EXTANT SEED PLANTS

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

MAGNOLIOPHYTA

Plant woody, evergreen; lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, cyanogenesis via tyrosine pathway, lignins derived from both coniferyl and sinapyl alcohols, containing syringaldehyde [in positive Maüle reaction, syringyl:guaiacyl ratio less than 2-2.5:1], and hemicelluloses as xyloglucans; root apical meristem intermediate-open; root vascular tissue oligarch [di- to pentarch], lateral roots arise opposite or immediately to the side of [when diarch] xylem poles; origin of epidermis with no clear pattern [probably from inner layer of root cap], trichoblasts [differentiated root hair-forming cells] 0; stem with 2-layered tunica-corpus construction; wood fibers and wood parenchyma +; reaction wood ?, with gelatinous fibres; starch grains simple; primary cell wall mostly with pectic polysaccharides; tracheids +; sieve tubes eunucleate, with sieve plate, companion cells from same mother cell that gave rise to the tube, the sieve tube with P-proteins; nodes unilacunar; stomata with ends of guard cells level with aperture, paracytic; leaves with petiole and lamina [the latter formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, vein endings free; flowers perfect, polysymmetric, parts spiral [esp. the A], free, numbers unstable, P not differentiated, outer members not enclosing the rest of the bud, A many, development centripetal, with a single trace, introrse, filaments stout, anther ± embedded in the filament, tetrasporangiate, dithecal, with at least outer secondary parietal cells dividing, each theca dehiscing longitudinally by action of hypodermal endothecium, endothecial cells elongated at right angles to long axis of anther, tapetum glandular, binucleate, microspore mother cells in a block, microsporogenesis successive, pollen subspherical, binucleate at dispersal, trinucleate eventually, tectum continuous, endexine compact, lamellate only in the apertural regions, pollen tube elongated, with callose plugs, penetrating between cells, growth rate moderate, siphonogamy occuring, nectary 0, G free, several, ascidiate, with postgenital occlusion by secretion, few [?1] ovules/carpel, ovules marginal, anatropous, bitegmic, micropyle endostomal, integuments 2-3 cells thick, megasporocyte single, megaspore lacking sporopollenin and cuticle, chalazal, female gametophyte ?type, stylulus short, stigma ± decurrent, wet [secretory]; P deciduous in fruit; seed exotestal; double fertilisation +, endosperm ?diploid, cellular [first division oblique, micropylar end initially with a single large cell, chalazal end more actively dividing], copious, oily and/or proteinaceous, embryo cellular ab initio; germination hypogeal, seedlings/young plants sympodial; Arabidopsis-type telomeres [(TTTAGGG)_n]; whole genome duplication, single copy of LEAFY and RPB2 gene, knox genes extensively duplicated [A1-A4], AP1/FUL gene, paleo AP3 and PI genes [paralogous B-class genes] +, with "DEAER" motif, SEP3/LOFSEP and PHYA/PHYC gene pairs.

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

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

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

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

BUXALES [GUNNERALES + CORE EUDICOTS]: ?

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

CORE EUDICOTS: Root apical meristem closed; flowers rather stereotyped: 5-merous, parts whorled, K and C distinct, K with 3 traces, A = 2x K, internal to the C whorl, (numerous, but then often fasciculate and/or centrifugal), pollen tricolporate, (nectary disc +), [G 5], [3] also common, compitum +, placentation axile, stigma not decurrent; endosperm nuclear; fruit dry, dehiscent, loculicidal [when a capsule]; euAP1 + euFUL + AGL79 genes [duplication of AP1/FUL or FUL-like gene], PLE + euAG [duplication of AG-like gene: C class], SEP1 + FBP6 genes [duplication of AGL2/3/4 gene].

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

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

ASTERIDS 1 + 2: Ellagic acid 0, proanthocyanidins not common; inflorescence cymose; C forming a tube, A epipetalous, = and opposite K 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:  Back to Main Tree
iridoids, myricetin, non-hydrolysable tannins usu. 0; nodes 1:1; K connate, anther sacs with placentoids; endothelium +.

Information on placentoids is needed for Hoplestigmataceae, Plocospermaceae, etc. Protein crystals in nuclei are apparently not known from Avicennia, etc. (Speta 1977, 1979), and information is needed for groups recently moved to Lamiales; Whether or not such crystals characterises both Lamiales and also Boraginaceae s. str. (see also Wagstaff & Olmstead 1997) needs to be confirmed. Although some Boraginaceae have protein bodies in their nuclei, they are of two very different kinds, and many lack them; there is also variation with Lamiales. Vahlia is sometimes placed sister to Lamiales, but with only 63% bootstrap support (Albach et al. 2001b), or is associated more specifically with Boraginaceae... (Lundberg 2001e). Lamiales and Boraginaceae have in common similar hydroxycinnamic acid derivatives (disaccharide esters or rosmarinic/lithospermic/caffeic acids: Mølgaard & Ravn 1988).

LAMIALES Bromhead  Main Tree, Synapomorphies.

Cornoside, verbascoside [caffeoyl phenylethanoid glycoside, caffeic acid ester, acteoside], methyl- and oxygenated flavones +; glandular hairs with vertical divisions in head +; leaves opposite; endosperm with micropylar haustoria; fruit a capsule; protein bodies in nuclei; mitochondrial coxII.i3 intron 0. - 23 families, 1059 genera, 23275 species.

Lamiales contain ca 12.3% eudicot diversity. Diversity is concentrated in families whose members are largely herbaceous and have monosymmetric flowers. Estimates of the age of this clade range from ca 97 to ca 74 million years before present (Bremer et al. 2004; Wikström et al. 2001 respectively). Parasitic and insectivorous Lamiales are nonmycorrhizal (Brundrett 2004 and references).

The occurence of cornosides and iridoids in this clade seem to be mutually exclusive, except in Martynia louisiana (Jensen 2000b). Verbascoside, a disaccharide derivate of caffeic acid, a hydroxycinnamic acid, and numerous variants, and also trisaccharide derivates, are notably prominent in this clade (Mølgaard & Ravn 1988). Intermediary cells with distinctive plasmodesmata associated with the ultimate leaf veins may be plesiomorphic in Lamiales; their presence is linked with the transport of raffinose and stachyose, oligosaccharides commonly found in phloem exudate in the order (Turgeon et al. 2001). Leaf teeth have a glandular apex, with one accessory vein proceeding into the tooth, the other going above it. Taxa with trinucleate pollen grains are scattered throughout the order. For integument thickness, for which I have no generally comparable information yet but which may be of systematic importance, see also Hjertsen (1997) and Fischer (2004b). An hypostase is common - e.g. Buddleja, some "scrophs" - but the level of this feature is unknown. Oleaceae seem to have a rather diferent embryo development from that of other Lamiales studied (Yamazaki 1974). A long, narrow suspensor may be common in Lamiales (di Fulvio 1979; Maldonado de Magnano 1987), but I do not know the general distribution of this character - it is certainly not found in Loganiaceae! Seed pedestals, variable in morphology and developed from the placenta, are scattered through the order, being known from Tetrachondraceae, Calceolariaceae, Plantaginaceae, Scrophulariaceae, Silbaceae, Orobanchaceae, Phyrmaceae and Paulowniaceae (Rebernig & Weber 2007).

Knowing the exact positions of Carlemanniaceae, placed sister to Oleaceae (rbcL data alone: Savolainen et al. 2000b; Lee et al. 2007, Plocospermataceae not included) or generally "basal" in Lamiales (K. Bremer et al. 2001, Plocospermataceae not included) and of Plocospermataceae themselves are important for understanding the evolution of the chemistry and floral morphology in particular of the group (cf. Endress 2001). Taxa with 4-merous or predominantly 4-merous flowers are common in the pectinations of the Lamiales tree below Gesneriaceae (see also Mayr & Weber 2006). If Carlemanniaceae are in this general part of the tree, as seems possible, one would expect them to lack iridoids - at least, to lack route II decarboxylated iridoids - and to have only a single endosperm haustorium. Note that Carlemanniaceae have both 4- and 5-merous flowers and that the lip of the apparently 4-merous flowers of Calceolariaceae may represent two completely connate members of the petalline whorl (Mayr & Weber 2006); evolution of floral meristicity in this part of the tree is clearly a complex matter, and exactly where changes of floral meristicity are to be placed is unclear.

The very provisional tree here is based in part on Oxelman et al. (1999) and Soltis et al. (2001). B. Bremer et al. (2002) analyse variation in three coding and three non-coding regions of the chloroplast genome; their sampling is sketchy, so the support for some family groupings in the part of the order is difficult to evaluate; Freeman and Scogin (1999) focus on the old Scrophulariaceae, but the pattern of relationships they find is unclear. Indeed, how family limits of those taxa with rather large, monosymmetric flowers (i.e., the bulk of the clade) will be drawn remains uncertain, and it is likely that characters distinguishing clades will be hard to come by, but an improvement over the old situation is likely. A tree in Müller et al. (2004) also suggests that at least a partial resolution of relationships is in sight, although sampling here is poor (this study focused on Lentibulariaceae); the three families known or suspected to be carnivorous (Byblidaceae, Lentibulariaceae and Martyniaceae) are not immediately related. Duplication of the FLO=LFY and DEF=AP3 genes within Lamiales suggests very interesting relationships, if confirmed; duplication occured in the representatives of Phrymaceae, Verbenaceae, Paulowniaceae and Orobanchaceae examined, but not in those of Plantaginaceae or Oleaceae (Aagard et al. 2005). It has been suggested that families such as Orobanchaceae, Lamiaceae and Acanthaceae form a clade with strongly monosymmetric flowers that mostly lack a staminode (Endress 2001). Calceolariaceae may be sister to Gesneriaceae (S. Andersson 2006: sampling!). Verbenaceae have been linked with Bignoniaceae and Paulowniaceae with Phrymaceae (Nie et al. 2006). Thomandersia, from tropical Africa, previously usually included in Acanthaceae, may go near Schlegeliaceae, however, support for any such association is weak (Wortley et al. 2005a and especially 2007). Few strongly supported relationships were evident in the recent study of Soltis et al. (2007a), indeed, although over 4600 bp have been sequenced, it has been estimated that at least 10000 bp more will need to be added if relationships within the clade are to be resolved (Wortley et al. 2005b). Savolainen et al. (2000b) place Carlemannia sister to Oleaceae (only 1 species in analysis) with moderate support, while Bremer et al. (2001) found the two genera to form a sister group that is part of a trichotomy at the base of Lamiales (Oleaceae [Ligustrum only] + all other Lamiales complete the trichotomy; Plocospermataceae were not included). A sister relationship to Oleaceae is also supported by Yang et al. (2007: 1.0 pp, Plocosperma included, but sampling still very poor), and that seems the best place to put the family.

The limits of families like Scrophulariaceae have long been problematic (Thieret 1967 for a summary). Rahmanzadeh et al. (2004), Albach et al. (2005) and Oxelman et al. (2005) are clarifying the contents of the separate clades that used to be subsumed in Scrophulariceae s. lato. (see also Tank et al. 2006).

R. Olmstead (pers. comm.) is compiling a synoptical classification of Lamiales from which some of the numbers of taxa included in the families below are taken. He also provided a readable account of some of the earlier changes in our ideas of relationships in the Scrophulariaceae s.l. in particular (Olmstead 2002; see Tank et al. 2006 for an update). Kadereit (2004) provided a summary of the order and its evolution.

Misplaced - even if there is little oxalate accumulation in Scrophulariaceae s.l. (although this may be more a sampling issue with other families in the clade).

For chemistry, see Harborne and Williams (1971: scutellarein, etc.), Zindler-Frank (1978: oxalate accumulation), Young and Siegler (1981: anthraquinones), Mølgaard and Ravn (1988: caffeoyl esters), Tomás Barberán et al. (1988: flavone glycosides), Scogin (1992: acteoside), and Grayer et al. (1999: general); note that cornoside is generally found in groups lacking iridoids (Jensen 1992, 2000a). For proteinaceous nuclear inclusions, see Bigazzi (1984, 1989a, 1989b, 1993, 1995) and Speta (1977, 1979), but unfortunately it is not known if they occur in Plocospermataceae and Tetrachondraceae in particular. For a useful general discussion, including suggestions of apomorphies for some clades, see Soltis et al. (2005b).

