EXTANT SEED PLANTS

Plant woody, evergreen; nicotinic acid metabolised to trigonelline; primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins rich in guaiacyl units; true roots present, xylem exarch; shoot apical meristem complex; arbuscular mycorrhizae +; stem with ectophloic eustele, endodermis 0, xylem endarch; vascular tissue in t.s. discontinuous by interfascicular regions; vascular cambium + [xylem ("wood") differentiating internally, phloem externally]; wood homoxylous, tracheids +; tracheid/tracheid pits circular, bordered; sieve tube/cell plastids with starch grains; phloem fibers +; stem cork cambium superficial, root cork cambium deep seated; nodes ?; leaf vascular bundles collateral; leaves spiral, simple, axillary buds?, prophylls [including bracteoles] two, lateral; plant heterosporous, sporangia eusporangiate, on sporophylls, sporophylls aggregated in indeterminate cones/strobili; true pollen [microspores] +, mono[ana]sulcate, pollen exine and intine homogeneous, ovules unitegmic, crassinucellate, megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development endo/exosporic, gametes two, with cell walls; 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 duplication, mitochondrial nad1 intron 2 and coxIIi3 intron present.

MAGNOLIOPHYTA

Plant woody, evergreen; lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, cyanogenesis via tyrosine pathway, lignins derived from both coniferyl and sinapyl alcohols, containing syringaldehyde [in positive Maüle reaction, syringyl:guaiacyl ratio less 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; stem with 2-layered tunica-corpus construction; wood fibers and wood parenchyma +; reaction wood ?, with gelatinous fibres; starch grains simple; primary cell wall mostly with pectic polysaccharides; tracheids +; sieve tubes eunucleate, with sieve plate, companion cells from same mother cell that gave rise to the tube, the sieve tube with P-proteins; nodes unilacunar; stomata with ends of guard cells level with aperture, paracytic; leaves with petiole and lamina [the latter formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, vein endings free; flowers perfect, polysymmetric, parts spiral [esp. the A], free, numbers unstable, P not differentiated, outer members not enclosing the rest of the bud, A many, development centripetal, with a single trace, introrse, filaments stout, anther ± embedded in the filament, tetrasporangiate, dithecal, 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, pollen subspherical, binucleate at dispersal, trinucleate eventually, tectum continuous, endexine compact, lamellate only in the apertural regions, pollen tube elongated, with callose plugs, penetrating between cells, growth rate moderate, siphonogamy occuring, nectary 0, G free, several, ascidiate, with postgenital occlusion by secretion, few [?1] ovules/carpel, ovules marginal, anatropous, bitegmic, micropyle endostomal, integuments 2-3 cells thick, megasporocyte single, megaspore lacking sporopollenin and cuticle, chalazal, female gametophyte ?type, stylulus short, stigma ± decurrent, wet [secretory]; P deciduous in fruit; seed exotestal; double fertilisation +, endosperm ?diploid, cellular [first division oblique, micropylar end initially with a single large cell, chalazal end more actively dividing], copious, oily and/or proteinaceous, embryo cellular ab initio; germination hypogeal, seedlings/young plants sympodial; Arabidopsis-type telomeres [(TTTAGGG)_n]; whole genome duplication, 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 PHYA/PHYC gene pairs.

Possible apomorphies are in bold. Note that the actual level to which many of these features, particularly the more cryptic ones, should be assigned is unclear, because some taxa basal to the [magnoliid + monocot + eudicot] group have been surprisingly little studied. Furthermore, details of relationships among gymnosperms will affect the level at which some of these characters are pegged.

NYMPHAEALES [AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]: vessels +, elements with scalariform perforation plates; pollen tectate-columellate, tectum reticulate [perforated]; nucleus of egg cell sister to one of the polar nuclei; ?genome duplication; "DEAER" motif in AP3 and PI genes lost, gaps in these genes.

PROTEALES [TROCHODENDRALES [BUXALES [GUNNERALES + CORE EUDICOTS]]]: ?

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

BUXALES [GUNNERALES + CORE EUDICOTS]: ?

GUNNERALES + CORE EUDICOTS: Ellagic and gallic acids common, cyanogenesis via phenylalanine, isoleucine or valine pathways; micropyle?; PI-dB motif +, small deletion in the 18S ribosomal DNA common.

CORE EUDICOTS: Root apical meristem closed; flowers rather stereotyped: 5-merous, parts whorled, K and C distinct, K with 3 traces, A = 2x K, internal to the C whorl, (numerous, but then often fasciculate and/or centrifugal), pollen tricolporate, (nectary disc +), [G 5], [3] also common, compitum +, placentation axile, stigma not decurrent; endosperm nuclear; fruit dry, dehiscent, loculicidal [when a capsule]; 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].

ASTERIDS - Sympetalae redux?: Nicotinic acid metabolised to its arabinosides; (iridoids +); tension wood decidedly uncommon; C sympetalous, if evident only early in development, petals free, anthers dorsifixed?, (nectary gynoecial), ovules unitegmic, integument thick, endothelium +, nucellar epidermis does not persist, style +, long; seed exotestal; endosperm cellular, embryo long.

ERICALES + ASTERIDS 1 + 2: (C tube in flower), ovules tenuinucellate.

This may be a synapomorphy for asterids as a whole.

Both B. Bremer et al. (2003) and Olmstead (2000) suggest strong support for Cornales being the sister to all other asterids. However, Hilu et al. (2003) reverse the positions of Cornales and Ericales, but the matK gene alone was sequenced. Note that in their study, Caiophora (Loasaceae) appears in Asterales, far separate from the other members of the family in the analysis - perhaps mistaken identity?

ERICALES Dumortier  Main Tree, Synapomorphies.

Nonhydrolysable tannins, triterpenoids incl. saponins +; nodes 1:1; leaf vascular bundles collateral; leaves spiral, teeth with single vein and opaque deciduous cap. - 25 families, 346 genera, 11,515 species.

The age of the stem group asterids may be ca 128 million years before present, mid Early Cretaceous, the Ericales diverging soon afterwards (K. Bremer et al. 2004). Interestingly, a fossil named Archaeamphora and assigned to Sarraceniaceae has been described from rocks about the same age as those in which Archaefructus was found, i.e. ca 124 million years before present (Li 2005), although this attribution needs to be confirmed. Sytsma et al. (2006) suggest diversification began 109-103 million years before present. However, Wikström et al. (2001) suggest a stem group age of 114-106 million years before present, divergence not beginning until 92-85 million years before present (if the clearly misplaced Roridulaceae are ignored: Ericales are also sister to Cornales in the reconstruction used there). Anderson et al. (2005: asterids other than Cornales and Ericales not sampled) suggest figures of ca 109 million years before present for stem group Ericales, 103-99 million years before present for the crown group. Almost all families had diverged by the early Eocene (50 million years before present: Sytsma et al. 2006). Indeed, in the late Cretaceous of E. North America there is a great diversity of fossil flowers that may belong to Ericales, the oldest being some 90 my old (Crepet et al. 2001, 2004; see also Herendeen et al. 1999). Schönenberger and Friis (2001) described Paradinandra from the Late Cretaceous of Sweden, and this has a number of Ericalean features, some suggesting relationships with Pentaphylacaceae in particular (perhaps the relationships could more accurately be described as being with Ericales minus the Balsaminaceae and Polemoniaceae clades). Its placentation was intrusive parietal, the pollen was tricolpate, and there was a nectary disc around the base of the ovary; there were paired stamens opposite the petals and single stamens opposite the petals, as in some Sapotaceae, Ebenaceae, Styracaceae, Pentaphylacaceae and perhaps even Actinidiaceae (see also Friis 1985 - ?Diapensiaceae; Keller et al 1996 - ?Actinidiaceae). Note that tricolpate pollen is uncommon in extant Ericales, being known only from Lecythidiaceae.

Today Ericales are an important component of the diversity of the understory in tropical rainforests, including ca 10% of the species and some 22% of the total stems (Davis et al. 2005a); families like Sapotaceae, Lecythidaceae and Ebenaceae are involved. However, this forest may have developed only early in the Tertiary (Burnham & Johnson 2004) whenever the clades now making it up initially diverged; members of Malpighiales are the other main component of this vegetation. Ericales contain ca 5.9% of eudicot diversity (Magallón et al. 1999), of which one third is made up of Ericaceae alone, not a noteworthy component of such forests. Lens et al. (2007b), however, suggest that the ancestors of Ericales-Cornales grew under more temperate and boreal-arctic conditions and moved into tropical lowland rainforest. Insectivorous members of Ericales are nonmycorrhizal (Brundrett 2004 and references).

Recent studies on the duplication of the RPB2 gene show that the I copy persists here almost alone in the eudicots + Trochodendrales + Gunnerales (and also in the asterid I clade: Oxelman et al. 2004). For leaf teeth that have a "?", their morphology is unknown. Schneider and Carlquist (2003) suggest that pit membrane remnants occur in some of this clade - perhaps mostly in some members of the terminal polychotomy.

Relationships within the order were for some time poorly understood, see R. J. Bayer et al. (1996) and Morton et al. (1997a - both largely molecular data) and Anderberg (1992 - morphological data). However, Polemoniaceae + Fouquieraceae, Myrsinaceae and relatives, Ericaceae and relatives, and Balsaminaceae and relatives formed distinct clades, and Styracaceae + Diapensiaceae were moderately (D. Soltis et al. 2000, 2007) or poorly (Albach et al. 2001b) supported, even if many relationships were unclear. Many taxa lack the mitochondrial coxII.i3 intron, but it is known from the Balsaminaceae and Maesaceae clades and also from Ebenaceae and Styracaceae (Joly et al. 2001). A study by Anderberg et al. (2002: five genes, both plastid and mitochondrial) suggested a beginning of resolution of basal relationships within the order; this forms the backbone of the tree here. B. Bremer et al. (2002) infered a similar set of relationships, although the resolution (and sampling) is less extensive.

Geuten et al. (2004) in a Bayesian analysis of some 13 kb of nucleotide sequences suggest a further clarification of relationships within the terminal polychotomy, they also suggest that Balsaminaceae and Marcgraviaceae may be sister taxa. Within the polychotomy, Theaceae s. str. may be sister to Symplocacaeae, this clade being in turn sister to Styracaceae and Diapensiaceae, in turn related to the Ericaceae-Sarraceniaceae clade - all these relationships had strong support in some analyses - while Pentaphylacaceae and the Primulaceae group were sister taxa (Geuten et al. 2004). However, only 16 terminals were included in their analysis, for instance, Ericaceae-Sarraceniaceae were represented by just two terminals. In a rather more extensive study employing some 59 terminals, nearly 20 kb of sequences, and a variety of analyses, Schönenberger et al. (2005) recovered a group differing only in some details from the Theaceae-Ericaceae-Sarraceniaceae clade just mentioned; they did not recover the [Pentaphylacaceae + Primulaceae] clade, rather, Primulaceae and relatives linked with Sapotaceae and Ebenaceae. Interestingly, Lecythidaceae, linked loosely with Sapotaceae in some earlier analyses (and earlier versions of this page) are largely unplaced. Details of the tree have been adapted to follow the relationships suggested by Schönenberger et al. (2005), however, caution is still in order when interpreting this (and other) phylogenies (the tree in Duangjai et al. 2006b shows rather weak [73% bootstrap] support for Lecythidaceae sister to most other Ericales - relationships in the order are not the focus of that study). Schönenberger et al. (2005) also looked at character evolution, which now perhaps makes a little more sense, although it still shows extensive homoplasy. The relationships just mentioned were largely recovered by Sytsma et al. (2006), and with strong support. Independently of such studies, Barkman et al. (2004, also 2007, there sampling poor) had suggested that Mitrastemonaceae belong to Ericales, a suggestion that is followed here (placing them next to Ericaceae and their immediate relatives in the tree is largely for convenience). The mitochondral genes cox1 and matR showed considerable divergence, but not the atp1 gene (Barkman et al. 2007).

