EXTANT SEED PLANTS

Plant woody, evergreen; nicotinic acid metabolised to trigonelline, (cyanogenesis via tyrosine pathway); primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins rich in guaiacyl units; true roots present, apex multicellular, xylem exarch, branching endogenous; arbuscular mycorrhizae +; shoot apical meristem multicellular, interface specific plasmodesmatal network; stem with ectophloic eustele, endodermis 0, xylem endarch, branching exogenous; vascular tissue in t.s. discontinuous by interfascicular regions; vascular cambium + [xylem ("wood") differentiating internally, phloem externally]; wood homoxylous, tracheids +; tracheid/tracheid pits circular, bordered; sieve tube/cell plastids with starch grains; phloem fibers +; stem cork cambium superficial, root cork cambium deep seated; nodes ?; stomata ?; leaf vascular bundles collateral; leaves spiral, simple, axillary buds?, prophylls [including bracteoles] two, lateral, veins -5 mm/mm² [mean for all non-angiosperms 1.8]; plant heterosporous, sporangia eusporangiate, on sporophylls, sporophylls aggregated in indeterminate cones/strobili; true pollen [microspores, i.e. no distal pore for release of gametes] +, grains mono[ana]sulcate, exine and intine homogeneous, ovules unitegmic, crassinucellate, megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development first endo- then exosporic, tube developing from distal end of grain, to ca 2 mm from receptive surface to egg, gametes two, with cell walls, with many flagellae; female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large", first cell wall of zygote transverse, embryo straight, endoscopic [suspensor +], short-minute, with morphological dormancy, white, cotyledons 2; plastid transmission maternal; two copies of LEAFY gene, PHY gene duplication [N/O//A/C and P//BE lines], 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 in ANITA grade?], 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 a sieve plate and cytoplasm with P-proteins, companion cells from same mother cell that gave rise to the sieve tube; nodes unilacunar [1:?]; stomata with ends of guard cells level with pore, paracytic, outer stomatal ledges producing vestibule; leaves with petiole and lamina [the latter formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, veins (1.7-)4.1(-5.7) mm/mm², endings free; flowers perfect, polysymmetric, parts spiral [esp. the A], free, development in general centripetal, numbers unstable; P not sharply differentiated, outer members not enclosing the rest of the bud, smaller than inner members; A many, 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, walls developing by centripetal furrowing; pollen subspherical, binucleate at dispersal, trinucleate eventually, tectum continuous or microperforate, ektexine columellar, endexine thin, compact, lamellate only in the apertural regions; nectary 0; G free, several, ascidiate, with postgenital occlusion by secretion, few [?1] ovules/carpel, ovules marginal, anatropous, bitegmic, [outer integument often largely subdermal in origin, inner integument dermal], micropyle endostomal, integuments 2-3 cells thick, megasporocyte single, megaspore lacking sporopollenin and cuticle, chalazal, female gametophyte four-celled [one-modular, nucleus of egg cell sister to one of the polar nuclei], stylulus short, hollow, stigma ± decurrent, dry [not secretory]; P deciduous in fruit; seed exotestal; pollen germinating in less than 3 hours, tube elongated, growing at 80-600 µm/hour, with callose plugs and callose-based walls, penetrating between cells, siphonogamy, penetration of ovules within ca 18 hours, distance to first ovule 1.1.-2.1 mm; 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, minute; 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 + C/PHYB + E gene pairs.

Evolution. Possible apomorphies for flowering plants are in bold. Note that the actual level to which many of these features, particularly the more cryptic ones, should be assigned is unclear, because some taxa basal to the [magnoliid + monocot + eudicot] group have been surprisingly little studied, there is considerable variation between families in particular for several of these characters, and also because details of relationships among gymnosperms will affect the level at which some of these characters are pegged. For example, if reticulate-perforate pollen is optimized to the next node on the tree (see Friis et al. 2009 for a discussion), it effectively makes the pollen morphology of the common ancestor of all angiosperms ambiguous....

NYMPHAEALES [AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]: vessels +, elements with scalariform perforation plates, axial parenchyma diffuse or diffuse-in-aggregate; tectum reticulate-perforate [here?]; ?genome duplication; "DEAER" motif in AP3 and PI genes lost, gaps in these genes.

ERICALES [ASTERID I + II]: ovules tenuinucellate.

ERICALES Dumortier  Main Tree, Synapomorphies.

Nonhydrolysable tannins, triterpenoids incl. saponins +; vessel elements with simple perforation plates; nodes 1:1; leaf vascular bundles collateral; leaves spiral, teeth with single vein and opaque deciduous cap; duplication of the PI gene. - 25 families, 346 genera, 11,515 species.

Evolution. 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) proposed that 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, while Janssens et al. (2009) date stem group Ericales to 123±10.5 million years ago and the crown group to 117±9.2 million years. Almost all families had diverged by the early Eocene (50 million years before present: Sytsma et al. 2006). Soltis et al. (2008: a variety of estimates) suggest an age of divergence of Ericales from the rest of 126-113(-85) million years ago.

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 million years old (Crepet et al. 2001, 2004; see also Herendeen et al. 1999), and some of these are quite unlike extant members of the clade, e.g. some have sepals with numerous huge abaxial and/or marginal glands (Crepet 2008). 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; Martínez-Millán et al. 2009 - ?Pentaphylacaceae). Note that tricolpate pollen is uncommon in extant Ericales, being known only from Lecythidaceae and Balsaminaceae.

Today Ericales are an important component of the diversity of the understorey 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. Currently 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).

Viaene et al. (2009) discuss the complex history of PI gene duplication, sub- and neofunctionalisation, and loss in the clade. For all taxa in which they found two copies of the PI gene, have connate filaments, however, Primulaceae-Theophrastoideae, with connate filaments, had only a single copy (Viaene et al. 2009).

Chemistry, Morphology, etc. 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.

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 ovary placentation, see Ng (1991); although true parietal placentation occurs in the order (e.g. Ericaceae - Pyroloideae), most other reports are incorrect.

Phylogeny. 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 other relationships were unclear. Many taxa lack the mitochondrial coxII.i3 intron, but it is known from the Maesaceae (and Balsaminaceae - plesiomorphic presence?) 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) suggested 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).

Previous Relationships. 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.

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, Aegiceratales Blume, Ardisiales J. Presl, Balsaminales Lindley, Barringtoniales de Candolle, Camelliales Link, Cyrillales Doweld, Diapensiales Engler & Gilg, Diospyrales Prantl, Ebenales Engler, Empetrales Rafinesque, Epacridales Berchtold & J. Presl, Fouquieriales Reveal, Gordoniales J. Presl, Halesiales Link, Lecythidales Cronquist, Lysimachiales Döll, Marcgraviales Doweld, Mitrastemonales Nakai, Monotropales Berchtold & J. Presl, 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; raphide sacs +, druses 0; lamina toothed; inflorescence racemose; stamens = and opposite sepals, free from C, anthers (near) basifixed, gynoecial nectary 0, ovules bitegmic, micropyle endostomal, style short, stigma little expanded; endosperm with micropylar haustorium.

The common ancestor of [Balsaminaceae + Marcgraviaceae] dates to middle Palaeocene ca 58.9 million years ago; the two families diverged ca 48.1 million years ago (Janssens et al. 2009).

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; the gene is normally expressed in the corolla and other inner whorls. Most taxa in this clade have more or less petal-like sepals, bracts or bracteoles - Tetramerista and Pentamerista, derived members of the clade, lack such expression patterns.

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).

Monophyly of the clade containing 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; Janssens et al. 2009: ML bootstrap support 98%).

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

Fleshy herbs; leucoanthocyanins, naphthoquinones +; 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 petals, 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 with 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 with the walls inrolling from the base.

Evolution. Impatiens is a genus that is most diverse in tropical and subtropical montane forests. Stem Balsaminaceae are ca 48 millions years old, crown diversification (Hydrocera diverged) began ca 31 million years ago, more diversification may have begun in the Early Miocene ca 22.5 million years ago, but the speciation rate much increased in the early Pliocene within the last 5 million years, when climate change caused much population fragmentation, isolation, and migration (Janssens et al. 2009).

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 gene has occured in this clade (Janssens et al. 2006b; Geuten et al. 2006).

Chemistry, Morphology, etc. 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.

Phylogeny. 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).

Previous Relationships. 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 perforation plates); 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.

Evolution. 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). 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).

Chemistry, Morphology, etc. 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). Johri et al. (1992) described the seeds as being arillate.

Phylogeny. Ward and Price (2002) suggest phylogenetic relationships within the family; Marcgravia, with its reversible heterophylly, two-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[list]/5. W. Malesia, Central and N. South America. Two groups below.

1. Pelliciera Bentham   Back to Ericales

Vessels in multiples; petiole bundle more or less flat where it joins the stem (?later annular); stomata ?; leaves involute, 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.

1/1: 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.

Evolution. From fossil pollen, Pelliciera was once much more widespread (A. Graham 1977), even being found in the Old World; for the evolution of the mangrove habitat, to which Pelliciera is restricted, see Rhizophoraceae.

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

2. Pentamerista + Tetramerista

Cork inner cortical; (vessel elements with scalariform perforation plates); wood fluorescing [1 sp. tested]; nodes 3:3; 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 = ?

2/4. 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; there are clearly three separate bundles at the base of the petiole. The flowers are rather small, the sepals and petals being rather similar and spreading; at least the sepals have glands on their adaxial surfaces.

Chemistry, Morphology, etc. 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; it is probably unilacunar. The products of different marginal glands of the one leaf may not be the same (Collins et al. 1977). The floral diagram of Pelliciera 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. Nodal anatomy of [Tetramerista + Pentamerista] was extrapolated from the petiole scars. In Tetramerista there are glistening dots on the adaxial surface of both calyx and corolla. The embryology, morphology and anatomy of Pellicieraceae s.l. are poorly known

Previous Relationships. Tetrameristaceae s. str. were placed in Theales by Takhtajan (1997). 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. .