Includes Acanthaceae, Bignoniaceae, Byblidaceae, Calceolariaceae, Carlemanniaceae, Gesneriaceae, Lamiaceae, Lentibulariaceae, Linderniaceae, Martyniaceae, Oleaceae, Orobanchaceae, Paulowniaceae, Pedaliaceae, Peltanthera, Phrymaceae, Plantaginaceae, Plocospermataceae, Rehmannia, Schlegeliaceae, Scrophulariaceae, Stilbaceae, Tetrachondraceae, Thomandersiaceae, Verbenaceae.

Synonymy: Acanthales Lindley, Bignoniales Lindley, Byblidales Reveal, Callitrichales Dumortier, Carlemanniales Doweld, Gesneriales Dumortier, Globulariales Dumortier, Hippuridales Thomé, Jasminales Dumortier, Lentibulariales Lindley, Ligustrales Bischof, Oleales Lindley, Pinguiculales Dumortier, Plantaginales Lindley, Rhinanthales Dumortier, Scrophulariales Lindley, Stilbales Doweld, Verbenales Horaninow - Lamianae Takhtajan, Oleanae Takhtajan (monotypic) - Lamiidae Reveal - Bignoniopsida Nees, Ligustropsida Meisner, Plantaginopsida Meisner, Selaginopsida Brongniart, Verbenopsida Brongniart

PLOCOSPERMATACEAE Hutchinson   Back to Lamiales

Shrubs or trees; cork?; vessels in radial multiples; petiole bundle annular; styloids +; hairs unicellular, calcified and/or with cystoliths, also bicellular, club-shaped, glandular; cuticle wax crystalloids 0; petiole articulated near base; inflorescences axillary, plant cryptically dioecious, bracteoles 0; flowers 5-6-merous, A extrorse, versatile, with parallel separate cells, nectary in carpellate flowers only, placentation parietal, 2 ovules/carpel, style divided twice, lobes stigmatic, not expanded, style, etc., 0 in staminate flower; seed with tuft of multicellular hairs at chalazal end; coat ?; endosperm ?development, slight; n = ?; protein crystals?

1[list]/1: Plocosperma buxifolia. Central America (Map: from Leeuwenberg 1967).

• Plocospermataceae are shrubby plants that may be recognised by their rather small, opposite leaves that are articulated near the base of the petiole, and their quite large, polysymmetric, campanulate-rotate flowers with versatile, extrorse anthers and a twice-divided style. The fruit is a few-seeded capsule and the seeds have a tuft of hairs at one end.

Plocospermataceae are poorly known. There are prominent groups of fibers in the outer cortex as the leaf leaves the stem. In each carpel, one ovule may be subapical and pendulous, the other subbasal and erect, or both may be in either position (Leeuwenberg 1967).

Plocospermataceae were included in Gentianales by Takhtajan (1997), probably because Plocosperma had long been associated with Loganiaceae.

See D'Arcy and Keating (1973: as Lithophytum, esp. anatomy), Jensen (1992: chemistry), M. Endress et al. (1996: general, and relationships), and Struwe and Jensen (2004: general) for information, and Oxelman et al. (1999a), Mueller et al. (2001) and Hilu et al. (2001) for their position as sister to other Lamiales.

[Carlemanniaceae + Oleaceae] [Tetrachondraceae [Calceolariaceae [Peltanthera + Gesneriaceae] [Plantaginaceae, Lamiaceae, etc.]]]: cells in heads of glandular hairs with vertical walls only; flowers 4-merous [?: reverses to 5-merous....].

Note that the arrangement of the sepals (and petals) is orthogonal in Oleaceae and Calceolariaceae, while that of Tetrachondraceae (and of 4-merous Veronica) is diagonal (Mayr & Weber 2006).

Carlemanniaceae + Oleaceae: A 2, stigma ± clavate; esotestal cells ± palisade.

CARLEMANNIACEAE Airy Shaw   Back to Lamiales

Perennial herbs or shrubs; chemistry?; vessel elements?; nodes?; no cuticle waxes; leaf margins toothed; inflorescences terminal and axillary; flowers weakly obliquely monosymmetric, 4- or 5-merous, heterostylous [Silvianthus], K unequal or not, ± linear, open?, C induplicate-valvate, anthers connivent and surrounding the style, pollen colpate, G inferior, nectary on top; fruit fleshy-capsular, loculicidal or 5-valved [valves correspond to calyx segments, and the fruit opens widely, exposing the placenta], K persistent; exotestal cells with radial walls thickened, unthickened cells interior to this, endothelium persistent [Silvianthus], or polygonal, all walls thickened [Carlemannia]; endosperm +, ruminate [Silvianthus], embryo small; n = 15, 19; protein bodies in nucleus?

2[list]/5. Thailand, Laos, Vietnam, S. China, Sumatra (Map: somewhat hypothetical).

• Carlemanniaceae can be recognised by their opposite, serrate leaves joined by a line across the stem; more or less swollen nodes; weakly monosymmetric four or five-merous flowers with only two stamens and inferior, bicarpellate ovary; and more or less fleshy but dehiscent fruit.

Characters such as stem anatomy, two stamens with connivent anthers and two carpels each with many ovules remove Carlemanniaceae from Caprifoliaceae; the toothed, exstipulate leaves, 2 stamens, anomocytic stomata, and absence of raphides (and crystal sand - Solereder 1893) from Rubiaceae (see Airy Shaw 1965; but included in Rubiales by Takhtajan 1997). The peltate, glandular hairs with unicellular stalks and the two stamens (their position is not entirely certain) suggest Lamiales. However, Carlemanniaceae are embryologically and chemically largely unknown.

Some information is taken from Tange (1999); Thiv (2004) provides a general account, and Yang et al. (2007) chromosome numbers, etc.

OLEACEAE Hoffmannsegg & Link, nom. cons.   Back to Lamiales

Woody; carboxycyclic iridoids, myricetin, orobanchin, mannitol +; (vessel elements with scalariform perforations); pits vestured; petiole bundle arcuate; sclereids + (0); hairs often peltate, secretory; cuticle deeply furrowed (waxes ribbons, platelets); branching from previous innovation; leaves odd-pinnate to simple, conduplicate [Chionanthus], margins entire to toothed, (2ndary veins palmate); (plant dioecious); flowers 4-merous, C tube formation early, often very short (0), (A 4), anther thecae ± back-to-back, ovules pendulous (hemitropous), (style short), stigma dry; testa often vascularised, exotesta moderately and evenly thickened, (endotesta fibrous; endothelium ± persistent); endosperm +/0; protein bodies in nuclear crystalline-globular; 9 bp deletion in ndhF.

24[list]/615 - five tribes below. More or less worldwide, especially East Asia (Map: from Meusel et al. 1975).

1. Fontanesieae L. Johnson

Secoiridoids +; pits vestured; 1 ovule/carpel; fruit a samara; n = 13.

1/2. Sicily, W. Asia, China.

2. Forsythieae L. Johnson

Cornosides +; 1-several ovules/carpel; fruit a samara or capsule; n = 14.

2/8. S.E. Europe, East Asia.

3. Myxopyreae Boerlage

Myxopyroside iridoid pathway +; cortical bundles in corners of angled stem (Dimetra not); 1(-3) ascending ovules/carpel; fruti a berry or schizocarp; n = 11, 12.

3 (Myxopyrum, Dimetra, Nyctanthes)/7. Indo-Malesia.

Synonymy: Nyctanthaceae J. Agardh (lamina surface rough; G transverse, 1 ascending ovule/carpel; exotesta little developed, mesotesta persistent, endotestal cells tangentially elongated, sclerotised; in the past, often placed in Verbenaceae)

Jasmineae + Oleeae: oleoside +; 2(-4) ovules/carpel.

4. Jasmineae Lamarck & Candolle

Secoirioidoids +; K and C to 14 or more, endothelium 0; fruit bilobed, berry or circumscissile capsule; seed coat multilayered, mesotesta with wholly thickened or band-thickened anticlinal walls; n = 11-13; 21kb chloroplast inversion.

1/225-450 (Jasminum: inc. Menodora). Tropical to warm temperate Old World, some in America. [Photo - Flower]

Synonymy: Bolivaraceae Grisebach, Jasminaceae Adanson

5. Oleeae (R. Brown) Dumortier

Secoiridoids, flavone glycosides +; vessels in multiples; pits vestured; libriform fibers +; (C free; 0), (A 4 - e.g. Nestegis); n = (20) 23.

17/415: Chionanthus (60-120: Linociera recently incorporated, but questionable?), Fraxinus (45-65: C may be 0, or free; leaves simple or compound, samara; wind pollination), Ligustrum (50), Noronhia (45), Olea (33). Tropical and subtropical, inc. New Zealand and Hawaii.

Schrebera looks superficially like Bignoniaceae.

Synonymy: Forestieraceae Endlicher, Fraxinaceae Vest, Ligustraceae G. Meyer, Nyctanthaceae J. Agardh, Schreberaceae (Wight) Schnizlein, Syringaceae Horaninow

• Oleaceae are woody plants that are recognisable by their opposite, exstipulate simple to odd-pinnately compound leaves; the twigs often dry pale and contrast with the dark-drying petioles, there are superposed buds, and the stem is also somewhat swollen at the nodes (note: there is no line across the node, cf. Gentianales, many other opposite-leaved Lamiales). The usually four-merous, sympetalous polysymmetric flowers with two stamens are also distinctive; even in Jasminum, with 5 or more sepals (and even spiral leaves), there are still only two stamens. The terminal bud quite often aborts.

Caterpillars of some Sphinginae are found on Oleaceae (and the same genera may also be on Solanaceae: Forbes 1958).

At least some species of Osmanthus have a lignified, torus-bearing, pit membrane (Coleman et al. 2004). Minute crystals in the wood are present (including in Nyctanthes), but are uncommon. The corolla tube is initiated early in a ring. The secretory hairs may be in groups forming extrafloral nectaries, while nectar is reported to be secreted from the ovary in Syringa and Ligustrum (Weberling 1989). Osmophores are common and their absence from the anthers may be of systematic interest (Nilson 2000: sampling?). Orbicules may be absent (Vinckier & Smets 2002a). Baillon (1891) illustrates both epitropous and apotropous ovules. The chloroplast gene accD (= ORF512, zpfA) has been lost (Doyle et al. 1995 and references) in at least some Oleaceae.

The route I secoiridoids are unlike other route I secoiridoids, e.g. those in Gentianaceae (Jensen 1992). Forsythia and Abeliophyllum (possibly sister to rest of the family) have cornosides, and some seem to lack iridoids; these features may be plesiomorphies.

Wallander and Albert (2000) suggest phylogenetic relationships within the family; the tribes they recognise (see above) have strong support, however, generic limits in Oleeae need much attention. Lee et al. (2006), however, found Myxopyreae to be sister to the rest of the family (100% bootstrap support), with Fontanesieae, Forsythieae and [Jasmineae + Oleeae] forming a trichotomy; they emphasized the complex pattern of chloroplast inversions in Jasminieae.

Tetrachondraceae [Calceolariaceae [Peltanthera + Gesneriaceae] [Plantaginaceae, Lamiaceae, etc.]: deletion in the matK gene.

For information, see Hilu et al. (2000); the sampling needs to be improved.

TETRACHONDRACEAE Wettstein   Back to Lamiales

Creeping to erect herb; sorbitol +; cork?; nodes split laterals; hair moniliform [Polypremum]; leaf bases connate or connected by membranaceous stipules; (inflorescence of 1-2 axillary flowers - Tetrachondra); flowers rather small, 4-merous, C with very short tube, anthers separate, pollen in tetrads, 6-sulcate, psilate, nectary 0, G 4 partite, 2 erect ovules/carpel, style gynobasic [Tetrachondra], or G slightly inferior, placentae peltate, many ovules/carpel [Polypremum], stigma small, subglobose; fruit with persistent green K, either schizocarpic, or a loculicidal (+ septicidal) capsule; endothelial cells with persistent thickened inner walls, testa thin; endosperm copious; n = 10, 11.

