EXTANT SEED PLANTS

Plant woody, evergreen; nicotinic acid metabolised to trigonelline, (cyanogenesis via tyrosine pathway); primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins rich in guaiacyl units; true roots present, apex multicellular, xylem exarch, branching endogenous; arbuscular mycorrhizae +; shoot apical meristem multicellular, interface specific plasmodesmatal network; stem with ectophloic eustele, endodermis 0, xylem endarch, branching exogenous; vascular tissue in t.s. discontinuous by interfascicular regions; vascular cambium + [xylem ("wood") differentiating internally, phloem externally]; wood homoxylous, tracheids +; tracheid/tracheid pits circular, bordered; sieve tube/cell plastids with starch grains; phloem fibers +; stem cork cambium superficial, root cork cambium deep seated; nodes ?; stomata ?; leaf vascular bundles collateral; leaves spiral, simple, axillary buds?, prophylls [including bracteoles] two, lateral, veins -5 mm/mm² [mean for all non-angiosperms 1.8]; plant heterosporous, sporangia eusporangiate, on sporophylls, sporophylls aggregated in indeterminate cones/strobili; true pollen [microspores, i.e. no distal pore for release of gametes] +, grains mono[ana]sulcate, exine and intine homogeneous, ovules unitegmic, crassinucellate, megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development first endo- then exosporic, tube developing from distal end of grain, to ca 2 mm from receptive surface to egg, gametes two, with cell walls, with many flagellae; female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large", first cell wall of zygote transverse, embryo straight, endoscopic [suspensor +], short-minute, with morphological dormancy, white, cotyledons 2; plastid transmission maternal; two copies of LEAFY gene, PHY gene duplication [N/O//A/C and P//BE lines], mitochondrial nad1 intron 2 and coxIIi3 intron present.

MAGNOLIOPHYTA

Plant woody, evergreen; lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, [cyanogenesis in ANITA grade?], lignins derived from both coniferyl and sinapyl alcohols, containing syringaldehyde [in positive Maüle reaction, syringyl:guaiacyl ratio less than 2-2.5:1], and hemicelluloses as xyloglucans; root apical meristem intermediate-open; root vascular tissue oligarch [di- to pentarch], lateral roots arise opposite or immediately to the side of [when diarch] xylem poles; origin of epidermis with no clear pattern [probably from inner layer of root cap], trichoblasts [differentiated root hair-forming cells] 0; stem with 2-layered tunica-corpus construction; wood fibers and wood parenchyma +; reaction wood ?, with gelatinous fibres; starch grains simple; primary cell wall mostly with pectic polysaccharides; tracheids +; sieve tubes eunucleate, with a sieve plate and cytoplasm with P-proteins, companion cells from same mother cell that gave rise to the sieve tube; nodes unilacunar [1:?]; stomata with ends of guard cells level with pore, paracytic, outer stomatal ledges producing vestibule; leaves with petiole and lamina [the latter formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, veins (1.7-)4.1(-5.7) mm/mm², endings free; flowers perfect, polysymmetric, parts spiral [esp. the A], free, development in general centripetal, numbers unstable; P not sharply differentiated, outer members not enclosing the rest of the bud, smaller than inner members; A many, with a single trace, introrse, filaments stout, anther ± embedded in the filament, tetrasporangiate, dithecal, with at least outer secondary parietal cells dividing, each theca dehiscing longitudinally by action of hypodermal endothecium, endothecial cells elongated at right angles to long axis of anther; tapetum glandular, binucleate; microspore mother cells in a block, microsporogenesis successive, walls developing by centripetal furrowing; pollen subspherical, binucleate at dispersal, trinucleate eventually, tectum continuous or microperforate, ektexine columellar, endexine thin, compact, lamellate only in the apertural regions; nectary 0; G free, several, ascidiate, with postgenital occlusion by secretion, few [?1] ovules/carpel, ovules marginal, anatropous, bitegmic, [outer integument often largely subdermal in origin, inner integument dermal], micropyle endostomal, integuments 2-3 cells thick, megasporocyte single, megaspore lacking sporopollenin and cuticle, chalazal, female gametophyte four-celled [one-modular, nucleus of egg cell sister to one of the polar nuclei], stylulus short, hollow, stigma ± decurrent, dry [not secretory]; P deciduous in fruit; seed exotestal; pollen germinating in less than 3 hours, tube elongated, growing at 80-600 µm/hour, with callose plugs and callose-based walls, penetrating between cells, siphonogamy, penetration of ovules within ca 18 hours, distance to first ovule 1.1.-2.1 mm; double fertilisation +, endosperm diploid, cellular [first division oblique, micropylar end initially with a single large cell, chalazal end more actively dividing], copious, oily and/or proteinaceous, embryo cellular ab initio, minute; germination hypogeal, seedlings/young plants sympodial; Arabidopsis-type telomeres [(TTTAGGG)_n]; whole genome duplication, single copy of LEAFY and RPB2 gene, knox genes extensively duplicated [A1-A4], AP1/FUL gene, paleo AP3 and PI genes [paralogous B-class genes] +, with "DEAER" motif, SEP3/LOFSEP and PHYA + C/PHYB + E gene pairs.

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

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

ERICALES [ASTERID I + II]: ovules tenuinucellate.

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

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

BORAGINACEAE + VAHLIACEAE + GENTIANALES + LAMIALES + SOLANALES: (8-ring deoxyflavonols +); vessel elements with simple perforation plates; C tube initiation late [sampling!]; [vascularised] nectary at base of G; style long.

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

Information on placentoid distribution is needed for Plocospermataceae, etc.

LAMIALES Bromhead  Main Tree, Synapomorphies.

Cornoside, verbascosides [caffeoyl phenylpropanoid glucosides (CPGs), caffeic acid esters, = acteosides], methyl- and oxygenated flavones +; glandular hairs with vertical divisions in head +; leaves opposite; endosperm with micropylar haustorium; fruit a septicidal capsule; protein bodies in nuclei; mitochondrial coxII.i3 intron 0. - 23 families, 1059 genera, 23275 species.

Evolution. Estimates of the age of Lamiales range from ca 97 to ca 74 million years before present (Bremer et al. 2004; Wikström et al. 2001 respectively); Janssens et al. (2009) date stem group Lamiales to 104±8.2 million years ago and the crown group to 95±11.9 million years. Lamiales contain ca 12.3% eudicot diversity. Most of this diversity is concentrated in families whose members are largely herbaceous to shrubby and have monosymmetric flowers and many small seeds. Both parasitic and insectivorous members of Lamiales are largely nonmycorrhizal (Brundrett 2004 and references).

Chemistry, Morphology, etc. The occurence of cornosides and iridoids in this clade seem to be mutually exclusive, except in Martynia louisiana (Jensen 2000b). Verbascoside, a disaccharide derivative of the hydroxycinnamic acid caffeic acid (= caffeoyl phenylethanoid glycoside), is common in this clade, it and and trisaccharide derivatives (over 100 altogether) are phenylpropanoid glycosides, a class of compounds usually with a central glucose, a C₆C₂ unit, commonly dihydroxyphenyl-ß-ethanol, and a C₆C₃ unit, hydroxycinnamic acid (Mølgaard & Ravn 1988). Such compounds are very rarely found elsewhere (an exception is Cassinopsis - see Icacinaceae s.l.: Cometa et al. 1993); they have not yet been recorded from Plocospermataceae. A great deal of work on characterising iridoids and understanding their distribution in Lamiales is being carried out by S. R. Jensen and collaborators.

Variation in nodal anatomy in the order is unclear. Neubauer (e.g. 1977, 1978) suggested that the single trace often divided immediately into three or more, and this nodal type is indeed common in the order. Bailey (1956) recorded 2:2 nodes in Lamiaceae, and other patterns of nodal vasculature occur as well. 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). A chalazal 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 here, being known from Tetrachondraceae, Calceolariaceae, Plantaginaceae, Scrophulariaceae, Stilbaceae, Orobanchaceae, Phyrmaceae and Paulowniaceae (Rebernig & Weber 2007).

Confirmation of the phylogenetic positions of Carlemanniaceae, placed sister to Oleaceae here (see below), and of Plocospermataceae, as well as studies of their anatomy, chemistry, etc., are important for understanding the evolution of the chemistry and floral morphology in particular of the whole Lamiales (cf. Endress 2001). Taxa with 4-merous or predominantly 4-merous flowers are common in the basal pectinations of the Lamiales tree (see also Mayr & Weber 2006). If Carlemanniaceae are in this general part of the tree, as seems lilkely, 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 merism in this part of the tree is clearly a complex matter, and exactly where changes of floral merism are to be placed is unclear.

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). For a useful general discussion, including suggestions of apomorphies for some clades, see Soltis et al. (2005b); Kadereit (2004) provided a summary of the order and its evolution.

Phylogeny. The very provisional tree here is based in part on the work of Oxelman et al. (1999a) and Soltis et al. (2001). Oxelman et al. (1999a), Mueller et al. (2001) and Hilu et al. (2001) among others suggested that Plocospermataceae is sister to other Lamiales. Savolainen et al. (2000b, rbcL data alone; see also Lee et al. 2007, Plocospermataceae not included) placed 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 was part of a trichotomy at the base of Lamiales (Oleaceae [Ligustrum only] + all other Lamiales complete the trichotomy; Plocospermataceae were again not included). A sister relationship to Oleaceae is also supported by Yang et al. (2007: 1.0 p.p., Plocosperma included, but sampling still very poor), and that seems the best place to put the family. The position of Hydrostachys has proved problematical. Here it is also included in Cornales, and there is a discussion of its relationships there. However, recent work (Burleigh et al. 2009) suggests that it should be included in Lamiales, with which its morphology is in general agreement. If it ends up in Lamiales, it is likely to be in the basal part of the tree. S. Andersson (2006, two genes, sampling poor) found 75% jacknife support for the clade [Calceolariaceae + Gesneriaceae], and 100% support for that clade as sister to remaining Lamiales, although Mayr and Weber (2006) did not think that the two families were particularly near each other. However, chemistry and morphology also suggest the relationship is close, and also that these families are basal or sister to (depending on details of their relationships) the remaining Lamiales.

For relationships in the "Scrophulariaceae"-Acanthaceae-Bignoniaceae-Lamiaceae area, see e.g. Wagstaff and Olmstead (1997), Olmstead et al. (2001), and Xia et al. (2009). B. Bremer et al. (2002) analysed variation in three coding and three non-coding regions of the chloroplast genome; their sampling was sketchy, so the support for some family groupings in the main part of the order is difficult to evaluate; Freeman and Scogin (1999) focussed on the old Scrophulariaceae, but the pattern of relationships they found was 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) 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) were not immediately related. 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. l. (see also Tank et al. 2006 for a summary). 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). 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). Petraea was sister to Bignoniaceae in some early molecular phylogenies (Wagstaff & Olmstead 1997). Amyloid is found in both Pedaliaceae and Acanthaceae, a family that is sometimes weakly associated with Pedaliaceae in molecular analyses (Soltis et al. 2005b and references). Both Martyniaceae and Pedaliaceae, probably not immediately related, have 10-hydroxylated carboxylic iridoids. Byblidaceae may be sister to Lentibulariaceae (e.g. Albert et al. 1992), although Müller et al. (2004) found no association between the two, nor of either with any Lamiales with viscid indumentum like that of the preceding two families, a feature which could perhaps be considered to be "precursory" to insectivory. On the other hand, Müller et al. (2004) found a weak association of Lentibulariaceae and Bignoniaceae.

Few strongly supported relationships in the bulk of the order were evident in the recent study of Soltis et al. (2007a), indeed, when over 4600 bp had been sequenced, it was estimated that at least 10000 bp more would need to be added if relationships within the clade were to be resolved (Wortley et al. 2005b). However, Refulio-Rodriguez and Olmstead (2008) have recently made substantial progress in disentangling relationships around Lamiacae-Verbenaceae and Scrophulariaceae s.l. (see below, also Xia et al. 2009). Although focusing on Triaenophora (ex "scroph", now Orobanchaceae s.l.), the relationships that Albach et al. (2009) found are broadly consistent with those suggested by others, and they, too, cast doubt on the monophyly of Phrymaceae as here delimited below...

Note that 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); this is consistent with the relationships discussed below. 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), although this is not so obviously consistent with these relationships.

Classification. 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. The limits of families like Scrophulariaceae have long been problematic (Thieret 1967 for a summary). Olmstead (2002) has provided a readable account of some of the earlier changes in our ideas of relationships in the Scrophulariaceae s.l. in particular.

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, Antirrhinales Döll, Aragoales D. Don, Bignoniales Lindley, Byblidales Reveal, Callitrichales Dumortier, Carlemanniales Doweld, Fraxinales Berchtold & J. Presl, Gesneriales Dumortier, Globulariales Dumortier, Hippuridales Thomé, Jasminales Dumortier, Lentibulariales Lindley, Ligustrales Bischof, Myoporales Berchtold & J. Presl, Oleales Lindley, Orobanchales Berchtold & J. Presl, Pedaliales Berchtold & J. Presl, Pinguiculales Dumortier, Plantaginales Lindley, Rhinanthales Dumortier, Scrophulariales Lindley, Selaginales Choisy, Stilbales Doweld, Utriculariales Döll, Verbascales Döll, Verbenales Horaninow, Viticales - 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, anthers extrorse, versatile, with parallel separate thecae, 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 bodies in nucleus?

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.

Chemistry, Morphology, etc. Plocospermataceae are poorly known. Jensen (1992) recorded verbascosides and cornoside from Plocosperma, but nor iridoids. There are prominent groups of fibers in the outer cortex where the leaf leaves the stem. In each carpel, one ovule may be subapical and pendulous, the other subbasal and erect, but both may also be in either position (Leeuwenberg 1967).

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.

Previous Relationships. Plocospermataceae were included in Gentianales by Takhtajan (1997), probably because Plocosperma had long been associated with Loganiaceae, although Cronquist included the genus in his Apocynaceae.

