EXTANT SEED PLANTS

Plant woody, evergreen; nicotinic acid metabolised to trigonelline; primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins rich in guaiacyl units; true roots present, xylem exarch, branching endogenous; arbuscular mycorrhizae +; shoot apical meristem complex; 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(-8) mm/mm²; plant heterosporous, sporangia eusporangiate, on sporophylls, sporophylls aggregated in indeterminate cones/strobili; true pollen [microspores] +, 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, mitochondrial nad1 intron 2 and coxIIi3 intron present.

MAGNOLIOPHYTA

Plant woody, evergreen; lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, cyanogenesis via tyrosine pathway [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 sieve plate, companion cells from same mother cell that gave rise to the tube, the sieve tube with P-proteins; nodes unilacunar; stomata with ends of guard cells level with aperture, paracytic; leaves with petiole and lamina [the latter formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, vein endings free; flowers perfect, polysymmetric, parts spiral [esp. the A], free, development in general centripetal, numbers unstable, P not differentiated, outer members not enclosing the rest of the bud, 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, pollen subspherical, binucleate at dispersal, trinucleate eventually, tectum continuous or microperforate, exine columellar, endexine thin, compact, lamellate only in the apertural regions, 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 , nectary 0, G free, several, ascidiate, with postgenital occlusion by secretion, few [?1] ovules/carpel, ovules marginal, anatropous, bitegmic, micropyle endostomal, integuments 2-3 cells thick, megasporocyte single, megaspore lacking sporopollenin and cuticle, chalazal, female gametophyte ?type, stylulus short, stigma ± decurrent, wet [secretory]; P deciduous in fruit; seed exotestal; double fertilisation +, endosperm ?diploid, cellular [first division oblique, micropylar end initially with a single large cell, chalazal end more actively dividing], copious, oily and/or proteinaceous, embryo cellular ab initio; germination hypogeal, seedlings/young plants sympodial; Arabidopsis-type telomeres [(TTTAGGG)_n]; whole genome duplication, single copy of LEAFY and RPB2 gene, knox genes extensively duplicated [A1-A4], AP1/FUL gene, paleo AP3 and PI genes [paralogous B-class genes] +, with "DEAER" motif, SEP3/LOFSEP and PHYA/PHYC gene pairs.

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

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

ERICALES [ASTERID I + II]: (C tube in flower), ovules tenuinucellate.

ASTERID I + II: ellagic acid 0, proanthocyanidins not common; inflorescence cymose; C forming a tube, A epipetalous, = and opposite sepals or P [polyandry (secondary) very uncommon indeed].

ASTERID I: G [2]; loss of introns 18-23 in d copy of RPB2 gene.

GENTIANALES + LAMIALES + SOLANALES: ?

LAMIALES + SOLANALES: iridoids, myricetin, non-hydrolysable tannins usu. 0; nodes 1:1; K connate, anther sacs with placentoids; endothelium +.

SOLANALES Dumortier  Main Tree, Synapomorphies.

Alkaloids, O-methyl flavonols (flavones) +; pollen tube usu. with callose; K persistent in fruit; endosperm development? - 5 families, 165 genera, 4080 species.

Stem group Solanales may date from the Capanian-Santonian 86-82 million years before present, diversifying 78-76 million years before present (Wikström et al. 2001: Sphenoclea not included).

Pyrrolizidine, tropane and pyrrolidine alkaloids are all synthesised from an ornithine precursor (Hegnauer 1973; Dahlgren 1988).

Montiniaceae are placed next to Solanaceae + Convolvulaceae (B. Bremer 1996, see also Soltis & Soltis 1997; cf. Takhtajan 1997 - in Hydrangeales). D. Soltis et al. (2000) found strong support for the association of Montinia and Hydrolea; Sphenoclea was not included. With the inclusion of the latter and broader sampling (three genera) in Montiniaceae, B. Bremer et al. (2002) found strong support for the association of Sphenoclea and Hydrolea, but only just above 50% for the association of Montiniaceae with that pair. The topology of the backbone of the tree here follows that of the latter paper.

Includes Convolvulaceae, Hydroleaceae, Montiniaceae, Solanaceae, Sphenocleaceae.