The order is made up largely of Sarracenianae, Ericanae, Primulanae, and some families in Theanae, all adjacent groups in the Dilleniidae of Takhtajan (1997). It is the asterid III group of some early phylogenetic studies.

For a summary of some chemical features of Ericales, see Grayer et al. (1999), for aluminium accumulation, see Jansen et al. (2004), and for wood anatomy, see Lens (2005) and Lens et al. (2007b: optimisation of characters on to a tree with rather different topology than that below). For details of the placentation, see Ng (1991); although true parietal placentation occurs in the order (e.g. Ericaceae - Pyroloideae), most other reports are incorrect.

Includes Actinidiaceae, Balsaminaceae, Cyrillaceae, Clethraceae, Diapensiaceae, Ebenaceae, Ericaceae, Fouquieriaceae, Lecythidaceae, Maesaceae, Marcgraviaceae, Mitrastemonaceae, Myrsinaceae, Pentaphylacaceae, Polemoniaceae, Primulaceae, Roridulaceae, Sapotaceae, Sarraceniaceae, Sladeniaceae, Styracaceae, Symplocaceae, Tetrameristaceae, Theaceae, Theophrastaceae.

Synonymy: Primulineae Burnett - Actinidiales Reveal, Balsaminales Lindley, Cyrillales Doweld, Diapensiales Engler & Gilg, Diospyrales Prantl, Ebenales Engler, Empetrales Rafinesque, Fouquieriales Reveal, Lecythidales Cronquist, Marcgraviales Doweld, Mitrastemonales Nakai, Myrsinales Spenner, Polemoniales Bromhead, Primulales Dumortier, Rhodorales Horaninow, Roridulales Nakai, Samolales Dumortier, Sapotales J. D. Hooker, Sarraceniales Bromhead, Styracales Bischoff, Ternstroemiales Doweld, Theales Lindley, Vacciniales Dumortier - Balsminanae Doweld, Diapensianae Doweld, Ericanae Takhtajan, Lecythidanae Reveal, Primulanae Reveal, Sarracenianae Reveal, Theanae Reveal - Ericidae C. Y. Wu, Theidae Doweld - Diospyropsida Brongniart, Ericopsida Bartling, Mysrinopsida Bartling, Primulopsida Brongniart, Styracopsida Bartling

Balsaminaceae + Marcgraviaceae + Tetrameristaceae: myricetin +, ellagic acid 0; vessel elements with simple perforations; raphide sacs +, druses 0; lamina toothed; inflorescence racemose; A = and opposite K, free from C, anthers (near) basifixed, disc 0, ovules bitegmic, micropyle endostomal, style short, stigma little expanded; endosperm with micropylar haustorium.

The raphide sacs are white pockets in the stem; they are visible under the dissecting microscope. Geuten et al. (2006) suggest that heterotopic SEP3-like gene expression - in bracteoles and calyx in extant members - was present in the common ancestor of the group.

Monophyly of these three families is well supported, and the clade is probably sister to rest of Ericales (e.g. Källersjo et al. 1998; Nandi et al. 1998; Soltis et al. 2000; Savolainen et al. 2000a; Geuten et al. 2004). If Balsaminaceae and Marcgraviaceae are sister taxa, there are no obvious synapomorphies for the family pair (Geuten et al. 2004).

For the wood anatomy of this group, see Lens et al. (2005b: that of Balsaminaceae is paedomorphic), for palynology, see Lens et al. (2005: Marcgraviaceae) and Janssens et al. (2005: the rest).

BALSAMINACEAE Berchtold & J. Presl, nom. cons.   Back to Ericales

Fleshy herbs; leucoanthocyanins, napthoquinones +; cork?; petiole bundle arcuate; mucilage sacs +; plant usu. glabrous; leaves (opposite), lamina involute, (extrafloral nectaries +, sometimes as paired glands or foliaceous lateral flaps on leaf base/stem); inflorescence axillary, (bracteoles 0); flowers vertically monosymmetric, inverting during growth; K 3 (5), functionally abaxial sepal with prominent spur (spur 0), nectary inside, C 5, when K 3 adaxial C often with a sepaloid keel, lateral petals connate in pairs (free), A forming cap over stigma, filaments stout, partly connate apically, anthers (with trabeculae in loculi), connate and forming cap over stigma, pollen intermixed with cellulose threads coming from cell walls and holding it to anther, starchy, 3- or 4-colpate or porate, endexine lamellated, G [(4-)5], opposite C, several (unitegmic) apotropous ovules/carpel, micropyle endostomal, stigma fairly broad, wet; fruit an explosive capsule, septifragal, walls inrolling from base, or a drupe/non-explosive schizocarp [Hydrocera]; seed pachychalazal, exotesta only thickened, ("hairs" with spiral thickenings; mucilaginous; sclerotic, testa 6-8 layers thickened cells and 5 layers unthickened - Hydrocera); endosperm scanty, also chalazal haustorium, cotyledons large, n = (3-)7-8(9)10+.

2[list]/1001: Impatiens (1000). Mostly Old World, Africa (esp. Madagascar) to mountains of S.E. Asia (Map: from Hultén 1971; Meusel et al. 1978; Grey-Wilson 1980a; Hultén & Fries 1986). [Photo - Flower.]

• Balsaminaceae are fleshy herbs with more or less translucent stems and usually spiral, toothed leaves. The flowers are strongly monosymmetric and nearly always have a spur on the adaxial sepal, but since the flower is held upside-down the spur is on the lower (abaxial) part of the flower. The anthers cap the gynoecium (until they are dislodged) and the style is at most short. The fruit is usually an explosively-dehiscent capsule.

The combination of leucoanthocyanins and raphides, both of which are found in Balsaminaceae, is rarely found in herbs (Fischer 2004). Indeed, Balsaminaceae are vegetatively rather uniform, if florally very diverse; duplication and probable subfunctionalisation of the class B DEF gener has occured in this clade (Janssens et al. 2006b; Geuten et al. 2006).

The flowers are protandrous. The abaxial-lateral sepal pair (non-inverted orientation) is often reduced, perhaps becoming fused with the abaxial petal (Caris et al. 2006; see also Grey-Wilson 1980c). The cellulose threads produced as the anther walls break down and then retract hold the exposed pollen in a basketwork of these threads over the anther (Vogel & Cocucci 1988). The integuments are quite thick and are free only at the micropyle (van Tieghem 1898); Narayana (1970) also illustrates more conventional ovules. A recent study by McAbee et al. (2007) shows considerable platicisty in integument development in the family, although many species show more or less well developed congenital fusion of the integuments (and bitegmy may be derived). There is variation in the embryo sac, Hydrocera and at least some species of Impatiens having an Allium-type embryo sac (bisporic, 8-celled); Hydrocera also has a much elongated embryo sac (Venkateswarlu & Lakshminarayana 1957). The fruit type of Hydrocera is unclear (see Wood 1975), and it may be septicidal or a several-seeded drupe, although the latter morphology is described by Grey-Wilson (1980a); see Leins (2000) for the fruit dehiscence of Impatiens. The micropylar haustorium is massive and may invade the funicle and even the placenta.

Hydrocera and Impatiens are clearly sister taxa (Yuan et al. 2004; esp. Janssens et al. 2006a). Taxa of Impatiens with five sepals are scattered through the genus, so that condition is apparently at least sometimes derived. The current infrageneric classification of Impatiens needs complete overhaul (Janssens et al. 2006a).

Balsaminaceae were included in Geranianae - Rosidae by Takhtajan (1997).

For information on floral anatomy, etc., see Grey-Wilson (1980b), for a revision and more of the African taxa, Grey-Wilson (1980c), for cellulose threads in the anthers, see Vogel and Cocucci (1988), ovule variation and seed development, Boesewinkel and Bouman (1991), for seed morphology, Utami and Shmizu (2005: variation considerable), for floral development, Caris et al. (2006a), and for general information, see Fischer (2004a).

Synonymy: Impatientaceae Barnhart

Marcgraviaceae + Tetrameristaceae: vessels in radial multiples; ± branched sclereids +; lamina supervolute, elongating in bud, with obscure abaxial lines; fruit indehiscent.

Beauvisage (1920) noted that both Pelliciera and Marcgraviaceae have large air spaces in the cortex.

MARCGRAVIACEAE Candolle, nom. cons.   Back to Ericales

Lianes, shrubs (small trees); (vessel elements with scalariform perforations); rays broad [Marcgravia]; stomata staurocytic; heterophylly common; lamina entire, with marginal to abaxial cavities (black dots); inflorescence a raceme, spike or umbel, flowers 4- or 5-merous; bracts abaxially ascidiate, nectariferous; K quincuncial, C ± connate, A 4-40, G [2-8], opposite ?, placentation very intrusive parietal, many ovules/carpel, stigma ?, dry; fruit dehisces more or less irregularly, placentae fleshy; seeds many, small, exotestal cells ± enlarged, inner walls much thickened; endosperm slight, cotyledons large to small; n = 18 [one count!].

7[list]/130: Marcgravia (60). New World tropics (Map: from Heywood 1978). [Photo - Flowers] [Photo - Fruits]

• Marcgraviaceae are often lianas and can be recognised by their often rather fleshy leaves with indistinct venation and minute abaxial cavities; their inflorescences have abaxially ascidiate bracts that function as nectaries. The fruits, with numerous small seeds borne on a fleshy, colored placenta, are distinctive. Heterophylly is common in Marcgravia, the climbing form of the plant often having orbicular leaves adpressed to the trunk.

The prominent inflorescences with nectar secreted in the cup-shaped (ascidiate) bracts attract a variety of large pollinators including humming birds and bats (e.g. Tschapka et al. 2006).

The black dots on the margin of the leaf blade make the leaf appear "serrate" - that character is not so much about serrations per se, as the marginal glands, etc., that terminate any serrations that are present. I do not know if the pollen grains are starchy (cf. Balsiminaceae). Although individual flowers of Marcgraviaceae are polysymmetric, the inflorescences of taxa like Margravia are monosymmetric from the point of view of their avian pollinators, which get pollen dusted on their heads as they take nectar from the modified bracts which are held beneath the ring formed by the flower (see also Westerkamp & Claßen-Bockhoff 2007). Johri et al. (1992) described the seeds as being arillate.

Ward and Price (2002) suggest phylogenetic relationships within the family; Marcgravia, with its reversible heterophylly, 2-ranked leaves, 4-merous flowers, calyptrate corolla, and nectaries adnate to abortive flowers, is distinct. In the rest of the family both synapomorphies and generic limits are unclear.

For general information, see Dressler (2004), for information on pollination (animals), see Dressler (1999).

TETRAMERISTACEAE Hutchinson   Back to Ericales

Evergreen trees; chemistry?; intervessel pitting opposite-alternate; petioles short; bracteoles rather large, ± caducous; K with adaxial glands, 1 ?orientation ovule/carpel.

3/5. W. Malesia, Central and N. South America. Two groups below.

1. Pelliciera Bentham   Back to Ericales

Vessels in multiples; nodes?; petiole bundle annular; stomata ?; leaf base asymmetrical, colleters?; flowers axillary, large, bracteoles petaloid; A extrorse, anthers very long, connective prolonged into a point, pollen strongly verrucate, G [2], ovule subapical, pendulous, campylotropous, style long, stigma bifid, punctate; fruit ± dry, indehiscent, K and C caducous; seed coat?; endosperm 0, cotyledons large; n = ?; germination phanerocotylar, hypogeal.