2[list]/3. Patagonia, Australia, New Zealand (Tetrachondra), S. U.S.A. to South America (Polypremum procumbens) (Map: approximate). [Photo - Polypremum Flower]

• Tetrachondraceae are perennial erect to prostrate herbs with small, opposite, more or less sessile leaves that are joined at the bases; the flowers are small, four merous, and polysymmetrical.

Chemistry (Harborne & Williams 1971 - scutellarein +, cf. Gelsemium!; Jensen 2000a), endothelium presence (absent in Loganiaceae), endosperm type, etc., of Polypremum are right for position in Lamiales, and the Polypremum + Tetrachondra pair is strongly supported (Oxelman et al. 1999a). Wagstaff et al. (2000) found that the sequences of the two species of Tetrachondra, from the Antipodes and from S. South America, were almost identical. Polypremum has both micropylar and chalazal endosperm haustoria, so the position of these features in this part of the phylogeny (see immediately below) is uncertain, also, its embryo sac protrudes through the nucellar epidermis.

Tetrachondra was placed in Boraginales by Takhtajan (1997) and in Lamiaceae by Cronquist (1981), while Polypremum has always been associated with Loganiaceae.

For general information, see Wagstaff (2004a), some additional information is taken from Moore (1948).

Synonymy: Polypremaceae Doweld & Reveal

Calceolariaceae [Peltanthera + Gesneriaceae] [Plantaginaceae, Lamiaceae, etc.]: shikimic-acid derived anthraquinones [see also Rubiaceae], 6- and/or 8- hydroxylated flavone glycosides + [? Tetrachondraceae], storage substances stachyose and other oligosaccarides; flowers vertically monosymmetric, C bilabiate, adaxial lobes outside the others in bud [ascending cochleate], tube formation late, many ovules/carpel; endosperm also with chalazal haustoria [?position in tree]; protein bodies in nuclei lamellar.

This clade has many members that are herbaceous or shrubby and have monosymmetric flowers and fruits with many small seeds. Westerkamp and Claßen-Bockhoff (2007) outline the morphological variation in monosymmetric, bilabiate flowers of the clade.

S. Andersson (2006, two genes) found 75% jacknife support for [Calceolariaceae + Gesneriaceae], and 100% support for that clade as sister to remaining Lamiales. A possible synapomorphy for this clade, if it is confirmed, is the presence of paired-flower cymes. The flower in front of the termianl flower is sometimes subtended by a "bracteole" which may represent the bract of that flower, the flower opposite it being totally suppressed (Weber 1973; Haston & Ronse De Craene 2007); what appears to be a rather strange dichasial cyme may be a modified "panicle". Both families also have at least some taxa with septicidal dehiscence of the capsule, but I don't know how the distribution of this character might appear on a combined tree of the two.

For the distribution of various flavone glycosides, see Tomás-Barberán et al. (1988: Mimulus and Orobanche lack the glycosides of Lamiaceae, Verbenaceae, Scrophulariaceae and Plantaginaceae, while those of Lentibulariaceae are somewhat different), for staminodes, see Endress (1998).

CALCEOLARIACEAE Olmstead   Back to Lamiales

Herbaceous to shrubs; cork?; leaves toothed (entire); cymes paired-flower; flowers 4-merous, K valvate, abaxial C lobe saccate, (adaxial "lip" strongly bilobed - Stemotria), elaiophores as pads of hairs on inside of abaxial lip (0), A 2 [adaxial pair] (3 - Stemotria), thecae (parallel) divergent, confluent on dehiscence or not, (theca 1), staminodes 0, nectary 0, (G semi-inferior), integument 3-4 cells across, stigma small or capitate or obscurely bilobed; capsule both septicidal and loculicidal; testa with anticlinal walls sinuous (straight); endosperm +; n = (8) 9.

2[list]/260: Calceolaria (240-270). Upland tropical and W. temperate South America, Brasil, also New Zealand (some Jovellana) (Map: from Sérsic 2004). [Photo - Habit, Flower.]

• Calceolariaceae are herbs or shrubs usually with opposite, serrate leaves that may be joined at the base. Their distinctive flowers are 4-merous, usually with two stamens, no nectary, and often a strongly saccate lower lip with oil-secreting glands as pads of hairs on inside of the lip.

Flowers with a closed mouth are visited by bumble bees, those with an open mouth by smaller bees. Visitors remove either oil from the oil glands, specialised hairs, or pollen if there are no oil glands (Vogel 1974; Sérsic 2004).

Each lip of the flower seems to be formed from two petals, judging by their vasculature, etc.; these primordium pairs may become connate only rather late in floral development (Mayr & Weber 2006).

Porodittia is a synonym of Stemotria, but neither name is needed as Stemotria is clearly derived from within Calceolaria (Mayr & Weber 2006), thus P. triandra = C. triandra (see S. Andersson 2006). The limits of the sections need adjusting.

Some information is taken from Weber (1973: inflorescence), Molau (1988: general, the abaxial lip appears to be bilobed and the odd stamen abaxial), Ehrhart (2000: general), Fischer (2004b: general, in Scrophulariaceae), and Mayr and Weber (2006: development - superb micrographs); see also Tank et al. (2006). I am grateful to Pamela Puppo for comments.

[Peltanthera + Gesneriaceae] + Plantaginaceae, Lamiaceae, etc.: monosymmetry of the 2:3 type, A 4 [often didynamous, with connivent anthers]; suspensor large [?level].

GESNERIACEAE Candolle, nom. cons.   Back to Lamiales

Usu. herbs or weak-stemmed trees (trees); hairs of stalked glands, or with thickened terminal cells; cork also deep seated; (cambium storied); (vessel elements with scalariform perforations); (nodes 1:1 with split laterals; 3:3 with split laterals; 5:5); stomata anisocytic; leaves joined at the base, (anisophyllous; 2-ranked; spiral), involute, rather fleshy and/or softly hairy, margins toothed or entire; inflorescence axillary (terminal), cymes paired-flower; abaxial C lobe outside others in bud [aestivation descending cochleate], (C spurred), A with staminode, (5, 2), anthers usu. didynamous and connivent in pairs, (thecae apically confluent), nectary vascularised, placentation intrusive parietal, placentae ± triangular, usu. covered by ovules, stigma broadly bilobed to trumpet-shaped, wet or dry; fruit a (septicidal) capsule (berry); exotestal cells variously elongated and thickened, endotestal cells at most simply persisting; GCyc duplication.

147(+)[list]/3200 - 3 main groups below. Largely tropical. [Photo - Flower]

Didymocarpoideae + Epithematoideae: 3-desoxyanthocyanins 0, chalcones, aurones +; ?stomata; ovary wall not richly vascularised, nectary vascularized from A traces; endosperm inconspicuous, cotyledons unequal, one accrescent. (Map: from van Steenis & van Balgooy 1966 [Malesia and Pacific]; Hillard & Burtt 1971 [Africa]; New World is only approximate.)

For anisocotyly - more accurately, one cotyledon is accrescent - and its development, see Burtt (1970) and Saueregger and Weber (2004).

1. Didymocarpoideae Arnott

Placentae lamelliform-recurved, ovules restricted to distal end, style usually not well set off from ovary; fruit elongated; testa cells ornamented, (with hairs); n = (4, 8) 9-11 (12, 13) 14-17, etc., polyploidy not uncommon.

82/2000: Cyrtandra (550), Aeschynanthus (180), Henckelia (155), Streptocarpus (155: A 2, inc. Saintpaulia, etc. - Möller & Cronk 2001, Imaichi et al. 2007 [growth patterns and the evolution of monophylly], Möller & Cronk 2002 [biogeography]; Harrison et al. 1999 [floral diversification]), Chirita (140), Agalmyla (100), Didymocarpus (100), Paraboea (90). S. Europe (scattered), Africa and Madagascar (possibly "basal" genera only), mostly Sri Lanka to Malesia and the Pacific (to Hawaii).

Haberlea and Ramonda, temperate, European, and with polysymmetric flowers and five stamens, may be sister to the rest of Didymocarpoideae (e.g. Mayer et al. 2003); they have dihydrocaffeoyl ester found nowhere else in flowering plants (Jensen 1996). The huge Didymocarpus has recently been dismembered (Weber & Burtt 1998), species that have been placed here are to be assigned to 27 genera (including two in Plantaginaceae); many species are in Henckelia.

Synonymy: Cyrtandraceae Jack, Didymocarpaceae D. Don, Ramondaceae Godron

2. Epithematoideae [no name]

Dihydroxyphenolics [e.g. acteoside] 0; secretory canals, medullary bundles +; cymes lacking bracteoles; (abaxial C lobe inside others in bud; placentation axile; nectary variously vascularized), ovary short, abruptly narrowed into the style; endosperm ?0; n = (8-)10(-12).

6/75: Monophyllaea (30+). India, South East Asia to Malesia, 1 sp. W. Africa, 1 sp. (Rhynchoglossum azureum) S. Mexico to Peru.

Epithematoideae perhaps include Cyrtandromoea (molecular data), but this is also sometimes placed in "Scrophulariaceae" - and it does have iridoids and is otherwise chemically similar to the latter; it also has endosperm and an exotesta with laminated, U-shaped thickenings in transverse section, the seeds are isocotylous, and the gynoecium bilocular (Burtt 1965, also a revision, he placed the genus in Scrophulariaceae and linked it with Leucocarpus). Epithematoideae are sister to Cyrtandroideae, see Smith (1996), Smith et al. (1997a, b) and especially Mayer at al. (2003). Branch lengths are long. Chemistry - Napeanthus (Gesnerioideae) is also similar!

3. Gesnerioideae Link

3-desoxyanthocyanins +, chalcones, aurones 0; seeds without surface ornamentation; endosperm conspicuous; GCyc2 gene lost.

63: 1250. Predominantly Neotropical, a few S.W. Pacific, East Asia.

3a. Coronathereae

Trees or ± shrubby (rotting from the nodes); stomata anomocytic; nectary embedded in G wall, vascularized from A traces; capsules septicidal (and loculicidal; fruit a berry); n = 37(-45).

9/20: Coronanthera (11). Solomon Islands, Antilles, New Caledonia, S. South America (Map: from Burtt 1998).

Titanotricheae + Gesnerieae: (plant with scaly rhizomes).

3b. Titanotricheae

Inflorescence racemose, with bulbils; testa striate-reticulate; n = 20.

1/1: Titanotrichum oldhamii. China, Japan, Taiwan, scattered in W. Malesia.

3c. Gesnerieae

(Nodes split-lateral - Episcieae); (leaves spiral); (flowers resupinate), G superior to inferior, numerous vascular bundles in ovary wall from which nectary is vascularized; (fruit with fleshy placentae or funicles - "display capsule"; berry); n = (8) 9 (10) 11 (12) 13-14 (16), polyploidy rare.

53/1500: Besleria (150), Drymonia (140+), Alloplectus (75+), Nautilocalyx (70+), Paradrymonia (70+), Gesneria (60), Sinningia (60), Columnea (s.l. = 270+, s. str., 75+, + 4 genera, inc. Dalbergaria [90], Tricantha [75+]). New World (Map: from Brummitt 2007, in part). [Photo - Leaves, Flower.]

This clade probably includes the Old World but more or less isocotylar Titanotrichum near the base (C.-N. Wang et al. 2004: substantial amount of molecular data; cf. D. Soltis et al. 2000; Albach et al. 2001), but it has also sometimes been placed in "Scrophulariaceae". See Pan et al. (2002) for its floral development. Kotarski et al. (2007) found 80% bootstrap support for the position of Coronathereae as sister to the other Gesnerioideae, and then Titanotrichum as sister to the remainder. For a phylogeny of Coronanthereae that suggests two dispersals E -> W across the Pacific, see Smith et al. (2006). Besleria and Napeanthus (n = 16) may also be near the base of the Gesnerioideae tree. For other relationships in the subfamily, see Smith (2001), Zimmer et al. (2002) and Smith et al. (2004a, b), for relationships around Alloplectus, see Clark and Zimmer (2003), for the phylogeny and biogeographic relationships of Gloxinieae, see Roalson et al. (2005 a, b; 2008b), the last paper focusing on relationships in Central America and the Antilles, and for diversification in Beslerieae, see Roalson and Clark (2006), and in Sinningieae, see Perret et al. (2006: the limits of Sinningia need adjusting, 2007: diversification). See also Skog (1976) for a revision of Gesneria and relationships in Gesnerieae.