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

Chemistry, Morphology, etc. 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; exotestal 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, thecae ?, pollen colpate, ovary 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, interior cells unthickened, 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.

Chemistry, Morphology, etc. The peltate, glandular hairs with unicellular stalks and the two stamens (their position is not entirely certain) suggest Lamiales. 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) information on chromosome numbers, etc.

Previous Relationships. 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. However, Carlemmanniaceae were included in Caprifoliaceae by Cronquist (1981) and in Rubiales by Takhtajan (1997).

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

Woody; carboxycyclic iridoids, myricetin, orobanchin, mannitol +; wood with minute calcium oxalate crystals; (vessel elements with scalariform perforation plates); fibre tracheids +; foliar crystals of various types, inc. styloids and raphides (0; druses); libriform fibres 0; petiole bundle arcuate; sclereids + (0); hairs (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, 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; C free, 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); (first 4 K diagonal, C contorted, early tube formation - Nyctanthes), 1(-3) ascending ovules/carpel; fruit 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, first 4 K diagonal, C tube formation early, 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 Dumortier

Secoiridoids, flavone glycosides +; vessel elements in multiples; (pits vestured); libriform fibers + (0); fibre tracheids 0 (+); marginal parenchyma +/0; (indumentum of peltate scales); C (free; 0), tube formation late (early - Ligustrinae), (A 4 - e.g. Nestegis); n = (20) 23.

17/415: Chionanthus (60-120: Linociera recently incorporated, but questionable?), Fraxinus (45-65), Ligustrum (50), Noronhia (45), Olea (33). Tropical and subtropical, inc. New Zealand and Hawaii.

Synonymy: Forestieraceae Endlicher, Fraxinaceae Vest, Ligustraceae G. Meyer, Nyctanthaceae J. Agardh, Schreberaceae 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. Schrebera (Oleeae) looks superficially like Bignoniaceae. Fraxinus is a distinctive genus; its corolla may be absent, or the petals are free, the leaves are simple or compound, and the fruit is a samara; pollination is by wind.

Evolution. Much diversification within Oleaceae is Tertiary (Besnard et al. 2009a).

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

Chemistry, Morphology, etc. 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 (but not if Myxopyreae are sister to the rest of the family - see below).

At least some species of Osmanthus have a lignified, torus-bearing, pit membrane (Coleman et al. 2004). The diversity of often minute crystal types in the vegetative plant (other than the wood) is very great; druses are uncommon (Lersten & Horner 2008a, 2009a, esp. b). Epidermal crystals often clustered in cells at the bases of trichomes, an unusual distribution pattern (Lersten & Horner 2009b). The corolla tube is initiated early in a ring. There may be groups of few-celled secretory hairs forming extrafloral nectaries (Zimmermann 1932), 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). There is infrageneric variation in the orientation of the two carpels; whether they are collateral or superposed, the two stamens are borne in the plane of the septum (Eichler 1874). 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.

For information, see Baas et al. (1988: wood anatomy), Bigazzi (1989a: protein nuclear inclusions), Kiew and Baas (1984) and Rohwer (1994b: both Nyctanthes), Rohwer (1993b, 1996: fruit and seed), Jensen et al. (2002: iridoids), Green (2004: general), and Sehr and Weber (2009: floral ontogeny).

Phylogeny. Wallander and Albert (2000) suggested phylogenetic relationships within the family; the tribes above have strong support. 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. Franzyk et al. (2001) note that Myxopyrum and Nyctanthes, both in Myxopyreae, have similar iridoids. Besnard et al. (2009a) looked at relationships in some Oleeae.

Tetrachondraceae [[Calceolariaceae [Peltanthera + Gesneriaceae]] [Plantaginaceae [Scrophulariaceae [Stilbaceae [[Lamiaceae [Paulowniaceae, Phrymaceae, Orobanchaceae]] [Thomandersiaceae + Verbenaceae] etc.]]]]]: deletion in the matK gene.

For information on the matK deletion, 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, thecae ?, 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, protein bodies in nucleus?

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, Morphology, etc. 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.

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

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

HYDROSTACHYACEAE Engler, nom. cons.   Back to Cornales

Annual to perennial submerged rosette herbs; primary root 0, adventitous roots +; kaempferol +, iridoids 0; vessels present, ?type; nodes ?; stomata 0; leaves deeply and complexly divided, surface with small enations, stipule single, intrapetiolar (two, lateral); inflorescence spicate, plants di(mon)oecious; P 0, nectary 0; staminate flowers: A 2, extrose, monothecal, pollen in tetrads, inaperturate; carpellate flowers: G [2], transverse, placentation parietal, many tenuinucellate ovules/carpel, styles separate, filiform, impressed in the apex of the ovary; fruit a septicidal capsule; seeds minute, exotestal, outer cell walls much thickened, mucilaginous; endosperm scanty or 0, micropylar haustorium +; n = 10-12.

1[list]/20. C. and S. Africa, Madagascar (map: from Rauh & Jäger-Zürn 1966b).

• Hydrostachyaceae are submerged, rosette-forming, aquatic herbs with complex leaves bearing enations, stipules that are usually intrapetiolar, dense, spicate inflorescence, and rather small flowers with extrorse anthers and free styles.

Chemistry, Morphology, etc. The caffeoyl ester chlorogenic acid is found here and in the Loasaceae-Hydrangeaceae clade (Rønsted et al. 2002). Another interpretation of the androecium is that is consists of one tetrasporangiate stamen. Vessels are reported (Jäger-Zürn 1998), but are not described. The integument is about 5 cells thick; there seems not to be an endothelium. The styles are more or less impressed into the apex of the ovary, a feature that Leins and Erbar (1988) noted was common in Lamiales, although I do not know the general distribution of this feature.

For floral development, see Leins and Erbar (1988), for general information, see Erbar and Leins (2004a), and for chemistry, Rønsted et al. (2002).

Previous Relationships. Hydrostachyaceae have variously been suggested as being sister to Decumaria (Hydrangeaceae) (e.g. Xiang 1999; Albach et al. 2001; Xiang et al. 2002), or close to Crassulaceae (Saxifragales), or - perhaps - close to Podostemaceae (near Clusiaceae, in Malpighiales). However, Takhtajan (1997) included Hydrostachyales in his Lamiidae, and Cronquist (1981) also put it in that general area.

[[Calceolariaceae [Peltanthera + Gesneriaceae]] [Plantaginaceae [Scrophulariaceae [Stilbaceae [[Lamiaceae [Paulowniaceae, Phrymaceae, Orobanchaceae]] [Thomandersiaceae + Verbenaceae] 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, (2:3), adaxial lobes outside the others in bud [ascending cochleate], tube formation late; A 4, didynamous, anthers connivent, many ovules/carpel; endosperm also with chalazal haustoria [?position in tree].

Biology This clade includes the bulk of the diversity within Lamiales; many members are herbaceous or shrubby and have monosymmetric, bilabiate flowers and fruits with many small seeds.

Chemistry, Morphology, etc. Westerkamp and Claßen-Bockhoff (2007) outline the morphological variation in the corolla. Monosymmetry of the 2:3 type is common and there are four stamens which are often didynamous and with connivent anthers; for staminodes, see Endress (1998). 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. I am not sure at what level to peg the character "embryo suspensor large".

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).

[Calceolariaceae [Peltanthera + Gesneriaceae]]: cymes paired-flower; endothelial cells in longitudinal rows, endosperm longitudinally furrowed [aulacospermous].

Chemistry, Morphology, etc. In paired-flower cymes the flower in front of the terminal 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". The two flowers of the flower-pair have the same orientation. Both Calceolariaceae and Gesneriaceae 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.

CALCEOLARIACEAE Olmstead   Back to Lamiales

Herbaceous to shrubs; cork?; leaves toothed (entire); flowers 4-merous, K valvate, abaxial C lobe saccate, (adaxial "lip" strongly bilobed - Calceolaria triandra), elaiophores as pads of hairs on inside of abaxial lip (0); A 2 [adaxial pair] (3 - C. triandra), thecae (parallel) divergent, confluent on dehiscence or not, (theca 1), staminodes 0; nectary 0; (G semi-inferior), stigma small or capitate or obscurely bilobed; ovules with integument 3-4 cells across; 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.

Evolution. 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).

Chemistry, Morphology, etc. 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). For floral development, see Endress (1999).

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).

Phylogeny. For a phylogeny of the family, see S. Andersson (2006).

Classification. Porodittia, with three stamens, is a synonym of Stemotria, but neither name is needed as Stemotria is clearly derived from within Calceolaria, thus P. triandra = C. triandra (S. Andersson 2006). The limits of the sections need adjusting. I am grateful to Pamela Puppo for comments.

GESNERIACEAE Candolle, nom. cons.   Back to Lamiales

Usu. herbs or weak-stemmed trees (trees); hairs of stalked glands, or with thickened terminal cells; (cambium storied); (vessel elements with scalariform perforation plates); petiole bundle arcuate; stomata anisocytic; leaves joined at the base, (anisophyllous; two-ranked; spiral), involute, rather fleshy and/or softly hairy, margins toothed or entire; inflorescence axillary (terminal); K connate, abaxial C lobe outside others in bud [aestivation descending cochleate], (C spurred); A with staminode (0), (5, 2), anthers didynamous, 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; 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.)

1. Didymocarpoideae Arnott

(Nodes 1:1 with split laterals; 3:3 with split laterals; 5:5); (A 2 [abaxial pair, the common condition in the clade], placentae lamelliform-recurved, ovules restricted to distal end, style usually not well set off from ovary; fruit ± elongated, twisted, or shorter, a berry; testa cells often little elongated, (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), Chirita (140), Agalmyla (100), Didymocarpus (100), Paraboea (90). S. Europe (scattered), Africa and Madagascar, mostly Sri Lanka to Malesia and the Pacific (to Hawaii).

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

2. Epithematoideae

Dihydroxyphenolics [e.g. acteoside] 0; secretory canals; (medullary bundles + - Rhynchoglossum); cymes lacking bracteoles; (abaxial C lobe inside others in bud; A 2 [adaxial pair]; nectary variously vascularized), ovary short, abruptly narrowed into the style, (placentation axile); 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.

3. Gesnerioideae Link

3-desoxyanthocyanins +, chalcones, aurones 0; seeds without surface ornamentation, cells much elongated, spirally arranged (ornamented; shorter; not spirally arranged); endosperm conspicuous; GCyc2 gene lost.

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

3a. Coronanthereae

Trees or ± shrubby (rotting from the nodes); stomata anomocytic (paracytic); (flowers polysymmetric0, (A 2 [adaxial pair]; 5), nectary embedded in G wall, vascularized from A traces; capsules septicidal (and loculicidal; fruit a berry); n = 37(-45); gcyc duplication.

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

Titanotricheae + Gesnerieae, etc.: (plant with scaly rhizomes).

3b. Titanotricheae

Scaly rhizomes +; (stomata anomocytic); inflorescence racemose, with bulbils; testa striate-reticulate; n = 20.

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

3c. Gesnerieae, etc.

(Raphides, styloids +); (nodes 3:3; split-lateral - Columneae, Episcieae); (petiole bundles deeply arcuate to annular); (stomata on raised mounds, usually single [widespread]); (leaves spiral); (flowers resupinate), (K ± free), ovary 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+]), Gesneria (50). New World (map: from Brummitt 2007, in part). [Photo - Leaves, Flower.]

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.

Evolution. Möller and Cronk (2002) discussed biogeographic relationships within the large African genus Streptocarpus. Within Coronanthereae there seems to have been a single dispersal E -> W across the Pacific (Smith et al. 2006). Cyrtandra, with its berry fruits, is a very diverse genus found throughout Malesia, being particularly speciose in places like New Guinea; it is widely distributed in the Pacific - the species there form a single clade - and has been aptly designated as a "supertramp" genus (Cronk et al. 2005).

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

Birds and bees are the major pollinators of Gesneriaceae. 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 since there is no twisting of the pedicel (Clark et al. 2006), they are not resupinate stictly speaking. 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).

Many Gesneriaceae have capsular fruits with wind dispersed seeds, but seed dispersal by birds, either of fleshy fruits in their entirety, of of the 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 common in the New World (Weber 2004b; Clark et al. 2006); in the Old World, the speciose Cyrtandra has fleshy fruits.

Although many taxa are rather succulent or sometimes quite delicate herbs, a surprising number 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). 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. (2007) discuss growth patterns and the evolution of monophylly in Streptocarpus. 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) suggested that in these leaves it was the blade, normally the last part to mature and stop growing, that kept on growing. The cotyledon that kept on growing was the one exposed to more light (Saueregger & Weber 2005), and the developmental pathways controlling meristem development became relocalised (Mantegazza et al. 2009). The plant body of monophyllous Gesneriaceae may become more complex by repetition of the cotyledonary unit. Monophylly in general seems to be an adaptation for life on rocks, the plant growing in cracks of the rock and the single leaf hanging down and covering the rock surface quite efficiently (plants whose stems consist of sprays of alternating [if opposite, then with strong anisophylly] leaves are also common in such situations). For anisocotyly - more accurately, one cotyledon is accrescent - and its development in [Didymocarpoideae + Epithematoideae], see Burtt (1970) and Saueregger and Weber (2004).

Morphology, Anatomy, etc. 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. Gesneria has spirally-inserted serrate leaves with an almost coriaceous texture - it looks quite "atypical".

For more information, see Trapp (1956b: androecium), Weber (1973: inflorescence), Wiehler (1970: vegetative anatomy, esp. Gesnerioideae), Wilson (1974a, b: nectary vascularization), Skog (1976: Gesnerieae s. str.; 1984: chromosomes), Beaufort-Murphy (1983: seed morphology under the S.E.M., 1984: response to growth hormones, etc.; Cyrtandroideae much more responsive than Gesnerioideae), Kvist and Pedersen (1986: phenolics), Citerne et al. (2000), Smith et al. (2004a), and Zhou et al. (2008: all 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). Pan et al. (2002) discussed the floral development of Titanotrichum (see below). Pollen variation is either uninformative or suggests problems in everything from species delimitation on up (Schlag-Edler & Kien 2001).