Synonymy: Convolvulales Dumortier, Nolanales Lindley, Sphenocleales Doweld - Solananae Reveal - Convolvulopsida Brongniart, Solanopsida Brogniart

Montiniaceae + Sphenocleaceae + Hydroleaceae: petiole bundle(s) arcuate; inflorescence terminal.

MONTINIACEAE Nakai   Back to Solanales

Shrubs, trees (lianes); plants with a peppery smell; route II secoiridoids +, slightly tanniniferous; cambium storied or not; pits vestured; young stem with a vascular cylinder (separate bundles; medullary bundles +); nodes 1:1(?-11:11 - Kaliphora); crystal sand, acicular crystals and styloids usu. all +; petiole arc of (rounded) bundles (+ additional strands); tuft of hairs at nodes; (stomata anisocytic - some Grevea); leaves also opposite; bracteoles 0; flowers imperfect, small, K small, C free [absolutely so - Montinia], valvate or not, nectary vascularised; staminate flowers: 3-4(-5)-merous, anthers extrorse, anthers ± basifixed, pollen grains large, pistillode minute; carpellate flowers: 4-merous, A staminodial (0), ovary inferior, placentation intrusive parietal-subaxile, 1-12 (campylotropous - Kaliphora) ovules/carpel, nucellus and endothelium?, style short, stout, hollow, stigma with 2 large lobes; fruit a capsule; seeds winged, exotesta lignified, periclinal walls thickened, adjacent wall of mesotesta also thickened [Montinia]; fruit indehiscent, placentae at least initially fleshy; testa thin-walled, ± pulpy when wetted [Grevea]; fruit a drupe; (exotesta not persistent Grevea); endosperm +/0, ?development, hemicellulosic, walls layered, cotyledons foliaceous, radicle oblique; cotyledonary petioles connate [Montinia]; n = 16 [Kaliphora], 34 [Montinia].

3[list]/5. Africa and Madagascar (Map: from Milne-Redhead & Metcalfe 1955; Verdcourt 1975; Bosser 1990; Brummit 2007 [C. and W. Africa]). [Photos - Kaliphora, Montinia Fruits © Serban Procheŝ.]

• Montiniaceae are woody plants that can be recognised by the persistent short style with large stigmatic lobes that cap the smooth fruit; the ovary is inferior. The often 4-merous flowers are small, with free, valvate petals and extrorse, more or less basifixed anthers. Crushed tissue may have a spicy or acrid smell.

Pericyclic fibers may be poorly developed; Grevea has vascular bundles in the pith. The axillary tufts of hairs are at best poorly developed in Kaliphora and are probably not apomorphic for the family. The pollen (Hideux & Ferguson 1976) is rather like that of some Araliaceae.

See Milne-Redhead and Metcalfe (1955) for general morphology and anatomy, Hegnauer (1973, 1990, as Saxifragaceae) for chemistry, Dahlgren et al. (1977) for germination and iridoids, Ramamonjiarisoa (1980), Carlquist (1989), and Gregory (1998) for vegetative anatomy, Krach (1976, 1977) and Takhtajan and Trifonova (1999) for testa anatomy, and Ronse Decraene (1992) and Ronse Decraene et al. (2000a) for floral details.

Synonymy: Kaliphoraceae Takhtajan

Sphenocleaceae + Hydroleaceae: placentae swollen [?level]; endosperm cellular, at most scanty, with multicellular micropylar and chalazal haustoria.

SPHENOCLEACEAE Baskerville   Back to Solanales

Rather fleshy annual herbs; fructose with isokestose linkages +; alkaloids?; cork ?mid-cortical; cortical air spaces +; stomata tetracytic; inflorescences spicate; early C-tube formation, C imbricate, pollen trinucleate, disc 0, G ± inferior, many ovules/carpel, style short, stigma capitate; fruit capsular, circumscissile; exotestal cells with inner walls thickened; n = 12, 16, 20, etc.

1/2. Pantropical. [Photo - Habit © B. Hammel]

• Sphenocleaceae are erect fleshy annuals of damp or flooded habitats that have dense spikes of small flowers with more or less inferior ovaries. The small fruits are circumscissile, the lid coming off just below the point of origin of the calyx. Older stems have a green reticulation marking the plates of cells in the tissue below bounding the cortical air spaces.