1[list]/1: Pelliciera rhizophorae. Central and N. South America (Map: from A. Graham 1977); mangroves. [Photo - Flower] [Photo - Fruit]

• Pelliciera is a tree in mangrove vegetation that may be recognised by its long, pointed terminal buds, more or less pseudoverticillate and toothed leaves with very short petioles and asymmetrical bases, and their large, axillary flowers with conspicuous petaloid bracteoles. The adaxial glands on the sepals and the large, sharply-pointed, single-seeded fruits are also distinctive. Fluted buttresses formed by vertical series of adventitious roots form a conical mantle at the base of the trunk.

From fossil pollen, Pelliciera was once much more widespread (A. Graham 1977).

There are no reports that Pelliciera accumulates aluminium, again unlike Theaceae s.l., in which it has often been included. Pelliciera was compared with Marcgraviaceae by Beauvisage (1920); details of wood anatomy suggest relationships with Tetrameristaceae (Baretta-Kuipers 1976). Nodal anatomy is extrapolated from petiole scars. The products of different marginal glands of the one leaf may not be the same (Collins et al. 1977). The floral diagram in Tomlinson (1986) suggests that either the two carpels are oblique, or the bracteoles are not in the lateral position and the carpels are transverse.

General information is taken in part from Kobuski (1951), Tomlinson (1986), and Maas and Westra (1993).

2. Pentamerista + Tetramerista   Back to Ericales

Cork inner cortical; (vessel elements with scalariform perforations); wood fluorescing [1 sp. tested]; stone cells [in stem] +, branched sclereids ?0; leaves with marginal glands; flowers 4- or 5-merous (bracteoles persistent); K and C similar, filaments slightly connate at the base, G [4, 5], ovule basal, ascending, ?epitropous, stigma punctate to minutely lobed; fruit a berry; testa several layers thick, walls thickened; endosperm copious, cotyledons small; n = ?

2[list]/4. Malesia (Tetramerista), Venezuelan Guyana (Pentamerista: leaves with abaxial domatia).

• The two genera are rather small woody plants that can be recognised by their spiral, short-petiolate leaves with indistinct venation and marginal glands. The flowers are rather small, the sepals and petals being rather similar and spreading; at least the sepals have glands on their adaxial surfaces.

In Tetramerista there are glistening dots on the adaxial surface of both calyx and corolla. Tetrameristaceae s. str. were placed in Theales by Takhtajan (1997).

The inclusion of Pellicieraceae in Tetrameristaceae is an option in A.P.G. II. Pellicieraceae and Tetrameristaceae formed a well-supported clade in the morphological analysis of Luna and Ochoterena (2004), but Macgraviaceae did not join them, nor were other Ericales part of the clade. The embryology, morphology and anatomy of Pellicieraceae s.l. are poorly known.

For general information, see Kubitzki (2004b).

Other Ericales: style long.

Polemoniaceae + Fouquieriaceae: cork outer cortical; vessel elements with simple perforations; C connate, nectary +, G [3], style hollow, stigma strongly lobed; K persistent in fruit; seeds winged; mitochondrial coxII.i3 intron 0.

Schönenberger (2006a) lists many other features occurring in this family pair, including free sepals, stomata on the calyx (also e.g. Ericaceae - what is the general distribution of this feature?), stamens adnate to the corolla, and details of gynoecial development.

POLEMONIACEAE Jussieu, nom. cons.   Back to Ericales

Sugars accumulated as kestose and isokestose oligosaccharides, cucurbitacins +, ellagic acid 0; cork also pericyclic; (vessel elements with scalariform perforations; stomata paracytic); leaves opposite to spiral (palmate, pinnate), conduplicate, margins entire to deeply lobed; inflorescence cymose, paniculate or thyrsoid; (flowers monosymmetric), K connate, lobes often with green midrib and colorless intermediate portion, C tube formation late, lobes usu. contortuplicate, A = and opposite K, inserted at different levels or filaments of different lengths, pollen pantoporate/zonocolporate, G [(2-4)], median member adaxial, 1-many ovules/carpel, stigma dry; seeds often mucilaginous when wetted, exotesta variously thickened, endotesta a pigment layer, radial walls ± thickened; endosperm nuclear, slight, embryo green or white.

Ca 18[list]/385 - two subfamilies below. N. temperate, W. North America, South America (Map: from Hultén 1971; Meusel et al. 1978).

1. Polemonioideae Arnott

Mostly ± desert-dwelling herbs (subshrubs); C veins usu. free or connected well above the base, filaments usu. merged with corolla; seeds not winged (narrowly winged - Loeselia); n = (6) 7 (8) 9, larger [not Loeselia].

13-22/350: Phlox (70), Linanthus (35), Navarretia (30), Polemonium (27), Gilia (25). W. North America, N. temperate, southern South America. There are several cases where predominantly western North American genera have a few species in the southern part of South America. [Photos - Collection (all except Cobaea).]

2. Cobaeoideae Arnott

Mostly mesic vines to small trees (herbs); flowers large; (K basally connate only), usu. herbaceous throughout, C veins connected at the base of lobe (and in upper lobe), A traces in two whorls, filaments often superficially adnate to C, (pollen 100-220 µm long), ovules with nucellar cap; fruit septicidal and/or loculicidal; seeds wing broad (narrow - Bonplandia), mesotestal cell walls thinly lignified; n = (9 - Acanthogilia), 15, 26, 27, small.

5/35: Acanthogilia (not mesic!), Cobaea, Cantua, Bonplandia, Huthia. Baja California, tropical America. [Photo - Flower & Fruits]

Synonymy: Cobaeaceae D. Don

• Polemoniaceae are herbs, or sometimes shrubs or lianas, that may be recognised by their often opposite leaves; the plant may smell unpleasant. The calyx is more or less connate, the units of which it is made up often having a green central portion, the rest being whitish. The corolla is strongly sympetalous and the five stamens are frequently inserted at different levels in the tube (not in Loeselia, Collomia, Microsteris, etc.) and/or are of different lengths. The style is usually three-lobed, the fruit is a capsule and the seeds are mucilaginous when wetted.

The cambium is sometimes storied; raylessness is frequent. The pollen tube has callose plugs. In Cobaea the leaves are tendrillar and the basal pair of leaflets is foliaceous-stipuliform.

Acanthogilia has been placed in its own subfamily (Porter et al. 2000 - Acanthogilioideae J. M. Porter & L. A. Johnson), and it may be sister to the Cobaea et al. clade (Prather et al. 2000) or even sister to the whole of the rest of the family (Schönenberger et al. 2005, only four taxa included). It has very dimorphic leaves and short shoots (cf. Fouquieraceae!), its branched spines are reduced leaves like those found scattered in Polemonioideae, and its sepals have a green midrib, as in Cobaeoideae. Seed wings?

Polemoniaceae were included as Polemoniales in Solananae by Takhtajan (1997).

For floral variation, see Grant and Grant (1965), for inflorscence morphology, see Weberling (1989), for general information, see Day and Moran (1986), Grant (1998), Johnson et al. (1996, 1999), Porter (1997), Porter and Johnson (1998) and Wilken (2004).

FOUQUIERIACEAE Candolle, nom. cons.   Back to Ericales

Woody and xeromorphic, with long and short shoots; flavonols only, ellagic acid, route I secoiridoids +, myricetin 0; cuticle wax crystalloids 0; leaves heteromorphic, margins entire, petiolar spines on long shoots; inflorescence variable; K ± scarious, A 10(-23), in a single whorl, free, nectary tissue in base of ovary, 6-20 ?apotropous bitegmic ovules/carpel, micropyle endostomal, stigmas punctate; fruit a capsule; seed winged, coat becoming crushed, testa and tegmen multiplicative, testa hypodermis with banded thickening; endosperm scanty, chalazal haustorium +; n = 12.

1[list]/11. S.W. North America (Map: Meusel et al. 1978). [Photo - Habit] [Photo - Branch] [Photo - Flowers]

• Fouquieriaceae are often little-branched shrubs with entire, spiral leaves, well developed short shorts, and petiolar spines on the long shoots. The flowers have free almost scarious sepals but a strongly connate corolla; the ten or more stamens are free from the corolla. The style is three-lobed.

Layers of fibrous cells alternate with layers of cork cells in the stem cork, while the cork cambium in the root is described as being superficial (Henrickson 1969), the unusual position for angiosperms, although perhaps commoner in desert plants. The perianth parts are borne in a distinct spiral, and the antepetalous stamens may be doubled (Schönenberger & Grenhagen 2005).

Fouquieriaceae were placed in Violales by Cronquist (1981). For relationships within the family, see Schultheis and Baldwin (1999).

See Kubitzki (2004b) for general information.

LECYTHIDACEAE A. Richard, nom. cons.   Back to Ericales

Trees (lianes); flavonols, ellagic acid +, kaempferol 0; vessel elements with simple (scalariform) perforations; cortical vascular bundles +; (wood siliceous and/or with SiO₂ grains); phloem stratified (with wedge-shaped rays); nodes 3 or more:3 or more; (cristarque cells +); petiole anatomy of numerous arcuate or annular bundles in arcs, etc.; stomata usu. anisocytic; leaf margins toothed or entire (tertiary veins subparallel, ± at right angles to midrib), cauline stipules small or 0, colleters +; inflorescence various, often racemose, pedicels articulated; flowers large; K (2-)4-6(-12), variously arranged, connate or not, valvate, C (0)3-10(-16), (free), A in concentric series or not, many [10-1200], centrifugal, connate or not, latrorse, filaments not articulated, tapetum amoeboid, pollen tricolpate, disc +, G [2-8], inferior, opposite K (C), 1-many (apical or basal) bitegmic ovules/carpel, integuments thick, micropyle endostomal, archesporium multicellular, style short/0, stigma ± capitate (punctate, divided), wet or dry; K persistent in fruit; seeds (hairy), often arillate, vascularised, testa multiplicative, exotestal cells variously thickened, palisade, or low with sinuous anticlinal walls, mesotesta sclerotic or not; endosperm nuclear, 0; mitochondrial coxII.i3 intron 0 [but sampling].

Ca 25[list]/310 - five groups below. Tropical, especially America and W. Africa.

1. Napoleonaeoideae Bentham

Secondary xylem with crystal chains; leaves distichous, supervolute, often with glands abaxially at the base of the lamina, margin serrate, stipules +/0; C 0, staminodial corona +, anthers long, (10, extrorse, monothecal - Napoleonaea), G opposite C, endothelium 0, style 0, stigma broad, pentagonal, flat; fruit a 1-several seeded drupe; embryo curved; n = 16.

2/11. W. tropical Africa (Map: from Liben 1971b). [Photo - Flower]

Crateranthus rbcL sequences have not been obtained, but it is placed here in joint analyses. In general morphology, including androecium, Asteranthos is similar (but cf. style, endosperm), yet its sequence data align it with Scytopetaloideae...

For general information, see Liben (1971a).

Synonymy: Napoleonaeaceae A. Richard

Scytopetaloideae + Lecythidoideae [Planchonioideae + Foetidioideae]: ?