For general information, see Kubitzki (2004b).

Other Ericales: corolla tube well developed, style long.

Polemoniaceae + Fouquieriaceae: cork cambium outer cortical; C connate, nectary +; G [3], style hollow, stigma strongly lobed; K persistent in fruit; 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 cambium also pericyclic; (vessel elements with scalariform perforation plates; stomata paracytic); leaves opposite to spiral (palmate, pinnate), conduplicate, margins entire to deeply lobed; inflorescence cymose, paniculate or thyrsoid; (flowers monosymmetric), K connate, lobes with green midrib and colorless intermediate portion, C tube formation late, lobes usu. contortuplicate, stamens = and opposite sepals, 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]; also sporophytic incompatibility system present.

13-22/350 [list]: Phlox (70), Linanthus (35), Navarretia (30), Polemonium (27), Gilia (25). Especially western North America, also a few 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); (short shoots +): 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 exine verrucate; 100-220 µm long), ovules with nucellar cap; fruit septicidal and/or loculicidal; seeds wing broad (narrow - Bonplandia), mesotestal cell walls thinly lignified; n = [x = 7-9?], n = 15, 26, 27, small.

4/34 [list]. Baja California, tropical America. [Photo - Flower & Fruits]

Synonymy: Cobaeaceae D. Don

3. Acanthogilioideae J. M. Porter & L. A. Johnson

Shrubby; leaves very dimorphic, persistent branched spines on long shoots, deciduous and unlobed on short shoots; pollen zonocolporate, exine coarsely verrucate; seeds few, winged; n = 9, chromosomes 2-4 .

1/1 [list]: Acanthogilia gloriosa. Baja California.

• 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.

Evolution. Gilisenium hueberi, perhaps close to Gilia, is known from the middle Eocene of Utah (Lott et al. 1998).

Chemistry, Morphology, etc. 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.

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

Phylogeny. Acanthogilia has been placed in its own subfamily (Porter et al. 2000), 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). However, its position relative to the other two subfamilies is still unclear (Johnson et al. 2008 and references). It has very dimorphic leaves and short shoots are in this is like Fouquieraceae, its branched spines are reduced leaves like those found scattered in Polemonioideae, and its sepals have a green midrib, as in Cobaeoideae. Johnson et al. (2008) suggest the following relationships within Polemonioideae - [[Polemonieae (one genus) + Phlocidae] [Gilieae + Loselieae]].

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

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, with micropylar and chalazal endosperm 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.

Chemistry, Morphology, etc. 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).

Phylogeny. For relationships within the family, see Schultheis and Baldwin (1999).

Previous Relationships. Fouquieriaceae were placed in Violales by Cronquist (1981).

See Kubitzki (2004b) for general information.

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

Trees (lianes); flavonols, ellagic acid +, kaempferol 0; (vessel elements with scalariform perforation plates); 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 sepals (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 petals, 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]

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), 2 or 4 ovules/carpel, (disc 0), style slender, relatively long, stigma punctate; ovules with endothelium; 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 two-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. Foetidioideae 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.

Evolution.

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 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 into 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: Note that monosymmetrtric flowers are not an apomorphy for Lecythidoideae.) Polysymmetric Lecythidoideae are pollinated by a variety of bees, other than euglossines. Napoleonaea vogelii pollination and floral morphology has been described in detail (Frame & Durou 2001). Despite the size of the flower, 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.

Chemistry, Morphology, etc. 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 Cariniana ianeirensis clade (Lecythidoideae) is described as having obliquely monosymmetric flowers (Mori et al. 2007). 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.

Phylogeny: [Napoleonaeoideae [Scytopetaloideae [Lecythidoideae [Planchonoideae + Foetidioideae]]]] - see Morton et al. (1998) and Mori et al. (2007). Crateranthus rbcL sequences have not been obtained, but it is placed with Napoleonaea in joint analyses. In Chemistry, Morphology, etc, including androecium, Asteranthos is similar to Napoleonaeaoideae (but cf. style, endosperm), yet its sequence data align it with Scytopetaloideae... 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.

Previous Relationships. Although a weakly supported relationship with Sapotaceae had been considered possible in earlier versions (8 and earlier) of this site, there is currently no evidence for particular sister group relationships between Lecythidaceae and other Ericalean clades, although they are clearly not sister to the Balsaminaceae clade (Schönenberger et al. 2005).

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: Napoleonaeaceae), Prance and Mori (2004: Lecythidaceae s. str.), and Scott Mori's The Lecythidaceae Pages (Lecythidoideae only).

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

Chemistry, Morphology, etc. Vessel elements 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 perforation plates; vessel-fiber pits bordered; nodes 1:1; petiole bundle arcuate; mucilage cells +; hairs unicellular; C ± campanulate, only basally connate, fairly small [petals 1³ 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.

Evolution. Pentapetalum trifasciculandricus is a fossil ca 91 million years old from New Jersey that is placed either with Theaceae or in the Pentaphylacaceae area depending on the analysis (Martínez-Millán et al. 2009).

Chemistry, Morphology, etc. The placenta is very well developed in Ficalhoa and many Ternstroemieae and Frezierieae. See Luna and Ochoterena (2004) and Martínez-Millán et al. (2009) for morphology.

Phylogeny. Luna and Ochoterena (2004) and Martínez-Millán et al. (2009) were unable to recover much in the way of strongly supported relationships in this area in morphological phylogenetic analyses; in some analyses in the latter paper Calophyllaceae were included in Theales, and adding morphology tended to reduce support measures, perhaps especially bootstrap support.

Classification. A.P.G. II (2003) 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, and so two families are recognised in A.P.G. III (2009).

Previous Relationships. See also Theaceae for another family that has been associated with Sladeniaceae and Pentaphylacaceae in the past.

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 [list]/3. S.E. Asia (Sladenia), tropical E. Africa (Ficalhoa) (map: see Verdcourt 1962; fossil Sladenia [blue], see Giraud et al. 1992).

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

Evolution. The wood of extant Sladenia is distinctive, and matches fossil wood from the ?Cretaceous-Albian/Cenomanian of northern Sudan remarkably closely (Giraud et al. 1992).

Chemistry, Morphology, etc. Sladenia is poorly known. 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 (especially Deng & Baas 1991). Li et al. (2003) have recently described a number of very distinctive embryological, etc., features for Sladenia, including monocot anther 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).

Phylogeny. Sladenia was sister to Pentaphylacaceae (Ternstroemiaceae) in rbcL studies (Savolainen et al. 2000b), albeit the DNA was rather degraded. Sladenia and Ficalhoa 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.

Previous Relationships. u>Sladenia has often been included in Theaceae, e.g. as Sladenioideae (see Takhtajan 1997).

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.

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 petals, 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.

Ternstroemieae + Frezierieae: ellagic acid +, iridoids 0; (pits vestured); pith often with diaphragms; sclereids +; stomata anomocytic; perulae 0; filaments to 2x longer than anthers, laterally variable in length, 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 petals, 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.).

3. Frezierieae

(Nodes 1:3, 3:3 [some Freziera]); sclereids usu. little branched; leaves scattered along shoot, two-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.]

• Pentaphylacaceae are shrubs or trees that often have two-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.

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

Chemistry, Morphology, etc. 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. When the shoot of Pentaphylax 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 Pentaphylacaceae. The pericylic sheath of Pentaphylax consists of fibers alternating with lignified parenchymatous cells (Beauvisage 1920). Overall, however, Pentaphylax is poorly known; details of the ovules are taken from Mauritzon (1936). Cleyera (Freziereae) 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. Freziera shows considerable variation in nodal anatomy, stomatal morphology, seed type and pollen surface (Weitzman 1987). Pentaphylacaceae such as Cleyera have quite long filaments. 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]. Ternstroemia itself, with its pseudoverticillate and usually entire leaves, can look rather unlike other Pentaphylacaceae.

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).

Previous Relationships. Theaceae often included Ternstroemia and relatives; Ternstroemioideae were a subfamily of Theaceae in Takhtajan (1997). However, 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. Note that the name Pentaphylacaceae is conserved... A position of Pentaphylacaceae in Ericales seems reasonable from the gross morphological point of view. 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, another member of Ericales - and found the two to be similar. 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.

Sapotaceae [Ebenaceae + Primulaceae]: ellagic acid 0.

Phylogeny. 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 scalariform perforation plates); 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 (two-ranked, opposite), conduplicate, margins entire (toothed), 2ndary veins often rather close, stipules +/0; (plants di- or monoecious), inflorescences cymose, fasciculate, pedicels not articulated; flowers (anisomerous), K ± connate at base, C 4-18 (variously lobed or divided), stamens = and opposite to 2x (-6x) C lobes, introrse to extrorse; tapetum multinucleate; pollen 3-6-colporate, infratectum ± granular, staminodes +; disc + (0); G 1[2-14(-30)], opposite sepals, hairs on the inside of the ovary, placentation axile to axile-basal, 1(-5) ascending ovules/carpel, (style short), stigma punctate or minutely lobed, dry; ovules apotropous; 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, (stipels +), 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).

Synonymy: Sarcospermataceae H. J. Lam

Sapotoideae + Chrysophylloideae: staminodes +, often ± petalline, opposite sepals (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.

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

3. Chrysophylloideae Luersson

Stipules 0; A high in the tube, (several stamens opposite each petal; staminodes outside/above the staminal whorl); endosperm copious, cotyledons foliaceous, radicle exserted.

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

• 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].

