2003a), Hoot et al. (1998), and Nandi et al. (1998), but support is rather weaker in Zhu et al. (2007). Within this large clade, although the rosid and asterid clades are well supported, other relationships have long been unclear, there being a six-radiate polytomy (see the sixth and earlier versions of this site) - the other clades involved were Crossosomatales, Berberidopsidales, Caryophyllales, and Santalales; Dilleniales and Saxifragales were also of rather uncertain positions. D. Soltis et al. (2003a) in a four-gene analysis suggested that Berberidopsidales are sister to the rest of the non-rosid core eudicots, but there was only 54% jacknife support for this position. Santalales were associated with asterids, while Saxifragales and Vitales linked with [Dilleniales + Caryophyllales], but with still less support (D. Soltis et al. 2003a). In some studies Dilleniaceae were placed sister to Caryophyllales, but with only very moderate support; D. Soltis et al. (2003a) provides rather stronger (83% jacknife) support for this position (see also Soltis et al. 2007a: 1.0 p.p.). It also seemed possible that Caryophyllales + Dilleniales and Santalales formed a clade (D. Soltis et al. 2000); Carlquist (2006) suggested that non-bordered perforation plates was a possible similarity between Santalales and Caryophyllales. Caryophyllales were linked with with asterids in a large 18S ribosomal DNA analysis (Soltis et al. 1997), albeit with only weak support (see also Hilu et al. 2003).

In studies using whole chloroplast genomes (Jansen et al. 2006a, esp. 2006b; Hansen et al. 2007; Cai et al. 2007; Ruhlman et al. 2006; Jansen et al. 2007; Moore et al. 2007; Logacheva et al. 2008) support for a [Caryophyllales + asterids] clade was stronger, however, Berberidospidales, Dilleniales, Santalales and Saxifragales were not included. [Caryophyllales + Santalales] were sister to asterids in some analyses in a study that focused on the position of Cynomoriaceae and Balanophoraceae (Nickrent et al. 2005), but again the sampling was only moderate. In a study using the mitochondrial gene matR, Caryophyllales repeated as sister to asterids, but with very little support; in other analyses including a reduced sampling and two chloroplast genes Santalales and Dilleniales were also in the area, but again with little support (Zhu et al. 2007). In the combined morphological and molecular study of Nandi et al. (1998) the position of Caryophyllales was uncertain, but this was perhaps partly because the ovules of Rhabdodendraceae, there sister to all other Caryophyllales, were interpreted as being unitegmic; recent work suggests that Rhabdodendraceae are not sister to all other Caryophyllales, but are embedded in the order (see Caryophyllales page). In any event, it was becoming increasingly likely that Caryophyllales, whether or not accompanied by a number of other taxa, were sister to asterids (but cf. Goloboff et al. 2009).

There are, however, major changes afoot. There is some support for placing Crossosomatales as sister to the core malvid group (Huerteales, Sapindales, etc.: see Zhu et al. 2007; Soltis et al. 2011, etc.). As we have seen, earlier studies often suggested that Caryophyllales, and perhaps also Santalales and Berberidopsidales, as well as Dilleniales, were closer to the asterids than to any other clade in the polytomy. In a two-gene study focussing of early-diverging eudicots, all these taxa grouped in a pectinate fashion with the asterids, although support was low (Hilu et al. 2008). In their 12-gene plus plastid inverted repeat study of the rosids, Wang et al. (2009) found Berberidopsidales did not group with rosids, but was sister to a clade made up of the few Caryophyllales and asterids included in the study. Moore et al. (2008) in a preliminary analysis of whole-chloroplast genome data, suggested that most members of the basal polytomy of the core eudicots could be placed as a series of pectinate branches immediately basal to the asterids (see Santalales) - suggested relationships are [Santalales [Berberidopsidales [Caryophyllales + Asterids]]], the position of Caryophyllales having the least support (Moore et al. 2010, see also Moore et al. 2011). Relationships in Bell et al. (2010) are [Berberidopsidales [Caryophyllales, Santalales, asterids]], in Soltis et al. (2011 - but little bootstrap support) [Santalales [[Berberidopsidales + Caryophyllales] + asterids]] and [Santalales [Berberidopsidales [Caryophyllales + Asterids]]] in Arakaki et al. (2011). These are all consistent with many of the relationships more tentatively suggested in the two paragraphs above.