Corolla tube formation is of the early type, and the corolla lobes are characteristically incurved; the lateral veins of adjacent lobes are fused producing commissural veins.

Sphenocleaceae, along with Hydrolea, another genus of uncertain position, were placed near Boraginaceae by Cosner et al. (1994). However, in morphological studies (e.g. Gustafsson & Bremer 1995) Sphenocleaceae seem to be satisfactorily positioned well within Asterales. Indeed, Sphenocleaceae have often been associated with Campanulaceae (e.g. they are placed in Campanulales by Takhtajan 1997), although they lack latex.

Some information is taken from Monod (1980).

HYDROLEACEAE Berchtold & J. Presl   Back to Solanales

Herb to shrubby; axillary-sublateral thorns or not; cork?; vessel elements?; stomata?; leaf margin toothed to entire; flowers 4-5-merous, K basally connate, C , tube formation late, A versatile, disc 0/+, G [2(-4)], diagonal, placentae bilobed, many mostly pleurotropous ovules/carpel, antipodals degenerating early, funicular bundle absent, styles +, stigma slightly funneliform or capitate; fruit a septi- (and loculi-)cidal capsule (irregularly dehiscent); seeds longitudinally ridged and ruminate, exotestal cells thin-walled, endotestal cells tanniniferous, with a cuticle; n = (9) 10 (12).

1/12. Tropical, warm temperate (Map: from Davenport 1988; FloraBase 2007). [Photo - Hydrolea Flower © B. Kenney]

• Hydroleaceae are more or less shrubby plants growing near or even in water. They have rather small leaves.

The axillary inflorescences may be cymose. Davenport (1988) suggested that the disc is absent. The two carpels are shown as being oblique by Schnizlein (1843-1870: fam. 147), and this is confirmed by Erbar et al. (2005), even for Hydrolea palustris, which has flowers with the median sepal abaxial. Di Fulvio (1997) notes that the four ventral bundles are all connate in the center of the ovary - cf. Hydrophyllaceae s. str., where there are two or four such bundles. There are no nuclear inclusions.

Hydrolea has usually been included in Hydrophyllaceae (e.g. Cronquist 1981; Takhtajan 1997). Not only molecular differences but also axile versus parietal placentation and embryological differences separate the two.

Details of embryology are taken from Svensson (1925) and di Fulvio (1989b); Davenport (1988) monographed the genus.

Convolvulaceae + Solanaceae: coumarins, caffeic acid esters, tropane [polyhydroxynortropanes] and pyrrolidine alkaloids, flavonol and flavone glycosides, acylated anthocyanins +, tannins 0; internal phloem +; leaves with conduplicate ptyxis; late C-tube formation, C usu. contorted or induplicate-valvate, G opposite petals, many ovules/carpel, (integumentary tapetum +); K persistent in fruit; young seeds starchy.

This clade may have diverged 78-76 million years before present, diversifying perhaps 66-65 million years before present (Wikström et al. 2001). Chrysomelidae-Cassidinae and -Criocerinae beetle larve like members of this clade, especially Convolvulaceae (Schmitt 1988; Jolivet 1988; Buzzi 1994; Vencl & Morton 1999).

Gemeinholzer and Wink (2001) discuss the sporadic distribution of tropane alkaloids in Solanaceae; they are known from Schizanthus and other clades; see Schimming et al. (1998) for the distribution of polyhydroxynortropanes (in most Convolvulaceae, not in Cuscuteae, unknown in Humbertia, scattered in Solanaceae. It is possible that flowers with oblique symmetry may be an apomorphy for this clade.

CONVOLVULACEAE Jussieu, nom. cons.   Back to Solanales

Plant laticiferous; stomata usu. paracytic; leaf margins entire; K quincuncial, large, free; placentoid 0, ovules apotropous, stigma dry; chloroplast gene rpl2 intron absent.

57[List]/1625 - two groups below. World wide (Map: from Meusel et al. 1978; Staples & Brummit 2007).

1. Humbertioideae Roberty

Large tree; chemistry?; vascular bundles collateral; petiole bundle annular; usu. articulated latex canals, or latex cells in the flowers alone; flowers single, axillary, strongly obliquely monosymmetric, A adnate to base of C, filaments bent in bud, style clavate; fruit a few-seeded drupe; endosperm copious; n = ?