Evolution. In Sideroxylon (Sapotoideae) 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.). Dating in Chrysophylloideae suggests that the largely New Caledonian Niemeyera clade reached that island in the latter part of the Oligocene (Swenson et al. 2008c, cf. Ladiges & Cantril 2007).

Chemistry, Morphology, etc. There is banded apotracheal parenchyma in Sapotaceae (cf. Lecythidaceae!). Some species of Sarcosperma have paired stipels at the apex of the petiole, a rather unexpected character for a member of Ericales. 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. Floral variation is considerable and most characters are very homoplasious (e.g. Swenson et al. 2008a, b, c). Swenson and Anderberg (2005) suggest that the basic floral morphology of the family is K5, C5; A 5 + 5 staminodes, however, anisomery is scattered in Sapotaceae (Swenson et al. 2008c), indeed, the basic floral construction of Sapotaceae is guite labile. Swenson and Anderberg (2005) suggest that the staminodes common in Chrysophylloideae, but derived within the clade, are perhaps not immediately comparable with those of other members of the family; the former are outside the staminal whorl while the latter are in the same whorl as the stamens. Amyloid is also known from the seeds of Omphalocarpum, a clade that is close to sister to the rest of Chrysophylloideae (see Kooiman 1960).

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).

Phylogeny. Sarcosperma is sister to the rest of the family. Its seedhas a shiny testa, albeit not as thick as that of most other Sapotaceae, and also a conspicuous hilar scar; the genus was placed in Sideroxyleae by Pennington.

Within the rest of the family there are 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. See Smedmark et al. (2006) for general discussion of relationships and character evolution in Sapotoideae. Swenson et al. (2007a, 2008a) discuss generic limits in Australasian members of Chrysophylloideae (the whole lot are monophyletic); for relationships among the monophyletic group of ca 80 species of the Pouteria complex on New Caledonia, see Bartisch et al. (2005); Pouteria sensu Pennington is polyphyletic (Triono et al. 2007). See Swenson et al. (2007b) for Planchonella and Swenson et al. (2008c) for its sister group, the largely New Caledonian Niemeyera complex. 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) helped clarify things somewhat and molecular data are providing much further information. Clade limits in e.g. New Caledonian Sapotaceae are now becoming clear enough so that species can be described in appropriate genera (Swenson et al. 2008b, esp. c).

Classification. For a checklist and bibliography, see Govaerts et al. (2001).

Ebenaceae + Primulaceae: ?

EBENACEAE Gücke, nom. cons.   Back to Ericales

Trees, bark and roots black; petiole bundle arcuate; sclereids +; leaves two-ranked, margins entire, flat glands on lower surface; pedicels articulated; flowers imperfect, ?4-merous; K connate, C contorted, connate, stamens adnate to corolla, 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 perforation plates); (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, 40-70 µm across, psilate, disc?; G [4], inferior, ?integuments, etc., stigma clavate, hairy apically; seeds 1-2; endosperm very hard; cotyledons foliaceous; n = ?

1/8. Tropical South America (map: from Wallnöfer 2004b).

Synonymy: Lissocarpaceae Gilg, nom. cons.

2. Ebenoideae Thorne & Reveal

Saponins, C-30 oxidised triterpenes, naphthoquinone derivatives of 7-methyljugone and plumbagin, flavonols, leucodelphinidin, myricetin +, ellagic acid 0; (cork pericyclic); cambium storied; (nodes 1:3); SiO₂ bodies + [not in Diospyros]; 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, pollen 25.9±6.4 µm across, infratectum granular, pistillode +; carpellate flowers: staminodes + (0); G [2-8], opposite petals 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 [not in Diospyros]), exotesta fibriform or mucilaginous, cells cuboid to palisade, endotesta crystalliferous or not, walls thickened or not; n = 15.

3/540: 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 two-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 naphthoquinones 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.

Evolution. Duangjai et al. (2009) focus on Diospyros in New Caledonia, where there are four separate lineages.

Chemistry, Morphology, etc. Ellagic acid may occur in Ebenaceae-Ebenoideae (Bate Smith 1962). Since I do not know of details the chemistry of Lissocarpoideae, whether the presence of ellagic acid or naphthoquinones is a synapomorphy of Ebenaceae as a whole or just part of them remains to be established. 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. The morphology of the inflorescence of Lissocarpoideae is unclear; one interpretation is that the flowers are axillary, whether on short or long shoots. The flowers seem to be imperfect. In Diospyros s.l., 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.

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). Some information on Lissocarpa is taken from Schadel (1978: leaf morphology) and Wallnöfer (2004a, b), but that genus is poorly known.

Phylogeny. Duangjai et al. (2006a and especially b), sequencing six plastid genes, found extensive phylogenetic structure in Ebenoideae; 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. Geeraerts et al. (2009) suggest apomorphies - especially palynological - for these genera, while Duanjai et al. (2009: eight genes, 119 species) provides a more detailed phylogeny of Diospyros itself with good Bayesian support for relationships along the backbone of the tree.

Previous Relationships. 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.

Classification. For a monograph of Lissocarpa, see Wallnöfer (2004b).

PRIMULACEAE Borkhausen, nom. cons.   Back to Ericales

(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, stamens = and opposite C, antesepalous whorl represented by 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.

58/2590. World wide.

• Primulaceae can be recognised by their usually spirally arranged leaves sometimes with dark dots or lines. Their flowers are sympetalous with stamens borne opposite to the petals and an ovary with several to many ovules and free-central placentation; the stigma is not lobed.

Evolution. 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.

Plants 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 Maesa but not so far on members of Theophrastoideae (Ehrlich & Raven 1964).

Chemistry, Morphology, etc. Leaves of Theophrasteae and Myrsinoideae are often described as being involute (?supervolute, cf. Cullen 1978) or conduplicate. There are common stamen/corolla primordia born on a ring primordium in this clade, but there is variation in the position/relative development of these primordia. In some cases such as Cyclamen the 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 especially Ma & Saunders 2003), whereas in other taxa the stamen primordia may initially be larger, as in Samolus (e.g. Sattler 1962). However, this is a tricky character, since there are really two variables, the relative positions of these primordia and how fast they initially develop, and, as with evicted terminal inflorescences, initial topological relationships between parts can speedily become disrupted by post-initiation growth. The number of carpels can be also difficult to ascertain, but five seems to be a common number; although their orientation is often unclear, they might be expected to be opposite to the sepals. However, the diagrams presented by Dickson (1936) mostly suggest that the carpels are opposite the petals, but in Primula, at least, the carpels appear to be opposite the sepals.

For the hollow style, see Guéguen (1901: is the condition in Maesa known?), for staminodes, see Saunders (1936) and Caris and Smets (2004: those of Samolus and Theoprasteae 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). For general morphology, see Anderberg et al. (2000) and especially Ståhl and Anderberg (2004).

Phylogeny. 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 Theophrasteae are reduced when morphological data are added to molecular.

Classification. This whole group was often recognised as Primulales in the past. 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 the old Theophrastaceae, the many herbaceous ex-Primulaceae that are sister to the old-style, woody Myrsinaceae rather than being in a clade with other Primulaceae, and the numerous features shared by the group as a whole, is whether it is worth recognising families at all... A broader circumscrioption was proposed in A.P.G. III (2009), and available subfamilial and tribal names fit well with the phylogeny here.

Previous relationships. Plumbaginaceae (see Caryophyllales here) were often associated with Primulaceae and related families because of apparently similar placentation and oppositipetalous stamens in common (see Cronquist 1981 for discussion).

1. Maesoideae de Candolle   Back to Ericales

Evergreen lianes or trees; vessel element type?; petiole bundles all annular; secretory canals well developed; leaves spiral or two-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).

• Maesoideae 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.

Chemistry, Morphology, etc. 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).

Synonymy: Maesaceae Anderberg, B. Ståhl & Kallersjö

Theophrastoideae [Primuloideae + Myrsinoideae]: herbs[?]; rays ³5-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.

Evolution. For the suggestion that rosette herbs might 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 to woody taxa, and variation in habit is very extensive in this whole clade. Smith and Donoghue (2008) found that the rate of mollecular evolution in the herbaceous taxa they examined was much greater than in the woody taxa.

2. Theophrastoideae A. de Candolle   Back to Ericales

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

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

1. 2A. Samoleae Reichenbach

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.]

Chemistry, Morphology, etc. 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. Theophrastoideae) the ovules are not embedded in placental tissue (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. 2B. Theophrasteae Bartling

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]

• Theophrasteae 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 extrorse anthers form a cone in the center of the flower when it opens, but later they spread. The subepidermal fibers are visible in the dried leaf as fine striations - use a hand lens!

Chemistry, Morphology, etc. The subepidermal fibres may lack lignification. For reports of glandular dots on calyx and corolla, see Mabberley (1997). Floral primordia may initially be quite strongly monosymmetric, as in Deherania (Sattler 1962), even if the flower at anthesis is polysymmetric. For general morphology, etc., see Ståhl (2004) and in particular Caris and Smets (2004).

Phylogeny. Phylogenetic relationships suggested by Källersjö and Ståhl (2003) imply that some generic realignments are needed.

Synonymy: Theophrastaceae Link, nom. cons.

Primuloideae + Myrsinoideae: two ndhF deletions.

3. Primuloideae Kostelesky   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, Primula flower.]

• Primuloideae 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.

Evolution. Heterostyly is common, although it is unlikely to be an apomorphy for the subfamily; it is sometimes lost, as in those Primula with buzz pollination, the erstwhile Dodecatheon (Mast et al. 2001, 2006).

Chemistry, Morphology, etc. 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 lactea. The corolla epidermal cells are isodiametric. Saunders (1936) suggested that some of the lobing of the corolla of Soldanella might be staminodial.

For pollen variation, see Mast et al. (2001), and for general information, see Anderberg (2004).