2. Scytopetaloideae O. Appel

Plants Al-accumulators; ?crystal chains; sclereids +; stipules +, minute; pedicels articulated; K connate, C 0, staminodes 6-28, connate, corolla-like, A basally connate, adnate to staminodes, tapetum cellular, (pollen tricolporoidate), G superior (half superior), (disc 0), 2 or 4 ovules/carpel, endothelium +, style slender, relatively long, stigma punctate; fruit a capsule or indehiscent; seeds often 1, hairy, ruminate (not - Oubanguia), endosperm +, hemicellulosic, (embryo J-shaped); n = 11, 18, 21.

6/21. Africa, South America (N.E. Brasil) (map: see Prance & Mori 1979; Heywood 1978). [Photo - Fruit]

The relationships of Asteranthos have long been uncertain (e.g. Prance & Mori 1979; Mori & Prance (1990). For stipules, see Breteler (2002).

Synonymy: Asteranthaceae R. Knuth, nom. cons., Rhaptopetalaceae Solander, Scytopetalaceae Engler, nom. cons.

Lecythidoideae [Planchonioideae + Foetidioideae]: ?

The food reserve in the embryo is usually starch.

3. Lecythidoideae Beilschmied

Secondary xylem with crystal chains; leaves 2-ranked or spiral, involute; (flowers monosymmetric via androecium), A with a basal ring, filaments contracted at the apex, pollen tricolp/oroid/ate, (fodder pollen +), disc 0/+, endothelium 0, style short (long); fruit operculate (indehiscent); seed with swollen funicle, or aril (= wing), or neither; embryo curved or not, hypocotylar or with long radicle and leaf-like (folded) or fat cotyledons; n = 17 (18).

10/215: Eschweilera (ca 100), Gustavia (40). Neotropical (see Planchonioideae below - New World only [blue]: from Prance & Mori 1979; Mori & Prance 1990). [Photo - Flower, Fruit, Flower, Fruits.]

Lecythidoideae have characteristically fibrous bark. Gustavia has inverted cortical bundles (Metcalfe & Chalk 1950). A few taxa have polysymmetric flowers, and monosymmetric flowers are not an apomorphy of the clade. Androecial variation is extreme; the pollen may be heteromorphic.

Synonymy: Gustaviaceae Burnett

Planchonioideae + Foetidioideae: cortical bundles inverted; leaves supervolute; fruit indehiscent.

In both subfamilies the nodal anatomy appears to be 3:3 if one looks only at the base of the petiole.

4. Planchonioideae Engler

Secondary xylem without crystal chains; leaves spiral, ptyxis?; A with a basal ring, pollen syntricolpate, strong colpus margin ridge; disc annular, endothelium 0 (+), style long; fruit usu. 1-seeded; embryo hypocotylar or with long radicle and leaf-like cotyledons; n = 13.

6/58: Barringtonia (40). Paleotropical (Map - Old World only [red]: from van Steenis & van Balgooy 1966; Payens 1967; Liben 1971b). [Photo - Flower]

In at least some species of Barringtonia there are little glands in the stipular position. Are there apotropous ovules (Baillon 1877)?

Synonymy: Barringtoniaceae F. Rudolphi, nom. cons.

5. Foetidoideae Engler

Secondary xylem with crystal chains; leaves elongating in bud; K woody, valvate, C 0, A free, introrse, disc indistinct, endothelium +, style 3- or 4-fid; n = ?

1/17. E. Africa (Pemba), Madagascar, Mauritius.

Synonymy: Foetidiaceae Airy Shaw

• Lecythidaceae are difficult to characterise. They are usually trees with spiral leaves in tufts at the ends of the branches, or the leaves are two-ranked and scattered along the stem. The leaves are minutely stipulate and/or the margins are serrate. The flowers are often large and have free petals or what appear to be petals. The numerous (up to 1200!) stamens are free to connate; when connate, they may form a complex, cochleate structure that obscures the style and top of the ovary. The ovary is inferior and crowned by the often persistent valvate calyx.

Polysymmetric Lecythidoideae are pollinated by a variety of beees, other than euglossines. Monosymmetric Lecythidoideae are pollinated largely by euglossine bees and several taxa have fodder pollen usually produced by the anthers in the hood, but sometimes by some of those in the ring; nectar secreted by staminodes in the hood is also found in some of these taxa (Prance & Mori 1979; Mori & Prance 1990); details of floral development, incuding the origin of monosymmetry, are to be found placed in a phylogenetic context in Tsou and Mori (2007). Monosymmetric flowers of Lecythidoideae are unlike those of any other angiosperm, with the monosymmetry primarily being evident in the massive development of the abaxial part of the staminal ring that leads to the production of the sometimes complexly coiled hood intio which bees force their way. A rather close evolutionary association between euglossine bees and these monosymmetric Lecthyidoideae has been suggested (e.g. Mori & Boeke 1987). The Cariniana ianeirensis clade is described as having obliquely monosymmetric flowers (Mori et al. 2007). Napoleonaea vogelii pollination and floral morphology has recently been described in detail (Frame & Durou 2001); pollination by thrips is suggested; there are also nectaries inside the flowers at the bases of some of the staminodes and also on the outside of the calyx. Seeds of Lecythidoideae are probably mostly dispersed by mammals, especially primates. Seeds of Lecythidoideae are probably dispersed mostly by mammals, esp. primates.

Although a weakly supported relationship with Sapotaceae had been considered possible in earlier versions of this site, there is currently no evidence for particular sister group relationships between Lecythidaceae and particular Ericalean clades, although they are clearly not sister to the Balsaminaceae clade (Schönenberger et al. 2005).

There is banded apotracheal parenchyma (cf. Sapotaceae!) and crystals in the axial parenchyma, the latter common in several other Ericales, but wood anatomy suggests little about groupings within Lecythidaceae (but see Mori & Prance 1990) and relationships of the family (cf. Lens et al. 2007b). Ditsch and Barthlott (1994) suggested that the rather dimorphic wax platelets of Asteranthos differ from those of Scytopetalaceae, but such platelets also occur in some species of Barringtonia (cf. their figs 26, 27, 29), so are not out of place in Lecythidaceae. The exact nature of the petal-like structures in the flower is still a matter of discussion. There is both centripetal and centrifugal androecial development in the family.

Relationships: [Napoleonoideae [Scytopetaloideae [Lecythidoideae [Planchonoideae + Foetidoideae]]]] - see Morton et al. (1998) and Mori et al. (2007). Scytopetalaceae were considered quite distinct until recently (e.g. Cronquist 1981, in Theales; Takhtajan 1997, in Ochnales [both Dilleniidae]) - C valvate, basally connate or nor, calyptrate and deciduous. The genera it included, plus Asteranthos, are placed in a subfamily in an extended Lecythidaceae, which can more or less be characterised; Lecythidaceae as restricted to the last three subfamilies in the summary phylogeny above cannot.

For more information, see Appel (1996: Scytopetaloideae), Letousey (1961: Scytopetaloideae), Tobe and Raven (1983a), Morton et al. (1997c, esp. 1998: phylogeny), Tsou (1994: embryology) and Tsou and Mori (2002: seed coat anatomy in Lecythidoideae; Takhtajan (1992) includes information on endothelium and testa vasculature. Recent general summaries are provided by Appel (2004: Scytopetalaceae), Prance (2004: Napoleonaceae), Prance and Mori (2004: Lecythidaceae s. str.), and Scott Mori's The Lecythidaceae Pages (Lecythidoideae only).

Sladeniaceae, etc., + Sapotaceae, etc., + [Theaceae + Symplocaceae, etc. + Ericaceae, etc.]: endothelium?

Vessels with vestured pits or walls are scattered, if uncommon, in this group - e.g. in some Symplocaceae, Theaceae, Ericaceae, Clethraceae, and Pentaphylacaceae (Ohtani 1983; Jansen et al. 1998 for general summary).

Sladeniaceae + Pentaphylacaceae: evergreen, woody; vessel elements with scalariform perforations; vessel-fiber pits bordered; nodes 1:1; petiole bundle arcuate; mucilage cells +; hairs unicellular; C ± campanulate, only basally connate, fairly small [petals 1> cm long], A basifixed, pollen 14-30 µm long, surface usu. little ornamented, nectary 0, placentae becoming ± swollen, bitegmic, both integuments ca 3 cells thick, micropyle endostomal; fruit a capsule, columella persistent, K persisting; endosperm +, embryo long.

A.P.G. II suggests as an option recognising three families in this clade, i.e. separating Pentaphylacaceae and Ternstroemiaceae, as well as Sladeniaceae. However, the first two are quite similar phenetically, far more so than they are to Sladeniaceae. See also Theaceae for another family that has been associated with Sladeniaceae and Pentaphylacaceae in the past.

The placenta is very well developed in Ficalhoa and many Ternstroemieae and Frezierieae.

Luna and Ochoterena (2004) were unable to recover any strongly supported relationships in this area in a morphological phylogenetic analysis.

SLADENIACEAE Airy Shaw   Back to Ericales

Exudate + [Ficalhoa]; chemistry?; cork pericyclic; vessels in radial groups [Sladenia] or not; intervessel pitting opposite-alternate; petiole also with wing bundles [Sladenia]; leaves toothed (not); inflorescences axillary, cymose; (C connate – Ficalhoa), A 10-15, anthers opening apically (sagittate – Sladenia), exothecium thickened, microsporogenesis successive [tetrads tetragonal], G [3, 5], 2 apical pendulous epitropous ovules/carpel, or placentation axile, placentae bilobed, many ovules/carpel, ovules tetrasporic, 8-nucleate [Adoxa type], style short, with pointed lobes; fruit also ?schizocarpic, endocarp crustaceous [Sladenia]; seeds winged [Sladenia], testa crustose, cells ± polygonal, little thickened [Ficalhoa]; n = 24 [Sladenia].

2/3. S.E. Asia (Sladenia), tropical E. Africa (Ficalhoa) (Map: see Verdcourt 1962).

• Sladeniaceae are evergreen trees with serrate, exstipulate leaves and distinctive axillary cymose inflorescences. The flowers are small, being less than 5 mm long.

Sladenia is poorly known. It was included in Theaceae (e.g. as Sladenioideae - Takhtajan 1997), and is sister to Pentaphylacaceae (Ternstroemiaceae) in rbcL studies (Savolainen et al. 2000b), albeit the DNA was rather degraded. Its pollen and wood anatomy is very much that of Pentaphylacaceae, but there are no sclereids. Ficalhoa is even more poorly known; it, too, lacks sclereids, but it was not associated with Sladenia in anatomical studies (see esp. Deng & Baas 1991). The two come out as sister taxa in some recent molecular analyses (Anderberg et al. 2002); note, however, that Schönenberger et al. (2005) did not find support for this clade. Li et al. (2003) have recently described a number of very distinctive embryological, etc., features for Sladenia, including monocot wall development; it will be interesting to see if Ficalhoa is similar in these respects. Sladenia has porose anthers, while in Ficalhoa the anthers open across the apex.

For general information, see Stevens and Weitzman (2004).

PENTAPHYLACACEAE Engler, nom. cons.   Back to Ericales

Plants Al-accumulators; parenchyma apotracheal, diffuse or in short tangential lines; intervessel pitting opposite-scalariform; lamina supervolute; inflorescence of axillary flowers or fasciculate; corolla greenish to yellowish [to orange-red in Balthazaria], crystals in the connective (?Pentaphlylax), ovules campylotropous to hemitropous, apotropous when few [?Symplococarpon], style hollow; mesotesta well developed; embryo U-shaped.

12[list]/337, three groups below. Tropical and subtropical, but few in Africa.

The flowers are basically single in the axils of reduced leaves; if the shoot on which they are borne is very much reduced, then the inflorescence is fasciculate.