Phylogeny. Epithematoideae are sister to Didymocarpoideae (Cyrtandroideae), see Smith (1996), Smith et al. (1997a, b) and especially Mayer at al. (2003), etc.. 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) or more likely near basal (Möller et al. 2009); they have dihydrocaffeoyl ester found nowhere else in flowering plants (Jensen 1996). Recent studies (Möller et al. 2009) place a number of small Asian and European clades all with four or five stamens basal in Didymocarpoideae. Of these, the odd Jerdonia, with its pollen in tetrads, four parietal placentae, large seeds with alveolate endosperm, and n = 14 (Burtt 1977b), may be sister to the rest of the subfamily (Möller et al. 2009). Although Didymocarpus was dismembered (Weber & Burtt 1998) and many species placed in Henckelia, the circumscription and relationships of that genus are still unclear (Möller et al. 2009). Within the diverse Cyrtandra, particularly speciose in places like New Guinea, all Pacific species studied are members of a single clade, and within this clade Hawaiian species are sister to the rest (Cronk et al. 2005). For relationships in Streptocarpus, to include Saintpaulia, see Möller and Cronk (2001).

Epithematoideae are perhaps to 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). Branch lengths are long. Chemistry - Napeanthus (Gesnerioideae) is also similar!

Kotarski et al. (2007) found 80% bootstrap support for the position of Coronanthereae as sister to the other Gesnerioideae, and then Titanotrichum as sister to the remainder. For a phylogeny of Coronanthereae, see Smith et al. (2006). Other studies also place the Old World but more or less isocotylar Titanotrichum in a similar position (C.-N. Wang et al. 2004: substantial amount of molecular data; cf. D. Soltis et al. 2000; Albach et al. 2001), but that genus has also sometimes been placed in "Scrophulariaceae". 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. (2003, 2006: the limits of Sinningia need adjusting, 2007: diversification); for relationships within Episcieae, see Clark and Smith (2009). See also Skog (1976) for a revision of Gesneria and relationships in Gesnerieae.

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 vessel elements with scalariform perforation plates like a few other Gesneriaceae (e.g. Kohleria, Carlquist 1997c). Secondary metabolites (lack of iridoids, presence of the caffeoyl phenylethanoid glycoside, sanangoside) suggest an association between Sanango and Gesnerioideae (Jensen 1996).

Classification. 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). In general all tribal classifications of Didymocarpoideae are decidedly unsatisfactory (Möller et al. 2009), and some genera, perhaps most notably Chirita, are polyphyletic. The huge Didymocarpus was fairly recently dismembered (Weber & Burtt 1998), species that had been included there being assigned to 27 genera (including two in Plantaginaceae); many species were placed in Henckelia, although this may still not be monophyletic.

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 [Scrophulariaceae [Stilbaceae [[Lamiaceae [Paulowniaceae, Phrymaceae, Orobanchaceae]] [Thomandersiaceae + Verbenaceae] etc.]]]]: route II decarboxylated iridoids [aucubin, catalpol widespread], 6- or 8-hydroxyflavones or 6 methoxyflavones +, cornosides 0; (embryo sac haustoria +).

Evolution. Caterpillars of Nymphalidae-Melitaeini and -"Kallimini" butterflies are quite common on plants in this group (see Plantaginaceae, Acanthaceae and Orobanchaceae below); they probably moved on to these families from Rosales (the Urticaceae group of families), and some Melitaeini in turn adopted members of Asteraceae as food plants (Nylin & Wahlberg 2008).

Chemistry, Morphology, etc. Flavonoid 7-O-glucosides and glucuroides are scattered in this clade (Lamiaceae, Pedaliaceae, Plantaginaceae: Noguchi et al. 2009). Extrafloral nectaries in this clade commonly consist of scattered multicellular trichomes (Zimmermann 1932).

PLANTAGINACEAE Jussieu, nom. cons.   Back to Lamiales

Herbs (shrubs; rooted aquatics); (mannitol, sorbitol +, iridoids 0, cornosides +), little oxalate accumulation; cork various; (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 (2 [adaxial pair]; 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), 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; Heide-Jørgensen 2008). [Photo - Callitriche Habit] [Photo - Hippuris Habit] [Photo - Flower]

• Hardly surprisingly, Plantaginaceae s.l. are very difficult to identify and distinguish from Scrophulariaceae s. str., Gesneriaceae, Stilbaceae, etc. However, the frequent absence of regular partitions in the heads of the glandular hairs is quite unusual in Lamiales, as is septicidal capsule dehiscence, both found here.

Evolution. For feeding preferences of a variety of insect groups that might suggest that the erstwhile Plantaginaceae s. str. and Scrophulariaceae s. l. are close, see Airy Shaw (1958) and Allen (1960, 1961); Allen (1960) found different insects eating Plantaginaceae and the immediately unrelated Scrophulariaceae s. str. (for which, see below). Larvae of Nymphalidae-Melitaeini butterflies are commonly found here and on Orobanchaceae, but not on Scrophulariaceae (Wahlberg 2001). There has been diversification of agromyzid dipteran leaf miners here (Winkler et al. 2009).

Since Hippuris and Callitriche (sister taxa) and genera like the more or less wind-pollinated Plantago are also in this clade, morphological diversification here has been very extensive (Leins & Erbar 1988: floral development; Olmstead & Reeves 1995; Reeves & Olmstead 1998). 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, but the family is predominantly pollinated by large insects and birds. Wilson et al. (2006, see also references) discuss shifts between bee and bird pollination in the speciose North American Penstemon clade; Penstemon is noted for is prominent bearded staminode. 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.

Sibthorpia has 5-8-merous, polysymmetric flowers, and polysymmetric flowers have been derived from monosymmetric flowers several times in this family. Linaria has flowers with a well-developed abaxial spur, but the well-known Peloria mutant has all five perianth members with spurs. Although we now know that it is under simple genetic control, Linnaeus was initially so impressed with this that he proposed to call it a separate genus, Peloria. 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, i.e. in bud the abaxial corolla lobes are outide the others.

Chemistry, Morphology, etc.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). Details of the distribution of these and other characters remain 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.

Both Digitalis and Isoplexis have cornosides. Iridoids with an 8,9 double bond - rather uncommon - are scattered in a number of genera (Jensen et al. 2007); at what level this character might be an apomorphy is unclear, although they are to be found in both Veronica and Plantago (Rønsted et al. 2000), and the two genera are close phylogenetically (Bello et al. 2002, 2004). Besseya and Plantago have a foliar endodermis. Veronica lyallii has successive subhypodermal phellogens (Gray 1937). Penstemon may have paired-flower cymes (elsewhere in Lamiales to be found in Gesneriaceae and Calceolariaceae); storied cambium is also reported from this genus. Bakker et al. (2006a) found major increases in the rate of evolution of the mitochondrial gene nad1 in Plantago and Littorella; 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).

For chemistry, see Jensen (2005), Taskova et al. (2006), and Jensen et al. (2009c), for Trapella, see Oliver (1888), and for a general survey, see Thieret (1967). Additional information is provided by Junell (1961: gynoecium), Leins and Erbar (2004a: general, as Hippuridaceae), Erbar and Leins (2004b: general, as Callitrichaceae), Schwarzbach (2004: general, as Plantaginaceae), Ihlenfeldt (2004: general, as Trapellaceae), Fischer (2004b: general, as Scrophulariaceae p. pte) and Wagenitz (2004: general, as Globulariaceae). For floral development, see Endress (1999). I thank Dirk Albach for comments.

Phylogeny. 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, summary, as Veronicaceae). Albach et al. (2005a) discuss the circumscription and phylogenetic relationships of the family and the variation that it encompasses. Gratiolaceae were recognised as a distinct family by Rahmanzadeh et al. (2004), who discuss the phylogeny of that family (only three species examined!) and give a list of included genera, etc. Gratiolaceae have an integument 3-6 cells across, with large, transversely elongated endothelial cells in vertical rows,\. This causes its seeds to have longitudinal ridges; the extotestal cells have hook-like thickenings. 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 Gratiolaceae; the relationships that they found can be represented as [Myoporum, etc. [Buddleja, etc. [Limosella, Manueleae, etc.]]]. Oxelman et al. (2005) also locate the majority of this clade in Plantaginaceae, although Limosella itself is again in Scrophulariaceae.

For the phylogeny of Antirrhineae, see Ghebrehiwit et al. (2003), and for that of Veroniceae, see Albach et al. (2004a, 2005c) and Taskova et al. (2004, 2006); the "new" molecular relationships are at least sometimes supported by other data such as chromosome number and iridoid type (Albach et al. 2004b, 2005c). Pedersen et al. (2007 and references) and Jensen et al. (2008a) report on some chemistry of ex-Hebe or Hebe s.l.; of the ca 125 species of this complex, all except for a few from New Guinea are found in New Zealand (Albach et al. 2005b). Albach (2008) discusses the limits of Veronica s.l., and Wolfe et al. (2006) outline phylogenetic relationships in Penstemon.

Classification. The circumscription of Plantaginaceae adopted here is because molecular studies suggest that a number of small but florally very distinctive families are to be included in a clade with relatively undistinguished monosymmetric flowers. Maintaining these as separate would entail the recognition of a number of other families. Since aquatic taxa like Callitriche and wind-pollinated taxa like some species of Plantago have very distinctive morphologies partly associated with different pollination syndromes, they have usually been segregated as separate families; they are briefly characterized below:

• Callitrichaceae - Water plant; hairs stellate; plant monoecious; staminate flowers; P 0; A 1(-3), pollen trinucleate, (exine, apertures 0 - see Cooper et al. 2000); carpellate flowers: G [2], transverse, subdivided, 2 ovules/carpel, integument thin, styles ± separate, slender; fruit schizocarpic, with 4 nutlets; exotesta perhaps persisting. For general evolution in Callitriche, see Philbrick and Les (2000). Pollination may be by wind, or under water, or self pollination through the plant, pollen grains germinating in the anthers and growing through the stem, etc., to the ovules of an adjacent pistillate flower.

• Globulariaceae - Shrublet to herbs; inflorescence capitate to racemose; gynoecium pseudomonomerous, 1 pendulous ovule/carpel, only the abaxial carpel developed; fruit nutlike, surrounded by the calyx; n = 8-10, 19. The pseudomonomerous gynoecium apparently distinctive; pollination by insects.

• Hippuridaceae - Water plant; flavones 0; hairs peltate, glandular; leaves whorled, linear; 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, style single, stigmatic its entire length, stigma dry; fruit achenial or drupaceous; testa structureless, endosperm thin, starchy; n = 16. Pollination by wind.

• Plantaginaceae s. str. - Herbaceous; nodes 3:3; leaves spiral; flowers polysymmetric, (3-)4-merous; G [2], placentation parietal, or unilocular, ovule 1, basal, epitropous, style long-branched, stigmatic its entire length, stigma dry; fruit circumscissile or nut-like, exotestal cells large, mucilaginous, endotestal cells persist; endosperm +, (embryo curved). Pollination by wind, insects. .

• Trapellaceae - Rhizomatous aquatic herb; flower quite large, slightly monosymmetric, A 2 [the adaxial pair], staminodes 2, connective apically peltate; ovary 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. Trapella sinensis. Pollination by insects.

Sutton (1988) monographed Antirrhineae.

Previous Relationships. Both Cronquist (1891) and Takhtajan (1997) recognise several of these smaller families, but they are in the same general part of the sequence; note Cronquist has a notably broad circumscription of Globulariaceae and includes a number of genera here placed in Scrophulariaceae. Trapella has been included in Pedaliaceae (e.g. Cronquist 1981), in part because its stoutly-spiny fruits appear to be so similar.

Synonymy: Antirrhinaceae Persoon, Aragoaceae D. Don, Callitrichaceae Berchtold & J. Presl, nom. cons., Chelonaceae Martynov, Digitalidaceae Martynov, Ellisophyllaceae Honda, Erinaceae Pfieffer, Globulariaceae Candolle, nom. cons., Gratiolaceae Martynov, Hemimeridaceae Doweld, Hippuridaceae Vest, nom. cons., Linariaceae Martynov, Littorellaceae Gray, Oxycladaceae Schnizlein, Plantaginaceae s. str., Psylliaceae Horaninow, Scopariaceae Trinius, Sibthorpiaceae D. Don, Trapellaceae Honda & Sakisaka, Veronicaceae Cassel

[Scrophulariaceae [Stilbaceae [[Lamiaceae [Paulowniaceae, Phrymaceae, Orobanchaceae]] [Thomandersiaceae + Verbenaceae] etc.]]]: ?

SCROPHULARIACEAE Jussieu, nom. cons.   Back to Lamiales

Herbs or shrubs; harpagide, harpagioside [8ß-8α-methyl substituted iridoids] +, little oxalate accumulation; (cork inner cortical); nodes also 1:3 + girdling bundle; (secretory cavities +); (indumentum stellate); (stomata anisocytic); leaves opposite (basally connate; ± foliaceous, stipuliform structures +) or spiral, (punctate), ptyxis flat; (inflorescence racemose; bracts recaulescent); (flowers polysymmetric; 4-merous), K unequal or not, (A 5 - Verbascum, Capraria; 2), thecae head to head and confluent, ± clavate, or parallel, (colpi diorate), staminodia +/0, nectary small or 0, ³1 ovule/carpel, apo/epi/pleurotropous, integument 5-11(-12) cells thick, stigma capitate (lingulate), dry; capsule septicidal (and apically loculicidal - Buddleja), (berry; drupe; schizocarp); seeds many, cushion-shaped scars on the placentae widespread; exotestal and endotestal cells with thickened inner walls, (testa multiplicative, exotestal cells ± longitudinally elongated, inner walls thickened - Buddleja); endosperm (ruminate because of inpushings of individual entotestal cells), copious to 0; n = 6-9, 12+ [18 - Myoporaceae s. str.], (protein crystal stacks in nucleus).