1/1: Humbertia madagascariensis. Madagascar.

The zygomorphy is largely positional; indeed, the flowers are drawn as being polysymmetrical by Pichon (1947). The sepals have five traces, but in Convolvuloideae there are fewer. Secretory cells are apparently restricted to the flower (Deroin 1993). Humbertia has hard wood with the odor of sandalwood.

Some other information is taken from Pichon (1947) and K. Kubitzki and H. Manitz (pers. comm.), but Humbertia is poorly known, esp. embryologically.

Synonymy: Humbertiaceae Pichon, nom. cons.

2. Convolvuloideae Burnett

Herbaceous right-twining vines (lianes to 30 m; trees); (ergoline alkaloids + - from clavicipitalean fungus); (cork pericyclic); secondary thickening anomalous; (fibers or sclereids +); unicellular T-shaped hairs common (hairs stellate); leaves (compound; margins lobed, toothed [dentate - Hyalocystis]), 2ndary veins pinnate to palmate; inflorescence usu. a dichasium; tapetum cells multinucleate, pollen pantoporate or 6-polycolpate, G [2(-5)], (false septum - Mina), (1-)2(-4) erect weakly crassinucellate ovules/carpel, placental obturator common, stigmas capitate and with multicellular papillae or punctate and smooth; fruit usu. a variously dehiscent capsule; testa with unbranched bundle, exotesta with papillae or hairs, usu. little thickened, outer hypodermis of small cells, little thickened, inner hypodermis elongated or not, 1+ palisade layers, thickened, or cells little elongated; endosperm nuclear, embryo green, curved, cotyledons often folded or coiled, bifid; n = 7-15+; chloroplast gene atpB with 6-15 bp deletion, trnF with 150 bp deletion, rpl2 intron 0.

56/1600: Ipomoea (500: paraphyletic, spiny pollen clade derived, some 50% of clade [Manos et al. 2001a] - I. batatas, the sweet potato), Cuscuta (145), Convolvulus (100), Argyreia (90), Jacquemontia (90), Erycibe (75), Merremia (70). World-wide. [Photo - Flower, Fruit.]

Within Convolvuloideae, part of Poraneae (Cardiochlamyeae: Porana itself is polyphyletic!), Erycibeae s. str., and a clade made up of all other Convolvuloideae form a basal trichotomy. Cardiochlamyeae have foliaceous bracts and fruits that are utriculate, i.e. they are one-seeded and with a papery pericarp. Erycibe itself (= Erycibeae) has broad, radiate, sessile stigmas and a berry-like fruit; it can look very unlike other members of the family and herbarium specimens are often misidentified. Ipomoea, Convolvulus, and their relatives form an embedded clade that is sister to a rather unexpected clade made up of Poraneae, Cresseae, Dichondreae (with gynobasic styles), some Erycibeae (Maripeae), etc., as well as Jacquemontia. Several members of this latter clade have distinct styles, or with only at most a short style and long styles (but not Jacquemontia, etc.), and leaf blades with more or less pinnate venation; Jacquemontia could be sister to the other taxa. Despite the sequencing of over 6800 bp, the position of Cuscuteae remains unclear (Stefanovic & Olmstead 2001, 2004; Stefanovic et al. 2002 and esp. 2003).

Synonymy: Cressaceae Rafinesque, Cuscutaceae Berchtold & J. Presl, nom. cons. (parasite; no internal phloem; C imbricate, with fimbriate corona especially well developed below and adaxial to A, ovules (incompletely) tenuinucellate, styles separate to connate, stigma capitate to elongated; embryo spirally coiled, almost acotyledonous, root 0), Dichondraceae Dumortier, nom. cons., Erycibaceae Meisner, Poranaceae J. Agardh

• Convolvulaceae are recognisable by their usually viny habit and leaves with palmate venation; the plants are often obviously laticiferous. The quincuncial calyx is often notably persistent; the sepals are free, and are rarely green in fruit, often being brown and woody - as in wood roses (Merremia tuberosa) - to scarious. The large, radially symmetric corolla has contorted or induplicate-valvate aestivation and the five stamens are adnate to the tube. The gynoecium is bicarpellate, each loculus usually having two ovules (you rarely need to check this since the calyx is so distinctive), and the seeds have a notably hard, shiny testa.