The general position of Dilleniales still remains uncertain, thus Bell et al. (2010) place it sister to Caryophyllales and Soltis et al. (2011) in a broadly similar position as sister to a clade they call superasteridae - it includes Santalales, Berberidopsidales, Caryophyllales and Asterids - with 87% ML bootstrap support; this position was also found by Arakaki et al. 2011), who included a larger sample of caryophyllalean chloroplast genomes. Similarly Qiu et al. (2010) found a weakly supported [Dilleniales + Berberidopsidales] as sister to an [Asterales [Santalales + Caryophyllales]] clade, but support for the broader group was also very weak. Moore et al. (2011) found a weakly supported position sister to a [rosids s.l. + asterids s.l.] clade. All in all, a position in this general part of the tree is becoming more probable, however, Dilleniaceae are left unplaced for now./p>

Note that in some analyses of four mitochondrial genes, Qiu et al. (2010) found that Fabidae were not monophyletic, there being quite strong support for a grouping [COM clade + rosid II/Malvidae] (see also Duarte et al. 2010; Burleigh et al. 2011). Similarly, in an analysis of 154 protein-coding genes Shulaev et al. (2011) found that Populus was sister to [Carica + Arabidopsis], rather than to four taxa from the nitrogen-fixing clade, so again, this clade was not monophyletic, while Burleigh et al. (2011) in a genome-level analysis found that Malpighiales were embedded in the malvid clade, although no representatives of Celastrales or Oxalidales were included in their study (see also Duarte et al. 2010); similar relationships were rejected by all tests in the combined analysis of Zhu et al. (2007), although they were found in the analysis of matR data alone. See also Soltis et al. (2011) for a discussion on the influence of the mitochondrial genome on relationships around here; in their study analysis of mitochondrial genes alone placed a weakly supported COM clade as sister to core Malvidae with quite strong support. Endress and Matthews (2006a) suggest some morphological characters that are consistent with such relationships.

The position of Saxifragales and Vitales with respect to rosids has been uncertain; although [Saxifragales, Vitales, rosids] form a strongly-supported clade, it is unclear whether Vitales are sister to Saxifragales or to rosids or to [Saxifragales + rosids] (Moore et al. 2010, 2011 - support for the latter position increased with reduced taxon sampling). Support for [Vitales + rosids] is only 72% ML bootstrap (Wang et al. 2009), but the topology in Bell et al. (2010) and Soltis et al. (2011) is similar, and this may well turn out to be the preferred relationships.

For further discussion, see Caryophyllales, asterids, Saxifragales, and Vitales - immediately preceding mention of these orders.

Classification. In Versions 8 and earlier of this site this was called the the core eudicot clade, largely because the evolution of the "typical" core eudicot flower can be pegged to this node; the current delimitation of core eudicots refers to a clade that is molecularly well supported but that is perhaps morphologically less interesting.

ROSIDS ET AL. = [DILLENIALES [SAXIFRAGALES [VITALES + ROSIDS]]]: nodes 3:3; stipules + [usually apparently inserted on the stem].

Evolution. Moore et al. (2010: 95% highest posterior density) suggest ages of (112-)108(-103) million years for this split.

DILLENIALES Berchtold & J. Presl  Main Tree, Synapomorphies.

Lamina with secondary veins proceeding straight to the teeth; A many, often centrifugal; G separate, compitum 0; ?micropyle; fruit a follicle; endotesta ± palisade, massively lignified, exotegmen usu. tracheidal. - 1 family, 10 genera, 300 species.