Synonymy: Belloniaceae Martynov, Besleriaceae Rafinesque

• Gesneriaceae are herbs or rather soft-stemmed shrubs that can be recognised by their opposite, serrate leaves that are often softly hairy and have arching, ascending secondary veins that do no join at the margin and their usually conspicuous monosymmetric flowers which have parietal placentation. In the distinctive cymose inflorescence there is an "extra" flower in front of the terminal flower.

Gesneriaceae are not often eaten by the caterpillar larvae of butterflies (Ehrlich & Raven 1964).

Variation in growth patterns in this family is considerable, and Weber (2004) provides a useful survey. The architecture of some Didymocarpoideae is distinctive, Streptocarpus in particular showing much variation in growth pattern, some species having only a single, ever-growing cotyledon (e.g. Hilliard & Burtt 1971); evolution of growth form in this clade shows many parallelisms and reversals, as well as being linked with other life history variables, such as age to flowering and flowering periodicity (Möller & Cronk 2001). Imaichi et al. 92007) discuss the evolution of monophylly. Plant architecture is notably diverse and complex in Epithematoideae, with anisophylly being common; taxa like Rhynchoglossum have "alternate" leaves. The plant body of many species of Monophyllaea is a single, ever-growing structure that is derived from a single cotyledon; there is a basal meristem and in some species the flowers arise along the midrib of the blade rather than from separate inflorescences at the base of the cotyledonary petiole. Tsukaya (2005) describes this growth in terms of the basal part of the leaf blade, normally the last to mature and stop growing, as keeping on growing.

Although many taxa are rather succulent, sometimes even delicate, herbs, there is a surprising number that grow on exposed rocks (Boea hygrometrica is an example) and are resurrection plants (Burtt 1998 for literature). Epiphytes are common, with well over 400 epiphytic species known from neotropical Episceae alone (Maddison 1977; Weber 1978). Polysymmetric flowers have arisen independently several times in the family, the ten or so genera involved not being immediately related (e.g. Burtt 1970; Smith et al. 2004a), indeed, polysymmetric flowers are notably abundant compared with some other families of the strongly monosymmetric Lamiales (Endress 1997). Harrrison et al. (1999) discuss floral diversification in Streptocarpus, which includes species with strongly monosymmetric flowers as well as Saintpaulia, with almost polysymmetric flowers, so encompassing very different flower morphologies and pollinators. Wiehler (1978) estimated that ca 60% neotropical Gesnerioideae (perhaps a thousand species) were hummingbird pollinated, and he divided the floral morphologies involved into three common and one less common "types"; ca 30% Gesnerioideae were pollinated by euglossine bees of both sexes (cf. e.g. Orchidaceae where it is only male bees seeking scents that are are involved). Pollination by birds is relatively less common in Old World Gesneriaceae. Flowers with inverted orientation are known from some Episcieae (Clark & Zimmer 2003). They seem to have evolved ca 3 times and this inverted orientation is evident from the very earliest stages of the ontogeny of the flower, and there is no twisting of the pedicel (Clark et al. 2006); they are not strictly resupinate. Many Gesneriaceae have capsular fruits with wind dispersed seeds, and seed dispersal by birds, either of fleshy fruits in their entirety, of glistening seeds exposed on a fleshy placenta, or of a number of the other variants of fleshy capsule/drupe fruit type found in the family, is also widespread (Weber 2004b; Clark et al. 2006) is also common; in the Old World, Cyrtandra has fleshy fruits.

There is quite a lot of anatomical variation which I have not integrated with the clades recognised here. Thus sclereids are common in the stem; Aeschynanthus has strongly U-thickened sclereids in the pericycle, other taxa lack fibers or sclereids in the pericyclic position; some taxa have lignified hairs; Gesneria has a U-shaped petiole bundle cradling a unmedullated circle of vascular tissue, and there are also two wing bundles; etc. In paired-flower cymes, there is a flower in front of the terminal flower and with the same orientation as it.

Gesneria has spirally-inserted serrate leaves with an almost coriaceous texture - it looks quite "atypical"!

As might be expected of a family in which there are conspicuous flowers and much obvious adaptation to pollinators, current generic limits, based as they are on floral characters, are unsatisfactory. However, much-needed changes are underway, and those in New World Gesneriaceae are clearly explained in a series of articles in Gesneriads 56(3). 2006. Also, see the World Checklist and Bibliography of Gesneriaceae (Skog & Boggan 2005 a, b).

Dickison (1994), Jensen (1994, 1996), Norman (1994), and Wiehler (1994) all deal with Sanango which had previously been placed in Loganiaceae, etc. Molecular studies (e.g. K. Bremer et al. 2001; B. Bremer et al. 2002) continue to place Sanango in this area, but exactly where it and Peltanthera (see below) go is unclear; the latter is very similar in wood anatomy to Buddleja, the former has vessels with scalariform perforation plates like a few other Gesneriaceae (e.g. Kohleria, Carlquist 1997c). Secondary metabolites (lack or iridoids, presence of the caffeoyl phenylethanoid glycoside, sanangoside) suggest an association between Sanango and Gesnerioideae (Jensen 1996).

For more information, see Trapp (1956b: androecium), Weber (1973: inflorescence), Wilson (1974a, b: nectary vascularization), Skog (1984: chromosomes), Beaufort-Murphy (1984: response to growth hormones, etc.; Cyrtandroideae much more responsive than Gesnerioideae), Kvist and Pedersen (1986: phenolics), Citerne et al. (2000) Smith et al. (2004b: both molecular details of floral development), Burtt and Wiehler (1995) and Wiehler (1983: both general), Möller and Kiehn (2004: cytology), and Weber (2004a: excellent general account, 2004b: history of classification, the four major groups with informal names).

PELTANTHERA   Back to Lamiales

Verbascosides, cornoside derivatives +; nodes 3:3; petiole bundle flattened annular, with (medullary and) rib bundles; hairs branched-moniliform; leaves involute, serrate, rather soft; inflorescence axillary, thyrsoid; flower slightly monosymmetric, K largely free, C valvate, A 5, thecae confluent, many ovules/carpel; n = ?

1/1: Peltanthera costaricensis. Central and W. South America.

Placed here, see Oxelman et al. (1999a), and perhaps to be included in Gesneriaceae. For information on general morphology, see Hunziker & Di Fulvio (1957), and for chemistry, see.

Plantaginaceae, Lamiaceae, etc.: route II decarboxylated iridoids [aucubin widespread], 6- or 8-hydroxyflavones or 6 methoxyflavones +, cornosides 0.

See Wagstaff and Olmstead (1997) and Olmstead et al. (2001) for relationships.

Sensitive stigmatic lobes are of sporadic occurence in this clade, and are found in some Phrymaceae, Bignoniaceae, Pedaliaceae, Lentibulariaceae, Linderniaceae and Acanthaceae at least (see also Endress 1994b). In many taxa the nectary is vascularised by branches from the main carpellary vascular traces - examples are Schlegeliaceae, some Pedaliaceae, Verbenaceae. On the other hand, the nectary is vascularised separately from the gynoecium in Bignoniaceae, Acanthaceae, other Pedaliaceae. There are septal vascular bundles - the gynoecial vascular system forms a sort of figure of 8 in transverse section - in Bignoniaceae and Schlegeliaceae. In other taxa like Acanthaceae there are no septal bundles, the gynoecial vasculature forming a sort of circle as a result (there are of course placental bundles: see Wortley et al. 2005a for details). Knowledge of the distributions of both these characters needs to be extended (somebody has probably already done this), and the variation in the vasculature of the androecium suggests that either the distinction between gynoecial and receptacular nectaries (Smets 1988; Smets et al. 2003) is overly simplistic and/or there is homoplasy in this feature.

PEDALIACEAE R. Brown, nom. cons.   Back to Lamiales

Annual to perennial herbs to deciduous trees; orobanchin, amyloid +; cambium storied; pericycle also with sclereids (fibers few); petiole bundle interrupted-annular; hairs broadly capitate, mucilaginous; leaf (spiral - Sesamum; venation palmate), margins toothed, lobed or entire; flowers usu. axillary (inflorescence dichasial), paired nectaries (modified flowers) at base of pedicel or not, A (5), didynamous, thecae ± confluent, at right angles to filaments, staminode + (0), pollen 5-13 zonocolpate, G [2-4], 2-many ovules/carpel (8 loculi, 1 ovule/loculus - Josephinia), stigma with 2 broad lobes, wet, often sensitive; fruit (schizocarp; nut; wind-dispersed), usu. with hooks or prickles of the endocarp exposed as mesocarp rots and fruit splits loculicidally, style base indurated; seeds winged or not, testa multiplicative, exotestal cells palisade or otherwise thickened, (mesotesta with crystals); slight staining for amyloid in cotyledons, endosperm thin; n = 8 (13); protein bodies in nucleus?

13[list]/70: Sesamum (19), Pterodiscus (13). Mostly tropical, in coastal or arid habitats, Old World (Map: from Ihlenfeldt & Grabow-Seidensticker 1979; FloraBase 2005; rather approximate esp. in Africa).

• Pedaliaceae are herbs or shrubs with mucilaginous hairs and large, monosymmetric flowers with a superior ovary. Placentation is axile, and the fruits often have hooks, prickles, or other emergences. Some taxa have glands at the base of the pedicel - these are modified flowers. [Photo - Flower]

The mucilage glands normally have four cells, and the apparently single axillary flowers of some taxa may represent reduced cymes (Manning 1991). Josephinia may have four carpels, each loculus being divided - a remarkable feature for a member of the asterid I + II group. Although Rogeria is reported to occur in Brasil, this seems to be a mistake (Volker Bittrich, pers. comm.).

Amyloid is also found in Acanthaceae, sometimes weakly associated with Pedaliaceae in molecular analyses (Soltis et al. 2005b and references). Both Martyniaceae and Pedaliaceae have 10-hydroxylated carboxylic iridoids.

Some information is taken from Stapf (1895), S. D. Manning (1991: U.S.A., general), and Ihlenfeldt (1967, 2004: general).

Synonymy: Sesamaceae Berchtold & J. Presl

MARTYNIACEAE Stapf, nom. cons.   Back to Lamiales

Annual herbs, roots often tuberous (perennials; woody); harpagide, harpagioside [8ß-8α-methyl substituted iridoids] +; petiole bundle deeply arcuate, also adaxial cortical and medullary bundles; plant sticky-hairy; leaves also spiral, margins toothed; inflorescence racemose; (K free), A (2), didynamous, staminode(s) +, pollen trinuclete, inaperturate, exine dissected into 20-40 platelets, placentation parietal, placentae bilobed, 2-many ovules/carpel, stigma bilobed; capsule with apical spurs or hooks [developing from sterile upper part of ovary], mesocarp ± fleshy, falling off, endocarp woody, with crests and spines; testa subgelatinous, or inner and radial walls with cellulosic bands, inner layers lignified [Proboscidea], or lignified exotesta only persistent; endosperm at most thin; n = 15, 16.

5[list]/16: Proboscidea (10). Tropical and subtropical America.

• Martyniaceae are herbs with sticky hairs and large, monosymmetric flowers with a superior ovary. The placentation is parietal, and the fruits have two prominent apical hooks as well as other hooks, spines or prickles. [Photo - Flower]

Insects may stick to the very viscid indumentum of Martyniaceae, although there is no evidence that the plants are carnivorous (but cf. Stylidiaceae [Asterales], which also have sticky hairs and for which there are recent suggestions that there may be carnivory). Martyniaceae are not immediately related to Lentibulariaceae and Byblidaceae, which are directly or indirectly carnivorous (Müller et al. 2004).

Martyniaceae and Pedaliaceae have often been combined, but there is currently no evidence that they form a monophyletic group. Differences in pollen and placentation best separate the two.

Some information is taken from Stapf (1895: general) and S. Singh (1970: embryology, etc.); Ihlenfeldt (2004) provides a general account of the family.

