LIGNOPHYTA

True roots +; lateral meristems: cork cambium producing cork abaxially, vascular cambium producing phloem abaxially and xylem adaxially.

EXTANT SEED PLANTS/SPERMATOPHYTA

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 derived from (some) sinapyl and particularly coniferyl alcohols, thus containing p-hydroxyphenyl and guaiacyl lignin units, (lignins derived from p-coumaryl alcohol, i.e. S [syringyl] lignin units); true roots present, apex multicellular, xylem exarch, and 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 and rays alone, tracheid/tracheid pits circular, bordered; mature sieve tube/cell lacking functioning nucleus, plastids with starch grains; phloem fibres +; stem cork cambium superficial, root cork cambium deep seated; leaves with single trace from sympodium ["nodes 1:1"]; stomata ?; leaf vascular bundles collateral; leaves megaphyllous [determinancy evolved first, then ad/abaxial symmetry], spiral, simple, lamina with vein density up to 5 mm/mm² [mean for all non-angiosperms 1.8]; axillary buds associated with at most some leaves; prophylls [including bracteoles] two, lateral; 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, developing after pollination, 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 duplications [three - [BP [A/N + C/O]] - copies], nrDNA with 5.8S and 5S rDNA in separate clusters; mitochondrial nad1 intron 2 and coxIIi3 intron and trans-spliced introns present.

MAGNOLIOPHYTA

Lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, [cyanogenesis in ANITA grade?], S [syringyl] lignin units common, positive Maüle reaction [syringyl:guaiacyl ratio more 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; shoot apex with tunica-corpus construction, tunica 2-layered; reaction wood ?, with gelatinous fibres; starch grains simple; primary cell wall mostly with pectic polysaccharides, poor in mannans; tracheid:tracheid [end wall] plates with scalariform pitting, wood parenchyma +; sieve tubes enucleate, sieve plate with pores (0.1-)0.5-10< µm across, cytoplasm with P-proteins, cytoplasm not occluding pores of sieve plate, companion cells from same mother cell that gave rise to the sieve tube; sugar transport in phloem passive; nodes unilacunar [1:?]; stomata with ends of guard cells level with pore, paracytic, outer stomatal ledges producing vestibule; leaves petiolate, lamina [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; most/all leaves with axillary buds; flowers perfect, pedicellate, polysymmetric, parts spiral [esp. the A], free, numbers unstable, development in general centripetal; P not sharply differentiated, with a single trace, outer members not enclosing the rest of the bud, often smaller than inner members; A many, filament not sharply distinguished from anther, stout, broad, with a single trace, anther introrse, tetrasporangiate, sporangia in two groups of two [dithecal], ± embedded in the filament, 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, 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, stylulus short, hollow, cavity not lined by distinct epidermal layer, stigma ± decurrent, dry [not secretory]; ovules few [?1]/carpel, marginal, anatropous, bitegmic, micropyle endostomal, outer integument 2-3 cells across, often largely subdermal in origin, inner integument 2-3 cells across, often dermal in origin, parietal tissue 1-3 cells across [crassinucellate], nucellar cap?; megasporocyte single, hypodermal, megaspore tetrad linear, functional megaspore chalazal, lacking sporopollenin and cuticle; female gametophyte four-celled [one module, nucleus of egg cell sister to one of the polar nuclei]; P deciduous in fruit; seed exotestal; pollen binucleate at dispersal, trinucleate eventually, germinating in less than 3 hours, pollination siphonogamous, tube elongated, growing at 80-600 µm/hour, with pectic outer wall, callose inner wall and callose plugs, growing between cells, penetration of ovules via micropyle [porogamous] within ca 18 hours, distance to first ovule 1.1.-2.1 mm, tube moves between nucellar cells; double fertilisation +, endosperm diploid, cellular [micropylar and chalazal domains develop diffently, first division oblique, micropylar end initially with a single large cell, divisions uniseriate, chalazal cell smaller, divisions in several planes], copious, oily and/or proteinaceous, embryo cellular ab initio, minute; germination hypogeal, seedlings/young plants sympodial; Arabidopsis-type telomeres [(TTTAGGG)_n]; whole genome duplication, ndhB gene 21 codons enlarged at the 5' end, 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 three copies of the PHY gene, [PHYB [PHYA + PHYC]].