The flowers often last for only a single day.

Glycine betaines are rather commonly accumulated in this family (Rhodes & Hanson 1993), perhaps surprising since it is not a family of halophytes. Wood fluorescence occurs, but not often. The bracts may be adnate to the pedicel and accrescent (wind dispersal: Neuropeltis), or the bracetoles may be much enlarged (e.g. Calystegia). The corolla tube of some Cuscuta and other members of the family is strictly speaking a corolla-stamen tube, both contributing integrally to the tubular structure (Prenner et al. 2002). Some Convolvuloideae have flowers with slight oblique disymmetry (Lefort 1951), while Weberling (1989) described the flowers as being fundamentally gynobasic, although with an apical septum. There are a few, mostly old records of protein crystalloids in the nucleus (Speta 1977; Thaler 1966).

For Cuscuta, see Kuijt (1969), the Parasitic Plants website (Nickrent 1998 onwards) and the Digital Atlas of Cuscuta (Costea 2007 onwards), for chemistry, see Hegnauer (1964, 1989), for ergoline alkaloids, see Markert et al. (2008), for seed reserves, see G. Dahlgren (1991), for the rpl2 intron, Downie et al. (1991) and Stefanovic et al. (2002), for a morphological phylogeny, see Austin (1998), for ovary morphology, see Deroin (1999b), for general information, including tribal groupings, see Staples and Brummitt (2007), and for relationships within the family, see Stefanovic and Olmstead (2000, 2001) and especially Stefanovic et al. (2002, 2003).

SOLANACEAE Jussieu, nom. cons.   Back to Solanales

Herbs to shrubs, branching sympodial; hygroline alkaloids, (withanolides [steroidal lactones]), oligosaccharides, (myricetin) +; roots diarch [lateral roots 4-ranked]; (hairs branched/stellate); wood commonly fluoresces; pits vestured; crystal sand +, esp. in stem; cystoliths +; stomata various; leaves simple to compound; (branching in inflorescence/leaf insertion apparently deviating from normal); (flowers 4 merous; obliquely monosymmetric), anthers often porose, or connate and pollen exiting communal apical hole, tapetum cells often 4-nucleate, G [(-5)], oblique, often pseudo-4-locular, usu. many often campylotropous ovules/carpel, style solid, stigma 2-lobed or peltate, wet; fruit a berry or septicidal capsule (drupe), K accrescent or not; exotestal walls thickened, endotesta persistent, walls ± lignified; endosperm (helobial, nuclear) +, cotyledons and radicle same width; n = 7-14, etc., chromosomes 1-3 µm long, protein bodies in nuclei.

102[list]/2460 - 7 subfamilies below, but treatment needs work. World-wide, but overwhelmingly tropical America (Map: from van Steenis & van Balgooy 1966; Meusel et al. 1978; van Balgooy 1984; Heywood 2007). [Photos - Iochroma Flower, Przewalkskia Fruiting Calyx, Schizanthus Flower.]

1. Schwenckioideae

Annual herbs; pericycle fibers +; flowers monosymmetric, A 4, didynamous, or 2 + 2 or 3 staminodes; embryo straight, short; n = 12.

4/31. South America.

The corolla lobes themselves each have three lobes in Schwenckia and Melananthus.

Schizanthoideae + Goetzeoideae + Cestroideae + Petunioideae [Nicotianoideae + Solanoideae]: ?

2. Schizanthoideae Hunziker

Annual herbs; tropane alkaloids +; cork pericyclic; pericycle fibers 0; flowers strongly monosymmetric, abaxial pair of C connate, forming a keel, A 2 [abaxial-lateral], staminodes 3; endosperm nuclear, embryo curved; n = 10.

1/12. Chile.

The flower is described as having oblique rather than inverted symmetry (Cocucci 1989b), and anther dehiscence is explosive (Cocucci 1989a); for floral evolution, see Pérez et al. (2006: midpoint rooting). Schizanthoideae have distinctive tropane alkaloids, hairs, and pollen (Hunziker 2001).