1. Pentaphylaceae

Chemistry?; druses 0; buds perulate; stomata mostly paracytic; lamina margins entire; flowers in axils of reduced leaves; A 5, filaments very broad, narrowed and incurved apically, theacae each opening by a valve that lifts up, pollen smooth, tectum thin, columellae poorly developed, endexine thick, G [5], opposite C, 2 apical crassinucellate ovules/carpel, stigmas shortly radiate; fruit a capsule, midrib ["teeth"] separating from rest of valves, endocarp cells transversely elongated; seeds flattened, exotestal cells slightly thickened, elongated, mesotestal cells large, thin-walled; endosperm development?, slight, cotyledons longer than the radicle; n = ?

1/1: Pentaphylax euryoides. Kwangtung and Hainan to Sumatra, scattered.

When the shoot does not develop expanded leaves after the flowers appear, it appears that the inflorescence is racemose. Although the leaves are entire, they, the bracts, and some sepals, are terminated by blackish, deciduous and probably glandular points, rather similar to those found in the rest of the Pentphylacaceae. The pericylic sheath consits of fibers alternating with lignified parenchymatous cells (Beauvisage 1920).

A position of Pentaphylacaceae in Ericales seems reasonable. In terms of overall similarity, the anthers are superficially like those of Diapensiaceae (Ericales), while Pentaphylax and Theacaeae s.l. are generally similar. The seed is Ericalean (Huber 1991), and there have been suggestions that Pentaphylacaceae link with Balsaminaceae, etc., also in Ericales (Nandi et al. 1998). Wei et al. (1999) compared the pollen of Pentaphylax with that of Clematoclethra (Actinidiaceae) - again, Ericales. On the other hand, Pentaphylacaceae were associated with Cardiopteridaceae and Gonocaryum in Savolainen et al. (2000a); the latter are strongly associated with Aquifoliales in a three-gene analysis (D. Soltis et al. 2000: see Kårehed 2001). More recently, however, Pentaphylax was placed sister to Ternstroemiaceae s. str. (Anderberg et al. 2001), and this seems to be its resting place for now.

Pentaphylax is poorly known; details of the ovules are taken from Mauritzon (1936).

Ternstroemieae + Frezierieae: ellagic acid +, iridoids 0; (pits vestured); pith often with diaphragms; sclereids +; stomata anomocytic; perulae 0; filaments to 2x longer than anthers, connective usu. prolonged; fruit ± fleshy; mesotesta lignified, ± crystalliferous; endosperm +, ?nuclear, radicle longer than cotyledons, incumbent.

2. Ternstroemieae Candolle

Sclereids much branched; leaves pseudoverticillate, often with black spots, margins entire to crenulate; flowers single from axils of reduced leaves; K opposite C, filaments shorter than anthers, G [2-3], (inferior - Anneslea), 4-12 apical ovules/carpel; fruit irregularly dehiscing; seeds few, 3< mm long, brown, sarcotestal [either exotesta or pockets of fleshy cells on either side of seed], exotesta 10< cells across, sclerified mesotesta 7-15 cells across; n = 20, 25.

2/103: Ternstroemia (100). Tropics, esp. Malesia and Central to South America (Map: from Camp 1947, approximate; M. Sosef, pers. comm.).

The reports of an aril in Ternstroemieae (e.g. Keng 1962) are incorrect; there is a sarcotesta which may, by its expansion, aid in the irregular rupture of the fruit. [Photo - Flowers & Fruits © Nick Turland]

Synonymy: Ternstroemiaceae Candolle

3. Frezierieae

(Nodes 1:3, 3:3 [some Freziera]); sclereids usu. little branched; leaves scattered along shoot, 2-ranked (spiral), lamina margins entire to serrate; plant dioecious or flowers perfect; inflorescence fasciculate or flowers single, at least some from axils of expanded leaves; (K connate; C urceolate [Freziera]), A 5-30(-60), from ring primordium, in a single whorl, (filaments to 5x longer than anthers [Cleyera]; connective not prolonged), G [(1-)3(-10)] (inferior - Symplococarpon), (placentation parietal), 4-many ovules/carpel, also styles +, separate; fruit a berry (drupe); seeds (1-)many, <4(-6) mm long, brown or black, inner walls of exotesta thickened and lignified or not, sclerified mesotesta 1-5 cells across; (embryo curved); n = 12, 13(?), 15, 18, 21(commonest)-23, etc.

9/233: Adinandra (80), Eurya (75), Freziera (57). Southeast Asia to Malesia, Hawaii, Central to South America, E. (Balthasaria) and W. (Adinandra) Africa, and Canaries (Visnea) (Map: from Camp 1947; Verdcourt 1962; van Balgooy 1975; Weitzman 1987). [Photo - Eurya Flower, Flowers & Fruits, Flower, Fruit.]

Cleyera lacks pericyclic fibers in the petiole. It, and also Eurya (for which see Brown 1938), secrete nectary from the basal part of the ovary wall.

• Pentaphylacaceae are shrubs or trees that often have 2-ranked, serrate leaves which may remain rolled up as they elongate, so the lower surface of the blade has longitudinal markings. The flowers are medium-sized to smallish, and the numerous stamens usually have short filaments and long, basifixed anthers. The fruit, which has a persistent calyx, is usually a berry, and the embryo is curved.

Pentaphylax and Visnea are reported fossil from the late Cretaceous Maastrichtian of Europe, while Eurya is known from the Santomasian there (Knobloch " Mai 1986).

Pentaphylacaceae are morphologically amply distinct from Theaceae. The former has pollen 14-28.5 µm long (versus 36.5-54.5 µm), vessel-fiber pits bordered (versus unbordered), etc. Note, however, that differences in the proportion of radicle:cotyledons in the embryo (long radicle in Pentaphylacaceae, short in Theaceae) are not so clear-cut given the inclusion of Pentaphylax and Sladeniaceae in the mix. Freziera shows considerable variation in nodal anatomy, stomatal morphology, seed type and pollen surface (Weitzman 1987). Pentaphylacaceae such as Cleyera have quite long filaments. Ternstroemia itself, with its pseudoverticillate and usually entire leaves, can look rather unlike other Pentaphylacaceae.

Some recent work (Geuten et al. 2004) suggests that [Pentaphylacaceae + Sladeniaceae] are sister to Maesaceae and related families.

Note that the name Pentaphylacaceae is conserved... Ternstroemioideae were a subfamily of Theaceae in Takhtajan (1997).

For general information, see Weitzman et al. (2004, as Ternstroemiaceae), for floral development, see Tsou (1995) and Zhang et al. (2007), for pollen, see Wei (1997), for a phylogeny that includes Theaceae s. str. and a few other Ericales, see Yang et al. (2006: relationships unclear).

Sapotaceae + Ebenaceae + Maesaceae + Theophrastaceae + Myrsinaceae + Primulaceae: ellagic acid 0.

Sapotaceae and Maesaceae and relatives were sister taxa (89% bootstrap) in a six-gene study focusing on Ebenaceae (Duangjai et al. 2006b); the latter was part of a polychotomy including many other Ericales.

SAPOTACEAE Jussieu, nom. cons.   Back to Ericales

Trees and shrubs; saponins, C-30 oxidised triterpenes, pyrrolizidine alkaloids, flavonols, leucodelphinidin, myricetin +; vessel elements with simple (scalariform) perforations; wood siliceous and/or with SiO₂ grains; nodes (1:1) 3:3; (medullary bundles +); petiole bundle arcuate, horizontal D-shaped or annular (wing bundles +); latex sacs +, secreting gutta; sclereids +; hairs brownish, T-shaped, arms unequal or not, unicellular (not in Delpyodon), (cauline stipules +); leaves (2-ranked, opposite), conduplicate, margins entire (toothed), 2ndary veins often rather close, stipules +/0; (plants di- or monoecious), inflorescences cymose, fasciculate, pedicels not articulated; flowers 4-12-merous, K ± connate at base, C 4-18 (variously lobed or divided), A = and opposite (to 6x) C lobes, introrse to extrorse, tapetum multinucleate, pollen 3-6-colporate, staminodes +, disc + (0), G 1[2-14(-30)], opposite K, hairs on the inside of the ovary, placentation axile to axile-basal, 1(-5) ascending apotropous ovules/carpel, (style short), stigma punctate or minutely lobed, dry; fruit a berry (drupe; capsule), K persistent; seeds large, hilar scar large, white, basal, coat multiplicative, hard, shiny, outer part with isodiametric much lignified cells; endosperm + or 0, nuclear; n = (10-)13(-14).

53[list]/1100. Pantropical.

1. Sarcospermatoideae Swenson & Anderberg

Leaves ± opposite, stipules cauline; inflorescence axis apparently well developed [actually a reduced branch]; G 1[-2], style stout; endosperm ?.

1/6. Indo-Malesian (Map: from Aubréville 1964).

The seed has a shiny testa, albeit not as thick as that of most other Sapotaceae, and hilar scar; the genus was placed in Sideroxyleae by Pennington. Some species have paired stipels at the apex of the petiole.

Synonymy: Sarcospermataceae H. J. Lam

Sapotoideae + Chrysophylloideae: staminodes +, ften ± petalline, opposite K (0).

Throughout the tropics (Map: from Aubréville 1964).

2. Sapotoideae Eaton

(K in two whorls of 2-4 valvate members in each; C with three segments); seed with axile hilum; endosperm ?, (amyloid +).

27/543: Palaquium (120), Madhuca (110), Manilkara (80: sapodilla, chicle), Sideroxylum (75), Mimusops (50). Pantropical.

See Smedmark et al. (2006) for general discussion of relationships and character evolution. In Sideroxylon there seems to have been ancient hybridisation 43-36.6. million years ago between a basically African clade and a basically American clade. The descendents, previously segregated as Nesoluma and found on very young islands in the Pacific, may have persisted hopping from island to island ever since (Smedmark & Anderberg 2007; for other taxa behaving similarly, see Hillebrandia [Begoniaceae], Psiloxylum [Myrtaceae], etc.).

Synonymy: Achradaceae Vest, Boerlagellaceae H. J. Lam, Bumeliaceae Barnhart

3. Chrysophylloideae Luersson

Stipules 0; (several stamens opposite each petal; staminodes outside/above the staminal whorl); endosperm copious, cotyledons foliaceous.

25/550: Pouteria (200-305: inc. Planchonella), Chrysophyllum (80), Micropholis (38). Pantropical.

Swenson and Anderberg (2005) suggest that the staminodes common in Chrysophylloideae, but derived within the clade, are perhaps not immediately comparable with the staminodes of other members of the family; the former are outside the staminal whorl, the latter in the same whorl as the stamens. See Swenson et al. (2007a) for generic limits in Australasian members, Swenson et al. (2007b) for Planchonella.

• Sapotaceae are woody plants that may be recognized by their rather small, naked terminal buds with adpressed, brownish, often T-shaped unicellular hairs, axillary branches often with prominent, basal prophyllar buds and then long internode(s), leaf blades that tend to have rather closely parallel secondary veins and entire margins, and petioles that are bottle-shaped when fresh. The twigs usually exude copious gutta. The flowers, with their persistent sepals and often protruding style, are distinctive, as are their seeds, which are large and have a thick, shiny, brown testa with a very large, pale-colored hilum.[Photo - Collection], [Photo - Fruit].

There is banded apotracheal parenchyma (cf. Lecythidaceae!). Anderberg and Ståhl (1995) suggest that bracteoles are absent, Wood and Channell (1960) that they are present. The flowers are sometimes described as being up to 6-merous, i.e. following the number of sepals in a single whorl, however, petals, androecium and gynoecium must then be considered to have doubled in number; see Pennington (2004) for a good summary of floral variation. Swenson and Anderberg (2005) suggest that the basic floral morphology of the family is K5, C5, A 5 + 5 staminodes. Amyloid is also known from the seeds of Omphalocarpum, a clade near sister to the rest of Chrysophylloideae (seee Kooiman 1960).