STILBACEAE Kunth, nom. cons.   Back to Lamiales

Ericoid shrubs, ordinary shrubs, or herbs; C8 iridoid glucosides +, (cornosides +); cork just outside pericycle; vessel elements also with scalariform perforations; nodes ?; petiole bundle?; stomata?, cuticle waxes as rods or threads; (leaf margins minutely toothed), revolute or not; inflorescences axillary or terminal (flowers axillary), flowers radial to monosymmetric, (4) 5(-7)-merous, bracteoles as long as K; K bilobed or not (free), C lobes equal to unequal, A = and opposite K, (one fewer; staminode +), anther thecae confluent apically or parallel with separate slits, ovary apically unilocular, 1-2 ovules/carpel, ascending and/or descending, apo/epitropous, unilocular [1 G infertile, or septum 0], or many ovules, bilocular, stigma sl. bifid or punctate; K and C persistent, fruit a loculicidal (and septicidal) capsule (indehiscent); embryo cylindrical [always?], endosperm +; n = 10, 12, 19; protein bodies in nucleus crystalline [Halleria].

11[list]/39: Nuxia (15). Most South Africa, the Cape Province, also to tropical Africa, Madagascar, the Mascarenes and Arabia (Map: from Leeuwenberg 1975). [Photo - Nuxia Inflorescence, Halleria Flower.]

The gynoecium is reminiscent of that of Scrophulariaceae - Manueleae, while the iridoids are like those of Loasaceae and Hydrangeaceae (e.g. unedoside is in common - Jensen et al. 1998). Thesmophora has G 2, tranverse, with 1 descending ovule/carpel (Rourke 1993) - perhaps the abaxial G with a false septum.

The C8 iridoid glucosides common in this family are extremely uncommon elsewhere; for the distribution of unedoside, present in at least some genera, see Oxelman et al. (2004a).

Retziaceae and Stilbaceae come out together in rbcL trees (Wagstaff & Olmstead 1997); for another early study on relationships, see B. Bremer et al. (1994). Rourke (2000) recognised two subfamilies, Retzioideae and Stilboideae, in Retziaceae, but these do not incorporate genera more recently moved to this clade. Thus Nuxia (ex Loganiaceae) is placed here in molecular phylogenies (Backlund et al. 2000; Wallick et al. 2002); for the association of Halleria with Stilbaceae, see Olmstead et al. (2001). Kornhall (2004) and Oxelman et al. (2005) have further clarified the limits of the family, the former dividing it into three tribes. Additional genera such as the almost gesner-like Charadrophila (the common name for this plant is "Cape gloxinia"!) and Scrophulariaceae-Bowkerieae (Bowkeria, Anastrebe and Ixianthes) are now included; Thesmophora was not included in these studies. The characterisation of the family is now greatly changed from what it was ten years ago, and it is unclear what might be apomorphies.

For anatomy and morphology, see Carlquist (1986), Dahgren et al. (1979), Engell (1987), for the general morphology of Charadrophila, see Weber (1989), and for general information, see Linder (2004: the family in a narrow circumscription), Fischer (2004b: some genera under "Scrophulariaceae"), and Tank et al. (2006: composition).

Synonymy: Hallieraceae (Link) Trinius, Retziaceae Bartling

PLANTAGINACEAE Jussieu, nom. cons.   Back to Lamiales

Herbs (shrubs; rooted aquatics); (mannitol, sorbitol +, iridoids 0, cornosides +), little oxalate accumulation; cork?; (leaf endodermis +); hairs with gland head not often vertically divided, (with cystolith); leaves spiral to opposite, simple to compound; (bracteoles 0 - Antirrhineae); corolla often bilabiate (spurred; 0; descending cochleate), stamens (5-8), thecae parallel, end-to-end, or sagittate, confluent [e.g. Penstemon] or not, (staminode + [esp. Cheloneae, Antirrhineae]/0 (2, adaxial pair of A), exine tectate and reticulate, (placentation intrusive parietal) (-1) campylotropous? ovules/carpel, integument 7-22 cells across, stigma (slightly) capitate or bilobed, dry (wet); fruit a septicidal capsule (loculicidal [Veronica]; poricidal [Antirrhineae]; circumscissile); seeds (1-)many, exotestal cells with inner walls ± thickened, when winged, cells with reticulate thickenings; endosperm +, embryo (green); n = 6-10 +; protein bodies in nucleus amorphous [excluding Angelonieae and Gratioleae].

Ca 90[list]/1700: Veronica (ca 450, inc. Hebe, Parahebe, Synthyris, etc.), Penstemon (275: cambium storied, bearded staminode, Wolfe et al. 2006 for a phylogeny), Plantago (275), Linaria (150: tubular protein bodies), Bacopa (55), Stemodia (55), Russelia (50). Mostly temperate (Map: from van Steenis & van Balgooy 1966; Hultén 1971; Meusel et al. 1978; Hong 1983). [Photo - Callitriche Habit] [Photo - Hippuris Habit] [Photo - Flower]

The frequent absence of regular partitions in the heads of the glandular hairs is quite unusual in Lamiales, as is septicidal capsule dehiscence. In a number of taxa in this clade the androecium is initiated before the corolla, but other patterns also occur, so it is perhaps unlikely to be a synapomorphy for Plantaginaceae (Bello et al. 2004, cf. Judd et al. 2002). However, details of the distribution of these and other characters need to be clarified. Thus Lindernieae were until very recently included in Plantaginaceae even though the heads of their glandular hairs are divided by vertical partitions, and taxa like Russelia and even some Penstemon, still in Plantaginaceae, also seem to have similar hairs (Raman 1991 and references). Morphological/developmental synapomorphies for Plantaginaceae may well yet be found.

For feeding preferences of a variety of insect groups suggesting a relationship between the erstwhile Plantaginaceae and Scrophulariaceae, see Airy Shaw (1958) and Allen (1960, 1961); Allen (1960) found different insects eating Plantaginaceae and Scrophulariaceae as distinguished here. Larvae of Nymphalidae-Melitaeini butterflies are commonly found here, and on Orobanchaceae, but not on Scrophulariaceae (Wahlberg 2001). Wilson et al. (2006, see also references) discuss shifts between bee and bird pollination in the speciose Penstemon clade. Some species of Callitriche self pollinate, the pollen tube growing through the tissue of the plant, sometimes even through the stem and then pollinating an adjacent flower (Philbrick & Les 2000). In other species, pollination occurs by pollen drifting through the water, yet others are perhaps wind pollinated.

Philcoxia, a white sand endemic from Brasil, is a remarkable plant, having small peltate leaves borne at about ground level and coming from underground stems; it has been suspected of being carnivorous (Fritsch et al. 2007).

Floral morphology is very variable, leaving aside the aquatic taxa like Callitriche and the wind-pollinated taxa like some species of Plantago. Sibthorpia has 5-8-merous, polysymmetric flowers. Collinsia has a remarkable papilionoid flower, with the distinctively-colored standard being almost lip-like in coloration and formed from the two adaxial petals, the three other petals are plane-coloured, the median abaxial petal forming the keel. Indeed, the overall colour scheme and functional floral morphology is very like that of some species of Lupinus. Aragoa has 4-merous, polysymmetric flowers (cf. Oleaceae and Tetrachondraceae!), but with five sepals. Veronica, to which it is close, has a 4-lobed corolla (but only two stamens); in some taxa there are two main veins in the adaxial lobe, perhaps suggesting that it is formed by the fusion of the two adaxial lobes so common in monosymmetric members of the family. Bello et al. (2004: see Bello et al. 2002 for a phylogeny; Muñoz-Centeno et al. 2006 for seed morphology and phylogeny) discuss floral evolution in this part of the family, emphasising the evolution of polysymmetry, and noting that these genera, as well as Plantago (sister to Aragoa) and Digitalis are members of a clade that has descending-cochleate aestivation. Hardly surprisingly, Plantaginaceae s.l. are very difficult to identify and distinguish from Scrophulariaceae s. str., Gesneriaceae, and even Stilbaceae.

Both Digitalis and Isoplexis have cornosides. Iridoids with an 8,9 double bond are scattered in a number of genera (Jensen et al. 2007); at what level this character might be an apomorphy is unclear. Besseya and Plantago have a foliar endodermis. Veronica lyallii has successive subhypodermal phellogens (Gray 1937). Penstemon may have paired-flower cymes (elsewhere in Gesneriaceae and Calceolariaceae).

Hippuris and Callitriche (sister taxa) and genera like Plantago are also in this clade, so morphological diversification has been very extensive (Olmstead & Reeves 1995; Reeves & Olmstead 1998). Bakker et al. (2006) also found major increases in the rate of evolution of the mitochondrial gene nad1 in Plantago and Littorella (see also Cho et al. 2004).

For the circumscription of the family, which initially had only rather weak support, see Olmstead et al. (2001, as Veronicaceae: inclusion of Cheloneae and Hemimerideae may be the problem, and for the latter, see Scrophulariaceae), Oxelman et al. (2005: support stronger), and Tank et al. (2006, as Veronicaceae). Gratiolaceae were recently recognised as a distinct family by Rahmanzadeh et al. (2004), who discuss phylogeny (only three species examined!) and give a list of included genera, etc. It is possible that Angelonieae (2/27: integument 5-12 cells across) should also be included in their Gratiolaceae, and the widespread Limosella was also part of it. However, the limits and correct placement of this clade clearly need confirmation. Note that Kornhall and Bremer (2004) place Limosella firmly in Scrophulariaceae, although they did not look at other members of this immediate clade - the relationships that they found can be represented as [Myoporum, etc. [Buddleja, etc. [Limosella, Manueleae, etc.]]]. Oxelman et al. (2005) locate the majority of this clade firmly in Plantaginaceae, although Limosella itself is in Scrophulariaceae.

For the phylogeny of Antirrhineae, see Ghebrehiwit et al. (2003), for Veroniceae, see Albach et al. (2004) and Taskova et al. (2004: the "new" molecular relationships are at least sometimes supported by other data such as chromosome number and iridoid type), while Pedersen et al. (2007 and references) and Jensen et al. (2008) report on some chemistry of ex-Hebe. Albach (2008) discusses the limits of Veronica s.l.

For chemistry, see Jensen (2005) and Taskova et al. (2006), for Trapella, see Oliver (1888), and for a general survey, see Thieret (1967). Additional information is provided by Junell (1961: gynoecium), Sutton (1988: monograph of Antirrhineae), Leins and Erbar (2004a: Hippuridaceae), Erbar and Leins (2004b: Callitrichaceae), Schwarzbach (2004: Plantaginaceae), Ihlenfeldt (2004: Trapellaceae), Fischer (2004b: Scrophulariaceae p. pte) and Wagenitz (2004: Globulariaceae). Albach et al. (2005) discuss the circumscription and phylogenetic relationships of the family and the variation that it encompasses. I thank Dirk Albach for comments.