3. Goetzeoideae Thorne & Reveal

Fruit often a drupe; endosperm at most slight, cotyledons large, fleshy [Goetzea, etc.], or embryo curved, cotyledons small [Duckeodendron]; n = 13.

4/7. Greater Antilles, but not Jamaica, South America. [Photo - Flower]

Includes Metternichia, with Duckeodendron sister to the rest (the latter with only moderately good support: Santiago-Valentin & Olmstead 2001, 2003).

Synonymy: Duckeodendraceae Kuhlmann ([genera list], (wood with large, open, radial canals [cf. Apocynaceae s. str.]; carpels oblique [Kuhlmann 1934]; 1 seed/fruit - Huchinson [1973] placed it in Boraginaceae, but doubtfully; Cronquist [1981] kept it as a poorly-known family; Takhtajan [1997] placed it in Solanales), Goetzeaceae Airy Shaw (odd growth pattern; leaves often xeromorphic; fruit a drupe)

4. Browallioideae Kosteletzky

(Steroid alkaloids - Cestrum); cork superficial or deep-seated; bordered pits +; pericyclic fibers +; A 4 or 5, often didynamous, staminode +/0; n = 7-13.

8/195: Cestrum (175). South and Central (and North) America.

Arabidopsis-type telomeres are absent from some of this clade (Sýkorová et al. 2003a), and Cestreae in particular (which lack these telomeres) have chromosomes that at 7.21-11.51 µm long are considerably larger than those of the rest of the family, which are much smaller, e.g. 1.5-3.52 µm long in Nicotianoideae (Acosta et al. 2006); Cestreae also have n = 8 (Las Penas et al. 2006). Metternichia has seeds with only slight endosperm and n = 13.

Synonymy: Cestraceae Schlechtendal, Salpiglossidaceae (Bentham) Hutchinson

Androgenesis, an uncommon condition, has been recorded for at least one member - Petunia, Capsicum and Nicotiana - of each of the last three subfamilies (Chat et al. 2003 for references).

Petunioideae + Solanoideae + Nicotianoideae: (tropane alkaloids [calystegines] +).

5. Petunioideae

Cork superficial (deep-seated); bordered pits +; pericyclic fibers +(0); druses 0(+); (flowers disymmetric), A 4(-5), usu. of two lengths; embryo also slightly curved; n = 7-9, 11.

13/160. Brunfelsia (45). Central and South America.

Nicotianoideae + Solanoideae: (nicotine [pyridine alkaloid] +); (cotyledons accumbent); x = 12.

6. Solanoideae Kosteletzky

(Steroid alkaloids +); pits not vestured; (crystal sand +) [level?]; A 5 (4), (of different lengths), base of the filament [stapet] often enlarged, with lobes, etc., (style gynobasic, G 3, or subdivision of carpels into 1-seeded units - "Nolanaceae"); fruit a berry (drupe; circumscissile capsule - Hyoscyameae; schizocarp); seeds flattened; endosperm cellular, embryo curved, often coiled; (n = 10-15).

61/1925: Solanum (1250-1700: inc. Cyphomandra, Lycopersicon; hairs often stellate, prickles common - see Bohs 2005, 2007; Levin et al. 2006; Weese & Bohs 2007 [three genes, S. thelopodium sister ro rest, or unresolved in Bayesian analysis], Botany 2008: Botany without Borders 120-121. 2008 for phylogenies), Lycianthes (200), Lycium (90: paraphyletic, to include Phrodus and Grabowskia - Levin & Miller 2005; Levin et al. 2007), Nolana (90: Tago-Nakawaza & Dillon 1999; Dillon et al. 2007 for a phylogeny), Physalis (80). World-wide, but esp. South America and others N. temperate. [Photo - Flower]

For branching patterns in Solanum, see Danert (1967). Lycium is recorded as accumulating glycine betaines, and some members at least are halophytic (Levin & Miller 2005). For alkaloids in Datureae, see Doncheva et al. (2006), for floral development, see Yang et al. (2002).