65[list]/1700: Verbascum (360), Eremophila (215), Scrophularia (200), Selago (190), Buddleja (125: inc. Nicodemia, Emorya, Gomphostigma), Jamesbrittenia (85), Manulea (75), Diascia (70), Nemesia (65), Zaluzianskya (55), Sutera (50). World wide (map: from Hultén 1958, 1971; van Steenis & van Balgooy 1966; Meusel et al. 1978; Leeuwenberg 1979; Hong 1983; Hilliard 1994; Norman 2000) [Photo - Flower, Flower, Myoporaceae s.str., again.]

Evolution. Mohrbutter (1937) notes both fungi and leaf miners that attack members of Scrophulariaceae s.l. (Buddlejaceae and Scrophulariaceae s. str.), thus the dipteran agromyzid miner Amauromyza verbasci has been found on Verbascum, Scrophularia and Buddleja (Spencer 1990). Some other insect herbivores seem to be able to distinguish between Plantaginaceae and Scrophulariaceae (e.g. Allen 1960). There are quite a few taxa that have oil-flowers with oil-secreting hairs (Vogel 1974; Vogel & Cocucci 1995 for a list). Details of the pollination mechanism of the remarkable two-spurred oil-flower Diascia are quite well known, the bee Redivia collecting oil from the oil-secreting hairs in the spurs by its sometimes remarkably elongated front pair of legs (Vogel 1984; Steiner 1990; Steiner & Whitehead 1991); flowers of some Orchidaceae from the same area are rather similar.

Chemistry, Morphology, etc. Harpagide and harpagioside, occuring here, are iridoids found elsewhere in Lamiales in Lamiaceae (inc. Caryopteris) and Pedaliaceae (Hegnauer & Kooiman 1978; Nicoletti et al. 1988), but Soltis et al. (2005b) suggest that such acylated rhamnosyl iridoids characterise the clade. The chemistry of Buddlejaceae (see Jensen 2000b) and Scrophulariaceae s. str. is in general similar (Houghton et al. 2003); for the chemistry of Myoporaceae s. str., see Ghisalberti (1994). Nicodemia (= Buddleja) is reported to have tannin (Bate-Smith & Metcalfe 1957). The wood anatomy of Buddleja is similar to that of Nuxia, Peltanthera, Androya, etc. (Carlquist 1997c), i.e. with taxa that are not immediately related to it. Some taxa have opposite leaves, an angled stem, and 1:3 nodes, however, I have not seen the little bundles of fibers that run along the ridges of otherwise similar stems in Linderniaceae.

Taxa with more or less polysymmetric flowers - sometimes rather like those of Silene, which some South African species may mimic - are common in almost all tribes, although the corolla tube of such flowers may be more or less bent and the androecium consits of two pairs of anthers borne at different heights in the tube (didynamy). There are also taxa with five corolla lobes and stamens (Capraria) and four lobes and stamens (some species of Buddleja), and in both cases the flowers are fully polysymmetric. Flowers of Verbascum s. str. have five stamens, but those of Celsia, included in Verbascum, have only four. Hemimeris may have inverted (and inversostylous!) flowers, but the patterning on the corolla is on the adaxial lobe, i.e. it is not really different from the normal condition in the family where the patterning is on the abaxial lobe and adjacent lateral abaxial lobes. Stamens with two thecae that are confluent apically are common (the anthers may be straight or U-shaped, but not sagittate); both Buddleia and Verbascum lack orbicules in their anthers.

A number of taxa have cushion-shaped scars, often with a central umbo, on the placenta marking the place where the seeds fell off; other taxa in e.g. the single-ovuled members of Manueleae have much thickened funicles that may be related to these scars. Similar scars are found in at least some Plantaginaceae and Stilbaceae (the latter, in Charadrophila), but details of the distribution of this feature are unclear.

Some information is taken from Hartl (1959: seed coat/rumination), Harborne and Williams (1971: chemistry) and Rogers (1986: as Loganiaceae, but including some genera belonging here), Maldonado de Magnano (1986b, 1987: embryology of Buddleja), Oxelman et al. (2004a: Buddlejaceae s. str., 2005), Theisen and Fischer (2004: as Myoporaceae), Fischer (2004b: Scrophulariaceae p. pte); , for orbicules, see Vinckier and Smets (2002a), for taxa included, see Tank et al. (2006). For floral development, see Armstrong and Douglas (1989), Endress (1999). F. Zapata made useful comments on the family.

Phylogeny. The old Selaginaceae/Selagineae with a single apical ovule per loculus link with Scrophulariaceae-Manueleae, although the latter have more ovules; Manueleae are very variable in both number and orientation of ovules in the loculus (see also Hilliard & Burtt 1977; Hilliard 1994). A number of these taxa have bracts that are adnate to the calyx (Kornhall et al. 2001). Also included here are Scrophulariaceae - Hemimerideae (Oxelman et al. 1999b), and recent work suggests that the cosmopolitan aquatic Limosella is to be placed with these southern African taxa (Kornhall & Bremer 2004).

Buddleja is very much paraphyletic and includes Nicodemia, Emorya, and Gomphostigma, but several lines of evidence place it here (e.g. Maldonado de Magnano 1986b); Teedia and Oftia have strong support as the sister group to Buddleja s.l. (Wallick et al. 2001, 2002). Other genera such as Androya that used to be in Loganiaceae also belong in Scrophulariaceae.

Myoporaceae in a somewhat expanded sense are usually shrubby plants that can be recognised by their more or less sessile and isobilateral leaves, sympetalous and often strongly monosymmetric flowers combined with leaves that have pellucid gland dots - but Leucophyllum has only a single pellucid gland at the apex of the lamina and Eremogeton has none. Core Myoporaceae have only a few epitropous ovules per loculus, the seeds have only slight endosperm, and the fruit is a drupe or schizocarp. The association of Leucophyllum with Myoporaceae s. str. seems quite well established (e.g. Schwarzbach & McDade 2002), and a distinctive pollen type the two have in common - tricolpate, with each colpus diorate - also agrees with this position (Niezgoda & Tomb 1975; Argue 1980). Capraria has glands in its leaves and pollen like that of other Myoporaceae; again, it fits nicely here (for leaf glands of these two genera, see Lersten & Beaman 1998; cf. also Henrickson & Flyr 1985; Lersten & Curtis 2001). Note that Scrophularia and Verbascum have distinctive cells (idioblasts) in their leaves (Lersten & Curtis 1997) perhaps similar to the glands of Myoporaceae s. str. Androya (used to be buddlejaceous) and Aptosimum may be around here; the former, however, has pollen that has been compared with that of Nicodemia (Loganiaceae s. str.). Oftia is unusual in that it has intraxylary phloem (not checked for Teedia); its inflorescence is a raceme, it has only four ovules/carpel, and its fruit is a drupe, the seeds haviving a very hard testa and copious endosperm. It, too, has been placed in Myoporaceae (see Takhtajan [1997: some information is taken from Dahlgren & Rao 1971). Olmstead et al. (2001) suggest that recognition of Myoporaceae may make Scrophulariaceae paraphyletic. Myoporaceae may be sister to Scrophulariaceae s.l., inc. Buddleja (Kornhall et al. 2001), or sister to Leucophylleae, in turn sister to Androya, the whole lot embedded in Scrophulariaceae (Oxelman et al. 2005). Chinnock (2007) monographed Myoporaceae s. str., but in a detailed discussion of its relationships suggested it could well be included in Scrophulariaceae.

For phylogenetic relationships, see also B. Bremer et al. (1994) and Nickrent et al. (1998).

Classification. The limits of Scrophulariaceae have long been problematic (Thieret 1967 for a summary; Olmstead 2002 for a readable account of the implications of the findings of molecular data). Rahmanzadeh et al. (2004), Albach et al. (2005a) and Oxelman et al. (2005) are clarifying the contents of the separate clades that used to be subsumed in Scrophulariaceae s. l. (see also B. Bremer et al. 2002; Tank et al. 2006); for further details see the introduction to Lamiales above. Members of the classical Scrophulariaceae are now also to be found in Plantaginaceae and Orobanchaceae (these contain the bulk of the taxa that have moved), as well as Stilbaceae, Phrymaceae, and Linderniaceae. Genera also associated with Scrophulariaceae, even if they were thought to be more or less links with other families, include Nelsonia and its relatives (see Acanthaceae) and Paulownia (see Paulowniaceae). Buddleja used to be included in Loganiaceae or placed in its own family; Takhtajan (1997) included both Polypremum (here Tetrachondraceae) and Sanango (here Gesneriaceae) in his Buddlejaceae.

Synonymy: Bontiaceae Horaninow, Buddlejaceae K. Wilhelm, nom. cons., Caprariaceae Martynov, Hebenstretiaceae Horaninow, Limosellaceae J. Agardh, Myoporaceae R. Brown, nom. cons., Oftiaceae Takhtajan & Reveal, Selaginaceae Choisy, nom. cons., Verbascaceae Rafinesque

[Stilbaceae [[Lamiaceae [Paulowniaceae, Phrymaceae, Orobanchaceae]] [Thomandersiaceae + Verbenaceae] etc.]]: ?

STILBACEAE Kunth, nom. cons.   Back to Lamiales

Ericoid shrubs, ordinary shrubs, or herbs; C-8 iridoid glucosides +, (cornosides +); cork just outside pericycle; vessel elements also with scalariform perforation plates; 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, stamens = and opposite sepals, (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 slightly 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.]

Chemistry, Morphology, etc. The C-8 iridoid glucosides common in this family are extremely uncommon elsewhere (Frederiksen et al. 1999); for the distribution of unedoside, present in at least some genera of Stilbaceae, see Oxelman et al. (2004a). Indeed, some iridoids are like those of Loasaceae and Hydrangeaceae, thus unedoside is common there (Jensen et al. 1998). The gynoecium is reminiscent of that of Scrophulariaceae - Manueleae. Thesmophora appears to have two tranverse carpels, with one descending ovule/carpel (Rourke 1993). This perhaps perhaps respresents the abaxial carpel that is divided by a false septum.

For anatomy and morphology, see Carlquist (1986) and Dahgren et al. (1979), for embryology, see 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).

Phylogeny. 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. Nuxia (ex Loganiaceae) is also placed here in molecular phylogenies (Backlund et al. 2000; Wallick et al. 2002), and phytochemically there are strong grounds for these associations (Frederiksen et al. 1999). 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 gesneriad-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 circumscription of the family is now greatly changed from what it was ten years ago, and it is unclear what might be apomorphies.

Synonymy: Hallieraceae Trinius, Retziaceae Bartling

[[Lamiaceae [Paulowniaceae, Phrymaceae, Orobanchaceae]] [Thomandersiaceae + Verbenaceae] etc.]: ?

[Lamiaceae [Paulowniaceae, Phrymaceae, Orobanchaceae]] : ?

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 uniseriate (unicellular; 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 ovules/carpel on inner side of placenta/carpel margin, style ?, (unequally) bifid, stigma inconspicuous, not expanded, dry (wet); ovules epitropous; 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; ?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 a drupe; ?endosperm.

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

Synonymy: Viticaceae Jussieu

3. Ajugoideae Kosteletzky

(Aromatic, no terpenoids, etc.); flowers (4 [Aegiphila] merous), also polysymmetric or 1-lipped [lobes 0:5 - Teucrium], exine with branched (simple, granular, etc.) columellae, disc slight-0 (+), style often ± terminal; endosperm several-layered, 0; n = 7, 10, 13, 14, 16+.

24/1115: Clerodendrum (400-500), Teucrium (250), 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 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 [CH₃CH₂)₁₀CH=C=CH(CH₂)₃COOH] +; 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.

Synonymy: Melittaceae Martynov

7. Nepetoideae Kosteletzky

Commonly aromatic [volatile terpenoids, rosmarinic acid], nepetoidin A and B [caffeic acid esters], (distinctive seed fatty acids) +, 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+: ?inc. Rosmarinus, Thymus, Mentha, Origanum), 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.

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

And 10 genera unassigned, including Callicarpa (140: hairs branched/stellate; flowers polysymmetric, 4-5(-7)merous), 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, but it is usually distinctively ridged and dry. Woody members often have strongly lenticillate twigs.

Evolution. 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. Gall-forming fruit flies of the Tephretidae-Tephrellini are found here (and on Acanthaceae and Verbenaceae: Korneyev 2005), as are agromyzid dipteran leaf miners (Winkler et al. 2009).

The very big, mostly New World Salvia has only two unithecate anthers, however, the connective is expanded forming a lever arm that is involved in pollination which, as in other members of the family, is predominantly by large insects and birds. Claßen-Bockhoff et al. (2004a, b) described stamen development in SalviaRosmarinus was part of the Salvia clade in this study) suggested that the distinctive stamen with its lever arm might have evolved three times. Reith et al. (2007) describe details of pollination in Salvia pratensis, and Wester and Claßen-Bockhoff (2007) focus on pollination by birds.

The calyx is conspicuous in fruit and is an integral part of the dispersal mechanism of the disseminule, whether being brightly coloured and helping to attract frugivores, as in Clerodendrum, having hooked hairs of being itself hooked (Priva and some species of Salvia respectively), forming a kind of catapult mechanism (Scutellaria) or a wing. Myxocarpy, the nutlets producing mucilage and so adhering to their disperser, is common in Nepetoideae (Ryding 1992).

The ca 60 species of mints endemic to Hawaii have diversified considerably and are placed in three separate genera (Lindqvist & Albert 2002). They are all polypoids and are derived from west North American Stachys (Lamioideae) and probably represent a single introduction to the islands (see also the silversword alliance - Asteraceae).

Chemistry, Morphology, etc. Trisaccharide esters alone of verbascoside are found in Lamiaceae, but along with with disaccharides they are also to be found in Verbenaceae, Oleaceae and Orobanchaceae in particular (Mølgaard & Ravn 1988). Bailey (1956) notes the vegetative nodes of Lamiaceae and "Verbenaceae" may be two trace, two gap; the extent of occurence of such nodes needs to be clarified. 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. 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.