Phylogeny. For ITS-based relationships within Primuloideae, 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).

Classification. Richards (2003) provides good general descriptions of the species of Primula s. str.

4. Myrsinoideae Burnett   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, many species in the Pacific), 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.

Evolution. 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 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 the bacteria might be playing (Miller 1990).

For the evolution of the mangrove habitat, to which Aegiceras is restricted, see Rhizophoraceae and Tomlinson (1986). 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, since seeds lack endosperm and contain a large, viviparous embryo (cf. Rhizophoraceae-Rhizophoreae, Acanthaceae-Acantheae-Acanthus ilicifolius, etc.).

Chemistry, Morphology, etc. The presence of coloured glands may well not be a synapomorphy of Myrsinoideae (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; nodal anatomy needs study. The epidermal cells of the corolla are often elongated; this is a derived feature within the family. Trientalis has anisomerous flowers (Swenson et al. 2008c). Cyclamen has one cotyledon and one integument. See Oh et al. (2008) for the seed morphology of herbaceous taxa around Lysimachia.

Coris is a particularly distinctive genus morphologically. It is a small ericoid sub-shrub with monosymmetric flowers that have a spine-tipped epicalyx. 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, but 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 or oblique). The rays of Aegiceras are relatively narrow, the ovules are unitegmic, etc. (Staåhl å Anderberg 2004; Lens et al. 2005).

Some information is taken from Otegui and Cocucci (1999) and from Ståhl and Anderberg (2004); Lens et al. (2005a) provide much information about wood anatomy.

Phylogeny. Myrsinaceae, previously circumscribed to include only woody taxa, were found to include Anagallis, Ardisiandra, Asterolinon (?= Lysimachia), Coris, Cyclamen, Glaux (= Lysimachia - it lacks a corolla), Lysimachia, Pelletiera, Stimpsonia and Trientalis (Anderberg et al. 2000, 2001), although the limits of the clade were not so clear in Martins et al. (2003: ITS data alone). Anderberg et al. (2007) was particularly interested in the relationships of the herbaceous taxa; the clade as a whole had moderate support as being monophyletic (72% jacknife), and Cyclamen, the herbaceous taxa, and the woody taxa then formed a trichotomy. Hao et al. 2004 also provide a phylogeny of much of the group, although focusing on Lysimachia.

Classification.Generic limits in the woody members in particular are unsatisfactory, but the limits of genera like Lysimachia are also unclear (Anderberg et al. 2007).

Synonymy: Aegicerataceae Blume, Anagallidaceae Adanson, Ardisiaceae Jussieu, Coridaceae J. Agardh, Embeliaceae J. Agardh, Lysimachiaceae Durande, Myrsinaceae R. Brown, nom. cons.

Mitrastemonaceae + Theaceae + Symplocaceae, etc. + Ericaceae, etc.: ?

MITRASTEMONACEAE Makino, nom. cons.   Back to Ericales

Endophytic root parasites; ?anatomy; leaf waxes hummocky; leaves opposite, scale-like; flowers single, terminal; P 4, connate, anthers 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.

Evolution. A mitochondrial gene has moved from the parasite to its host, Quercus (Systma et al. 2008).

Chemistry, Morphology, etc. Watanabe (1936: V) talks a lot about a "Mitrastemon-Pilz" (cf. ectomycorrhizae of Ericaceae?). 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).

For general information (including a more extensive list of hosts) and references, see Meijer and Veldkamp (1993), the Parasitic Plants website (Nickrent 1998 onwards) and also Heide-Jørgensen (2008).

Previous Relationships. Along with Cytinaceae and Rafflesiaceae, relationships with of Mitrastemonaceae to Malvales have also 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. Since its parietal placentation is found in many other parasitic angiosperms, as well as in the echlorophyllous hyperparasitic Ericaceae-Monotropoideae, this is not necessarily a taxonomically informative character. Cocucci and Cocucci (1996) suggested that Mitrastemonaceae had relationships with Annonaceae.

Theaceae + Symplocaceae, etc. + Ericaceae, etc.: cork?; vessel elements with scalariform perforation plates; leaves serrate.

THEACEAE Ker Gawler, nom. cons.   Back to Ericales

Trees or shrubs; plants Al-accumulators; myricetin, ellagic acid +; cork pericyclic (subepidermal); (pits vestured); intervessel pitting opposite-scalariform; pericyclic fibers +/0; petiole bundle arcuate; sclereids and mucilage cells common; stomata paracytic, anisocytic or cyclocytic; hairs unicellular; leaves also two-ranked, lamina involute or supervolute (conduplicate), margins toothed (entire); flowers single, axillary; C ± free; A usu. 40<, ± basally connate, centrifugal, from 5 primordia opposite petals or ring primordium in 2-5 whorls, laterally variable in length,anthers articulated, connective usu. not prolonged, pollen tricolporoidate, pseudopollen produced from connective, nectar from base of filaments or ovary; G [(3-)5(-10)], opposite petals, 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, 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.

1. Theeae Szyszylowicz

Pedicels multibracteolate, K and C intergrading, A in 2 whorls, embryo sac bisporic (the spores chalazal) and 8-celled [Allium-type], outer integument vascularized; 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], connective with stomata, inner integument vascularized; 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, embryology and pseudopollen?; capsule lacking columella; seeds narrowly winged or not, ?vascularization; n = 15, 17, 18.

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

Evolution. The function of the pseudopollen is unknown, but it does not appear to be nutritious (Tsou 1997; Iqbal & Wijesekara 2002).

Chemistry, Morphology, etc. The cotyledons of (?all) Theaceae have three or more traces from a single gap. The stomata are often described as being "gordoniaceous", i.e. cyclocytic to anisocytic. In some species of Camellia, at least, a cyclocytic arrangement of cells may be surrounded by anisocytic arrangement (Lu et al. 2008). 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).

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 (1997, 1998: embryology), Stevens et al. (2004b: general), Wang et al. (2006: Apterosperma, chromosomes and morphology) and Zhang et al. (2009: sclereids in Camellia); for pollen, see Wei (1997).

Phylogeny. Major relationships within the family are still poorly understood. 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. Above is a suggestion of how the family might be broken down.

For relationships within Camellia, see Vijayan et al. (2009); current sectional limits need overhauling.

Classification. Stewartia is to include Hartia (e.g. Prince 2002), and generic limits in other Theaceae are difficult; for useful notes on the genera, see Prince (2007).

Previous Relationships. 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.

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; true tracheids +; crystal sand +; stomata usu. paracytic, very large "water stomata" also present; (leaves two-ranked), lamina ± supervolute; (inflorescence branched), pedicels articulated (not Cordyloblaste); K basally connate; A (= and opposite sepals)-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 +, style hollow [?all], stigma ± capitate, wet or dry; 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.

2[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.

Evolution. Symplocos is locally very abundant as both pollen and fruits in the Tertiary fossil record of Europe, but is also known from the southern USA amnd East Asia (Krutzsch 1989).

There are subapical lobes on the style just below and alternating with commissural "stigmatic" lobes in the ca 145 species of the New World Symplocos subg. Symplocos sect. Symplocastrum; the papillae on these lower lobes are rich in lipid that may help the pollen stick to the pollinators (Kriebel et al. 2007). However, pollen germinates on the subapical lobes which are thus (and from their position) the true stigmatic lobes (Kelly & Nicholson 2009).

Chemistry, Morphology, etc. 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).

Phylogeny. For a phylogeny of Symplocos sensu lato, see Y. Wang et al. (2004) and Fritsch et al. (2006, 2008); section Cordyloblaste appears to be sister to the rest. The infrageneric taxonomy needs reworking, but it has been suggested that two genera in the family should be recognised (Fritsch et al. 2008).

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

Phylogeny. 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 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 small calyx and a more or less inferior ovary. The filaments are stout and not sharply distinguished from the anthers.

Evolution. For the early Tertiary fossil history of Styracaceae that are now East Asian endemic, see Manchester et al. (2009).

Van Steenis (1949b) illustrates the remarkable galls found on Malesian species of Styrax. There is a rather close association between the aphids involved (Cerataphidinae) that cause some of these galls and individual species of Styrax; the morphology of the galls is ultimately determined by the aphids (Cerataphidinae also produce soldiers - Stern 1995; Stern & Foster 1996). Cecidomyids also produce galls on Styrax.

Chemistry, Morphology, etc. The floral vasculature suggests that although the stamens are in a single whorl, they are basically obdiplostemonous (Dickison 1993). Van Tieghem (1898) shows Halesia as having two ascending epitropous ovules and two descending apotropous ovules. Pterostyrax (and Styrax?) lack endothelium. There are no septal bundles, as in many Ericales (but details of the distribution of this character?).

Phylogeny. For relationships within Styracaceae, see Fritsch et al. (2001) and within Styrax Fritsch (2001). The main phylogenetic structure in the family is [[Huodendron + Styrax] [[Alniphyllum + Bruinsmia] The Rest]]; both main clades, especially the second, are well supported. Members of the former clade have entire leaf blades, members of the latter have dentate blades, an inferior ovary, and bud scales, with the exception of the Alniphyllum + Bruinsmia clade which differs from The Rest on all three counts. For possible additional synapomorphies for Styracaceae, see Fritsch et al. (2001).

For general information, see Fritsch (2004).