Synonymy: Atropaceae Martynov, Daturaceae Rafinesque, Hyoscyamaceae Vest, Lyciaceae Rafinesque, Nolanaceae Dumortier, nom. cons. (2 long 2 short stamens - Nolana)

7. Nicotianoideae Miers

Cork superficial; pericyclic fibers +/0; A 4 (staminode +), 5, (of two lengths); embryo straight (curved), radicle short; n = (7-11).

8/125: Nicotiana (95). Mostly Australian, also North and South America, Africa.

Synonymy: Nicotianaceae Martynov

• Solanaceae are quite often rather foetid herbs or weak shrubs with polysymmetric or weakly, rarely strongly, monosymmetric sympetalous flowers - when monosymmetrical, the flowers are held so that the odd or median petal is in the adaxial position. Vegetatively many Solanaceae - especially Solanoideae, the bulk of the family - are distinctive because the insertion of leaves, branches and flowers often does not seem to follow any regular sequence; paired leaves adjacent on the stem are common. The flowers often have porose anthers or the anthers as a group are connivent at the apex. The fruit is usually bicarpellate and is a septicidal capsule or a berry with flattened seeds; the calyx consists of connate sepals that persist and even enlarge in fruit.

Stem group Solanaceae date from 66-65 million years before present, crown group (including Schizanthus and Duckeodendron) from 41-36 million years before present (Wikström et al. 2001). A genome duplication event in Solanaceae has been dated from ca 50-52 million years before present (Schlueter et al. 2004). New World Solanaceae are eaten by larvae of Nymphalidae-Ithomiinae butterflies (Ehrlich & Raven 1964; Drummond et al. 1987; Willmott & Freitas 2006: Schizanthoideae, Goetzeoideae and Schwenkioideae may not be eaten). Detailed coevolution seems not to be involved; Ithomiinae probably moved on to Solanaceae from Apocynaceae, and Solanum is especially important as a food source (ca 70% records of neotropical Solanaceae food sources, ca 89% those of all Ithomiinae: Willmott & Freitas 2006; see also Brower et al. 2006). Although some larvae may be distasteful, the adult butterflies are distasteful not because of the alkaloids, etc., that may be in Solanaceae that the larvae eat but because of the dihydroxypyrrolizidines that the adults obtain especially from Boraginaceae-Heliotropioideae and Asteraceae-Asteroideae (esp. Eupatorieae, see Brown 1987). The mimicry rings in which Ithomiinae are involved may be associated with particular host plants (Willmott & Mallet 2004). Tobacco hornworm caterpillars prefer members of the Solanoideae + Nicotianoideae clade as food sources, although they didn't like Nicandra much; they died on Petunia, and didn't grow on Browallia and Brunfelsia. Other plant feeders show similar distinctive patterns (e.g. Fraenkel 1959), thus other sphingids are found here and on Oleaceae (Forbes 1958). Phytophagous Chrysomelidae beetles (perhaps especially Criocerinae) are notably more common on New World than Old World Solanaceae, perhaps suggesting an origin of the use of the family as a food source in the former area (Jolivet & Hawkeswood 1995; see also Hsiao 1986), while Chrysomelinae and Megalpodinae are also found on New World Solanaceae (Jolivet 1988). Criocerinae may have moved onto Solanaceae from monocots; the larvae are covered by faecal shields (Vencl & Morton 1999). In general, however, Solanaceae have multiple lines of defence and are avoided by most insect herbivores (Harborne 1986; Hsiao 1986).

Petunia and Hyoscyamus, in different subfamilies, can be intergrafted (Taiz & Zeiger 2006).

Unusual stomata with degenerate guard cells seem often to have been reported in the family (Cammerloher 1920; D'Arcy & Keating 1973). Leaves in the fertile part of the stem of Solanaceae, perhaps especially in Solanoideae, are often geminate and/or branching is apparently not axillary; Petunia can have ordinary-looking cymose inflorescences, but Schwenkia, Schizanthus and many other taxa have more or less recaulescent bracts, only one branch of the inflorescence is developed each node, or they develop in different ways, etc., making interpretation of the construction of the plant very difficult (see especially Danert 1958; Child & Lester 1991 for a brief summary; Bell & Dines 1995). Physalis has notably inflated calyx surrounding the fruit, in the development of which heterotopy of a foliar gene may be involved (He & Saedler 2007; cf. Hu & Saedler 2007); inflated calyces occur in some nine genera in total, although details of the pattern of evolution - and perhaps also loss - are unclear.