For gynoecial morphology and embryology, see Junell (1934: note that there is variation in ovule attachment within the family), for seedlings, see Vassilczenko (1947: cotyledons in Lamiaceae s. str. usu. cordate to hastate), for pollen, ovules and seeds, see Wunderlich (1967b), for variation in the proteinaceous inclusions in the nucleus, see Speta (1979), for fatty acids in the seed, see Badami and Patil (1981), 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), for leaf anatomy in Mentheae, see Moon et al. (2009), and for a comprehensive general treatment, see Harley et al. (2004). For floral development, see Endress (1999).

Phylogeny. 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. For relationships in Viticoideae, see Bramley et al. (2009); Vitex is paraphyletic. for the phylogeny of the Australian-centered Chloantheae (Prostantheroideae: Conn et al. 2009). Species of Lamioideae and Scutellarioideae, but not Nepetoideae, tend to have relatively massive ammounts of fibrous tissue associated with the veins in the calyx, e.g. with the tertiary veins (Ryding 2007). Wenchengia has spiral leaves; it is unclear where it should be placed (Cantino & Abu-Asab 1993). In Ajugoideae, Clerodendrum is being divided (Steane et al. 1999, 2004), while within Nepetoideae, the large New World-centred Salvia, with over 900 species, is possibly para- or polyphyletic, Rosmarinus, Thymus, and Origanum also being involved (Walker et al. 2004; Walker & Sytsma 2007) - alas for "Scarborough fair". Nepetoideae include the large tribe Ocimeae with synthecous, dorsifixed anthers (Paton et al. 2004). Wunderlich (1967b) suggests that there is no endosperm in the subfamily. Moon et al. (2008) surveyed pollen morphology especially of Salviinae. For phylogenetic relationships in Lamioideae, see Wagstaff et al. (1995). For relationships of the ca 60 species of mints endemic to Hawaii, see Lindqvist and Albert (2002); recognition of the three genera in which they are placed makes Stachys paraphyletic, but even without this problem the limits of that genus is difficult to determine; Leucas is also highly paraphyletic (Scheen & Albert 2009).

Classification. 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).

The classification here is based on that of Harley et al. (2004). Note that the circumscription of Viticoideae is more narrowly drawn than in Cantino et al. (1992), some genera included there not being assigned to subfamilies here. Generic limits will need some attention; in both Stachys and Leucas characters associated with pollination prove unreliable indicators of clades, and there are clearly other places where rearrangements are to be expected; generic limits also need adjusting in Chloantheae (Prostantheroideae: Conn et al. 2009).

Previous Relationships. 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 were often placed fairly close to each other (e.g. Cronquist 1981, where both are in a rather narrowly drawn [compared to here] Lamiales). However, there are numerous differences (chemistry, leaf insertion, floral symmetry, ovule morphology, etc.) between the two, and the radicle in Boraginaceae points upwards in fruit while in Lamiaceae it points downwards.

[Paulowniaceae, Phrymaceae, Orobanchaceae] : (protein crystal stacks in nucleus).

Chemistry, Morphology, etc. For nuclear protein crystals, see Albach et al. (2009).

PAULOWNIACEAE Nakai   Back to Lamiales

Trees; cork cambium outer cortical; nodes 1:1; hairs uniseriate-branched; petiolar bundle annular; leaves entire; flowers large, K deeply lobed, anther thecae head-to-head, ?staminode, style hollow, head expanded or not, stigma punctate; seeds winged; exotesta cells broad, with complex reticulate thickenings; endosperm +; n = 19, 20.

1/6. (Warm) temperate East Asia (map: from Hu 1959). [Photo - Flower]

• Paulowniaceae may be recognised by their opposite leaves and terminal inflorescences with large, monosymmetric flowers; the calyx is densely covered with brown indumentum.

Chemistry, Morphology, etc. For some information, see Schilling et al (1982: verbascoside, etc.) and Fischer (2004b: as Scrophulariaceae; he includes Wightia, etc., in the same immediate group - "Paulowniaceae" - but in a separate tribe).

Phylogeny. There is moderate support for a position sister to Lamiaceae (Olmstead et al. 2000), but rather more for a position sister to Orobanchaceae (Olmstead et al. 2001; Mueller et al. 2001; Hilu et al. 2003; Müller et al. 2004: Wortley et al. 2005a [80% bootstrap]), or with Phrymaceae interpolated between them and Orobanchaceae (some analyses in Albach et al. 2009). The phylogenetic significance of the wood anatomical differences between Catalpa and Paulownia (Dos Santos & Miller 1993) is unclear.

Previous Relationships. Paulownia is superficially like Catalpa (Bignoniaceae) and both have been shuttled back and forth between "Scrophulariaceae" and Bignoniaceae. Paulownia has endosperm and lacks the distinctive ovary and seed anatomy of Bignoniaceae (Armstrong 1985; Manning 2000; Lersten et al. 2002).

PHRYMACEAE Schauer, nom. cons.   Back to Lamiales

Annual or perennial herbs (woody); (iridoids 0); 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<)many near-basal to axile (parietal) straight 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.]

Evolution. 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.).

Chemistry, Morphology, etc. 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.

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: as Scrophulariaceae p. pte). For floral development in Mazus, see Rawat et al. (1988).

Phylogeny. 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; Beardsley & 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). Xia et al. (2009) and Albach et al. (2009) found some support for the paraphyly of Phrymaceae as delimited here, with Mazus and Lancea forming a clade separate from that containing the rest of the family, and dismemberment may well be in order. There may also be parallel chemical differences within Phrymaceae s.l., thus Mazus has iridoids while Mimulus does not (Hegnauer & Kooiman 1978), although sampling is poor.

OROBANCHACEAE Ventenat, nom. cons.   Back to Lamiales

Most (hemi)parasitic herbs, many turn black on drying, (shrubs); stomata do not close; (mannitol), orobanchin +, little oxalate accumulation, 6- and/or 8-hydroxylated flavone glycosides 0; cork?; head of hair lacking vertical partitions; leaves spiral to opposite, often toothed to deeply lobed; inflorescence often racemose; (K ± free), C (tube development intermediate), abaxial lateral lobes outside others in bud [quincuncial], (otherwise); A (free from C - Eremitilla), 4, didynamous, (2), staminode 0 (1), anther thecae parallel or ± confluent sagittate to inverted U-shaped, (unequal or single: Castilejinae), often hairy, with tails or basal awns, (thecae unequal), pollen often starchy, commonly colpate with a retipilate surface, (polyporate), ([G 5]), (-1 ovule/carpel), placentation parietal, (placentae [2, bilobed] 4 [-6]), integument 4-12 cells across, stigma clavate to capitate; capsule loculicidal to septicidal; (seeds with elaiosomes); (cells of seed wings with reticulate thickenings on radial walls), exotestal cells variously thickened on the inner walls, (cells of other layers thickened and lignified); endosperm + (starchy; 0), (embryo minute, undifferentiated; germination via germination tube); n = 7+.

99[list]/2060: Pedicularis (600-?800 - Mill 2001), Castilleja (160-200), Euphrasia (170-350), Orobanche (150), Buchnera (100), Bartsia (50), Agalinis (45), Rhinanthus (45), Alectra (40), Harveya (40), Sopubia (40). World wide, but especially N. (warm) temperate and Africa-Madagascar (map: from van Steenis & van Balgooy 1966; Hultén 1971; Meusel et al. 1978; Hong 1983). [Photo - Plant, Collection.]

• Orobanchaceae are herbs, rarely shrubs, mostly with racemose inflorescences. In the flower, the abaxial lobes of the corolla are outside the others in bud. Nearly all members of the family have a holo- or hemiparasitic life-style. Some Plantaginaceae have similar corolla aestivation, but they are autotrophic.

Evolution. Orobanchaceae may be some 50-40 million years old (Wolfe et al. 2005). Euphrasia has a North Temperate distribution but is also circum-Pacific, i.e., it is basically bipolar; much dispersal seems to have been involved in attaining this range (Gussarova et al. 2008).

There has been diversification of agromyzid dipteran leaf miners on the hemiparasitic members of this clade (Winkler et al. 2009).

The diversification of the speciose of Castilleja is becoming better understood; there is a speciose West North American/Caentral/South American perennial clade - some 120 species - derived apparently quite recently from an annual ancestor; polyploidy is common in the polyploids, but not in the diploids (Tank & Olmstead 2008, 2009). Annuals have dispersed more than once to South America (Tank & Olmstead 2009).

Larvae of Nymphalidae-Melitaeini butterflies are commonly found here (also on Plantaginaceae, not on Scrophulariaceae: Wahlberg 2001).

Holoparasites have evolved from hemiparasites more than once in this clade (dePamphilis et al. 1997; Nickrent et al. 1998; Young et al. 1999; Schneeweiss et al. 2004a; Bennett & Mathews 2006, etc.). Indeed, the hemiparasitic Harveya obtusifolia is well embedded in a holoparasitic clade of the genus; whether there has been reversion in habit, or several independent acquisitions of the holoparasitic habit in that part of the family alone is unclear (Morawetz & Randle 2009). Within the family there are several different types of haustoria, some forming a connection with xylem only, others with both xylem and phloem; nevertheless, haustoria may have but a single origin (Fischer 2004b for a summary). Stomata in this family seem to be perpetually open (Stewart & Press 1990; Smith & Stewart 1990), even in the apparently autotropic Lindenbergia, sister to most other Orobanchaceae; the situation in Rehmannia and relatives (see below) is unknown. Perpetually-open stomata are common in parasitic plants because they increase the transpiration flow in the parasite so faciltating movement of water, etc., from the host to the parasite.

Some Orobanchaceae, particularly the hemiparasitic taxa with chlorophyll, may take up largely water, nitrogen, etc., from their hosts, others take up organic material as well; iridoid glucosides, pyrrolizidine and quinolizidine alkaloids, etc., may also move from host to parasite (e.g. Adler & Wink 2001; Hibberd & Jaeschke 2001; Shen et al. 2005 [also host selection]; Rasmussen et al. 2006 and references). Alder (2000, 2002, 2003) found a complex relationship between host and parasite, the the annual Castilleja indivisa. Association with Lupinus in particular led to a decrease in herbivores eating the parasite (sometimes), more visitors by pollinators, an increased seed set, etc., when compared with other hosts of the parasite. These effects were mediated by the movement both of alkaloids (?anti herbivore?) and nitrogenous compounds (increase in growth s.l.) from the lupin to the parasite. There may also be movement of material from parasite to host (Rank et al. 2004 for references). Indeed, some of the severe effects on the host caused by orobanchaceous parasites may be due in part by the release of the cytotoxic iridoid aglucone; the breakdown of the iridoid glucoside of the parasite is perhaps caused by the host's ß-glucosidases, themselves common because they are involved in the host's cyanogenic defence pathway (Rank et al. 2004). For other information on parasitism in Orobanchaceae, see Irving and Cameron (2009 and references).

Variation in floral morphology in Pedicularis is very great, some species having a corolla tube ca 10 cm long or more, and others having asymmetric flowers with an asymmetrical, proboscis-like extension of the upper lip, the two adaxial corolla members that form the galea. For comments on the floral evolution of the genus, see Ree (2005). Pollen morphology - there is quite extensive variation - is linked with corolla morphology and pollinator type (Wang et al. 2009). For myrmecochory (Melampyrum, Pedicularis), see Lengyel et al. (2009).

For the evolution of genome size in the family, see Weiss-Schneewiess et al. (2005); genome size is reduced after polyploidization.

Economic Importance. A number of Orobanchaceae, e.g. Striga and Alectra species of the tropical clade (see below), are very serious parasites especially on legume and grain crops in warmer and drier areas, especially in sub-Saharan Africa, where they are still spreading. Striga affects ca 40% of the cereal producing areas, causing average losses in yield of 30-90%, especially on poorer soils; a single plant of the parasite can produce up to 100,000 seeds which can remain viable for about 20 years (Scholes & Press 2008; see also Yoshida & Shirasu 2009; Irving & Cameron 2009); Alectra vogelii can cause the complete loss of legume crops it infects (Morawetz & Wolfe 2009).

Chemistry, Morphology, etc. Orobanchaceae have orobanchin, a phenylpropanoid ester of caffeic acid, and silicic acid, and their iridoids are derived from the aucubin pathway (Thieret 1971; Rank et al. 2004); cf. Gesneriaceae. For seed fatty acids in Orobanche, see (Velasco et al. 2000). Fischer (2004b) notes that a collar-like base of the corolla tube persists after the rest has fallen off - is this a family character? Corolla aestivation is interesting in this clade. The abaxial-lateral pair of corolla lobes commonly envelops the adaxial-lateral lobes, while in Euphrasia and its relatives the abaxial lobe also envelops this latter pair of lobes - both forms of quincuncial aestivation; in a number of other Orobanchaceae, the abaxial lobe envelops all other lobes, i.e. ascending cochleate aestivation (Armstrong & Douglas 1989). For floral development, see Armstrong and Douglas (1989), Endress (1999). Greilhuber (1974) observed endomitotic polyploidization in the cells of the inner tapetum in some genera - but not in Pedicularis, Melampyrum, and Plantaginaceae.

The recently-described Eremitilla is very distinctive morphologically, i.a. the stamens are free from the corolla tube and the anther thecae are more or less embedded in the expanded filament apex (Yatskievych & Jiménez 2009).

For embryology, see Tiagi (1963), for seed morphology, see Musselman and Mann (1976), for pollen, see Minkin and Eshbaugh (1989) and Lu et al. (2007), for corolla aestivation, see Eichler (1875) and Armstrong and Douglas (1989), for a general treatment, see Terekhin and Nikitcheva (1981) and Fischer (2004b: as Scrophulariaceae p. pte). Demissew (2004) provides a treatment of Cyclocheilaceae and Harley (2004) one of Nesogenaceae. See the Parasitic Plants website (Nickrent 1998 onwards) and also Heide-Jørgensen (2008) for general information. Robert Mill caught a number of mistakes around here.