Evolution. Divergence & Distribution. The age of stem Dilleniales has been suggested as being some ca 114.4 or 115.1 million years (relaxed and constrained penalized likelihood datings) and that of crown Dilleniales as only ca 52.2 or 52.7 million years (relaxed and constrained estimates again) by Magallón and Castillo (2009).

Phylogeny. Some evidence, including seed coat anatomy, suggests a relationship between Dilleniales and Vitales, but relationships between Dilleniales and Caryophyllales have also been suggested (e.g. D. Soltis et al. 2003a; Soltis et al. 2007a). Horne (2006) lists a number of features suggesting a relationship between Dilleniaceae and Rhabdodendraceae, in particular, which was then thought to be sister to the rest of Caryophyllales; some of these could be features (?synapomorphies?) of [Dilleniales + Caryophyllales], and the status of the others depended on an improved resolution of relationships. These features include absence of tension wood; successive cambia present; vessel elements with simple perforation plates; wood with SiO₂ bodies; nodes 3 or more:3 or more; leaves spiral; K persistent in fruit.

However, recent work suggests that Rhabdodendraceae are sister to the core Caryophyllales and immediately associated families rather than sister to all Caryophyllales (Drysdale et al. 2007; Brockington et al. 2007). Caryophyllales themselves are sister to asterids (e.g. Hansen et al. 2007; Jansen et al. 2007; Logacheva et al. 2008), while Dilleniales may be associated with the extended rosid clade (see below for details).

From the discussion above, a postion of Vitales as sister to other rosids could be seen as probable (see that page for more details and characters), but, other than that, everything seemed somewhat up in the air until very recently. However, an analysis of all 79 protein-coding plastid genes and four mitochondrial genes suggest that Dilleniales are sister to rosids, although support could be stronger (Moore et al. 2008). Wang et al. (2009: Dilleniales not included) in an analysis of 43,000 bp, largely chloroplast sequences, found substantial resolution within rosids s.l., and the relationships they suggest are followed here (they analysed a twelve gene and inverted repeat data sets separately and combined, preferring ML over MP analyses). Vitales and Saxifragales are successively sister to core rosids ([malvids + fabids]), and can be included in rosids, although the position of the former is only moderately supported (72% bootstrap in a ML analysis).

Includes Dilleniaceae.

Synonymy: Dillenianae Takhtajan - Dilleniidae Reveal & Takhtajan

DILLENIACEAE Salisbury   Back to Dilleniales

Trees and shrubs (lianes, perennial herbs); distinctive flavonols, myricetin, ellagic acid +; hairs ± stellate [esp. Hibbertia] and sclerified; primary stem with continuous cylinder; (successive cambia +); cork cambium deep-seated; (vessel elements with simple perforation plates); true tracheids +; raphides +, also common in wood; epidermis silicified; branching from previous flush; hairs unicellular; leaves spiral, lamina vernation conduplicate, surface often scabrid, margins toothed (entire), 2ndary veins parallel, percurrent, tertiary venation ± scalariform, fine veins areolate, teeth with clear glandular expanded apex, base rather broad, stipules 0, or long petiolar flanges; inflorescence?; pedicels articulated; (flowers [horizontally] monosymmetrical); K (3-)5(-20), (with 1 trace), C (2-)5, often crumpled in bud; androecium often asymmetrical, A (2-)many, from a ring primordium or fasciculate, fascicles opposite K, supplied by trunk bundles, (staminodes +), connective often well-developed, anthers basifixed, (dehiscing by pores), exodermis well developed; (pollen colpate); nectary 0; G (1-3)4-8[-20], (opposite petals, or odd member adaxial), styluli long, stigmas capitate to punctate, wet [1 record]; ovules 1-many/carpel, apotropous, often campylotropous (straight), micropyle zigzag or exostomal, outer integument 2(-3) cells across, inner integument 2-6 cells across, parietal tissue 6-14 cells across, nucellar cap ca 2 cells across, chalazal area massive; (megaspore mother cells several); (fruit a nut; berry), (K enclosing fruit, ± fleshy); funicular aril, often laciniate, exotesta often fleshy, exotegmen with spiral or annular thickenings, endotegmen tanniniferous; zygote with distinctive wall and protrusions into the endosperm ["mantle"]; n = 4, 5, 8-10, 12, 13; germination phanerocotylar.