In floral development, petal and stamen primordia together are lifted by zonal growth and the carpel primordia develop on a flat apex; in this respect there are some similarities between Solanaceae, Scrophulariaceae and Gesneriaceae, few with Montiniaceae (Huber 1980: 66-69; Ronse Decraene et al. 2000). Endothecial thickening in the family is very diverse (Carrizo García 2002).

The two carpels so common in Solanaceae (but Nicandra has 3-5) are often in the plane of the first K initiated; this is one of the abaxial pair. Zygomorphy and heteranthy (which in this context is really a kind of zygomorphy) have evolved several time in Solanum (Bohs et al. 2007). The basic plane of symmetry in flowers like Salpiglossis and Schizanthus may be monocotyledonous, and the "abaxial" one or three stamens are sterile (see Eichler 1875; Robyns 1931; Cocucci 1989; Knapp 2002; Ampornpan & Armstrong 2002). Floral symmetry in both Convolvulaceae and Solanaceae needs study, especially given the suggestion by Ampornpan and Armstrong (2002) that in the latter the gynoecium is in the median plane, the parts being initiated slightly off the vertical plane of symmetry and the whole flower slightly rotated, the result being the odd petal is in the adaxial position. For a detailed study of the frequent loss of gametophytic incompatability in Solanaceae, see Igic et al. (2006).

The grouping [Petunioideae [Solanoideae + Nicotianoideae]] is well supported in Olmstead et al. (1999), although less so in Olmstead and Santiago-Valentin (2003), while in the summary tree of Olmstead and Bohs (2007), although there is a clade [Solanoideae + Nicotianoideae], immediately below it is a polychotomy including Petunioideae, Cestroideae and Schwenkioideae. Indeed, the relationships between the branches basal to this group have only weak support, Schwenkia may be sister to the rest of the family (Olmstead et al. 1999). However, using the nuclear gene SAMT (salicylic acid methyl transferase), Martins and Barkman (2005) found Schizanthus to be sister to the rest of the family, and with rather strong support, with Schwenkia weakly linked with Cestroideae (see also Olmstead & Bohs 2007). Furthermore, Wu et al. (2006) found a strongly supported grouping of [Solanoideae [Petunioideae + Nicotianoideae]]. Although in this case no other clades of the family were included, Wu et al. (2006) noted that the sequences they included came from ten orthologous loci each on a different chromosome. (Wu et al. [2006] also dismissed the possibility that there had been a genome duplication either on the branch leading to Solanaceae or within Solanaceae themselves [cf. Blanc & Wolfe 2004].) Clearly there remain uncertainties about relationships between major groupings within the family, but until these are resolved I follow the infrafamilial classification of Olmstead et al. (1999).

For relationships, see Olmstead & Palmer (1992), Fay et al. (1998b), and especially Olmstead et al. (1999). For general chemistry, see Hegnauer (1973, 1990, also 1966, 1990 as Nolanaceae), for hairs, esp. of Solanum, see Seithe (1962), for wood anatomy, see Carlquist (1987, 1988a) and Jansen and Smets (2001: vestured pits - do Petunioideae and Nicotianoideae have them?), for floral and inflorescence morphology, see Huber (1980), for branching patterns, see Danert (1958), for pollen morphology of zygomorphic taxa, but not yet integrated with phylogeny, see Stafford and Knapp (2006), for evolution of secondary metabolites, see Wink (2003 and references), for evolution of fruit and flower types, etc., see Knapp (2002) and Knapp et al. (2004), for calystegines (tropane alkaloids), see Dräger (2004), for the evolution of floral scent, see Martins et al. (2007), and for fruit anatomy, see Pabon Mora and Litt (2007). For generic descriptions and much else, see Hunziker (2001), although his classification differs from that used here; Solanaceae Source includes information currently mostly about Solanum, but its coverage will expand.

Unplaced: Sclerophylacaceae Miers (crystal sand +; flowers sessile, G 2, oblique, inferior, ovules 2-3, pendulous from upper part of loculus); K accrescent, spinescent, surrounding the 1-3-seeded fruit - generally placed near Solanaceae).