Phylogeny. For the delimitation and composition of the family, see Young et al. (1999), Wolfe et al. (2005), Bennett and Mathews (2006), etc. Lindenbergia, perhaps sister to the rest of the family (e.g. Wolfe et al. 2005; Albach et al. 2009, but sampling limited) or linking more particularly with a small group of parasitic taxa (Bennett & Mathews 2006: support weak), is autotrophic (Hjertsen 1995). Lindenbergia has tricolporate pollen rather like that common in Lamiales, while the pollen of many other members is triporate and retipilate (Bennett & Mathews 2006). Other than that, a summary of relationships is [holoparasitic clade [Castilleja, Pedicularis, etc. [Euphrasia, Rhinanthus, etc. + tropical clade]]] (Bennett & Mathews 2006).

R. G. Olmstead (pers. comm.) notes that the inclusion in this clade of Nesogenes, Cyclocheilon and Asepalum - ex Cyclocheilaceae and Nesogenaceae and all poorly known - is likely (see also B. Bremer et al. 2002 for Cyclocheilon. There was strong support for Nesogenes (the only taxon of this group included) being sister to the shrubby Radamea (Bennett & Mathews 2006), and these two genera belonged to a strongly supported tropical clade relationships between whose members are poorly known (Bennett & Mathews 2006). In a more comprehensive analysis, ex Cyclocheilaceae and Nesogenaceae are sister to this tropical clade of Orobanchaceae, within which there was some resolution (Morawetz & Randle 2009).

These erstwhile Cyclocheilaceae and Nesogenaceae considerably increase the diversity of Orobanchaceae. Nesogenes has a verticillate inflorescence. Cyclocheilon and Asepalum, the two genera in Cyclocheilaceae, lack much in the way of a calyx but have large bracteoles enveloping the flower bud. They are also shrubs with red roots [?always]; the flowers are solitary, the calyx being at most a minute rim; A didynamous, pollen is 3-colpate, the exine being thickened near the apertures, the placentation is axile or parietal, with 1-5 ovules/carpel, endothelium?, the funicles are long and the stigma is lingulate. The fruit is a capsule or schizocarp; there is no endosperm. Although Harley (2004) notes similarities between the pollen of Cyclocheilaceae, Nesogenaceae (both have tricolpate pollen, that of Nesogenes is perhaps also pilate) and Orobanchaceae, nothing is known of the stomatal closure and parasitism - or lack of them - in these putative Orobanchaceae. There are other shrubby orobanchs, including Brandisia, an isolated genus of uncertain relationships (Bennet & Mathews 2006).

Rehmannia is a small genus of ca 6 species from China and Korea and the related Trianeophora includes two to three species from China. Oxelman (2005) found that Rehmannia linked very weakly with Phryma, Paulownia, Mazus and Lancea, as well as with genera of Orobanchaceae. 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; in a rather more extended analysis, Jensen et al. (2008b) found that Rehmannia was sister to Orobanchaceae. Albach et al. (2007) recorded the presence of iridoids in Rehmannia, although these are at best very uncommon in Gesneriaceae, and also at least some mannitol, a polyol not occuring in Scrophulariaceae s. str. but found i.a. in some Orobanchaceae, while in a more extended study Xia et al. (2009) placed both Rehmannia and Trianeophora in a strongly supported clade sister to Orobanchaceae (see also Albach et al. 2009), so they are provisionally included here. Rehmannia is not a hemiparasite and has a racemose inflorescence the flowers of which lack bracteoles and have quincuncial aestivation in which the two abaxial-lateral lobes are outside the others, as is common in Orobanchaceae; Trianeophora has bracteoles, there may be a staminode, but floral aestivation is similar (Wang & Wang 2005). Phytochemistry links Triaenophora closely with Rehmannia but Oreosolen is unrelated, linking with Verbascum and relatives, a north temperate group in Scrophulariaceae (Jensen et al. 2008b); the first two genera also show more particular similarities with Orobanchaceae (Xia et al. 2009). Rehmannia has the 1:3 nodes and petioles with arcuate + wing bundles so common in Lamiales (pers. obs.).

For a phylogeny of Pedicularis, see Ree (2005), for that of Euphrasia, see Gussarova et al. (2008), for that of Orobanche, see Park et al. (2008), and for that of Castilleja, see Tank and Olmstead (2008, 2009). For further details of relationships, see dePamphilis (1995) and Olmstead and Reeves (1995). Tank et al. (2006) summarize ideas on relationships within the family.

Classification. See Hjertsen (1995) for a monograph of Lindenbergia. Tank et al. (2009) provide a phylogenetic classification of Castillejinae.

Previous Relationships. In the past, the argument has usually been over whether or not the holoparasitic Orobanchaceae s. str. were distinct from the Scrophulariaceae s.l., which then included hemiparasitic genera like Euphrasia and Pedicularis (e.g. Boeshore 1920). Rehmannia has often been linked with Titanotrichum and included in Gesneriaceae (Xia et al. 2009 for references).

Synonymy: Aeginetiaceae Livera, Aragoaceae D. Don, Buchneraceae Lilja, Cyclocheilaceae Marais, Euphrasiaceae Martynov, Lindenbergiaceae Doweld, Melampyraceae Hooker & Lindley, Nesogenaceae Marais, Pedicularidaceae Jussieu, Phelypaeaceae Horaninow, Rhinanthaceae Ventenat

[Thomandersiaceae + Verbenaceae]: inflorescence racemose; 3³ ovules/carpel.

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; K with nectaries on the outside, staminode +; pollen 5-6-colpate; nectary vascularised by carpellary traces; gynoecial vasculature 8-shaped, 1-3 ovules/carpel; ovules hemianatropous; 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.

Chemistry, Morphology, etc. 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 with their radially-segmented heads which also often occur on the abaxial surface of the lamina. 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 endosperm has a distinct outer periclinal cell wall. I am not sure exactly how the cotyledons are folded. Study of the development of the ovule, embryo, endosperm and seed anatomy might well be profitable.

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.

Phylogeny. Thomandersia may also go somewhere near Schlegeliaceae, from tropical America. Characters like the vasculature of the floral nectary and petiole, also the nectaries on the outside of the calyx, link it with Schlegeliaceae, however, support for any Thomandersiaceae-Schlegeliaceae association is currently weak (Wortley et al. 2007).

Previous Relationships. Thomandersia was previously usually included in Acanthaceae.

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; leaves simple, margins toothed to deeply lobed; (inflorescence 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, (1 erect ovule/carpel - Lantata), 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 (+ - Lantata); 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 asymmetrically swollen and glandular and the ovary has four ovules.

Evolution. Gall-forming fruit flies of the Tephretidae-Tephrellini are found here (and on Acanthaceae and Lamiaceae: Korneyev 2005).

Chemistry, Morphology, etc. 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).

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).

Phylogeny. Within Verbenaceae, Petraea and Duranta, both woody, are successively sister to the rest of the family (Marx & Olmstead 2007); Petraea (and Nashia) have polysymmetric flowers (Jabbour et al. 2008). For relationships around Verbena, see Yuan and Olmstead (2008), while Lu-Irving et al. (2009) found that within the Lantana-Lippia complex, Aloysia formed a basal grade and members of the animal-dispersed Lantana with their pyrene-type fruits were polyphyletic.

Previous Relationships. Verbenaceae as currently circumscribed (especially 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.

Classification. The whole Lantana-Lippia complex, speciose athough it may be (ca 400 species?), may be best reduced to a single genus, the larger genera currently recognised being hopelessly para- or polyphyletic (Lu-Irving et al. 2009).

Synonymy: Durantaceae J. Agardh, Lantanaceae Martynov, Petraeaceae J. Agardh

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]

Chemistry, Morphology, etc. The mucilage glands normally have four apical cells. The apparently single axillary flowers of some taxa appear to represent reduced cymes, the paired nectaries at the base of the pedicel representing modified flowers (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.).

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) +, connective with apical gland; pollen trinuclete, inaperturate, exine dissected into 20-40 platelets; G with parietal placentation, placentae bilobed, 2-many ovules/carpel, stigma bilobed; capsule with paired apical spurs or hooks [developing from sterile upper part of ovary], mesocarp ± fleshy, falling off, endocarp woody, with crests and spines; exotesta 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]

Evolution. Insects may stick to the very viscid indumentum of Martyniaceae, although there is no evidence that the plants are carnivorous (see Plachno et al. 2009; 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).

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

Phylogeny. For relationships within Martyniaceae - rather poorly supported and varying somewhat according to the marker used - see Gutierrez (2008).

Previous Relationships. Martyniaceae and Pedaliaceae have often been combined (as Pedaliaceae, e.g. Cronquist 1981), but there is currently no evidence that they form a monophyletic group. Differences in pollen and placentation best separate the two.

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, bracteoles 0; glandular hairs on the inside of the C; A 4, staminode +/0, or A 2, the adaxial pair, also 2 large abaxial Z-shaped staminodes with an appendage, or staminodes much reduced, or A 2, the abaxial pair only, thecae parallel to ± head to head; pollen 3(-5)-colpate, stigma bilobed, sensitive; ovule with spathulate embryo sac; capsule septicidal or -fragal; seeds with ruminate endosperm; n = 8, 9, 12-14, etc. [x = 7-9?].

Ca 13[list]/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.

Evolution. 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, while in other species the long, curved abaxial filaments are joined by the connate anthers and form a sort of balustrade across the mouth of the corolla. Various hairs develop on the abaxial anther knees, and also inside the corolla, and the latter may have projections, flanges, etc.; all in all, a complex little flower (see e.g. Magin et al. 1989). It would be interesting to know details of pollination mechanisms for such flowers, although small bees have been recorded as visitors (Magin et al. 1989). In Torenia fournieri, which has a less obviously distinctive floral morphology, the adaxial stamen pair elongate quickly; the abaxial pair has anthers which, when touched on lever-like lateral flanges, open and forcibly extrude their pollen (Armstrong 1992).

Chemistry, Morphology, etc. 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. For the floral development of Torenia, see Armstrong (1988). Lewis (2002) suggests that the stamens are extrorse and the ovules are straight; 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 sometimesprotrudes beyond the micropyle in some species of both Torenia and Lindernia, at least (Wardlaw 1955; Yamazaki 1955); 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.

For more information, see Fischer (1989, 1992, 2004b - the latter Scrophulariaceae pro parte: general).

Phylogeny. 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. 2005a). 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. In another study (Oxelman et al. 2005), Micranthemum was sister to Torenia, the two in turn were sister to Stemodiopsis, the only three Linderniaceae included in the analysis. See Tank et al. (2006) for a summary of our ideas of relationships within this clade, and also of its composition (the generic list here is rather notional). If Micranthemum belongs in this clade, it is possible at least some other Scrophulariaceae-Microcarpeae will also have to be included. Microcarpeae include aquatic herbs whose flowers usually have only the abaxial stamen pair (in the rest of the family it is the adaxial pair that is likely to be fertile); the filaments have "clavate geniculations at base" (Fischer 2004b).

BIGNONIACEAE Jussieu   Back to Lamiales

Trees or shrubs; C-4 carboxyl and ecarboxylated iridoids +; cork also cortical; cambium storied; (vessel elements with scalariform perforation plates); nodes 1:1-3 or more; petiole bundles annular (also rib or adaxial bundles); stomata helicocytic [?level]; leaves bicompound, 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, 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; 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 - eight groups and unassigned below. Mainly tropical, esp. South America (map: from van Steenis 1977). [Photos - Amphitechna Flower, Distictella Flower.]

1. Jacarandeae Seeman

K ± free, staminode large, bearded; G with parietal placentation; fruit orbicular, angustiseptate; n = 18.

1(?2)/55: Jacaranda (50). Tropical America.

[Tourrettieae [Tecomeae [Bignonieae [[Catalpeae + Oroxyleae] [Crescentieae + Coleae]]]]]: ?

2. Tourrettieae G. Don

Tendrillar vines; inflorescence bracteate, racemose; staminode 0.

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

[Tecomeae [Bignonieae [[Catalpeae + Oroxyleae] [Crescentieae + Coleae]]]]: leaves once compound; (staminodes +, simple).

3. Tecomeae Endlicher

Distinctive C-4 formyl iridoids.

12/55. Worldwide, not Arctic.

[Bignonieae [[Catalpeae + Oroxyleae] [Crescentieae + Coleae]]]: ?

4. Bignonieae Dumortier

Tendrillar lianes; anomalous secondary thickening + [phloem discontinuous, the 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].

21/380: Adenocalymma (78), Arrabidea (69), Anemopaegma (42), Amphilophium (41). America, largely tropical.

[[Catalpeae + Oroxyleae] [Crescentieae + Coleae]]: ?

[Catalpeae + Oroxyleae]: ?

5. Catalpeae Meisner

Leaves simple; (A 2).

2-3/11. North America, the Greater Antilles, East Asia.

6. Oroxyleae A. H. Gentry

(Leaves bicompound); (flowers polysymmetrical; A 5); fruits septicidal.

4/6. Indo-Malesian.

[Crescentieae + Coleae]: ?

7. Crescentieae G. Don

Leaves palmate (unifoliate; spiral, simple, phyllodinous); (flowers bat-pollinated, ± cauliflorous; fruits ± indehiscent; seeds not [barely] winged).

12/147: Tabebuia (70). Central and South America and the Greater Antilles.

Synonymy: Crescentiaceae Dumortier

8. Coleae Bojer

(Leaves phyllodinous, articulated); flowers bat-pollinated, ± cauliflorous; fruits ± indehiscent; seeds ?not winged.

4/60. Madagascar and surrounding islands.

• 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.

Evolution. The divergence time of Bignoniaceae and Verbenaceae has been estimated at ca 49.5 million years before present (Nie et al. 2006). The family is probably New World in origin, with some five shifts to the Old World (and one back to the New World) (Olmstead et al. 2009).