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. This is because some taxa basal to the [magnoliid + monocot + eudicot] group have been surprisingly little studied, there is considerable homoplasy as well as variation within and between families of the ANITA grade 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... For other features such as details of sugar transport in the phloem, their placement on the tree is frankly speculative. Finally, for features such as parietal tissue/a nucellus only one (Nymphaeales) to three cells thick above the embryo sac and a stylar canal lacking an epidermal layer, although plesiomorphous for basal grade angiosperms (Williams 2009), I am unsure where on the tree a thicker nucellus and a stylar epidermal layer are acquired.

[NYMPHAEALES [AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]]: vessels +, elements with elongated scalariform perforation plates; wood fibres +; axial parenchyma diffuse or diffuse-in-aggregates; 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] / CORE ASTERIDS: ellagic acid 0, non-hydrolysable tannins not common; sugar transport in phloem active; inflorescence basically cymose; C forming a distinct tube; A epipetalous, = and opposite sepals or P, polyandry associated with increased numbers of C or G, very uncommon; (pollen with orbicules); duplication of the PI gene.

Evolution. Estimates of the age of diversification within core asterids ranges from (109-)100, 93(-85) million years (see Bell et al. 2010 for details).

ASTERID I / LAMIIDAE: G [2], superposed; 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!]; [vascularized] 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 +.

Chemistry, Morphology, etc. Information on placentoid distribution is needed for Plocospermataceae, etc.

LAMIALES Bromhead  Main Tree, Synapomorphies.

Cornoside, verbascosides [caffeoyl phenylethanoid glucosides (CPGs), caffeic acid esters, = acteosides], methyl- and oxygenated flavones +; eglandular hairs multicellular; leaves opposite; endosperm with micropylar haustorium; fruit a septicidal capsule; cotyledons incumbent; protein bodies in nuclei; mitochondrial coxII.i3 intron 0. - 23 families, 1059 genera, 23810 species.

Evolution. Divergence & Distribution. 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. Magallón and Castillo (2009) offer estimates of ca 80 and 63 million years for both relaxed and constrained penalized likelihood datings for stem and crown group Lamiales respectively - but note topology. However, Martínez-Millán (2010), using a rather scanty fossil record, suggested an Oligocene age for diversification within Lamiales - this would be the herbaceous Lamiales, since Oleaceae are known from the Eocene.

Lamiales contain ca 12.3% eudicot diversity. Most of this diversity is concentrated in families whose members are largely herbaceous to shrubby and have rather large, monosymmetric flowers and many small seeds.

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. Endress (2011a) suggested that a key innovation somewhere in Lamiales was tenuinucellate ovules.

Bacterial/Fungal Associations. Both parasitic and insectivorous members of Lamiales are largely non-mycorrhizal (Brundrett 2004 and references).

Confirmation of the phylogenetic positions of Carlemanniaceae, placed sister to Oleaceae (see below), and of Plocospermataceae, as well as studies of their anatomy, chemistry, etc., and also resolution of relationships within Oleaceae, are important for understanding the evolution of the chemistry and floral morphology in particular of 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 likely, 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 petal-like whorl (Mayr & Weber 2006); evolution of floral merism in this part of the tree is obviously a complex matter, and exactly where changes in this feature are to be placed on the tree is unclear.

Chemistry, Morphology, etc. The occurrence 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 trisaccharide derivatives (over 325 structures altogether - S. R. Jensen, pers. comm.) 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). 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; Turgeon 2010a). 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).

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), Jensen (1992), 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).

Phylogeny. The 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 [Carlemanniaceae + 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 peltate, glandular hairs with unicellular stalks and the two stamens (their position is not entirely certain) also suggest Lamiales, and anatomical features that are known (see below) are consistent with this relationship.

Finding the position of Hydrostachys in the asterid part of the tree has proved difficult. Here it is included in Cornales, and there is a discussion of its relationships there. Recent work (Burleigh et al. 2009) had suggested that it should be included in Lamiales, with which its morphology is in general agreement. If it should ends up in Lamiales, it is likely to be towards 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.