Synonymy: Halesiaceae D. Don

DIAPENSIACEAE Lindley, nom. cons.   Back to Ericales

Shrublets or herbs, mycorrhizae ecto- and endotrophic [?ectendomycorrhiza]; plants Al-accumulators; ellagic acid +, iridoids?; (cork superficial); vessel elements with simple (scalariform) perforations; secondary wood rays 0; pericyclic fibers 0 (+ - Shortia); nodes (1:1 - Pyxidanthera) - 3:3; petiole bundle(s) arcuate to annular (medullary bundles +); (stomata anisocytic); leaf margins toothed or entire, 2ndary veins subpinnate to palmate; (flowers axillary); K free or connate, C serrate or not, forming tube with flattened filaments, stamens = and opposite sepals (connate - Galax), anthers ± incurved, thecae horizontal, staminodes + (0); disc 0; G [3], median member adaxial, endothelium 0, stigma shortly 3-lobed, wet; exotesta with inner walls thickened; endosperm copious, embryo terete; n = 6.

6[list]/18. Arctic and N. temperate, esp. East Asia and E. U.S.A. (map: from Diels 1914; Wood & Channel 1959; Hultén 1971).[Photo - Diapensia Flower © J. Maunder] [Photo - Diapensia Fruit © J. Maunder]

• Diapensiaceae are herbs or subshrubs with moderately-sized and weakly sympetalous flowers; the five stamens with their flattened filaments alternate with the petals and the anthers are incurved. There are usually staminodes, the stigma is shortly three-lobed, and the fruit is a loculicidal capsule with small seeds.

Evolution. Friis (1985) described Actinocalyx from the Upper Cretaceous of Sweden. It has a number of similarities with extant Diapensiaceae, although the anthers are rather different, the pollen is smaller (7-9.5 µm, versus 17-40 µm), and the styles are separate.

Chemistry, Morphology, etc. The mycorrhizal association in Diapensiaceae may be a distinctive ectendomycorrhiza as is found in many Ericaceae (see Asai 1934, although at that time the distinctiveness of the ericaceous mycorrhizal association was not fully understood). The integument is 5-7 cell layers thick, and seems to consist of an outer and inner part in some taxa. An endothelium may not always develop (Samuelsson 1913; Diels 1914; Kapil & Tiwari 1978). Schnizlein (1843-1870: fam. 160) shows Galax with the median G abaxial.

Phylogeny. Galax and Pyxidanthera are successively sister taxa to the rest of the family (Rönblom & Anderberg 2002); if this series of relationships hold, the presence of staminodes is a derived feature within the family.

Previous relationships. Diapensiaceae have often been considered close to Ericaceae, but the anthers of some genera of the former which appear to be inverted, are not.

For general information, see Scott (2004), for pollen and a morphological phylogeny, see Xi and Tang (1990).

Synonymy: Galacaceae D. Don

[Sarraceniaceae [Actinidiaceae + Roridulaceae]] [Clethraceae [Cyrillaceae + Ericaceae]]: inflorescence racemose; anthers extrorse, inverting during development, opening by pores or short slits, pollen ± rugulate ["cerebellar"], tectum and foot layer solid, infratectum with granular elements; G [3], median member adaxial, also [5], opposite petals, many ovules/carpel, endothelium +, style impressed; fruit a capsule; testa with much thickened inner wall [?higher level], endosperm copious; mitochondrial coxII.i3 intron 0.

Evolution. Wikström et al. (2001) suggest a stem group age of 74-71 million years before present, with crown group divergence beginning 67-59 million years before present, however, relationships between members of the group are other than those shown here, and Roridulaceae are sister to all other Ericales...

Chemistry, Morphology, etc. There are pit membrane remnants in the perforations of vessels in several families of this clade (Schneider & Carlquist 2003, 2004; Carlquist & Schneider 2005). For a summary of pollen variation, see Zhang and Anderberg (2002).

Phylogeny. The whole clade may be sister to ((Theaceae s. str. + Symplocacaeae) (Styracaceae + Diapensiaceae)) (Geuten et al. 2004). Bracteole presence is rather variable (Anderberg & Xiaoping (2002).

Sarraceniaceae [Actinidiaceae + Roridulaceae]: route I secoiridoids +; nectary 0, hypostase +, stigma dry.

SARRACENIACEAE Dumortier, nom. cons.   Back to Ericales

Insectivorous acaulescent rosette herbs; O-methyl flavonols only +; cork?; vascular bundles separate; nodes ?; leaves with broad bases, ascidiate; (flowers solitary), bracteoles + [Heliamphora]; K (3-6), ± petaloid, C (0 - Heliamphora, 4), free; A 8-10 [Heliamphora] or many, centrifugal, anthers with slits, pollen 5+ colporoidate, verrucose-vermiculate, with small surface granules, (nectar at base of style - Sarracenia), integument ca 4 cells across, style hollow [?always], (peltate; apical styles), stigmas small; seeds small, with wings or hairs, exotesta ± thickened; endosperm haustoria?, embryo medium; n = 13, 15, 21.

3[list]/ca 15. E. and W. U.S.A. and the Guayana Highlands (map: from Uphof 1931; Schnell 2002).

• Sarraceniaceae are herbaceous, insectivorous pitcher plants. The pitchers are long, with short petioles, and are borne in a rosette. The flowers are quite large, with free sepals and petals, the former being petaloid, and there are usually numerous stamens. The capsules have many small seeds. [Photo - Inflorescence © J. Maunder, Flower, Flower.]

Evolution. Archaeamphora (Sarraceniaceae) has been described from rocks ca 124 million years old (Li 2005) - this is perhaps an unlikely identification.

There are nectar glands on the pitcher which attract insects that fall into the pitcher and drown - alternatively, the nectar may take up water increasing the possibility of an insect's hydroplaning into the pitcher (see Bauer et al. 2008); the colouring on the flap of the pitcher may also attract insects; it is a kind of pseudoflower (Cresswell 1993). The pitcher varies in the amount of digestive enzymes it contains. In Sarracenia purpurea, for example, there are few enzymes and nutrients from the entrapped animals are made available to the plant by the activity of detritivores that break up the prey that is further decomposed by bacteria, in turn eaten by rotifers and protozoa and ultimately by mosquito larvae - all forming a microcosm in the liquid of each pitcher (Kitching 2000; Ellison et al. 2003; Butler & Ellison 2007). For general information on carnivory, see especially Lloyd (1942) and Juniper et al. (1989).

Caterpillars of the moth Exyra fax drain the pitchers by opening up a hole at the base; they then eat the pitcher; species of Exyra occur throughout the range of Sarracenia. Flowers of Heliamphora lack a nectary and are buzz pollinated; Sarracenia has ten nectaries on the ovary wall above the stamen fascicles.

Chemistry, Morphology, etc. Jensen (1992) suggests the family has route I iridioids.In Sarracenia, at least, the leaves have an adaxial flange, but the pitcher develops from the midib area. When there are many stamens, development is centrifugal from initially 10 primordia. The ovules may be unitegmic or bitegmic; the integuments are thin. Is the style hollow?

For general information, see Kubitzki (2004b), McPherson (2006) and the Carnivorous Plants Database, for perforation plates, see Schneider and Carlquist (2004). I thank D. Hoekman for information.

Phylogeny. R. J. Bayer et al. (1996) and Neyland and Merchant (2006) provide more information about relationships within the family; the topology [Darlingtonia [Sarracenia + Heliamphora]] seems well supported.

Synonymy: Heliamphoraceae Chrtek, Slavikova & Studicka

Actinidiaceae + Roridulaceae: ?

ACTINIDIACEAE Gilg & Werdermann, nom. cons.   Back to Ericales

Trees, shrubs or lianes; (vessel elements with simple perforation plates); (nodes 3:3); petiole bundle deeply arcuate with wing bundles [Actinidia] or annular (medullary bundles +); raphide sacs +; hairs often ± (flattened) setose; leaves (opposite), conduplicate, apex of tooth expanded, clear, not deciduous, (2ndary veins subpalmate); plant usu. dioecious; C basally connate or not, (nectar at base of petals), A 10-many, centrifugal, (in groups opposite petals), inflexed in bud (not - Saurauia; ± connate), inverting rather late, porose, or ± with short slits, rugulate and transversely striate; G [(-20)], (10< ovules/carpel, style +, or styles separate (grooved [Actinidia]; stigma peltate, lobed); fruit usu. a berry; seeds embedded in placental pulp; endosperm haustoria?; n = 20, 29, 30; horizontal transfer of mitochondrial rps2 gene [Actinidia].

3[list]/355: Saurauia (300), Actinidia (30), Clematoclethra (25). Largely tropical, esp. South East Asia to Malesia, but not Africa (map: from Soejarto 1980).

• Actinidiaceae are usually shrubs or lianes with spiral, serrate, exstipulate leaves and often rather flattened setose hairs. The flowers have apparently free sepals and petals and many stamens that invert just before anthesis; the anthers dehisce by a slit that is much broader towards the apex. The styles are short and separate and the fruit is a berry. [Photo - Flowers, Collection.]

Evolution. Parasuarauia was described from flowers of Campanian (Late Cretaceous) age from the eastern USA. It has impressed styles, and both separate styles and numerous stamens are probably derived within the family (Keller et al. 1996; Herendeen et al. 1999); Parasuarauia itself is probably belongs to crown group Actinidiaceae.

Chemistry, Morphology, etc. Anthers of staminodes in Saurauia contain sterile pollen; in general, dioecy in the family is cryptic. For androecium development in Actinidia, see van Heel (1987), the synascidiate carpels are in a single whorl, and there is a large, flat, residual floral axis. There is paternal transmission of the plastid genome in Actinidia at least (Chat et al. 2003).

For information on the floral anatomy of Actinidia, see Schmid (1978), for general information, see Dressler and Bayer (2004).

Synonymy: Saurauiaceae Grisebach, nom. cons.