Synonymy: Antirrhinaceae Persoon, Aragoaceae D. Don, Callitrichaceae Berchtold & J. Presl, nom. cons. (water plant; hairs stellate; staminate flowers; P 0, A 1(-3), pollen trinucleate; carpellate flowers: G [2], transverse, subdivided, 2 ovules/carpel, integument thin, styles ± separate, slender; fruit schizocarpic, with 4 nutlets; exotesta perhaps persisting), Chelonaceae Martynov, Digitalidaceae Martynov, Ellisophyllaceae Honda, Erinaceae (Duvau) Pfieffer, Globulariaceae Candolle, nom. cons. (cork epidermal to pericyclic; leaves conduplicate; 1 or 2 ovules/carpel, one pendulous and one erect, or [Globulariaceae s. stricto] only the abaxial carpel developed, stigma dry; n = 8-10, 19), Gratiolaceae Martynov (integument 3-6 cells across, endothelial cells large, transversely elongated, in vertical rows; seeds with longitudinal ridges; extotestal cells with hook-like thickenings), Hemimeridaceae Doweld, Hippuridaceae Vest, nom. cons. (water plant; flavones 0; hairs peltate, glandular; leaves whorled; plant monoecious, flowers polysymmetric ; K a 2-4-lobed or entire rim, C 0, A 1, pollen trinucleate, G 1, inferior, ovules 4, pendulous, apotropous, chalazogamy occurs, stylodium stigmatic its entire length, stigma dry; fruit achenial or drupaceous; testa structureless, endosperm thin, starchy; n = 16), Linariaceae Martynov, Littorellaceae Gray, Oxycladaceae (Miers) Schnizlein, Plantaginaceae s. str. (sorbitol +; cambium storied; nodes 3:3; leaves spiral, margins toothed to entire; flowers polysymmetric, (3-)4-merous; G [2], placentation parietal, or unilocular, ovule 1, basal, epitropous, stylodium +, stigmatic its entire length, stigma dry; fruit circumscissile or nut-like, exotestal cells large, mucilaginous, endotestal cells persist; endosperm +, (embryo curved). Plantago has substitution rates at synonymous sites in the mitochondrial genome that are 3,000-4,000 times those of nearly all other angiosperm clades [Cho et al. 2004]), Psylliaceae Horaninow, Scopariaceae (Link) Trinius, Sibthorpiaceae D. Don, Trapellaceae Honda & Sakisaka (rhizomatous aquatic herb; flower slightly monosymmetric, A 2 [the adaxial pair], staminodes 2, connective apically peltate, G inferior, the abaxial loculus much reduced, 2 pendulous ovules/carpel, stigma unequally bilabiate; fruit indehiscent, K persistent, with an appendage from below the apex of the sepals; testa with thin-walled cells; n = ca 25), Veronicaceae Cassel

LINDERNIACEAE Borsch, K. Müller, & Eb. Fischer   Back to Lamiales

Ephemerals to suffruticose perennials; iridoids 0; cork?; nodes 1:3; leaves opposite (basally connate), venation also pamate, margins entire or serrate; inflorescence racemose or flowers from the axils of leaves; glandular hairs on the inside of the C, A 4, or A 2 + 2 large abaxial Z-shaped staminodes with an appendage, or 2 much reduced staminodes, or A 2, pollen 3-colpate, embryo sac exserted from the ovule, stigma bilobed, sensitive; capsule septicidal or -fragal; seeds with ruminate endosperm; n = ?

Ca 13/195: Lindernia (100), Torenia (40). Pantropical to warm temperate, mostly New World (Map: based on Fischer 1992; Lewis 2000).

• Many Linderniaceae are rather small herbs with opposite leaves, square stems, and monosymmetric flowers with a sensitive stigma and the lower stamens usually very different from the upper, either Z-shaped or long and curved, with a projection, or simply very much reduced.

In some species the anthers of the abaxial stamens are yellow and lie on the abaxial lip; they appear to contribute to the attractive aspect of the lip. In other species the long, curved filaments are joined by the connate anthers and form a sort of balustrade across the mouth of the corolla. It would be interesting to know details of pollination mechanisms for such flowers.

The nodes appear to be 1:3, rather than 3:3 as I originally thought. Small strands of lignified tissue are associated with the sharp ridges of the stems. The glandular heads of the hairs on the corolla and the vegetative plant have vertical partitions, as is common in Lamiales. Lewis (2002) suggests that the stamens are extrorse and the ovules are straight [atropous]; Fischer (1992), however, gives a floral diagram showing introrse anthers and describes the ovules as being anatropous to hemitropous. The integument is up to about four cells thick. The embryo sac protrudes beyond the micropyle in both Torenia and Lindernia, at least (the synergids can then be ablated easily in studies of fertilization: Higashiyama et al. 2006). The rumination of the endosperm is caused by inpushings of endothelial cells; these may become confluent and the seeds may then have longitudinal ridges. The seed coat is otherwise pitted or smooth.

Of four genes analysed, only one did not suggest separation of this group from Plantaginaceae when analysed separately; the joint analysis also supported separation (Albach et al. 2005). Micranthemum, with only two stamens, was the rather unexpected sister taxon to Lindernieae, whose members make up the rest of this clade, but it was included neither in the two gene analysis of Rahmanzadeh et al. (2004), nor in the list of genera that they included in their Linderniaceae. However, Micranthemum is sister to Torenia, the two in turn are sister to Stemodiopsis, the only three Linderniaceae included in the analysis (Oxelman et al. 2005); see Tank et al. (2006) for a summary of our ideas of relationships within this clade, and also of its composition.

Information is taken from Fischer (1989, 1992: general), Rahmanzadeh et al. (2004: relationships, genus list, etc.), and Albach et al. (2005: relationships, etc.).

BIGNONIACEAE Jussieu   Back to Lamiales

Trees, shrubs or vines; iridoid aldehydes +; cork also cortical; cambium storied; (vessel elements with scalariform perforations); nodes 1:1-3 or more; petiole bundles annular (also rib or adaxial bundles); stomata helicocytic [?level]; leaves bicompound, (spiral; simple), conduplicate (involute - Pyrostegia), margins entire (toothed); flowers large, K often with nectaries, A 4 (5, 2), thecae sagittate or head-to-head, usu. not confluent, tapetum amoeboid, staminode usu. +, pollen tricolpate, psilate, nonperforate, bundles in the ovary wall and also opposite septum, ovules in two groups in each loculus, (placentae lobed), nucellar endothelium +, stigma lobes broad, often sensitive, wet; fruit often with nectaries; seeds many, winged (wingless); cells in wings with helical or annular (none; reticulate) thickenings; endosperm 0; n = 20; seedings epigeal and phanerocotylar (cryptocotylar), cotyledons obcordate, lobed, persistent.

110[list]/800 - four tribes below. Mainly tropical, esp. South America (Map: from van Steenis 1977). [Photos - Amphitechna Flower, Distictella Flower.]

1. Jacarandeae Fenzl

Trees and shrubs; staminode large, bearded; n = 18.

1/50: Jacaranda. Tropical America.

2. Tourretieae K. Schumann

Tendrillar vines; inflorescence a bracteate raceme; staminode 0 [Tourretia].

2/6. Andes in South America and N. to Mexico. [Photo - Eccremocarpus Flower.]

Tourretia has sub four-locular ovaries each with a single rank of ovules, while Eccremocarpus has parietal placentation.

3. Bignonieae Dumortier

Tendrillar lianes; anomalous secondary thickening + [basal condition is for the xylem cylinder to be 4-lobed]; leaves usu. ternate; fruit usu. septifragal, with persistent septum and separate whip-like strands of woody tissue [= vascular bundles opposite septum].

21/360: Adenocalymma (78), Arrabidea (69), Anemopaegma (42), Amphilophium (41). America, largely tropical.

Perianthomega is sister to the rest of the tribe. It has biternate leaves, also, it has robust unbranched tendrils that represent petioles, and three small scars (leaflets!) can be seen at their ends. A number of Bignonieae with septifragal dehiscence also have cracks in the loculicidal position along the backs of the valves.

For a comprehensive (2-gene + morphology) phylogeny of the tribe, see Lohmann 2006a). Major clades are supported by a mixture of floral and vegetative characters, and generic limits have been extensively reworked (Lohmann 2002, 2003, esp. 2006a, b; the latter lists all species accepted in the tribe).

Bignonieae may be close to Oroxylum and relatives (leaves bicompound; capsules septicidal) and Catalpa (Olmstead et al. 2002). Catalpa has only two stamens.

4. The Rest

Trees and shrubs, lianes; (leaves palmate - Tabebuia; simple, phyllodinous or not); (A 5; 2 + 3 staminodes), (fruit indehiscent).

66?: 384: Tabebuia (100: good wood). Tropical.

Synonymy: Crescentiaceae Dumortier

• Bignoniaceae can be recognised quite easily. Vegetatively, they are woody and with lenticillate stems, and most have opposite, compound leaves; the New World vines and lianes often have leaf tendrils. The often rather large flowers are monosymmetric, and there are often nectaries on the leaf, stem (at the nodes), calyx and even the ovary surface. The calyx is often spathaceous (Old World taxa), there are two series of ovules in each carpel, and the broad, almost spathulate stigmatic lobes are often sensitive. The seeds are usually flattened and winged.

The divergence time of Bignoniaceae and Verbenaceae has been estimated at ca 49.5 million years before present (Nie et al. 2006). Bignoniaceae are, along with Sapindaceae, the most important neotropical group of lianes; Bignonieae in particular are nearly all lianes.

Palmate leaves may have arisen independently within Bignoniaceae, but are known only from New World taxa. Amphitecna has spiral, simple leaves as does Crescentia; their petioles are short and the lamina has distinctive venation. Such leaves may be phyllodinous, indeed, in some species of Crescentia palmately-arranged leaflets are borne on the end of a lamina-like petiole. Relationships between the New World Tabebuia and its relatives have recently been clarified. Amphitecna and Crescentia, both with spiral, simple leaves, are probably derived; Crescentieae are embedded in a paraphyletic Tabebuia, with palmately compound leaves (Grose & Olmstead 2002, 2007a [phylogeny], b [nomenclature]). Associated with these two genera are Parmentiera and Spirotecoma, also with palmately compound leaves; all four genera have bat-pollinated flowers. Amphitecna and Crescentia (calabash) have indehiscent fruits, Parmentiera has fleshy fruits, although its seeds still have a small wing, Spirotecoma dehiscent fruits. The petiolate leaves of Catalpa (opposite or whorled leaves)/Chilopsis (spiral leaves), which hybridise, have a very different morphology; they appear to be more conventionally simple.

There has been much discussion of pollination and seed dispersal in Bignoniaceae, indeed, over-reliance on characters associated with pollination and dispersal syndromes as markers of generic distinctness has caused serious problems with generic limits in the past (see Lohmann 2003, 2006a, b). The large flowers of Bignoniaceae are animal pollinated, and show considerable variation in details of floral morphology and flowering phenology which affects the behaviour and type of visitor (Gentry 1974a, b, 1990). One of the coomonest flower types in the New World is the Anemopaegma type, visited by euglossine bees (along with anthophorids), and the sometimes nectarless Cydista type, also visited by euglossines. Parmentiera has bat-pollinated flowers and much-elongated fleshy fruits, although its seeds have a small wing; Crescentia has a rather similar complex of floral and fruit characters, although its fruits are spherical. The pinnately-compound Kiggelaria also is bat-pollinated and with massive, sausage-shaped, indehiscent fruits; Oroxylon has three to four times compound leaves and is also pollinated by bats, its flowers being almost polysymmetrical and with five stamens; however, it has capsules and wind-dispersed seeds. Dispersal syndromes are also quite diverse (Gentry 1983; 1990). A number of taxa have seeds dispersed by water, including Dolichandrone, a mangrove plant; the modified seed wing is corky and becomes a flotation device.

Aliform-confluent xylem parenchyma is common. There are four main carpel bundles, but only two in the "Scrophulariaceae" (Armstrong 1985), Gesneriaceae, etc.

The basic phylogenetic structure within the family seems to be [[[Jacarandeae [Tourrettieae [Bignonieae + the rest]]] (Olmstead et al. 2002).

For information on pollen, which is very variable, see Gentry and Tomb (1979), for wood anatomy, see Gasson and Dobbins (1991: lianes and the rest compared), for protein bodies in the nucleus, see Bigazzi (1995), for tapetum, Huysmans et al. (1998), for iridoids, von Poser et al. (2000), for general information, Manning (2000) and Fischer et al. (2004a: the classification, including the generic limits recognised, is very "classical", cf. e.g. Lohmann 2006b), and for seed anatomy, including that of Schlegliaceae and Paulowniaceae, see Lersten et al. (2002). There is a species level checklist for the family. I am grateful to L. Lohmann for comments.

VERBENACEAE Jaume Saint-Hilaire, nom. cons.   Back to Lamiales

Vines, trees, or herbs; 4-carboxy-iridoids +, (ethereal oils + - Lantaneae); (pits vestured); petiole bundles arcuate (also medullary, associated with median bundle); needle crystals common; stomata dia(anomo)cytic; stems often square; eglandular hairs unicellular hairs; leaves simple, margins toothed to deeply lobed; inflorescence racemose (capitate); flower weakly bilabiate, A didynamous, (or of two lengths, but free; A 5 - Verbena), staminode 0 (+), tapetal cells 2-4-nucleate, pollen (colpate, por(or)ate), exine thickened near apertures, G [2 (4 [Duranta])], transverse (only 1 fertile), 2 erect (descending) apotropous ovules/carpel, stigma bilobed, with conspicuous stigmatoid tissue, wet; fruit a schizocarp or drupe with 1, 2 or 4 stones, K persistent; testa thin-walled; endosperm ± 0 (+); n = 5-12+.