Relationships in the "Scrophulariaceae"-Acanthaceae-Bignoniaceae-Lamiaceae area have been uncertain for some time, 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 is difficult to evaluate; Freeman and Scogin (1999) focussed on the old Scrophulariaceae, but the pattern of relationships they found was unclear. A tree in Müller et al. (2004) suggested that at least a partial resolution of relationships is in sight, although sampling 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) began to clarify 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 2007a). 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); the former position is more likely (Schäferhoff et al. 2010). Petraea (Verbenaceae) was sister to Bignoniaceae in some early molecular phylogenies (Wagstaff & Olmstead 1997, consistent with its position in Schäferhoff et al. 2010). 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 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) suggested that substantial progress in disentangling relationships around Lamiacae-Verbenaceae and Scrophulariaceae s.l. might be on the horizon (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. Indeed, the recent results of Schäferhoff et al. (2010) are consistent with the outline suggested by Refulio-Rodriguez and Olmstead (2008). The bulk of the free-living Scrophulariaceae are paraphyletic at the base of the iridoid-containing clade of Lamiales, there can be little further doubt that Lamiaceae and Verbenaceae are to be separated, the insectivorous and putatively insectivorous clades in Lamiales are all unrelated, etc. (Schäferhoff et al. 2010 - and another immediately unrelated carnivorous clade has recently been added - see Pereira et al. 2012). Note that Albach et al. (2009) has cast doubt on the monophyly of Phrymaceae (and see also Schäferhoff et al. 2010) while Qiu et al. (2010) and Soltis et al. (2011) suggest that Peltanthera may fall outide the [Calceolariaceae + Gesneriaceae] clade. Taxon limits in these two areas, narrowly drawn below, will probably have to be adjusted, but I wait for better sampling... As might be anticipated, there is little morphological support for internal nodes of this iridoid-containing clade and even for some of the families.

The pattern of duplication of the FLO=LFY and DEF=AP3 genes within Lamiales suggests very interesting relationships, if confirmed; duplication occurred 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), 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), but Olmstead (2002) has provided a readable account of some of the recent changes in our ideas of relationships in the Scrophulariaceae s.l. in particular.

Despite the lack of morphological support for some of the families, little is to be gained and more lost if their limits are much expanded.

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

Synonymy: Acanthales Berchtold & J. Presl, Antirrhinales Döll, Aragoales D. Don [?status], Bignoniales Berchtold & J. Presl, Byblidales Reveal, Callitrichales Link, Carlemanniales Doweld, Fraxinales Berchtold & J. Presl, Gesneriales Berchtold & J. Presl, Globulariales Dumortier, Hippuridales Link, Jasminales Berchtold & J. Presl, Lentibulariales Berchtold & J. Presl, Ligustrales Bischof, Myoporales Berchtold & J. Presl, Oleales Berchtold & J. Presl, Orobanchales Berchtold & J. Presl, Pedaliales Berchtold & J. Presl, Pinguiculales Dumortier, Plantaginales Berchtold & J. Presl, Rhinanthales Dumortier, Scrophulariales Lindley, Selaginales Martius, Stilbales Martius, Utriculariales Döll, Verbascales Döll, Verbenales Berchtold & J. Presl, Viticales Link {?status]

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; plant cryptically dioecious; inflorescences axillary, bracteoles 0; flowers 5-6-merous; staminate flowers: anthers extrorse, versatile, with parallel separate thecae; nectary 0; style 0; carpellate flower: nectary +; placentation parietal, style divided twice, branches stigmatic, not expanded; ovules 2/carpel; 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 fibres 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 (1981) included the genus in his Apocynaceae, probably because of the hairs on its seeds.