Generic limits have been notoriously fickle in Sapotaceae: "it is difficult to understand how two authors working on the same family could have come to such widely different conclusions" (Pennington 1990, p. 29), but Pennington himself (1991) has helped clarify the situation.

Sarcosperma is sister to the rest of the family; there are then two major clades, the (Isonandreae + Mimusopeae + Sideroxyleae) and (Chrysophylleae + Omphalocarpeae). Xantolis may be sister to the latter clade (Anderberg & Swenson 2003). In a combined molecular + morphological analysis and after successive weighting the same three clades were recognised, the latter still with only moderate support (79% jacknife) because of the inclusion of Xantolis (the rest of that clade minus Xantolis had 97% support); these were recognised formally as the subfamilies listed above (Swenson & Anderberg 2005). Morphological characters are highly homoplasious, more so than the molecular data, and characters for the subfamilies are hard to come by.

The mitochondrial coxII.i3 intron is absent in Chrysophyllum, at least (Joly et al. 2001).

For more information on Sapotaceae, Ebenaceae, etc., see Franceschi (1993), and Ng (1991). For general information, see Pennington (1991, 2004), and for a checklist and bibliography, see Govaerts et al. (2001).

Ebenaceae + Maesaceae + Theophrastaceae + Myrsinaceae + Primulaceae: ?

EBENACEAE Gücke, nom. cons.   Back to Ericales

Trees, bark and roots black; petiole bundle arcuate; sclereids +; leaves 2-ranked, margins entire, flat glands on lower surface; pedicels articulated; flowers imperfect, ?4-merous; K connate, C contorted, connate, A adnate to C, in two series, basifixed, anthers long, 2 pendulous apotropous ovules/carpel, style ± divided; fruit a berry, K persistent; testa vascularized; endosperm copious, radicle long.

4[list]/548, two subfamilies below. Tropical (to temperate).

1. Lissocarpoideae Wallnöfer

Iridoids, etc.?; hairs 0; cork?; (vessel elements with scalariform perforations); (petiole bundle arcuate but with recurved edges and wing bundles); stomata anomocytic and cyclocytic; plant glabrous; flowers axillary, or inflorescences subfasiculate; bracteoles large, apical; flowers 4(-5)-merous, C with an 8-lobed corona, A 8, filaments connate, anther connective prolonged, pollen 3-porate, psilate, disc?, G [4], inferior, ?integuments, etc., stigma clavate, hairy apically; seeds 1-2; endosperm very hard; cotyledons foliaceous; n = ?

1[list]/8. Tropical South America (Map: from Wallnöfer 2004b).

The morphology of the inflorescence is unclear; one interpretation is that the flowers are axillary, whether on short or long shoots. The flowers seem to be imperfect.

For general information, including a monograph, see Wallnöfer (2004b).

Synonymy: Lissocarpaceae Gilg, nom. cons.

2. Ebenoideae Thorne & Reveal

Saponins, C-30 oxidised triterpenes, napthoquinone derivatives of 7-methyljugone and plumbagin, flavonols, leucodelphinidin, myricetin +, ellagic acid 0; (cork pericyclic); cambium storied; (nodes 1:3); SiO₂ bodies +; secretory cells common; cuticle wax crystalloids 0; stomata usu. paracytic; hairs (T-shaped), unicellular; leaves (opposite, spiral), conduplicate; inflorescence cymose, axis short; flowers 3-7-merous, (C valvate), nectary 0; staminate flowers: A (3-)12-20(-many), extrorse, anthers often hairy, pistillode +; carpellate flowers: staminodes + (0), G [2-8], opposite C or K, loculi often divided, often 1 ovule/carpel, ovules bitegmic, micropyle endostomal, endothelium +, stigmas little expanded, dry; K often accrescent; seed pachychalazal, often ruminate, testa multiplicative (not vascularised), (radicle surrounded by ingrowth of coat), exotesta fibriform or mucilaginous, cells cuboid to palisade, endotesta crystalliferous or not, walls thickened or not; n = 15.

3[list]/540: Diospyros (500+). Tropical (to temperate) (Map: from Morley & Toelken 1983; White 1988 [Africa] - still imprecise). [Photo - Carpellate flower, Fruit, Collection.]

Synonymy: Diospyraceae Vest, Guaiacanaceae Jussieu

• Ebenaceae are woody plants that may be recognised by their usually 2-ranked, entire leaves that may have flat, dark-colored glands on the lower surface of the blade. The corolla is contorted. In Diospyros and relatives the flower buds have adpressed brown indumentum (sometimes of T-shaped hairs) and are distinctively pointed; the calyx is usually much accrescent in fruit. The bark (including that of Lissocarpus), even of twigs, is black, its undersurface is yellow; the heartwood is also black and the leaves, too, may dry blackish - because of the napthoquinones that are common in the family. The flowers of Lissocarpa are glabrous and have a rather narrow sympetalous corolla and an inferior ovary below which are two well-developed bracteoles.

Ellagic acid may occur in the family (Bate Smith 1962). Vessels sometimes occur in radial multiples. Both Massart's model (rythmic monopodial branches) and variants (e.g. Roux - continuous branching) occur in Diospyros. The terminal bud of each innovation frequently aborts. Both integuments appear to be very thick, although the inner is only three cells across at the endostome (van Tieghem 1898). There is variation in germination - foliaceous cotyledons and alternate subsequent leaves vs thick cotyledons and opposite leaves.

Lissocarpaceae have often been placed in or close to Ebenaceae, but they were unassigned in A.P.G. (1998). Rather degraded rbcL sequences suggested that Lissocarpa was to be included in Sapindales - Rutaceae (Savolainen et al. 2000a), however, it is well supported (rbcL only) as sister to Ebenaceae s. str. (Berry et al. 2001), with which it also has much morphologically in common, so it is reasonable to combine the two. However, I do not know of details of its chemistry, so whether the presence of napthoquinones is a synapomorphy of Ebenaceae as a whole or just part of them remains to be established. Duangjai et al. (2006a and especially b), sequencing six plastid genes, found extensive phylogenetic structure in the group; the African(-Arabian) Euclea and Royena were sister to Diospyros, and within Diospyros there were a number of well-supported clades, although relationships between them are unclear.

For Diospyros and relatives and carpel orientation, see Baillon (1891), Eichler (1875) and Le Maout and Decaisne (1868). For general information, see Wallnöfer (2001, 2004a), and on this and putatively related families, see Francheschi (1993), for possible generic characterisations, phylogeny, etc., see Duangjai et al. (2006b). Some information on Lissocarpa is taken from Schadel (1978: leaf morphology) and Wallnöfer (2004a, b), but the genus is poorly known.

Maesaceae [Theophrastaceae [Myrsinaceae + Primulaceae]]: (schizogenous secretory canals [material yellow, red, brown: tannins, etc.]); nodes ?3:3; (stomata anisocytic); small ± immersed often peltate glandular hairs +; inflorescence racemose; C and A from common primordia, C connate, A = and opposite C, antesepalous whorl at least a vascular trace [?Maesa], nectary +, G [5], opposite C, placentation free-central, ovules at least partly immersed in swollen placenta, apotropous, bitegmic, micropyle bistomal, endothelium +, tanniniferous, style short, hollow, stigma ± capitate; seeds angled; endotesta crystalliferous; endosperm nuclear, copious, cell walls thick, with amyloid or hemicellulosic.

Wikström et al. (2001) suggest a stem group age of 75-72 million years before present, with crown group divergence beginning 49-46 million years before present. Members of this group are not often eaten by butterfly larvae, but Lycaenidae-Riodininae-Hamearini and a few Riodinini (see also Abisara) are found on them, especially on Maesaceae but not so far on Samolus and Theophrastaceae (Ehrlich & Raven 1964).

Leaves of Theophrastaceae and Myrsinaceae are often described as being involute (?supervolute, cf. Cullen 1978) or conduplicate. The number of carpels can be difficult to ascertain, but five seems to be a common number, however, the orientation is unclear. The diagrams presented by Dickson (1936) mostly suggest that the carpels are opposite the petals, but in Primula, at least, they suggest that the carpels are opposite the sepals.

The monophyly of the group is not in doubt (see Anderberg & Ståhl 1994; Anderberg et al. 1998; and especially Källersjö et al. 2000: note that support values for Samolus as sister to Theophrastaceae s. str. are reduced when morphological data are added to molecular), and it was often recognised as Primulales in the past (Plumbaginaceae [see Caryophyllales] were also often associated with it because of similar placentation and stamen arrangement). Perhaps the only question, particularly in light of the break-up of Primulaceae, the removal of Maesa from Myrsinaceae, the placement/addition of Samolus as sister to/within Theophrastaceae, the many herbaceous taxa perhaps sister to the old-style, woody Myrsinaceae, and the many features shared by the group as a whole, is whether it is worth recognising families at all... Subfamilial names are already available. For general morphology, see Anderberg et al. (2000) and especially Ståhl and Anderberg (2004).

For the hollow style, see Guéguen (1901: is Maesaceae known?), for staminodes, see Saunders (1936) and Caris and Smets (2004: those of Samolus and Theoprastaceae are developmentally rather different), for nectar secretion, see Vogel (1986, 1997) and Caris and Smets (2004), for embryology especially of the herbaceous taxa, i.e. Primulaceae in the old sense, see Dahlgren (1916), for wood anatomy, see Lens et al. (2005a), and for floral morphology and ontogeny, Dickson (1936: esp. gynoecial arrangement), Sattler (1962), Sundberg (1982), Ronse Decraene (1992), Ronse Decraene et al. (1995) and especially Ma and Saunders (2003).

MAESACEAE Anderberg, B. Ståhl & Kallersjö   Back to Ericales

Evergreen lianes or trees; vessel element type?; petiole bundles all annular; secretory canals well developed; leaves spiral or 2-ranked, induplicate, margin toothed to entire; inflorescence often branched; flowers small; C induplicate-valvate, stamen primordium smaller than petal primordium, A basally connate, attached at the middle of the C tube, nectary on G, G [3-4], half inferior, ovules apotropous, endothelium +, stigma truncate or capitate and lobed; fruit a many-seeded drupe, K persistent; testa 2-layered, inner layer with rhombic crystals; n = 10.

1/150. Old World tropics to Japan, the Pacific, and Australia (map: from Palgrave 2002).

• Maesaceae are woody plants that can be recognised by their serrate, exstipulate leaves that often lack much venation, even when dry, and which have well-developed and conspicuous canals in the leaf blades, sepals and petals. The small, obviously bracteolate flowers have a connate corolla, stamens opposite the petals and an inferior ovary; the many-seeded fruit is drupaceous.

Vessels are in radial multiples (as quite commonly in woody Theophrastaceae and Myrsinaceae); there may be groups of druses in the abaxial epidermis; the fibers are septate; and the lateral bundles arise about half an internode below the leaf they supply.

Information on floral development is taken from Caris et al. (2000); the ovules are separated by and partly sunken in placental tissue (see also Utteridge & Saunders 2001).

Theophrastaceae [Myrsinaceae + Primulaceae]: herbs[?]; rays >5-seriate, uniseriate rays 0 [not herbaceous taxa]; bracteoles 0; C imbricate, arising abaxially on common primordium [i.e. stamen primordium > petal primordium], subrotate, tube rather short.