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

Shrub; unspecified iridoids +, ellagic acid?; cork?; pericyclic fibers 0; hairs numerous, glandular; leaves curved, sessile, linear, margins entire or with narrow lobes; inflorescence with a terminal flower [?always; sometimes looking racemose]; C free, quincuncial, stamens = and opposite sepals, connective swollen at base of anther, conspicuous and nectariferous, anthers early inverting, fibrous endothecium 0, pollen densely and minutely spinose, placentation apical, 1-4 ovules/carpel, stigma capitate, ?surface; testa also mucilaginous; micropylar haustorium +; n = 6.

1[list]/2. Southern Africa.

• Roridulaceae are small shrubs with linear leaves that have resin-secreting capitate hairs especially along the margins and abaxial midrib; the leaves are neither circinate in bud nor are the hairs sensitive (cf. Droseraceae, Drosophyllaceae, and Byblidaceae). The flowers are medium-sized with free sepals and petals, only five stamens that invert early in development and that have a visually attractive, yellow, apparently nectariferous basal portion, and three carpels with an undivided style. [Photo - Roridula Flower © M. Schmidt].

Evolution. The age of Roridulaceae is ca 90 million years, suggesting that they are very much a relictual element in the Cape flora (Warren & Hawkins 2006).

Although the family may not be carnivorous in a conventional sense, digestive enzymes not having been recorded from it (e.g. Hartmeyer 1997), its two species live in very close mutualistic association with two species of the hemipteran, Pameridea. These eat the insects that get stuck to the plant, and the plant absorbs nutrients from their excreta (Ellis & Midgley 1996; Anderson 2005). However, Plachno et al. (2009) suggest that Roridula is fully carnivorous, recording mineral uptake from Drosophila stuck on the leaves.

Pameridea is also involved in pollinating Roridula (Ellis & Midgley 1996; Anderson 2005).

Chemistry, Morphology, etc. Whether or not the roots are mycorrhizal is disputed (Conran 2004 for literature). There are bracteoles.

For general information, see Vani-Hardev (1972), Dahlgren and van Wyk (1988), Wilkinson (1998), Conran (2004), McPherson (2008) and the Carnivorous Plants Database.

Previous Relationships. Roridula was included in Byblidaceae by Cronquist (1981); for further information on relationships, see that family (Lamiales!).

Clethraceae [Cyrillaceae + Ericaceae]: ellagic acid +; cork pericyclic; pericyclic fibers absent; leaves spiral; bracteoles 0; flowers pendulous, stamens = 2x K, nectariferous basal part of ovary wall +, style hollow; endosperm with micropylar and chalazal haustoria; embryo terete.

Chemistry, Morphology, etc. It is possible that the accumulation of sugars as ketose and isokestose oligosaccharides is of systematic significance. Bracteoles have to be regained somewhere in this clade, but I have not worked out where.

CLETHRACEAE Klotzsch, nom. cons.   Back to Ericales

Mycorrhiza as modified ectendomycorrhiza?; sugars accumulated as kestose and isokestose oligosaccharides, iridoids?; (pits vestured); petiole bundle arcuate or annular with medullary bundle; stomata also paracytic and actinocytic; hairs stellate [sect. Clethra]; leaves conduplicate-subplicate, margins toothed (entire); inflorescences terminal, branched or not, (bracts conspicuous - Purdiaea); K quincuncial, C basally connate or free; A ?obdiplostemonous, adnate to C or not, inverting late, anthers ± sagittate, with pores or short slits, pollen <20µm, oblate, psilate to rugulate, (nectary 0); G ?orientation, (one apical straight ovule/carpel - Purdiaea), stigma lobed or not; K persistent; seeds winged or not, (or fruit dry, indehiscent, testa undistinguished, ± disappearing - Purdiaea); endosperm hemicellulosic; n = 8.

2[list]/75: Clethra (65). E. Asia to Malesia, S.E. U.S.A. (sect. Clethra), Mexico southwards, Cuba, 1 sp. on Madeira (Clethra sect. Cuellaria); largely tropical montane or subtropical (map: from Sleumer 1971d; Good 1974; Heywood 1978).

• Clethraceae are woody plants that can be recognised by their medium-sized, spiral, often toothed leaves that tend to be clustered at the ends of branches, their racemose inflorescences, and their rather small flowers with quincuncial sepals, free or basally connate petals, and ten late-inverting stamens with distinctive often sagittate and porose anthers. [Photo - Fruits & Flowers © A. Gentry], [Photo - Inflorescence], [Photo - Purdiaea Inflorescence].

Chemistry, Morphology, etc. Iridoids were described as being absent but scored as being present in Hufford (1992); they are not mentioned by Schneider and Bayer (2004). In Clethra, there is a prominent endodermis in the stem and the pith tends to be heterogeneous.

Some information is taken from Sai 1934 (mycorrhiza), Thomas (1960: Purdiaea), Sleumer (1967: Clethra) and especially Anderberg and Zhang (2002: pollen) and Schneider and Bayer (2004: general).

Phylogeny. See Fior et al. (2003) for a phylogeny of Clethra; they suggest that the Macaronesian C. arborea may be sister to the E. North American C. alnifolia.

Cyrillaceae + Ericaceae: myricetin +; colleters +; C connate, stigma wet.

CYRILLACEAE Endlicher, nom. cons.   Back to Ericales

Iridoids?, sieve tube plastids with protein crystalloids and fibers; petiole bundle annular, complex or deeply concave; leaf supervolute, margins entire, petiole obscure; (flowers 6-7-merous), K connate basally, C connate basally; A diplostemonous, or = and opposite sepals [Cyrilla], anthers ellipsoid, introrse, not inverting, dehiscing by slits, pollen >16µm, spherical, smooth; G [2-5], 1-3 apical mostly apotropous ovules/carpel, style short, stigma lobed; fruit a dry 1-4-seeded drupe or one-seeded, 2-5-winged samara; testa undistinguished, ± disappearing; endosperm moderate; n = 20.

2[list]/2. S. U.S.A. to N. South America (map: from Thomas 1960).

• Cyrillaceae are trees or shrubs that can be recognised by their spirally-arranged entire leaves with short petioles and often prominent, black colleters, and their long-racemose inflorescences with rather small flowers. The flowers appear to be polypetalous, the style is short, and the stigma is often three-lobed. [Photo - Cliftonia Habit].

Chemistry, Morphology, etc. Goldberg (1986) notes the presence of small, scarious stipules; I have not seen them. However, some taxa have slight excavations in the leaf base enclosing the buds and the colleters may be sublateral, hence suggesting the presence of stipules; Thomas (1960, p. 15) described them as being bright red, ligulate and glandular, although it is not clear to what genera he was referring. Goldberg (1986) also shows a floral diagram in which the median K is abaxial. The sepals are small and do not overlap, except perhaps very early in development. Anderberg and Zhang (2002: see also Copeland 1953) draw the anthers as being introrse and also suggest that the stamens do not invert during development. Cyrilla is described as having its five stamens opposite the petals by Thomas (1960). There are stomata in the nectary, but apparently not in nectaries in Ericaceae (Brown 1938). The integument is 4-5 cells across.

Previous Relationships. The old separation between Clethraceae and Cyrillaceae was based on fruit type (dehiscent versus indehiscent fruits), the new limits correlate better with general floral morphology. Clethra is sometimes included here (e.g. Mabberley 1997).

For more information, see Copeland (1953: general floral morphology and anatomy), Thomas (1960: monograph, 1961), and Vijayaraghavan (1970: ovule morphology, etc.), also Anderberg (1993), Zhang and Anderberg (2002: pollen), Anderberg and Zhang (2002: general) and Kubitzki (2004b: general).

ERICACEAE Jussieu, nom. cons.   Back to Ericales

Woody; benzo- and naphthoquinones, route I secoiridoids +, ellagic acid 0; (vessel elements with simple perforation plates); pericyclic fibers poorly developed; petiole bundle arcuate; leaves involute, margins entire to toothed, teeth associated with multicellular hairs; inflorescence terminal; K connate basally, C connate; A obdiplostemonous, inverting late, tapetal cells uni-, bi- or multinucleate, pollen >26µm, surface ± rugulate; G [5], stigma expanded; K persistent; testa with outer wall unthickened; chloroplast infA gene defunct.

Ca 126[list]/3995 - eight groups below. World-wide, but rare in lowland tropics (map: N. part of range, see Hultén 1971; Meusel et al. 1978; Luteyn 1995).

1. Enkianthoideae Kron, Judd & Anderberg

Pith with small, thick-walled and lignified and larger and thin-walled cells mixed [heterogeneous]; leaves pseudoverticillate; anthers with paired awns, pollen grains trinucleate, surface ± graulate; megagametophyte with "ears"; n = 11.

1/16. South East Asia: China, Japan and environs (map: from Kron & Luteyn 1995). [Photo - Habit.]

Monotropoideae + Arbutoideae [[Cassiopoideae + Ericoideae] [Harrimanelloideae [Styphelioideae + Vaccinioideae]]]: ectendomycorrhizal, hair roots present [consisting of endodermis, exodermis, tracheid, sieve tube + companion cell], invested by hyphae [Hartig net, usually formed by basidiomycetes], fungal hyphae with complex coiled intrusions into the exodermal cells of hair roots [pegs only in Monotropeae, Pterosporeae], clade A Sebacinales involved; anthers with exothecium; no chalazal vascular bundle in seed.

2. Monotropoideae Arnott

Herbs, some echlorophyllous, mycoheterotrophic and hyperparasitic; sugars accumulated as kestose and isokestose oligosaccharides ; multicellular hairs 0 [+ - Pterosporeae]; inflorescence a raceme (bracteoles + - Monotropeae, Pterosporeae); C often free, anther dehiscence various, pollen usu. in tetrahedral tetrads [Pyroleae]; testa variously winged or not, walls thickened or not; embryo short to minute and undifferentiated; n = 8, 13, 19, 23, etc.; protein crystals in nuclei.