Bignoniaceae are, along with Sapindaceae, the most important neotropical group of lianes, and Bignoniaceae are the second most speciose family in drier tropical forest types in America (Gentry 1988). Bignonieae in particular are nearly all lianes with branched tendrils and distinctively rayed xylem (Lohmann 2006 for a phylogeny). Perianthomega has biternate leaves, also, it has robust unbranched tendrils that represent petioles, and three small scars (leaflets!) can be seen at their ends. Within Bignonieae, variation in the detail of the ray-like fluting of the xylem can be interpreted as complexity increasing by terminal addition and is mirrored by ontogenetic increases within an individual; shrubby Bignonieae (polyphyletic) show a simple pattern that results from a heterochronic reversal (Pace et al. 2009).

Palmate leaves may have arisen independently within Bignoniaceae, but are known only from New World taxa. The New World Tabebuia s.l., which has opposite, palmate leaves, is polyphyletic (Grose & Olmstead 2007b); a number of taxa - some apparently very different vegetatively - are derived from this basic morphology. Amphitecna has spiral, simple leaves as does Crescentia; their petioles are short and the lamina has distinctive, widely spreading venation. Such leaves are phyllodinous, indeed, in some species of Crescentia palmately-arranged leaflets are borne on the end of a lamina-like petiole. These genera are derived from within a clade of palmately-leaved taxa (Grose & Olmstead 2002, 2007a) all included here in an expanded Crescentieae. Associated with these two genera are Parmentiera and Spirotecoma, both with opposite palmately compound leaves; all four genera have bat-pollinated flowers. The simple and clearly petiolate leaves of Catalpa (opposite or whorled leaves) and Chilopsis (spiral leaves: the two genera hybridise), have a very different morphology from those of Crescentia, etc.; 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 commonest 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.

Dispersal syndromes are also quite diverse (Gentry 1983; 1990) but are not particularly correlated with pollination syndromes. Wind dispersal is common in the family, and the seeds have broad, papery wings. 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. Of the Crescentieae just mentioned, Amphitecna and Crescentia (calabash) have spherical indehiscent fruits, Parmentiera has elongated fleshy fruits, although its seeds still have a small wing, and Spirotecoma and Tabebuia and relatives have elongated, dehiscent fruits and winged seeds. Kigelia africana has pinnately-compound leaves; it is also bat-pollinated and has 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.

Chemistry, Morphology, etc. Aliform-confluent xylem parenchyma is common. There are four main carpel bundles, but only two in the "Scrophulariaceae" (Armstrong 1985), Gesneriaceae, etc. In Tourrettieae, Tourrettia has sub four-locular ovaries each with a single rank of ovules, while Eccremocarpus has parietal placentation. A number of Bignonieae with septifragal dehiscence also have cracks in the loculicidal position along the backs of the valves.

For information on pollen, which is very variable, see Gentry and Tomb (1979) and Burelo-Ramos et al. (2009: Pithecocteniinae), 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.

Phylogeny. The basic phylogenetic structure within the family is [[[Jacarandeae [Tourrettieae [Bignonieae + the rest]]] (Olmstead et al. 2002), and this has been further amplified by Olmstead et al. (2009: ca 3/4 of the genera sampled, three genes), although a number of relationships between major groups, e.g. of Tecomeae, remain poorly supported. For a comprehensive (2-gene + morphology) phylogeny of Bignonieae, see Lohmann (2006a); Perianthomega is sister to the rest of the tribe. Major clades are supported by a mixture of floral and vegetative characters, and generic limits have been reorganized accordingly (Lohmann 2002, esp. 2006). Bignonieae may be close to Oroxylum and relatives - which have bicompound leaves and septicidal capsules - and Catalpa - which has only two stamens (Olmstead et al. 2002). Coleae as narrowly delimited here are restricted to Madagascar, and their phylogeny and fruit evolution has been examined by Zjhra et al. (2004); they are part of a larger and well supported clade that includes Kigelia, Spathodea, etc. Relationships between the New World Tabebuia, with opposite, palmately-compound leaves and its relatives have recently been clarified. Amphitecna and Crescentia, both with spiral, simple leaves, are probably derived (Grose & Olmstead 2007a, b).

Classification. The tribes above are those recognised by Olmstead et al. (2009). Note, however, that their tribal classification is not exhaustive in that quite a number of genera are not assigned to tribes, partly because their phylogenetic position is still ambiguous (e.g. Argylia, Delostoma) or simply because; extension of the circumscription of some tribes and the addition of new ones will be needed in a classification such as that used here. Note that Coleae and Crescentieae, with similar flowers and fruits and both with "simple" leaves, but of different morphologies, are not sister taxa; Crescentieae have been expanded to include Tabebuia, etc. (the Tabebeuia alliance of Grose & Olmstead 2007a, b), the expanded clade being characterized by palmate leaves. Coleae, too, could well be expanded to include genera like Kigelia, Spathodea, etc. Generic limits in Bignonieae have been extensively reworked (Lohmann 2006, esp. 2009); in Lohmann and Ulloa (2007) all the species accepted in Bignoniaceae are listed.

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.

Chemistry, Morphology, etc. 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 of 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 rather improbable, unless it occurs in the Cuban Synapsis, 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.

Previous Relationships. Schlegelia and relatives are usually included in Bignoniaceae. The two may indeed be close (Olmstead et al. 2001), and there are no obvious differences in wood anatomy between them (Gasson & Dobbins 1991) - or in Scrophulariaceae s.l.

ACANTHACEAE Jussieu, nom. cons.   Back to Lamiales

Quaternary methylammonium compounds, amyloid +; (cork cambium deep seated); (intraxylary phloem +); stomata diacytic; leaf margins entire to toothed; inflorescence indeterminate thyrse [i.e. racemose], bracts large, conspicuous; K free or connate, often sharply pointed, adaxial lobes of C outside others in bud [aestivation descending cochleate], (lobes narrow); A didynamous (2; + 2 staminodes; 5), staminode +/0; G lacking septal bundles; ovule with "thin" integument; capsule dehiscence explosive, walls cartilaginous, K persistent; endosperm development asymmetrical [the two haustoria come to lie close to each other], embryo often ± curved.

229[list]/ca 4000 - eleven groups below. Mostly tropical.

1. Nelsonioideae Sreemadhavan

Herbs; gland-headed hairs with 2-celled heads; leaves opposite to spiral, margins?; inflorescence a raceme, (branches cymose - Saintpauliopsis), bracts spiral, (bracteoles 0 - Nelsonia); stamens 2 or 4, anthers variable (e.g. thecae ± separate), (placentation parietal - Elytraria), many (-4) ovules/carpel, funicular obturator +, stigma broadly (unequally) lobed, (sensitive - Elytraria); seeds 2-many, ruminate, testa ± disorganised; endosperm +, oily; n = 9.

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

Synonymy: Nelsoniaceae Sreemadhavan

Acanthoideae [Thunbergioideae + "Avicennioideae"]: (inverted vascular bundles in the pith); acicular fibers +; (adaxial C lobes outside others - [descending cochleate]), pollen other than tricolpate or -colporate common, 2 collateral ovules/carpel, endothelium 0; amyloid in cotyledons +, endosperm 0.

2. Acanthoideae Link

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

217/3220: Asystasia (70). World-wide; the bulk of the family. There are more species in the New World, more genera in the Old World. [Photo - Habit, Flower.]

2A. Acantheae

Nodes not swollen; anthers monothecous, pollen tricolpate.

21/500: Aphelandra (170), Blepharis (130).

[Ruellieae + Justicieae] [BAWN clade]: cystoliths +; pollen porate.

Ruellieae + Justicieae: ?

2B. Ruellieae

Filament curtain +, pollen often reticulate, (3+ ovules/carpel); seeds with hygroscopic trichomes.

Strobilanthes (250), Ruellia (250).

2C. Justicieae

Parallel ridges on upper lip of corolla [rugula] holding style (0); A (2), thecae displaced, not opposite, pollen tricolporate, hexapseudocolpate.

Justicia (600), Ptysiglottis (60).

BAWN clade: ?

2D. Barlerieae

C quincuncial: seed with hygroscopic trichomes.

/420: Barleria (300). Pantropical.

2E. Andrographidae

Pollen colporate, ornamented and thickened exine surrounding or over apertures; (3+ ovules/carpel).

/75. Asia.

2F. Whitfieldieae

C contorted/abaxial lobes outside others in bud [aestivation ascending cochleate]; pollen biporate, lenticular, granular around apertures; seeds with concentric rings of ridges, (also hygroscopic trichomes + - Lankesteria).

Andrographidae + Barlerieae: ?

2G.Nemacanthus

K united 3 + 2, pollen tricolporate, intercolpal regions psilate/foveolate; seed with hygroscopic trichomes.

1/30. Africa, Madagascar, Arabia to Vietnam.

Synonymy: Justiciaceae Rafinesque, Meyeniaceae Sreemadhavan

Thunbergioideae + Avicennioideae: cotyledons folded.

3. Thunbergioideae Kosteletzky

Twining vines (erect); (C-8 iridoid glucosides [unedoside] +); petiole bundles arcuate or annular; leaf ptyxis strongly curved; inflorescence with 2 or more flowers in the median plane of the leaf/inflorescence bract axil, adaxial flowers opening first; bracts 0, bracteoles very large, connate, K a rim, (up to 16 lobes), C often contorted, anthers with lignified unicellular hairs (multicellular awns), sagittate, (thecae slightly displaced), (elongated) porose/slits, endothecium 0, pollen 8-colpate or spiraperturate, connective elongated, (adaxial carpel aborting - Mendoncia), stigma wet, small and sub-bilobed to trumpet-shaped, with broad and often unequal papillate lobes; capsule also septifragal, (fruit a 1-2-seeded drupe - Mendoncia); (cotyledons twice folded - Mendoncia, etc.); n = 9, 28.

Ca 5/150: Thunbergia (90), Mendoncia (60: M. belizensis has rather boraginaceous hair bases). Tropical America, Africa and Madagascar, fewer in South East Asia - Malesia. [Photo - Flowers.]

Synonymy: Mendonciaceae Bremekamp, Thunbergiaceae Lilja

4. Avicennioideae Miers

Trees; betaines +, tanniniferous; wood with successive cambia, phloem islands occuring in bands of conjunctive tissue; nodes 3:3; petiole bundle annular; sclereids +; leaves with salt glands on both sides, colleters +; inflorescence in dense thyrsoid spicate units; flowers (polysymmetric), 4(-6)-merous; K ± free, C with nectary glands[?], stamens = and alternating with C, disc small [nectaries as tufts of hairs?], (false septae +), loculi apically confluent, ovules apical on partitions, pendulous, ± straight[?], no micropyle, stigma with 2 blunt lobes; fruit an achene, K persistent, green; embryo green, viviparous, cotyledons induplicate-reduplicate; n = 18, 32.

1/8 (species limits need attention). Mangroves in tropics, but also warm temperate (map: from Moldenke 1960; Tomlinson 1986). [Photo - Flower]

Synonymy: Avicenniaceae Endlicher, nom. cons.

• Acanthoideae are usually herbs, and can be recognised by their often swollen nodes, the stem immediately above them collapsing on drying, and the opposite leaves that have cystoliths (but not in Acanthus and immediate relatives). The bracteoles are often prominent and the calyx (and corolla) lobes may be narrow. The obovoid, almost woody capsules have few, relatively large, flattened seeds cradled by hook-like jaculators; the latter are hardened funicles. The seeds are shot out as the capsule dehisces explosively.

• Thunbergioideae are vines. There are no bracts, but large bracteoles envelop the flower. The calyx is reduced to a rim or is represented by a number of linear lobes. The flowers are large, the anther connective is elongated and there are two ovules per carpel.

• Avicennioideae are mangrove trees with pneumatophores that can be recognised by their opposite, entire, somewhat fleshy leaves with invisible secondary and finer venation, clavate hairs covering the lower surface, and salt glands on both surfaces. The flowers have as many stamens as corolla lobes and a persistent green calyx, and the fruits are achenes with a single rather large seed, as befits a mangrove plant.

• Nelsonioideae are rather undistinguished!

Evolution. Gall-forming fruit flies of the Tephretidae-Tephrellini are found here (and on Verbenaceae and Lamiaceae: Korneyev 2005). Mass defoliation of Avicennia by lepidopteran larvae seems to be not uncommon (Fernandes et al. 2009).

Pollination has been extensively studied in the family. The filament curtain, formed from decurrent filament ridges in the corolla tube and more or less connate filaments immediately above the adnate portion of the filaments, is probably involved in pollination in Ruellieae and perhaps other taxa, too. The curtain divides the corolla tube vertically into two apartments, and so the nectar can be enclosed in a separate chamber (Mantkilow 2000; see also Moylan et al. 2004b). Tripp and Manos (2008) studied the pollination systems in the speciose Ruellia. They found that although flowers specialised for bird or bee pollination may reverse pollinators, sphingid-adapted flowers do not reverse, perhaps because thay have entirely lost their floral pigments. For bird pollination in Aphelandra, see McDade (1992). Full (180^o) or partial resupination has evolved several times independently in Acanthoideae, and this is sometimess caused by the twisting of the corolla tube rather late in development (Daniel & McDade 2005); this is a rather unusual mechanism. Elytraria (Nelsonioideae) may also have inverted flowers.

Avicennia has numerous features associated with the mangrove habitat, including large, green, more or less viviparous embryos that are the unit of dispersal, pneumatophores, and salt glands on both surfaces of the fleshy leaf (Tomlinson 1986: for the evolution of the mangrove habitat, see Rhizophoraceae). Thes salt glands have largely radially-arranged cells in their heads (Fahn 1979), and appear to be variants of the common glandular hair type in Lamiales. Capsules open explosively in all taxa except Avicennioideae and some Thunbergioideae, and Witztum and Schulgasser (1995) discuss in detail capsule dehiscence in Acanthoideae with their distinctive retinacula.