For the suggestion that rosette herbs may be the plesiomorphic condition for this part of the clade, see Anderberg et al. (2001); however, Lens et al. (2005a) find no evidence from wood anatomy that this is likely (apart from in a few Myrsinaceae). Note that in Myrsinaceae, herbaceous taxa such as Stimpsonia, Ardisiandra and Coris are basal, and variation in habit is very extensive in this clade as a whole.

THEOPHRASTACEAE Link, nom. cons.   Back to Ericales

Bracts displaced up the pedicels; staminodes +, petaloid, endothelium?

6-9[list]/105 - two groups below. Mostly New World and tropical, some also more temperate and Old World (Map: from Hultén 1971).

1. Samolus

Nodes ?1:1; leaves entire; K connate, nectary on ovary, G [5], semi-inferior, style impressed; fruit a 5-valved capsule, seeds many; coat undistinguished, exotesta and endotegmen tanniniferous, the latter crystalliferous; endosperm cell walls thin; n = (12) 13.

1/15. America, the Antipodes, Europe, tropical to temperate (Map: from Hultén 1971; Meusel et al. 1978; FloraBase 2005). [Photo - Flowers.]

Ståhl (2004) suggests that a secretory system is present, if not always conspicuous. The stomata are anomocytic. There are several petiole bundles forming an arc, and these seem to diverge very soon after the leaf trace departs from the central stele. The ovules completely cover the placenta, but fingers of placental tissue may poke up between them (but not seen in the material examined by Caris & Smets 2004); Ma and Saunders (2003) suggest that in this whole clade (i.e. Theophrastaceae s.l.) the ovules are not embedded (which would then be a synapomorphy for it...). The valves of the capsule are opposite the calyx (Caris & Smets 2004).

For general information, see Ståhl (2004: as Samolaceae).

Synonymy: Samolaceae Rafinesque

2. The Rest (Theophrastaceae s. str.)

Woody, tending to be pachycaul; rays broad; nodes also 1:1 [Jacquinia, dividing into three], 5:5 [Clavija]; secretory system?; petiole bundle deeply arcuate or annular, with small adaxial inverted bundles; scale leaves +; leaves conduplicate, margins spiny-toothed to entire, subepidermal fibers +; plant dioecious or flowers bisexual; anthers extrorse, with calcium oxalate, (nectariferous hairs +), style long, stigma dry or wet; fruit a (rather dry) berry, placentae ± pulpy, (drupe); seeds 1-few, rounded, exotestal cells flattened, thick-walled, hypodermal cells (with thickened anticlinal walls), often crystalliferous; endosperm cell walls pitted, cotyledons usu. foliaceous; n = 18, 20, 24.

4/90: Clavija (50), Jacquinia (35 - perhaps to be divided). New World tropics (Map: from Ståhl 1989, 1991, 1995). [Photos - Collection]

• Theophrastaceae have sympetalous flowers with stamens opposite the petals and five, petaloid staminodes opposite the sepals. Most taxa are woody plants with broad rays and often pseudoverticillate leaves with toothed or spiny margins or a pungent apex. The flowers are sympetalous and have five stamens. The extrorse anthers form a cone in the center of the flower when it opens, but later spread; the staminodes are more or less petaloid. The subepidermal fibers are visible in the dried leaf as fine striations - use a hand lens!

The subepidermal fibres may lack lignification. For reports of glandular dots on calyx and corolla, see Mabberley (1997). Källersjö and Ståhl (2003) suggest some generic realignments within the family. For morphology, etc., see Ståhl (2004) and in particular Caris and Smets (2004).

Primulaceae + Myrsinaceae: two ndhF deletions.

PRIMULACEAE Borkhausen, nom. cons.   Back to Ericales

Cucurbitacins +; ?cork; glands 0, trichomes articulated; leaves involute or revolute, margins entire to dentate or serrate; inflorescence scapose; K often connate, C hypocrateriform, A attached at or above middle of C tube, pollen syn- or polycolpate, nectary on ovary, ovules not immersed in placenta (immersed - Dionysia), (inner integument ca 4 cells across), style usu. long, (heterostyly +); fruit a capsule; seeds many, angled, exotesta ± persistent, walls thickened or not, (endotesta with inner walls thickened [Primula]), endotegmen often crystalliferous; (endosperm cell walls thin); n = 8-12.

9[list]/900: Primula (490-600: inc. Cortusa, Dionysia [some chasmophytes, for which see Trift et al. 2004, relationships, biogeography; Lidén 2007, revision), Dodecatheon), Androsace (160: inc. Douglasia, Vitaliana, see Schneeweiss et al. 2004b). Northern hemisphere, scattered elsewhere (Map: from Hultén 1971; Meusel et al. 1978). [Photo - Dodecatheon flower © R. Kowal] [Photo - Primula flower]

• Primulaceae are usually rosette herbs with a scapose inflorescence and medium-sized flowers with connate sepals, obviously connate petals, hypocrateriform corolla and stamens borne opposite the corolla lobes; the placentation is free-central and the capsular fruits have numerous, angular seeds.

The involute leaves can be sharply bent rather than incurved (for ptyxis, see Conti et al. 2000; Mast et al. 2001). Solereder (1908) reports that secretory tissues occur in Androsace. Heterostyly is common, although it is unlikely to be an apomorphy for the family; it is sometimes lost, as in those Primula with buzz pollination, the erstwhile Dodecatheon (Mast et al. 2001, 2006). The corolla epidermal cells are isodiametric. Saunders (1936) suggested that some of the lobing of the corolla of Soldanella might be staminodial.

For ITS-based relationships within the family, see Martins et al. (2003), and for relationships within Primula, see also Trift et al. (2002) and Mast et al. (2004, 2006), for relationships within Androsace, see Wang et al. (2004) and Schneeweiss et al. (2004b), for pollen variation, see Mast et al. (2001), and for general morphology, see Anderberg (2004). Richards (2003) provides a good general account of Primula s. str.

MYRSINACEAE R. Brown, nom. cons.   Back to Ericales

Also trees to shrubs or lianes; benzoquinones +; (vessel elements with scalariform perforations); (nodes 3:3 - unnamed taxon from Atlantic Forest; Ardisia densiflora); glands/canals throughout the plant (0); leaves (opposite), also supervolute (curved), margins entire (crenate to serrate, teeth cartilaginous); (plant dioecious), inflorescence often fasciculate/corymbose; flowers (3-)4-5(-7)-merous, C often contorted, (nectariferous hairs +), A dorsifixed or basifixed, sagittate, (porose), (micropyle endostomal - Coris; endothelium 0; style 0; long), stigma (punctate), dry or wet; fruit a berry, drupe or capsule [latter in herbaceous taxa], placentae ± pulpy; seeds 1-few, rounded (ruminate; hilum depressed) [woody taxa] or many, small, angular, seed coat undistinguished, (endotesta crystalliferous - Cyclamen), tegmen thickened before becoming crushed, (endotegmen crystalliferous); endosperm walls pitted, (embryo slightly curved; medium); n = 10-13, 15, 17, 23.

41[list]/1435: Ardisia (450), Myrsine (155: inc. Rapanea, Suttonia), Lysimachia (150: sometimes staminodes?, some woody; white flowers with nectariferous hairs, some yellow oil flowers [Macropis the pollinator], or selfers - see Vogel 1986), Discocalyx (115: inc. Tapeinosperma), Embelia (100), Parathesis (85), Stylogyne (60). Pantropical and N. Temperate (Map: from Hultén 1958, 1971; FloraBase 2008: S. Hemisphere a bit notional). [Photos - collection woody members, Cyclamen flower © H. Schneider, fruit © H. Schneider], collection of ex Primulaceae.]

• Myrsinaceae are trees to herbs, and many have distinctive yellowish to blackish dots or streaks on the often spiral leaves and also often obvious on the persistent calyx and on the fruit - I have not seen them on genera like Trientalis, Anagallis (but there may be pigmented spots on the abaxial surface of the lamina), Cyclamen, or some species of Lysimachia. The sympetalous corolla is often contorted, the five stamens are antepetalous, the ovary is superior, and the ovules and/or seeds are immersed in the placenta. In many woody taxa the point of insertion of the branches on the main stem is vertically elongated; growth is often sylleptic.

Vogel (1986) discusses pollination, which in a group of yellow-flowered Lysimachia in particular is by oil-collecting Macropis (Mellitidae) bees (see also Simpson et al. 1983). The oil is secreted by trichomes. Anderberg et al. (2007) suggested that Lysimachia with buzz-pollinated flowers and those with nectar-producing hairs both formed separate clades and were both derived from oil-producing ancestors. Some species of Ardisia have pustules along the edge of the leaf blade; although inhabited by bacteria, it is unclear what role they might play (Miller 1990).

Myrsinaceae include Anagallis, Ardisiandra, Asterolinon (?= Lysimachia), Coris, Cyclamen, Glaux, Lysimachia, Pelletiera, Stimpsonia and Trientalis (Anderberg et al. 2000, 2001; the limits of Myrsinaceae are not so clear in Martins et al. 2003, but ITS data alone in the latter study). Generic limits in the woody members are unsatisfactory. Anderberg et al (2007) studied the relationships of the herbaceous taxa; the family had moderate support as being monophyletic (72% jacknife), and Cyclamen, the herbaceous taxa, and the woody taxa then formed a trichotomy.

Coris is a particularly distinctive genus. It is a small ericoid sub-shrub with monosymmetric flowers that have a spine-tipped epicaly. There is nectary at the base of the ovary and there are only 5-6 ovules. Monosymmetry is expressed early in development by the calyx, monosymmetry of the corolla becomes evident only later (Ronse Decraene et al. 1995: they suggest that the median sepal is abaxial, i.e. that the orientation of the flower is inverted). The mangrove Aegiceras has a number of anomalous anatomical and morphological features, the seed characters in particular are those that might be expected from a mangrove plant. Its rays are relatively narrow, the seeds lack endosperm and contain a large, viviparous embryo, and the ovules are unitegmic, etc. (Staåhl å Anderberg 2004; Lens et al. 2005).

The presence of coloured glands may well not be a synapomorphy of the family (Hao et al. 2004). There are breakdown areas in the rays of woody members, and these may be filled with dark contents (Lens et al. 2005). Discocalyx has three traces in the petiole base, and some other taxa may be trilacunar. The epidermal cells of the corolla are often elongated (Glaux - to be included in Lysimachia - lacks a corolla); this is a derived feature within the family. Cyclamen has one cotyledon and one integument. Note that its stamens are initiated as adaxial outgrowths of a common primordium, i.e. the petal primordia are early larger than the stamen primordia, as also in Myrsine and Aegiceras (see esp. Ma & Saunders 2003).

Some information is taken from Otegui and Cocucci (1999) and from Ståhl and Anderberg (2004); Hao et al. (2004) provide a phylogeny of the family, although focusing on Lysimachia, while Lens et al. (2005a) provide much information about wood anatomy.

Synonymy: Aegicerataceae Blume, Anagallidaceae Adanson, Ardisiaceae Jussieu, Coridaceae J. Agardh, Embeliaceae J. Agardh, Lysimachiaceae Durande

MITRASTEMONACEAE Makino, nom. cons.   Back to Ericales

Endophytic root parasites; ?anatomy; leaf waxes hummocky; leaves opposite, scale-like; flowers single, terminal; P 4, connate, A extrorse, completely connate and surrounding G except for small apical pore, polythecate, pollen binucleate, 2-porate [?colpate], ektexine reduced to tuberculae, G with 8-20 intrusive parietal placentae, ovules many, unitegmic, integument 2 cells thick, with a funicular obturator, style stout, stigma hemispherical; fruit berry-like, circumscissile; funicle sticky; exotestal cells with massive U thickenings; endosperm 1-layered, embryo undifferentiated, 4-celled; n = 20.