15/50. N. hemisphere, largely temperate (map: from Meusel et al. 1978; Hultén & Fries 1986; Kron & Luteyn 2005; the distribution in E. Asia is rather unclear). [Photo - Chimaphila Flower, Monotropa Habit, Pterospora Habit, Flower, Pyrola Flower.]

Sieve tube plastids in at least some of the echlorophyllous members lack both starch and protein inclusions. Monotropa uniflora has a two-celled embryo (Olson 1991); much smaller than this you cannot get.

For variation in proteinaceous nuclear inclusions, see Speta (1979), for the phylogeny of Pyrola and its relatives, see Freudenstein (1999).

Synonymy: Hypopityaceae Link, Monotropaceae Nuttall, nom. cons., Pyrolaceae Lindley, nom. cons.

Arbutoideae [[Cassiopoideae + Ericoideae] [Harrimanelloideae [Styphelioideae + Vaccinioideae]]]: (petiole bundle annular); bracteoles + [but see Monotropoideae], anthers without endothecium, porose, pollen in tetrahedral tetrads.

3. Arbutoideae Niedenzu

Ellagic acid, C-8 iridoid glucosides +; corolla urceolate; anthers with paired awns, 10³ ovules/carpel, style continuous; fruit berry or drupe; testa cells rather thick-walled; n = 13.

1-6/ca 80: Arctostaphylos (60). Warm (cold) temperate, esp. S.W. North America, Mediterranean (map: from Meusel et al. 1978; Hultén & Fries 1986; Kron & Luteyn 2005).[Photo - Flower (x-sec)] [Photo - Inflorescence]

Phylogeny. Arbutus appears to be paraphyletic with respect to the other genera (Hileman et al. 2001; see also Kron et al. 2002b), but broader sampling with the ITS gene yields a topology compatible with conventional delimitations of genera, in particular, Arctuos is not sister to Arctostaphylos (Greg Wahlert, pers. comm.).

Synonymy: Arbutaceae Bromhead, Arctostaphylaceae J. Agardh

[[Cassiopoideae + Ericoideae] [Harrimanelloideae [Styphelioideae + Vaccinioideae]]]: (toxic andromedane diterpenes +); no Hartig net, mycorrhizae-forming fungi commonly ascomycetes; cauline pericyclic fibers well-developed, fibers associated with leaf bundles; stamens early inverting.

Cassiopoideae + Ericoideae: leaves opposite, revolute [ericoid].

Chemistry, Morphology, etc. There is a variety of morphologies subsumed by the term "ericoid leaf".

4. Cassiopoideae Kron & Judd

Pith with large thin walled cells surrounded by smaller thick-walled and lignified cells [Calluna-type]; bud scales 0; inflorescences axillary; anthers with awns, embryo sac bisporic.

1/12. Circumboreal (map: from Meusel et al. 1978; Hultén & Fries 1986; Kron & Luteyn 2005). [Photo - Habit.]

The leaf midrib may lack associated fibers (Kron et al. 2002b), but this may be a plesiomorphy here and then apomorphic at higher levels.

5. Ericoideae Link

Leaves also spiral, flat (convolute); (pedicel articulated); flowers held erect to ± pendulous, (C free), anthers lacking appendages (present - Ericeae), (pollen with viscin threads); capsule septicidal.

19/1790: Rhododendron (850: inc. Azalea, Ledum, Menziesia, Tsusiophyllum), Erica (765+, 600+ in the S. Cape region: inc. Philippia, etc., see Oliver 2000). Most diverse in South Africa and Malesia to S.E. Asia, also general N. hemisphere to S. South America, inc. Tristan da Cuhna and Falkland Islands (map: from Meusel et al. 1978; Hultén & Fries 1986; Kron & Luteyn 2005). [Photo - Flower, Habit, Flower, Empetrum Fruit © J. Maunder.]

Synonymy: Azaleaceae Vest, Diplarchaceae Klotzsch, Empetraceae Berchtold & J. Presl, Ledaceae Gmelin, Menziesiaceae Klotzsch, Rhododendraceae Jussieu, Rhodoraceae Ventenat, Salaxidaceae J. Agardh

Harrimanelloideae [Styphelioideae + Vaccinioideae]: K in fruit persisting, not withering.

6. Harrimanelloideae Kron & Judd

Leaves acicular; flowers single, ebracteolate; anthers porose, with spurs.

1/2. Interruptedly circumboreal (map: from Hultén & Fries 1986; Kron & Luteyn 2005).

Styphelioideae + Vaccinioideae: ?

7. Styphelioideae Sweet

Epidermis lignified; leaf bundles with well-developed abaxial fibrous vascular sheaths, no adaxial cap; multicellular hairs and leaf serrations 0 (+ - Prionoteae); leaves xeromorphic, pungent, midrib not evident (it is in Prionoteae), veins parallel or palmate; inflorescences often axillary, usu. spikes or multibracteolate axillary flowers; A = and opposite sepals, epipetalous (not), anthers bisporangiate/monothecal, with 1 (2) slits, appendages 0, (pollen in pseudomonads), (ovules apotropous - Lysinema); C persistent in fruit.

35/545: Leucopogon (230, but perhaps to be split), Dracophyllum (50), Epacris (45). Australasia, Chile (Lebetanthus) (map: Sleumer 1964; Kron & Luteyn 2005; FloraBase 2006). [Photo - Flower, Fruit & Flowers.]

For wood anatomy, see Lens et al. (2003).

Synonymy: Epacridaceae R. Brown, nom. cons., Prionotaceae Hutchinson, Stypheliaceae Horaninow

8. Vaccinioideae Arnott

(Hyphal sheath with inter/intracellular hyphae in cortex, basidiomycetes involved); apical bud aborting; stomata often paracytic; inflorescence usually axillary; pedicel often articulated; anthers with spurs, 2 or 4 awns or appendages 0, stigma truncate; n = 12.

Ca 50/1580. Inferior-ovaried taxa: Vaccinium (450), but in Southeast Asia Agapetes (ca 400, inc many Vaccinium), Dimorphanthera (ca 85) sister to Paphia (18), used to be part of Agapetes, but not immediately related to it, while in tropical America Cavendishia (155), Psammisia (60), Thibaudia (60), Macleania (55), and Gaylussacia (50). The rest: Gaultheria (235, inc. Pernettya, Diplycosia, Tepuia). N. hemisphere, Malesia and montane Central and South America, Australia (Queensland), few in Africa (map: from Meusel et al. 1978; Hultén & Fries 1986; Kron & Luteyn 2005, still incomplete). [Photo - Psammisia Flowers, Gaultheria Flowers, Vaccinium Flower © J. Maunder.]

Chemistry, Morphology, etc. A group of genera around Lyonia has a lignified epidermis, bands of fibers in the secondary phloem, anomocytic stomata, etc.

For general information about New World taxa, see Luteyn (2000, 2002), for wood anatomy of superior-ovaried taxa, see Lens et al. (2004a) and including many inferior-ovaried taxa, see Lens et al. (2004b: ecological correlations are mainly with latitude, also life form and precipitation).

Synonymy: Andromedaceae Schnizlein, Oxycoccaceae A. Kerner, Vacciniaceae Perleb, nom. cons.

• Ericaceae are usually woody and shrubby and may be recognized by their flowers, which are often urceolate or tubular and are sympetalous; the ten stamens often dehisce by pores and sometimes have appendages on the anthers or the tops of the filaments. Vegetatively the family is very variable, but the leaves are always exstipulate and often spiral and rather mesomorphic, or they are xeromorphic, in that case being either opposite, ericoid, narrow and with a channel on the abaxial surface, or variable in size, thick, and with close parallel venation.

Evolution. For Paleoenkianthus, an interesting Late Cretaceous fossil from some 90 million years before present, see Nixon and Crepet (1993), for diverse but much later (early Pleistocene) Stypheliodeae fossils, see Joran et al. (2007).

Kron and Luteyn (2005) discuss the historical biogeography of Ericaceae and give useful distribution maps for the subfamilies (but Cassiopoideae are all over Greenland...); an Eurasian origin of the family is likely. Outlines of interesting biogeographical groupings in tropical Vaccinieae are developing (Kron et al. 2002a) and these for the most part cut across the limits of the larger genera in particular (see also Powell & Kron 2003; Pedraza-Peñalosa 2009), but may be correlated with variation in wood anatomy (Lems et al. 2004c).

Ericaceae are most commonly found in open, more or less acidic habitats in cold to warm temperate climates, being most diverse in Mediterranean climates (e.g. the 600 or more spp. of Erica in the Cape region of South Africa alone) and in the mountains of South East Asia-Malesia and tropical America. There are four (partly overlapping) main ecological lines of diversification in the family: taxa with fleshy fruits of one sort or another, ca 1500 spp., taxa with xeromorphic leaves living in more or less Mediterranean or dry habitats, 1300+ spp., taxa that are epiphytic (or epilithic), ca 400 spp., and taxa with viscin threads, ca 900 spp. Many taxa in Mediterranean climates in particular (the South African Cape - Ericoideae, Australia - Styphelioideae) form starch-rich lignotubers that allow the plants to resprout after fires, or they regenerate by seeding; germination is enhanced by heat and/or smoke (Bell & Ojeda 1999; Cairney & Ashford 2002 - cf. Restionaceae). Fleshy-fruited taxa predominate in Vaccinieae, and in both the Indo-Malesian and the Andean regions they may be epiphytes. Some Vaccinieae in both the Old and New Worlds have seeds with a mucilaginous testa and a green embryo; plants with such seeds are generally epiphytic or epilithic. Finally, bird pollination is particularly common in the Andean Vaccinieae (Stiles 1981 and references) and may also occur in the large-flowered Indo-Malesian members of the tribe.