34[list]/1175: Verbena (200-250), Lippia (200), Lantana (150), Citharexylum (130), Glandularia (100), Stachytarpheta (90), Junellia (50). Pantropical (to warm temperate), but mostly New World (Map: from van Steenis & van Balgooy 1966; Hultén 1971; Meusel et al. 1978; Brummitt 2007). In Europe, Verbena officinalis may be native only from S. Europe eastwards; is Phyla nodiflora native to Australia? [Photo - Flower]

• Verbenaceae are shrubs or trees that may be recognised easily because the plants are often aromatic, the opposite leaves are serrate, the stem is often angled and with a line across the node, and the often rather weakly monosymmetric flowers with a trumpet-shaped corolla and more or less included anthers are borne in racemes or heads. The stigmatic area is conspicuously asymmetically swollen and glandular and the ovary has four ovules.

The endothelium seems to be poorly developed (Johri et al. 1992). For the position of the carpels, see Sattler (1973); the ovules are described as being attached to the false septae (see Junell 1934). Two-chambered mericarps or stones may contain ovules from both carpels... (Sanders 2001).

Verbenaceae as currently circumscribed (see esp. Cantino 1992a, b) are much reduced compared to a decade or so ago, and many genera have been placed in Lamiaceae. The two families are now more easily distinguishable than before! For Avicennia, also once included in Verbenaceae, see Acanthaceae; Phrymaceae are also separate and are not immediately related, although included in Verbenaceae by Cronquist (1981) and others in the past, in part because they have a similar racemose inflorescence. Verbenaceae s. str. may be in a different part of Lamiales from Lamiaceae, or they may be sister to them (e.g. Olmstead et al. 2001, but only one member of each sampled: see Wagstaff & Olmstead 1997 and Cantino 2004 for more information). Within Verbenaceae, Petraea and Duranta, both woody, are successively sister to the rest of the family (Marx & Olmstead 2007).

For hairs and stomata, see Cantino (1990), for the megagametophyte, see Rudall and Clark (1992), for exine thickening, see Chadwell et al. (1992), for iridoids, see von Poser et al. (1997 - also Soltis et al. 2005b), and for general information, see Sanders (2001), Atkins (2004) and Brummitt (2007).

Synonymy: Durantaceae J. Agardh, Lantanaceae Martynov (exine thickened near apertures, 1 erect ovule/carpel; endosperm + - does this really belong?), Petraeaceae J. Agardh (in some molecular phylogenies sister to Bignoniaceae - Wagstaff & Olmstead 1997)

LAMIACEAE Martynov, nom. cons.//LABIATAE Jussieu, nom. cons. et nom alt.   Back to Lamiales

Herbs (trees, vines); diterpenoids [Symphorematoideae?], betaines, C4-decarboxylated iridoids +; cork also deep-seated; (pits vestured); (nodes 1:2); petiole bundles arcuate (annular); stomata dia(anomo)cytic; stem often square; eglandular hairs (unicellular), uniseriate (stellate); leaves simple or palmately compound, ptyxis variable, margins toothed; A didynamous (2 [e.g. Salvia]), staminode 0 (+), tapetal cells multinucleate, pollen 3-colpate, 2-nucleate, exine not thickened near apertures, G [2(-5)], 2 ± erect epitropous ovules/carpel, ovules on inner side of placenta/carpel margin, style (unequally) bifid, stigma inconspicuous, not expanded, (wet) dry; fruit a schizocarp, berry or drupe, K persistent or accrescent; exotestal cells elongated or not, thickened on radial and often inner walls (hypodermal cells sclerenchymatous).

236[list]/7173 - 7 subfamilies below. World-wide (Map: from Vester 1940; Hultén 1971; Van Balgooy 1975). [Photos - Collection] [Photo - Fleshy fruit]

1. Symphorematoideae Briquet

Lianes; inflorescences in 3-7-flowered capitate cymes with an involucre of bracts; flowers polysymmetrical, K 5-8, C 5-16, A 4-18, disc 0, G imperfectly 2-locular, ovules apical, straight [atropous]; ?endosperm; n = 12, 14, 17, 18.

3/27. India, Sri Lanka, South East Asia, Malesia.

Synonymy: Symphoremataceae Reveal & Hoogland

2. Viticoideae Briquet

Often woody; (hairs branched); (leaves compound); disc 0 or poorly developed; fruits drupes.

10/376-526: Vitex (250), Premna (50-200). Tropical, esp. South East Asia-Australia.

Note that the circumscription of this subfamily is more narrowly drawn than in Cantino et al. (1992), some genera included there not being assigned to subfamilies.

Synonymy: Viticaceae Jussieu

3. Ajugoideae Kosteletzky

(Aromatic, no terpenoids, etc.); flowers (4 [Aegiphila] merous), polysymmetric or 1-lipped, exine with branched (simple, granular, etc.) columellae, disc slight-0 (+); endosperm several-layered, 0; n = 7, 10, 13, 14, 16+.

24/1115: Clerodendrum (400-500, but to be divided [Steane et al. 1999, 2004]), Teucrium (250: lip sometimes 5-lobed), Aegiphila (120), Rotheca (50-60), Ajuga (40-50). Cosmopolitan, but many temperate, and esp. South East Asia to Australia.

Synonymy: Aegiphilaceae Rafinesque, Siphonanthaceae Rafinesque

4. Prostantheroideae Luersson

(Aromatic); flowers polysymmetric (monosymmetric), 4-8 merous, (staminodes 2), (disc 0); endosperm +; n = ?

16/317: Prostanthera (100), Hemigenia (50), Pityrodia (45). Australia.

Synonymy: Chloanthaceae Hutchinson

5. Scutellarioideae (Dumortier) Caruel

(Aromatic, no terpenoids, etc.); stem endodermis +; K two-lipped (not Holmskioldia), lobes rounded; seeds tuberculate; endosperm various; n = 12+.

5/380: Scutellaria (360). ± Cosmopolitan.

Synonymy: Salazariaceae F. Barkley, Scutellariaceae Caruel

6. Lamioideae Harley

(Aromatic), laballenic fatty acid +; stem endodermis +; (stamens 4, about the same length - Pogostemon and relatives), embryo sac with micropylar lobe longer and broader than chalazal lobe, style gynobasic; endosperm several-layered; n = 6+.

63/1210: Stachys (300), Sideritis (140), Leucas (100), Phlomis (100), Pogostemon (80), Eremostachys (5-60). Esp. Europe to Asia, some cosmopolitan, but v. few Antipodean.

For phylogenetic relationships, see Wagstaff et al. (1995), and for relationships of mints endemic to Hawaii, see Lindqvist and Albert (2002).

Synonymy: Melittaceae Martynov

7. Nepetoideae Kosteletzky

Commonly aromatic [volatile terpenoids, rosmarinic acid], nepetoidin A and B [caffeic acid esters] +, betaine concentration low, iridoid glycosides, acteosides 0 (+); stem endodermis +; pollen trinucleate, hexacolpate, style gynobasic; exocarp with mucilaginous cells producing hygroscopic spiral fibrils; endosperm 1-layered, cotyledons investing embryo; n = 6+.

105/3675: Salvia (900+: A 2, unithecate, connective expanded forming a lever arm involved in pollination - but possibly para- or polyphyletic, Rosmarinus, Thymus, Mentha and Origanum also involved [Walker et al. 2004] - alas for "Scarborough fair"; see Claßen-Bockhoff et al. [2004a, b] for stamen development. Walker and Sytsma [2007: only Rosmarinus was part of the clade in this study] suggested that the distinctive stamen with its lever arm might have evolved three times.), Plectranthus (300: inc. Coleus), Hyptis (280), Thymus (220), Nepeta (200+), Clinopodium (100), Isodon (100), Micromeria (70), Ocimum (65), Platostoma (45), Aeollanthus (40), Hedeoma (40), Lepechinia (40), Origanum (40), Pyconostachys (40). World-wide, but esp. (warm) temperate.

Nepetoideae include the large tribe Ocimeae with synthecous, dorsifixed anthers (Paton et al. 2004). Wunderlich (1967b) suggests that there is no endosperm. Myxocarpy is common (Ryding 1992). Moon et al. (2008) surveyed pollen morphology especially of Salviinae.

Synonymy: Glechomaceae Martynov, Mellitidaceae Martynov, Menthaceae Burnett, Nepetaceae Horaninow, Salviaceae Rafinesque

And 10 genera unassigned, including Callicarpa (140: hairs branched/stellate), Tectona (4).

• Lamiaceae are herbs, also shrubs and lianes, that may be recognised by their opposite, usually serrate leaves, more or less square stems, and monosymmetric flowers borne in often clearly cymose clusters. The plants are often aromatic. The often gynobasic style is apically bifid, the two lobes being sharply pointed, and the four ovules of the bicarpellate gynoecium are borne in uniovular compartments. The calyx is an integral part of the dispersal mechanism and is or more or less accrescent; it may be fleshy, or distinctively ridged and dry. Woody members often have strongly lenticillate twigs.

The leaf beetle Phyllobrotica (Chrysomelidae) eats plants from Scutellarioideae, Lamioideae and Viticoideae, but not members of Nepetoideae or Verbenaceae (Farrell & Mitter 1990). Larvae eat the roots, adults the above-ground parts, which they can decimate.

Bootstrap support for the family as circumscribed is 100% (Wagstaff et al. 1998); Congea may be sister to the rest, but some relationships are still in a state of flux. Wenchengia has spiral leaves; it is unclear where it should be placed (Cantino & Abu-Asab 1993).

Many Lamiaceae have a single layer of sclerenchymatous, bone-shaped cells on the inside of the mesocarp, others have thicker pericarp walls, and the cells are often crystalliferous. Pericarp anatomy of Verbenaceae is more complex (Ryding 1995). There may be differences in seed coat anatomy: the testa of at least some Verbenaceae has the hypodermal layer(s) thickened, while in Labiatae it is the exotestal cells that are thickened, particularly on their inner periclinal and anticlinal walls (Rohwer 1994a). The ovules are described as being attached (just) to the false septae.

Only trisaccharide esters of verbascoside are found in Lamiaceae, with disaccharides they are found especially in Verbenaceae, Oleaceae and Orobanchaceae (Mølgaard & Ravn 1988). Some Lamiaceae have asymmetric development of the endosperm such that the two haustoria come to lie very close to each other (Ram & Wadhi 1964 for references). This pattern of development may be restricted to Nepetoideae (further studies are needed), but it is also to be found in many Acanthaceae.

As their distinctive alternative name Labiatae implies, Lamiaceae have always been considered as an "eminently natural" family, being immediately recognisable because of their paired, serrate leaves, square stems, monosymmetric flowers, gynobasic style, and four nutlets. However, the gynobasic style and the four nutlets have evolved more than once (Cantino 1992a), and a considerable number of ex-Verbenaceae must now be included in Lamiaceae (Junell 1934; Cantino et al. 1992a, b: see Verbenaceae for further discussion). Lamiaceae and Boraginaceae have always been considered distinct, but their similar gynobasic styles and fruits with four separate nutlets (and also some chemistry) have invited comparisons between the two, and they are often placed fairly close to each other (e.g. Cronquist 1981: both are in a rather narrowly drawn [compared to here] Lamiales). However, there are numerous differences (oher chemistry, leaf insertion, floral symmetry, ovule morphology, etc.) between the two, and the radicle in Boraginaceae points upwards in fruit, while in Lamiaceae it faces downwards.

For gynoecial morphology and embryology, see Junell (1934: note that there is variation in ovule attachment within the family), for pollen, ovules and seeds, see Wunderlich (1967b), for variation in the proteinaceous inclusions in the nucleus, see Speta (1979), for hairs and stomata, see Cantino (1990), for megagametophyte, see Rudall and Clark (1992), for betaine distribution, see Blunden et al. (1996: widespread, but what about Verbenaceae and other Lamiales?), for secondary metabolite evolution, see Grayer et al (2003) and Wink (2003), and for a comprehensive general treatment, see Harley et al. (2004).