1/2. South East Asia, Malesia, Central America, N.W. South America, scattered (Map: from van Steenis and van Balgooy 1966; Meijer & Veldkamp 1993). [Photo - Habit © S. Hsiao]

• Mitrastemonaceae are root parasites, often found on Fagaceae, with opposite, scale-like leaves. The single, terminal flowers are surrounded at the base by the perianth, the stamens are completely connate, forming a conspicuous cone entirely surrounding the gynoecium except for a terminal opening, and the anthers are extrorse. The ovary has parietal placentation and the fruit contains many minute seeds.

Watanabe (1936: V) talks a lot about a "Mitrastemon-Pilz". The pollen may have three or four pores - see Watanabe (1936: III). Cronquist (1981) and Meijer and Veldkamp (1993) describe the fruit as being a berry or berry-like and opening via a transverse slit - i.e., it is also some sort of circumscissile capsule - and the latter both described the ovule as being unitegmic and the seed as as being formed from the inner integument (the latter following Watanabe 1937: VII).

Along with Cytinaceae and Rafflesiaceae, relationships with Malvales have been suggested (Nickrent 2002). Barkman et al. (2004) use mitochondrial sequences to place Mitrastemonaceae in Ericales, a position that appeared in most analyses in Nickrent et al. (2004: for further discussion, see Rafflesiaceae). Its cellular endosperm is certainly compatible with a position in Asterids, and its extrorse anthers are perhaps comparable with those of Ericaceae and their relatives. Its parietal placentation is found in many other parasitic angiosperms, as well as in the echlorophyllous hyperparasitic Ericaceae-Monotropoideae.

For general information (including a more extensive list of hosts) and references, see Meijer and Veldkamp (1993), also the Parasitic Plants website (Nickrent 1998 onwards).

Theaceae + Symplocaceae, etc. + Ericaceae, etc.: cork?; leaves serrate.

THEACEAE Ker Gawler, nom. cons.   Back to Ericales

Trees or shrubs; plants Al-accumulators; myricetin, ellagic acid +; cork pericyclic (subepidermal); vessel elements with scalariform perforations; (pits vestured); intervessel pitting opposite-scalariform; pericyclic fibers +/0; petiole bundle arcuate; sclereids and mucilage cells common; stomata paracytic or cyclocytic; hairs unicellular; leaves also 2-ranked, lamina involute or supervolute (conduplicate), margins toothed (entire); flowers single, axillary; C ± free, A usu. 40<, ± basally connate, centrifugal, from 5 primordia opposite C or ring primordium in 2-5 whorls, anthers articulated, connective usu. not prolonged, pollen tricolporoidate, pseudopollen produced from connective, nectar from base of filaments or ovary, G [(3-)5(-10)], opposite C, 2-few (basal) bitegmic ovules/carpel, outer integument 5-7 cells thick, inner integument ca 4 cells thick, micropyle endostomal, (styles +, separate), stigma wet; fruit a capsule, central axis often persistent, K persistent or not; seeds few, often >4 mm long, flattened, testa and tegmen vascularised, exotesta lignified or not, mesotesta lignified (fibrous; with sclereids), endotesta lignified or not; endosperm nuclear, usu. slight, cotyledons longer than radicle, accumbent.

Ca 7[list]/195(-460!): Camellia (120), Pyrenaria (42). Mostly South East Asia-Malesia, also S.E. U.S.A. [Photo - Flower, Fruit, Collection.]

• Theaceae are shrubs or trees that are recognised by their quite thick and serrate exstipulate leaves, they often turn red just before falling. The plant has sclereids that may be visible as minute punctations, e.g. in the leaf blades. The flowers are usually medium-sized to large, and the bracteoles, calyx and corolla are quite often in a single spiral and are not clearly distinguishable one from another. The numerous stamens have quite long filaments and short, dorsifixed anthers. The fruit is usually capsular and with a columella, and the seeds, often flattened, have straight embryos.

The cotyledons have three or more traces from a single gap. The stomata are often described as being "gordoniaceous".

Generic limits in Theaceae are difficult, and family relationships are still poorly understood (for useful notes on the genera, see Prince 2007). An analysis of two chloroplast genes by Prince and Parks (2001) suggests that there are three major clades within the family and that Polyspora and Laplacea should be separated from Gordonia (see also Airy-Shaw 1936; Yang et al. 2004: genes from all three genomes, 2006: mitochondrial gene only, family in broad sense, including Pentaphylacaceae). However, the relationships between the three clades is unclear. An analysis of matK data alone suggested that Theeae were sister to the other two tribes, but there was a polychotomy in the combined analysis (including rbcL data: Prince & Parks 2001), while Yang et al. (2004) found Stewartieae to be sister to the rest, although not with very strong support. Below is a suggestion of how the family might be broken down.

1. Theeae Szyszylowicz

Pedicels multibracteolate, K and C intergrading, A in 2 whorls; capsule with columella; seeds winged or not; n = 15.

5/Apterosperma, Camellia, Laplacea, Polyspora, Pyrenaria. Southeast Asia, Malesia, tropical America (Map: from Camp 1947, approximate).

2. Gordonieae de Candolle

Pedicels bibracteolate; K 5, K and C dissimilar, A in 3-5 whorls [?is this a character]; capsule with columella, dehiscence also septicidal; seeds apically winged (not – Franklinia]; n = (15), 18.

3/4-30. Franklinia, Gordonia, Schima. Southeast Asia, West Malesia, S.E. United States (Map: from Camp 1947; Bloembergen 1952).

3. Stewartieae Choisy

Pedicels bibracteolate; K 5, K and C dissimilar; capsule lacking columella; seeds narrowly winged or not; n = 15, 17, 18.

1/9. East Asia, E. North America (Map: from Hong 1993).

Stewartia includes Hartia (e.g. Prince 2002).

Camellia has bisporic, eight-nucleate Allium-type embryo sacs. Although the carpels seem to be opposite the sepals in Camellia, this may be connected with the arrangement of the perianth, not to that of the gynoecium, since the basic orientation of the gynoecium is clearly the same as that of Gordonia, where the carpels are clearly opposite the petals (Eichler 1878).

Theaceae s. str. may be sister to Symplocacaeae, this clade being in turn sister to Styracaceae and Diapensiaceae, in turn related to the Actinidiaceae/Clethraceae clade (Geuten et al. 2004).

Theaceae s.l. have in the past been associated with Asteropeiaceae (e.g. Takhtajan 1997), for which, see Lecythidaceae. Pentaphylacaceae, Sladeniaceae and Pellicieraceae, erstwhile Theaceae, are all separate families here; Pentaphylacaceae, often recognised as a monotypic family (see e.g. A.P.G. 1998; /APweb/ versions 1-3) are here expanded to include Ternstroemiaceae, also part of or placed adjacent to the old Theaceae.

For further information about Theaceae s.l., see Keng (1962), Grote and Dilcher (1989: fossil record), Liang and Baas (1991), Yang and Min (1995a, b: embryology and systematics), Tsou (1995, 1997, 1998), Stevens et al. (2004b: general), and Wang et al. (2006: Apterosperma, chromosomes and morphology); for pollen, see Wei (1997).

Synonymy: Camelliaceae Candolle, Gordoniaceae Sprengel

Symplocaceae + Styracaceae + Diapensiaceae: inflorescence racemose; endosperm copious.

SYMPLOCACEAE Desfontaines, nom. cons.   Back to Ericales

Trees; plants Al-accumulators, O-methyl flavonols, route II decarboxylated iridoids, ellagic acid +, myricetin 0; vessel elements with scalariform perforations; true tracheids +; crystal sand +; stomata usu. paracytic, very large "water stomata" also present; (leaves 2-ranked), lamina ± supervolute; (inflorescence branched), pedicels articulated; K basally connate, A (= and opposite K)-many, in bundles, (connate), adnate to C, anthers globose, pollen angular, spinuliferous, G [2-5], (half) inferior, median member abaxial, disc +, 2-4 pendulous epitropous ovules/carpel, endothelium +, stigma ± capitate; fruit drupaceous, with as many pores as fertile carpels, K persistent; seed usu. 1, exotestal cells with inner walls thickened or thin; embryo large, (curved); n = 11 (12); mitochondrial coxII.i3 intron 0.

1[list]/320. Tropical to subtropical, inc. New Caledonia, not Africa, subgenus Hopea common as fossils in Europe (Eocene) (Map: see Nooteboom 1975). [Photo - Symplocos chinensis Flowers]

• Symplocaceae are shrubs or trees that can be recognised by their serrate, exstipulate leaves which often dry yellowish and racemose inflorescences with rather small flowers. The petals are only basally connate, there are often many stamens, and the ovary is inferior - indeed, the flowers look superficially rather like those of Rosaceae-Maloideae. The fruit is drupaceous and the stone has germination pores.

Symplocos is locally very abundant in the Tertiary fossil record of Europe. The style just below the stigma secretes oil in the ca 145 species of Symplocos sect. Symplocastrum which probably helps the pollen stick to the pollinators (Kriebel et al. 2007); this seems to be unique in flowering plants.

For a phylogeny of the genus, see Y. Wang et al. (2004) and Fritsch et al. (2006); section Cordyloblaste appears to be sister to the rest. The infrageneric taxonomy needs reworking.

Although the placentation is described as being fully axile, in material seen it is parietal at the apex. The androecium is basically obdiplostemonous (Caris et al. 2002).

For testa anatomy, see Corner (1976) and Huber (1991), and for general information, see Nooteboom (2004).

Styracaceae + Diapensiaceae: cork pericyclic; glandular hairs 0; leaves spiral, (margins entire); A basifixed, style continuous, hollow; fruit a capsule.

There is also fairly good support for this clade in B. Bremer et al. (2002). Scott (2004) and Fritsch (2004) suggest that there are embryological features in common between the two families; I do not know if any of them are really synapomorphies.

STYRACACEAE Candolle & Sprengel, nom. cons.   Back to Ericales

Trees or shrubs; ellagic acid, myricetin 0, iridoids?; vessel elements with scalariform (simple) perforations; wood siliceous; resin canals often +; petiole bundle arcuate or D shaped (medullary and/or wing bundles +; complex - Parastyrax); indumentum stellate or scaly; leaves conduplicate-plicate or supervolute; bracteoles 0, (pedicels articulated); flowers (4-)5(-7)-merous; K ± completely connate, open, C valvate or not, A 2(3)x or = and alternate with K, adnate to C, often basally connate, (filaments as broad as anther - Styrax sect. Pamphilia), connective produced or not, pollen spinuliferous, nectary 0, G [2-5], ± inferior, alternate with K, median member ?abaxial, often with hairs inside, 1 basal apotropous [Pamphilia]-many (bitegmic - Styrax) ovules/carpel, micropyle endostomal, placental obturator +, (endothelium + - Alniphyllum), (long style branches), stigma punctate or lobulate, dry; fruit also drupaceous; testa vascularised, crushed; n = 8.

11[list]/160: Styrax (120 - benzoin, gum bejamin [sic], pedicels not jointed). Warm N. temperate to tropical (Map: from van Steenis 1949b; Sales & Hedge 1996; Fritsch 1999). [Photo - Flower, Fruit.]

• Styracaceae may be recognised by their stellate or lepidote indumentum; spiral, serrate, exstipulate leaves; and sympetalous flowers with a more or less completely connate and/or