Endophytes are common (Petrini 1988); Ngugi and Scherm (2006) discuss the pseudoflowers formed by some fungal associates of Vaccinium.

Ericaceae are particularly noted for their distictive ectendomycorrhizae; organic nitrogen and phosphorus is taken up by the mycrorhizal fungus associated with the plant, perhaps an element in the success of the family in the often rather acidic and nitrogen-poor habitats in which many of the family grow. This nitrogen and phosphorus then moves to the ericaceous associate (Cairney & Ashford 2002 and references). For ericoid mycorrhizae, in which the fungi are usually ascomycetes, see Cullings (1996) and Smith and Read (1997); some of these fungi also form mycorrhizal associations with Jungermanniales, leafy liverworts (Presel et al. 2008). Massicotte et al. (2008 and references) discuss mycorrhizal associations in North American Pyroleae. Bougoure et al. (2007) detail the diversity of fungal associated with Vaccinium and Calluna - some may be ectomycorrhizal. Selosse et al. (2007) describe the association of Sebacinales, a "basal" order of basidiomycetes with Ericaceae; they find group A to be largely associated with Arbutoideae and Monotropoideae, while group B are found associated with members of Ericoideae Styphelioideea and Vaccinioideae (Harrimanellloideae and Cassiopoideae are unknown) (Selosse & Weiß 2009; Weiß et al. 2009), while basidiomycetes may be proportionally particularly common in Vaccinoiodeae (Bougoure et al. 2007). Recent work shows that several clades of Sebacinales are involved in the distinctive mycorrhizae formed on the hemiepiphytic Cavendishia nobilis (Setaro et al. 2006), and this association has recently been described as a new mycorrhizal type, the cavendishioid type (Setaro et al. 2008); how widespread it is in Ericaceae is unclear. For the mycorrhizae of Styphelioideae in particular, see Cairney and Ashford (2005). Interestingly, Enkianthus has arbuscular mycorrhizae of the Paris type (Abe 2005), i.e. it is endomycorrhizal, and it apparently lacks hair roots; I have seen few accounts of mycorrhizae in Clethraceae or Cyrillaceae.

The association of particular basidiomycete fungi with individual species of echlorophyllous and mycoheterotrophic Monotropoideae is often close and specific (Bidartondo and Bruns 2001, 2002; Bidartondo 2005: note that problems with the identity of Monotropeae studied make interpretation of the earlier literature difficult), and both carbon and nitrogen move from the fungal associate to chorophyllous and echlorophyllous Monotropoideae alike (Zimmer et al. 2007; Tedersoo et al. 2007). The mycoheterotrophic habit (and probable hyperparasitism) has arisen at least twice in Monotropoideae; Pyrola aphylla, more closely related to other heterotrophic Monotropoideae-Pyroleae than to mycoheterotrophic Monotropoideae (= [Monotropeae + Pterosporeae]), is also a mycoheterotroph at times (Zimmer et al. 2007; Hynson et al. 2009b), although it may have small green leaves and the fungi associated with it show no particular specificity, as in other more photosynthetically conventional species of Pyrola (Hynson & Bruns 2009). Monotropoideae are mixotrophic or fully heterotrophic (Hynson et al. 2009b; Selosse & Weiß 2009).

Host preferences of the rust fungi Chrysomyxa and Exobasidium link Empetreae with other Ericaceae, perhaps Ledum with Rhododendron, etc.; Exobasidium is also found on Theaceae and Symplocaceae (Savile 1979b; see Jackson 2004 for possible codivergence). However, many characters other than rust preferences link Empetreae with Ericoideae in particular, e.g. both Rhodoreae and Empetreae have the flavonoid gossypetin. Molecular data strongly associate Empetreae with Ericaceae, and the ericoid leaves of the former are common (?independently derived) in other Ericoideae.

Chemistry, Morphology, etc. The best developed pit membrane remnants in Ericaceae occur in Enkianthus of the genera sampled; they are more poorly developed in other genera, but are well developed in other families in this part of Ericales (Carlquist & Schneider 2005) - a plesiomorphy? The hair roots found in many (?nearly all) Ericaceae are distinctive, being barely wider than a root hair and consisting merely of endodermis, exodermis, tracheid, sieve tube, and companion cell; they are relatively long-lived. Note that even quite low-order root branches in Vaccinium corymbosum, at least, are quite thin (Valenzuela-Estrada et al. 2008: the ultimate root branches are described there as first-order branches). For the development of the distinctive pollen of many Styphelioideae in which only a single cell of the tetrad persists, see Furness (2009). A common surface morphology of pollen grains in Ericaceae is faintly cerebellar, although there are some notable exceptions, as in Vaccinium japonicum - indeed, pollen is somewhat more variable than one perhaps might have thought (Sarwar et al. 2006 [Vaccinium], 2008 [Arbutoideae]; Sarwar & Takahashi 2006a [Vaccinioideae excl. Vaccinieae], 2006b [Enkianthus], 2007 [Vaccinieae], 2009 [Cassiopoideae and Harrimanelloideae]; Lu et al. 2009 [Gaultheria and relatives]). Viscin threads are well known as occuring in Rhodoreae, and have recently been reported from Gaultheria (Lu et al. 2009). Carpels are opposite the calyx in Vaccinium, Dracophyllum and Monotropa (Schnizlein 1843-1870: fams 160, 161). There is considerable variation in integument thickness in Ericaceae, but I have not attempted to collate all the literature on this feature: Pyrola may have an integument only two cells across, while that of Epacris is three cells thick and of Vaccinium, Arctostaphylos, etc., ca. 5 cells (Samuelsson 1913; Diels 1914).

For oligosaccharide storage, Fouquieriaceae, Diapensiaceae, and Cyrillaceae (and Lennoaceae) also sampled, see Pollard and Amuti (1981); for protein crystals in the nucleus, see Speta (1977); and for a phenetic analysis of some staminal characters, see Vander Kloet and Avery (2007). For general information on the family, see Kron et al. (2002b) and Stevens et al. (2004a).

Phylogeny. Early studies are summarized by Kron et al. (2002b). For the phylogeny of Ericoideae-Ericeae, see McGuire and Kron (2005); the African species are probably monophyletic and are derived from a lineage that originated within taxa currently found in Europe. Azalea, Ledum, Menziesia, Tsusiophyllum are to be included within Rhododendron (Kurashige et al. 2001, Gao et al. 2002; Milne 2004 [subsection Pontica paraphyletic, other subgenus Hymenanthes embedded in it; Kron 2003 [limits of genus, sections]; Brown 2003; Brown et al. 2006a, b, c [section Vireya, subsections need recircumscribing; biogeography]; and especially Goetsch et al. 2005 for phylogenies; Argent 2006 [account of species of subgenus Vireya] and Craven et al. (2008) classification of the Vireya group).

In Ericoideae-Rhodoreae, polypetaly is sporadic. The wind pollinated Empetreae are highly derived. Not only are their leaves opposite or whorled, small, and strongly revolute, but the plants are monoecious or dioecious, flower parts in 2s, or 4 tepals, free, the anthers lack appendages, there is one basal apotropous ovule/carpel, the stigma is expanded, and the fruit is a few-seeded drupe. Ericeae usually have similar small, opposite or whorled, strongly revolute leaves - the prototypical ericoid leaves - and their flowers are 4-merous, the bracteoles are often recaulescent, the calyx and corolla are both more or less scarious, they wither in fruit, but do not fall off, and the anthers are often with spurs.

For the phylogeny of Styphelioideae, see Crayn and Quinn (2000) and Quinn et al. (2003: Styphelieae). Within Styphelioideae, Prionoteae and Archerieae may be successively sister to remaining Styphelioideae. The former has such plesiomorphic features as multicellular hairs, leaves with serrate margins, pedicellate flowers, and anthers dehiscing by two slits. Styphelieae have drupes and the pollen grains are single, three cells of the meiotic quartet not developing (Furness 2009). Richeeae have 3<:3< nodes and sheathing leaf bases. Relationships in Styphelieae are being disentangled (e.g. Cherry et al. 2001; Quinn et al. 2005).

Within Gaultheria s.l. the epiphytic Diplycosia with some 100 species and Tepuia (Powell & Kron 2001; Bush et al. 2006; Bush & Kron 2008) may form a clade along with a few species of Gaultheria; the majority of Gaultheria s. str. forms a sister clade (Bush et al. 2009: see also optimisation of fruit and inflorescence characters).

For additional information on relationships, see Anderberg (1993), Cullings (2000), Judd and Kron (1993), Kron and Chase (1993), Kron et al. (1999a, b), and Crayn et al. (1998).

Classification. The infrafamilial classification outlined by Kron et al. (2002b) is largely followed here. Generic limits in Styphelieae and some other Styphelioideae are difficult (e.g. Cherry et al. 2001; Quinn et al. 2005 suggest some realignments). Still more substantial changes to generic limits are occuring in Vaccinioideae. Gaultheria is to include Pernettya, Diplycosia as well as Tepuia (Powell & Kron 2001; Bush et al. 2006; Bush & Kron 2008; Bush et al. 2009). Generic limits in the epiphytic Vaccinieae in particular are also in a mess (Powell & Kron 2003; Pedraza-Peñalosa 2009), and Vaccinium itself is probably pretty wildly paraphyletic. In particular, the "Tethyan" Vaccinium section Hemimyrtillus, with species from the Mediterranean area, etc., may be sister to the rest of the Vaccinieae, although with only weakish support (Powell & Kron 2002), while in Southeast Asia the Agapetes clade probably also includes 250< species of Vaccinium and 90< species of Agapetes s. str., all having superficial phellogen and a falsely 10-locular ovary, both probably derived features. Vander Kloet and Dickinson (2009) provide a sectional classification for Vaccinium - they recognize thirty sections.