SCHLEGELIACEAE Reveal   Back to Lamiales

Large trees, woody shrubs, vines or epiphytes; pericyclic sheath sclereidal; nodes 1:3; petiole bundle solid-(almost)annular, with wing bundles, no pericyclic lignification; sclereids +; stomata variable; leaves entire or serrate; flowers quite large, nectaries on outside of K, staminode +/0, nectary +/0, vascularised from carpellary bundles; fruit a berry, K persistent; exotestal cells with scalariform thickenings on the inner periclinal wall or mucilaginous with outer periclinal wall absent; endosperm +/0, embryo stout, cotyledons slightly over half its length; n = 20; seedings epigeal and phanerocotylar, cotyledons lobed.

4[list]/28: Schlegelia (15), Gibsoniothamnus (11). Mexico to tropical South America, Cuba (Synapsis) (Map: from Gentry 1980). [Photo - Flower, Flower, Fruit.]

• Schlegeliaceae may be recognised by their woody habit, white bark, opposite, often thick leaves at least sometimes with small surface glands on the lower surface, quite large and monosymmetric flowers, and berries.

Schlegelia and relatives are usually included in Bignoniaceae - and the two may indeed be close (Olmstead et al. 2001), although there are no obvious differences in wood anatomy (Gasson & Dobbins 1991) - or in Scrophulariaceae s.l. Schegelia may have anomocytic or paracytic stomata, while the stomata of Gibsoniothamnus are anisocytic or cyclocytic. The glands on the lower surface of the lamina are hairs with the normal lamialean structure or radially-arranged cells in the head. Gibsoniothamnus may be anisophyllous (cf. Thomandersia!). Winged seeds have been reported for the family, but the combination of winged seeds and baccate fruits seems at improbable, unless in the Cuban genus, which I have not seen.

Some information is taken from Leinfellner (1973: gynoecium of Schlegelia), Armstrong (1985), Burger and Barringer (2000), Barringer (2004: Gibsoniothamnus), and Fischer (2004b: under Scrophulariaceae). Gibsoniothamnus parvifolius: Herrera 672, - leaf, stem, seed; G. allenii: McPherson 11069 - leaf, seed; Schleglia darienensis: Neill et al. 11411 - seed.

THOMANDERSIACEAE Sreemadhavan   Back to Lamiales

Shrub or small tree; 2-indolinone alkaloids +; phloem stratified; pericyclic fibers ?short, massively thickened; nodes 1:3; petiole bundles forming a ring or incurved C-shaped; stomata anisocytic; lamina (deeply lobed), with flat glands abaxially, petiole swollen apically and basally; inflorescence racemose; K with nectaries on the outside, staminode +, pollen 5-6-colpate, nectary vascularised by carpellary traces, gynoecial vasculature 8-shaped, 1-3 hemianatropous ovules/carpel; fruit with "jaculators" [cup-shaped expansion of funicle], K accrescent; seed with rather large hilum; seed coat with ascending-imbricate scales or warts, exotesta palisade, not lignified, up to 6 layers of cells in the warts; endosperm 0, embryo strongly curved, cotyledons thin-foliaceous, complexly folded; n = ?.

1/6. W. and C. Africa (Map: from Wortley et al. 2007).

• Thomandersiaceae are woody plants with opposite, usually entire leaves the lower surface of which often have flat, black-drying glands; the inflorescence is a raceme of rather small flowers and there are extrafloral nectaries on the outside of the calyx. The septicidal, capsular fruits are distinctive with their accrescent calyx and few seeds each sitting on a thin, cup-shaped expansion of the funicle; the surface of the seed is warty or scaly, the hilum is large, and the embryos are strongly curved.

The flat glands in the characterization above are dark-drying, rounded, and up to 3 mm across, and are quite different from the lamialean glands which also often occur on the abaxial surface of the lamina. I am not sure exactly how the cotyledons are folded. Despite the presence of structures described as jaculators, fruit dehiscence is not explosive, unlike Acanthaceae; the seed, with its prominent hilum, sits in a thin, cup-like expansion of the funicle. The organised part of the seed coat described above is above a layer of apparently much crushed cells, in turn above a layer of a few less densely-crushed cells; the outer layer of the endoperm has a distinct outer periclinal cell wall. Ovule and seed anatomy need further study.

Thomandersia, from tropical Africa, previously usually included in Acanthaceae, may go somewhere near Schlegeliaceae, from tropical America. If this is to be recognised as a separate family, the name Thomandersiaceae is available, but if there are characters linking it with Schlegeliaceae (see the vasculature of the floral nectary and petiole, also the nectaries on the outside of the calyx) then a single family may be more appropriate - and I think Thomandersiaceae is the earlier name. However, support for any Thomandersiaceae-Schlegeliaceae association is currently weak (Wortley et al. 2007).

For alkaloids, see Ngadjul et al. (1995), and for general details, see Wortley et al. (2005a and especially 2007). Thomandersia hensii: de Wilde & Jongkind 9400, seed, stem; Ngok Bamak et al. 1263, leaf; T. laurifolia: Dibata 30, seed; Thomandersia sp.: Reitsma et al. 1819, leaf, stem.

PHRYMACEAE Schauer, nom. cons.   Back to Lamiales

Annual or perennial herbs (woody); cork ?; vessel elements ?; leaf margins toothed; inflorescence racemose; K tubular, toothed, subplicate-ribbed (4-, 3-lobed), C 2 + 3 (polysymmetric; 2 + 0 - Mimulus douglasii), A (2) 4, anthers subreniform, thecae confluent, (pollen <10-colpate; each colpus with 2 orae; spiraperturate, etc.), nectary +/0, (1<) near-basal to axile (parietal) straight [atropous] ovules/carpel, integument 3-5 cells across, stigma broadly 2-lobed (1-lobed; shortly 2-fid), sensitive; (fruit indehiscent), K persistent; endosperm +, cotyledons folded?; n = 7-10, 14, 22, etc.

Ca 19, but probably fewer[list]/234: Mimulus (150-170), Mazus (30). ± World-wide, esp. temperate and W. North America and Australia, but few humid tropics (Map: from Meusel et al. 1978; Barker 1982; Hong 1983, 1993). [Photos - Collection, Mimulus Flower.]

Phryma diverged from other Phrymaceae 52.5-28 million years before present (Nie et al. 2006), although its well-known East Asian E. North American disjunction is much more recent, ca 6-2 million years before present.

The Australian Glossostigma is scarcely bigger than Lemna, while small plants of Mimulus jepsonii may consist only of cotyledons, a pair of foliage leaves, and a flower (T. Livschultz, pers. comm.).

Phryma itself has opposite, serrate leaves and spicate inflorescences bearing small, monosymmetric flowers. The calyx persists in the secund fruit, the three adaxial members forming recurved spines, and the fruit has but a single seed - a very derived morphology. Whipple (1972) described the nodes as having three traces coming from a single gap; the ovules were described as being apotropous and hemitropous.

Phryma and Mimulus, into which should perhaps be placed about six genera or more, and its relatives, makes up this unexpected clade (Beardsley & Olmstead 2000, esp. 2002; Beardsley et al. 2001, 2004; Beardley & Barker 2004). However, there is still some doubt as to whether genera like Mazus are to be included, or not (Oxelman et al. 2005; Tank et al. 2006). There may also be chemical differences within Phrymaceae s.l., thus Mazus has iridoids while Mimulus does not (Hegnauer & Kooiman 1978).

For pollen, see Argue (1980, 1981) and Chadwell et al. (1992), for a monograph of Mimulus sensu stricto, see Grant (1924) and Thompson (2005), for Phrymaceae s. str., see Whipple (1972), Ramana et al. (2000: embryology), and Cantino (2004: general), for other genera included here, see Fischer (2004b: Scrophulariaceae p. pte).

Rehmannia   Back to Lamiales

This is a small genus of ca 6 species from China and Korea. Although previously included in Gesneriaceae or Scrophulariaceae, Oxelman (2005) found that it linked very weakly with Phryma, Paulownia Mazus and Lancea, and genera of Orobanchaceae. Albach et al. (2007) found that Rehmannia had iridoids, at best very uncommon in Gesneriaceae, and at least some mannitol, not occuring in Scrophulariaceae. In a restricted phylogenetic analysis, Rehmannia linked with Oreosolen (Albach et al. 2007), earlier placed in the Scrophulariaceae s. str. clade (Oxelman et al. 2005), but this may be a rooting problem.

ACANTHACEAE Jussieu, nom. cons.   Back to Lamiales

Quaternary methylammonium compounds, amyloid +; (cork deep seated); (intraxylary phloem +); stomata diacytic; leaf margins entire to toothed; (inflorescence racemose), bracts and/or bracteoles often large; K free or connate, often sharply pointed, (C lobes narrow), A didynamous (2, + 2 staminodes; 5), staminode +/0, G lacking septal bundles, integument thin; K persistent; endosperm development asymmetrical [the two haustoria come to lie close to each other], embryo often ± curved.

229[list]/3500 - four groups below. Mostly tropical.

1. Nelsonioideae (Nees) Sreemadhavan

Herbs; gland-headed hairs with 2-celled heads; leaves opposite to spiral, margins?; inflorescence racemose, (bracteoles 0 - Nelsonia); adaxial lobes outside others in bud [aestivation ascending cochleate], stamens 2 or 4, anthers variable, (placentation parietal - Elytraria), (-4) ovules/carpel, funicular obturator +, stigma broadly (unequally) lobed; seeds 2-many, ruminate, testa ± disorganised; endosperm +, oily; n = 9.

6-7/170: Staurogyne (140). Tropical.

Nelsoniaceae have often been placed in Scrophulariaceae s.l. or considered "intermediate" between Scrophulariaceae and Acanthaceae, but they are placed sister to rest of Acanthaceae s.l. in Hedren et al. (1995). Characters of the group suggested by Scotland and Vollesen (2000) - no retinacula or cystoliths, descending cochlear aestivation (i.e. the adaxial petals overlapping the abaxial petals in bud) - are likely to be plesiomorphies. There is some dispute as to whether Nelsonioideae have jaculators, but even if present, they certainly are not functional.

Some information is taken from Bremekamp (1955) and in particular Johri and Singh (1959).

Synonymy: Nelsoniaceae Sreemadhavan

Acanthoideae + Thunbergioideae + "Avicennioideae": (inverted vascular bundles in the pith); acicular fibers +; (abaxial C lobes outside others), pollen other than colporate [?level], ovules collateral, endothelium 0; amyloid in cotyledons +, endosperm 0.

This group appears to lack a funicular obturator, but I am uncertain as to the polarity of this feature.

2. Acanthoideae Link

Herbs (to shrubs); cystoliths + (0); petiole bundles arcuate, arranged in a circle (annular); (C contorted, abaxial lobe outside others in bud [descending cochleate], etc.), anthers sagittate, or thecae displaced and not opposite, (one theca ± reduced), pollen hideously variable, often porate, stigma dry, usu. bifid; capsules explosive, obovoid; seeds 2-few, flattened, (hairy), borne on hook-like hardened funicles [jaculators, retinaculae]; exotesta palisade, often mucilaginous when wetted [hygroscopic hairs], (hypodermal cells thickened); cytologically very variable.

217/3220: Justicia (600), Barleria (300), Strobilanthes (250), Ruellia (250: defined by pollen, many taxa are cleistogamous, includes things like Blechum, etc. - Tripp & Manos 2006; Tripp 2007), Aphelandra (170 - for bird pollination, see McDade 1992); Blepharis (130), Asystasia (70), Ptysiglottis (60). World-wide; the bulk of the family. There are more species in the New World, more genera(!) in the Old World. [Photo - Habit, Flower.]

[Acantheae [[Whitfieldieae (lenticular biporate pollen with a distinctive circumferential band ["Gürtelpollen"]) [Barlerieae (quincuncial corolla aestivation) + Andrographidae]] [Justicia + Ruellia)) - the individual genera here in fact represent groups of genera - seems a possible set of relationships within Acanthoideae (McDade & Moody 1999; McD