Larvae of Nymphalidae-Melitaeini butterflies commonly feed on Acanthaceae (Wahlberg 2001).

Chemistry, Morphology, etc.Features characterizing Nelsonioideae mentioned 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, as is their rather undistinguished tricolpate or tricolporate pollen. There is some dispute as to whether Nelsonioideae have jaculators, but even if present, they certainly are not functional.

Mendoncia lacks iridoids. Inverted vascular bundles in the pith, or anomalous secondary thickening where an internal and inverted cambium develops, are scattered in the family, but neither have yet been found in Nelsonioideae or Avicennioideae (Schwarzbach & McDade 2002 for literature). Thunbergia has extrafloral nectaries on the calyx, while in Avicennia nectar is secreted from glands on the corolla tube (for details, see Tomlinson 1986). There is discussion as to the nature of corolla tube initiation, which is probably usually more or less late, rarely early (cf. Leins & Erbar 1997; Schönenberger & Endress 1998; see also Endress 1999 for floral development). In Avicennia there may be fewer corolla than sepal lobes ("connation" of a pair of the former?). Bravaisia (Acanthoideae) is distinctive in that it has small bracteoles and rounded calyx and corolla lobes (the former are more or less scarious); the anthers have short basal appendages. Pollen variation is extensive, but shows extensive homoplasy (e.g. Kiel et al. 2006). Indeed, the variation in pollen morphology in the family is spectacular: for variation within Strobilanthes sensu lato, see Carine and Scotland (2000) and Wang and Blackmore (2003), for that within Acanthoideae as a whole, see Daniel (1998) and Scotland and Vollesen (2000) and references. Many Isoglossinae (Justicieae) have distinctive "Gürtelpollen" (Kiel et al. 2006) - lenticular biporate pollen with a prominent circumferential band. The ovule of Avicennia is sometimes reported to be straight, and the endosperm at least is extra-ovular (Padmanabhan 1970). Acanthoideae [Thunbergioideae + "Avicennioideae"] group appears to lack a funicular obturator, but I am uncertain as to the polarity of this feature. The fruit of Avicennia is a capsule, according to Takhtajan (1997) and Schatz (2001), but it may split only as the seed germinates. In Mendoncia and relatives only one carpel is functional and the fruit is a drupe, while in Thunbergia and relatives the stigma is more or less trumpet-shaped and the fruit is a capsule, usually with four seeds.

The distinctive asymmetrical endosperm development found in Acanthaceae is also to be found in Lamiaceae-Nepetoideae (a parallelism). In Acanthoideae other than Acantheae, other details of endosperm development show considerable variablity. There is often a central area in which divisions are free nuclear, and walls may be laid down subsequently, but in some taxa there is what is known as a "basal apparatus", an area in which walls are not laid down; this pattern of endosperm development occurs in no other angiosperms (Ram & Wadhi 1964; Johri et al. 1992 and references). For cotyledon folding, see Schwarzbach and McDade (2002).

For general anatomy, capsule dehiscence, see van Tieghem (1908), for embryology, etc., see Mauritzon (1934), Johri and Singh (1959), and Wadhi (1970), and for stomata, see Rohweder et al. (1971). Some information on Nelsonioideae is taken from (1955) and in particular Johri and Singh (1959). For information on Thunbergia, etc., see Schönenberger (1999). For embryology, etc., of Avicennia, see Padmanabhan (1970, as Verbenaceae), for general information, see Tomlinson (1986) and Sanders (1997). Within Acanthoideae, for information on acicular fibres, see Bremekamp (1965: = "raphidines"), for chemistry, see H. F. W. Jensen et al. (1988), for corolla aestivation, which shows interesting variation, see Scotland et al. (1994), and for floral morphology, see Endress (1994b).

Phylogeny. The erstwhile Nelsoniaceae were placed sister to rest of Acanthaceae in Hedren et al. (1995), and this position seems quite firm. Within Nelsonioideae, Elytraria may be sister to the rest (two genera not included), although the position of Nelsonia was uncertain (Wenk & Daniel 2009); for further details of relationships here, see McDade et al. (2009). For the phylogeny of Acanthaceae, especially Acanthoideae, see McDade et al. (2008: see also McDade & Moody 1999; McDade et al. 2000a; McDade et al. 2006). Acantheae have neither cystoliths nor swollen nodes; see McDade et al. (2005) for a phylogeny. Taxa with one- and two-lipped corollas form separate clades, Old World and largely New World respectively. For Justicieae, mostly New World, see McDade et al. (2000b) and Kiel et al. (2009), and for relationships in the Tetramerium group, see Daniel et al. (2008). Ruellia is defined by pollen morphology, and it includes genera like Blechum, etc.; many taxa are cleistogamous (Tripp & Manos 2006; Tripp 2007). For more on phylogenetic relationships, see Scotland et al. (1995).

The position of Avicennia (Avicenniaceae) within Acanthaceae s.l. is fairly well established (note also the distinctive endosperm development that they have in common); it shows a rather weakly supported sister group relationship with Thunbergioideae (Schwarzbach & McDade 2002; Hilu et al. 2003). Support for the [Thunbergioideae + Avicennioideae] clade has remained rather weak and comes mostly from the chloroplast genes (McDade et al. 2008). Like many other Acanthaceae, Avicennia has swollen nodes. Borg et al. (2006) provide a phylogeny for Thunbergioideae and discuss their biogeography and the evolution of some characters.

Classification.Generic limits in Acanthoideae are difficult, as in other groups where the genera are often based on variation in corolla morphology that represent adaptations to particular pollinators (e.g. Daniel et al. 2008 and references); Bremekamp (1944) dismembered Strobilanthes into some 54 genera. Genera in Justicieae will need recircumscribing (Kiel et al. 2009). For a listing of genera in Acanthoideae, obviously needing amendment now, see Scotland and Vollesen (2000). The tribal classification of Acanthoideae follows that in McDade et al. (2008).

Previous Relationships. Nelsonioideae have often been placed in Scrophulariaceae s.l. or considered "intermediate" between Scrophulariaceae and Acanthaceae. Avicennia was often included in Verbenaceae, largely because of it more or less cymose inflorescence and gynoecium with two ovules per carpel, but the similarity is only superficial. For a discussion about Thomandersia, the fruits of which have a structure described as a retinaculum (cf. Acanthoideae), although they do not dehisce explosively, see Thomandersiaceae.

BYBLIDACEAE Domin, nom. cons.   Back to Lamiales

Rhizomatous and woody to ephemeral herbs; cork?; young stem with separate bundles; nodes 1:1 or 1:3; stomata paracytic; leaves spiral, linear, abaxially curved, veins parallel; flowers single, axillary, polysymmetric, bracteoles 0; K connate only basally, C contorted, connate only basally, margins fimbriate, stamens = and opposite sepals, shortly epipetalous, anthers with short slits or pores, epidermal cells ephemeral, nectary 0, 2-several ± apical ovules/carpel, stigma punctate to capitate (slightly bilobed); exotestal cells tangentially elongated, walls thickened, mesotesta sclerenchymatous; endosperm copious, starchy, with aleurone; n = 8, 9; proteinaceous inclusions in the nucleus?

1[list]/6. W. and N. Australia, S. New Guinea (map: from van Steenis 1971; FloraBase 2004). [Photos - Collection]

• Byblidaceae are rhizomatous to ephemeral herbs with linear leaves that are abaxially curled in bud and are densely covered by glands. The larger glands, with their heads formed from a single layer of cells radially divided, are conspicuous and look like little parasols; the sessile glands are smaller versions of these. The axillary, polysymmetric flowers have a broadly-spreading, contorted corolla that is only shortly connate at the base, anthers that are porose and whose apices form a cone, and a long style.

Evolution. Although there is no evidence that the plant absorbs nutrients from the insects that may stick to it (Hartmeyer 1997, 1998; Mueller et al. 2001), Conran and Carolin (2004) note that mirid bugs are associated with the genus, and so there may be a relationship similar to that in Roridula (Ericales) where the plant absorbs nutrients from the excreta of the bug.

Although the flowers are basically polysymmetric, the stamens are held to one side of the flower. Buzz pollination is likely.

Chemistry, Morphology, etc. Byblis linifolia has leaves that are abaxially curled in bud - in that respect like those of Drosophyllum (Drosophyllaceae, Caryophyllales) The glandular hairs of Byblidaceae have the typical structure of those of core Lamiales and look like little parasols; they are not vascularised and with irregularly arranged cells in the head as are those of Drosophyllum. Diels (1930b) draws the flower with the odd sepal abaxial. Byblidaceae are often described as being bitegmic, but cf. Diels (1930b) and Vani-Hardev (1972).

See Conran (1996: embryology), Cutler and Gregory (1998: anatomy), Conran et al. (2002: chromosome numbers), Conran and Carolin (2004: general), McPherson (2008: general), and the Carnivorous Plants Database (general) for more information, and Lloyd (1942) and Juniper et al. (1989) for details of the morphology of the plant.

Previous Relationships. Roridula (see Roridulaceae - Ericales) has hitherto often been placed in the same family as Byblis and included in Rosales, as by Cronquist (1981), or the two split, and both placed in Byblidales, a member of Aralianae, as by Takhtajan (1997).

LENTIBULARIACEAE Richard, nom. cons.   Back to Lamiales

Insectivorous rosette-forming herbs, also other derived growth forms; primary root reduced immediately after germination, lateral roots [?always] lacking a root cap; plants often Al-accumulators, little oxalate accumulation; ?cork; vessel elements?; stomata also anisocytic; hairs variously secreting mucilage and digestive enzymes; leaves spiral, usu. modified, or 0 and vegetative plant body not readily categorizable; inflorescence racemose; K 5-partite, or divided into 2 lobes, C abaxial lobe outside the others in bud [descending cochleate], with an abaxial nectar-secreting spur; A 2 [the abaxial pair], filaments stout [always?], thecae (superposed), confluent, epidermal cells ephemeral, pollen trinucleate, staminode 0, disc 0; G free-central or basal, integument 3-6 cells across, style often 0, stigma with broad lobes, wet, (sensitive); fruit a capsule of various types; exotestal cells variously thickened; endosperm 0, starchy, embryo green [1 record], cotyledons present [1 or 2, Pinguicula] or embryo minute, undifferentiated; n = 7-12+.

3[list]/320: Utricularia (220: great variation in testa), Pinguicula (80). World-wide, introduced into Hawaii? (map: from Hultén 1958, 1971; Taylor 1989). [Photo - Flower]

• Lentibulariaceae are vegetatively diverse insectivorous herbs of wet habitats. Pinguicula has flat leaves that are sticky on the adaxial surface, Genlisea has tubular leaves and forked, subsurface traps with the opening spiralling along the branches of the fork, while Utricularia may have leaves or not, but in either case the rest of the vegetative body is impossible to equate with conventional plant parts and there are often little animal-collecting bladders with trap-door entrances that open on stimulation of one of the four sensitive hairs. The flowers are much more uniform. They are strongly bilabiate, with a nectar-secreting spur, and there are only two anthers.

Evolution. Details of the morphology of these plants as it relates to their carnivorous proclivities are given by Lloyd (1942) and Juniper et al. (1989) in particular. It can be particularly difficult to understand plants of some species of Utricularia in ordinary morphological terms. Whole plants commonly develop from small fragments of leaves (Merl 1915), conventional roots do not occur, and the traps are remarkable structures without parallel in other flowering plants. Reifenrath et al. (2006 and references) discuses details of trap architecture of Utricularia (see also Merl 1915); many species have sensitive hairs the stimulation of which opens the trap. Jobson et al. (2004) and Laakkonen et al. (2006) suggest a possibly associated change in cytochrome c oxidase that may increase respiratory capacity so providing the energy needed for the rapid changes involved in the movements of the suction bladder-traps. It is possible that some Utricularia may eat aquatic algae, especially if the water is very acid; algae may predominate in traps in such environments (Peroutka et al. 2008). Indeed, there is a diverse microbial community in the traps - perhaps a mutualistic association - that may aid in the uptake of phosphorus by the plant (Sirová et al. 2009). Some species of Utricularia have non-suction traps rather more like those of Genlisea; there animals get passively trapped in the spiraling branches of the trap. Pinguicula, which alone among Lentibulariaceae has roots and embryos with cotyledons (the latter, not all species), has fly-paper traps. Throughout the family, each secretory gland is attached to a single epidermal cell and has no contact with vessels. Lentibulariaceae are notably prominent in acid habitats including those of the ephemeral flora of inselbergs (Seine et al. 1996).

Mutation rates in the matK gene in Genilsea in particular, and also Utricularia, are about the highest in all angiosperms (Müller et al. 2004), and that of other genes is also high (Jobson et al. 2003). At the same time, some species of Genlisea, e.g. G. margaretae have the smallest genomes known from angiosperms (Greilhuber et al. 2006).

Chemistry, Morphology, etc. Roots are lost in Genlisea and Utricularia. Utricularia kuhlmannii is described as having odd pinnate leaves by Merl (1915). "Nutritive tissue" is described from the ovule, funicle, or placenta, but not in Pinguicula, while in some taxa the embryo sac more or less escapes from the ovule and apparently takes nutrients from the placenta (Khan 1970 and references). The integument may be multiplicative in Genlisea (see Merl 1915).

For vegetative morphology, see Brugger and Rutishauser (1989) and Rutishauser and Isler (2001), for seeds and embryos, see Khan (1970) and G. Degtjareva (2004), for general information, see Fischer et al. (2004b), McPherson (2008: Pinguicula) and the Carnivorous Plants Database.

Phylogeny. For general phylogeny of Lentibulariaceae, see Jobson et al. (2003), Müller et al. (2004, 2006b), and also Müller and Borsch (2005a), while Cieslak et al. (2005) and Degtjareva et al. (2006) discuss the phylogeny and evolution of Pinguicula.

Classification. For a classic revision of Utricularia, see Taylor (1989).

Synonymy: Pinguiculaceae Dumortier, Utriculariaceae Hoffmannsegg & Link