10[list]/300-410 - four groups below. Tropical and warm temperate (map: from van Steenis & van Balgooy 1966; van Balgooy 1975; Heywood 1978; Horn 2009, which see for more detail). [Photos - Collection.]

1. Delimoideae Burnett

Lianes; stomata paracytic.

1/44 (Tetracera). Pantropical.

[Doliocarpoideae [Hibbertioideae + Dillenioideae]]: ?

2. Doliocarpoideae J. W. Horn

(Compitum +), stigma infundibular; ovules 2/carpel, collateral, one epitropous, the other apotropous.

5/65: Doliocarpus (40). Neotropical.

[Hibbertioideae + Dillenioideae]: ?

3. Hibbertioideae J. W. Horn

Nodes 1:1, 3:3; (hairs stellate); (lamina entire), tertiary venation ± random, areoles at most weakly developed; (flowers monosymmetric); A 1-200< (outer staminodes +; obdiplostemonous, the outer A basally connate).

1/115-225. Madagascar to Fiji, but nearly all endemic to Australia.

4. Dillenioideae Burnett

Cork cambium superficial; leaf base surrounding stem; (flowers monosymmetric); A very numerous [200+]; G many, (compitum +); (exotestal cells large, tanniniferous, becoming flattened - Dillenia).

4/75. Dillenia (60). Madagascar to Fiji, most Indo-Malesian, few Australia.

• Dilleniaceae are recognisable by the often strong and parallel secondary veins that proceed straight into the teeth; scalariform tertiary venation is common, and the lamina is often rough; the leaves tend to elongate when they are still rolled up. The bark is often rich brown; prop roots are common, at least in Malesia. The pedicels are articulated near the apex and persist after the flower falls off; the flowers themselves are often conspicuous and have crumpled petals and numerous stamens that are reflexed in bud and often have anthers dehiscing by pores. The fruits are shiny follicles containing arillate seeds; the calyx is persistent, sometimes accrescent, and the filaments are also persistent.

Evolution. Plant-Animal Interactions. Caterpillars of the tortricid Phricanthini are known only from Dilleniaceae (Powell et al. 1999).

Floral Biology & Seed Dispersal. Buzz pollination is likely to be common in Dilleniaceae (Endress 1997b).

The dispersal unit is generally the seed, which may be arillate and then endozoochorous (Dillenia) or myrmecochorous, as in most of the rest of the family (Lengyel et al. 2009, 2010).

Chemistry, Morphology, etc. Hibbertia is very variably both vegetatively and florally. Some taxa have very much reduced leaves and winged, photosynthetic stems, floral symmetry varies, and stamen number varies from one to well over 150. The monosymmetric flowers of Didesmandra aspera have two bundles of stamens on the functionally upper side of the flower, in each there is a single fertile stamen longer than the rest; the flower is drawn as if the plane of symmetry in horizontal (Stapf 1900); the monosymmetric flowers of Schumacheria have only a single staminal bundle in which all stamens are about the same lengths. There are often sclereids in the pith.

See Kubitzki (1971) and especially Horn (2006, 2009) for general information, Paetow (1931), Sastri (1958) and Swamy and Periasamy (1955) for floral morphology, embryology, etc., and Tucker and Bernhardt (2000) for floral development in Hibbertia.

Phylogeny. Fpr relationships, I follow Horn (2002, 2009). Note that Tetracera has reticulate perforation plates in its smallest vessels and the endotesta seems to be in general poorly differentiated (Horn 2006); the latter in particular, if confirmed, may require some adjustments to clade characterizations.

Classification. See Horn (2009).

Synonymy: Delimaceae Martius, Hibbertiaceae J. Agardh, Soramiaceae Martynov