[[Carlemanniaceae + Oleaceae] [Tetrachondraceae [Peltanthera, [Calceolariaceae + Gesneriaceae], [Plantaginaceae [Scrophulariaceae [Byblidaceae, Linderniaceae, [Stilbaceae [[Lamiaceae [Mazus etc. [Phrymaceae [Paulowniaceae + Orobanchaceae]]]] [Thomandersiaceae, Verbenaceae, Pedaliaceae, [Schlegeliaceae + Martyniaceae] Bignoniaceae, Acanthaceae, Lentibulariaceae]]]]]]]]]: 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; pollen tricolpate; stigma ± clavate; exotestal cells ± palisade, endothelium persistent.

CARLEMANNIACEAE Airy Shaw   Back to Lamiales

Perennial herbs or shrubs; chemistry?; vessel elements?; nodes?; no cuticle waxes; lamina margins toothed; inflorescences terminal and axillary; flowers weakly obliquely monosymmetric, 4- or 5-merous, heterostylous [Silvianthus]; K members unequal or not, ± linear, open?, C induplicate-valvate; anthers connivent and surrounding the style, introrse, thecae ?; 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. 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, lamina vernation conduplicate [Chionanthus], margins entire to toothed, (2ndary veins palmate); (plant dioecious); flowers 4-merous; K valvate; anther thecae ± back-to-back; pollen (trinucleate); (style short), stigma dry; ovules apical, (hemitropous), epitropous or apotropous, integument ca 7 cells across, (postament +); testa often vascularized, exotesta moderately and evenly thickened, (endotesta fibrous; endothelium ?not 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 [secologanoside] +; pits ± vestured; C free, imbricate; ovule 1/carpel; fruit a samara; n = 13.

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

2. Forsythieae L. Johnson

Pith chambered; C valvate; ovules 1-several/carpel; fruit a samara or capsule; n = 14.

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

3. Myxopyreae Boerlage

Myxopyroside iridoid pathway +; inverted cortical bundles in corners of angled stem (Dimetra not); (K diagonally oriented, C contorted, early tube formation - Nyctanthes); placentation ± basal; ovules 1(-3)/carpel, integument to 20 cells across [Nyctanthes]; (megaspore mother cells several, embryo sac bisporic, 8-nucleate [Allium type]); fruit a berry or schizocarp; n = 11, 12.

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

Synonymy: Nyctanthaceae J. Agardh

Jasmineae + Oleeae: secoirioidoids [oleoside] +; ovules 2(-4)/carpel; fruit fleshy.

4. Jasmineae Lamarck & Candolle

K and C to 14 or more, first 4 K diagonal, C quincuncial-imbricate, tube formation early; endothelium 0; (megaspore mother cells several, several elongated embryo sacs developing); 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 Jussieu

5. Oleeae Dumortier

Flavone glycosides +; vessel elements in multiples; (pits vestured); libriform fibres + (0); fibre tracheids 0 (+); marginal parenchyma +/0; (indumentum of peltate scales); K (open), C valvate, (imbricate; 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 Meisner, 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. Divergence & Distribution. Samaras of Fraxinus have been reported from Eocene rocks of that are some 44 million years of age (Call & Dlicher 1992). Much diversification within Oleaceae is Tertiary (Besnard et al. 2009a).

Plant-Animal Interactions. Caterpillars of some Sphinginae are quite common 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). Damtoft et al. (1995) noted that the secoiridoids of Fontanesia (loganic acid, etc., and 5-hydroxylated derivates like swertiamarin) were produced by a somewhat different biosynthetic pathway than the oleoside-type secoiridoids common elsewhere in the family. Forsythia and Abeliophyllum (possibly sister to rest of Oleaceae) 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: Dute et al. 2010b). 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), and epidermal crystals often clustered in cells at the bases of trichomes, an unusual distribution pattern (Lersten & Horner 2009b).

The calyx is sometimes diagonally oriented (Sehr & Weber 2009). There is variation in corolla tube initiation, both early and late initiation being known in the family (Sehr & Weber 2009). 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; however, whether they are collateral or superposed, the two stamens are always borne in the plane of the ovary 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 Andersson (1931), Kapil and Vani (1966), and Maheshwari Devi (1975), all embryology, 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) and dadpur et al.(2011), both floral ontogeny, latter also some inflorescence.

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 tritomy; they emphasized the complex pattern of chloroplast inversions in Jasminieae. Kim and Kim (2011) suggested quite well supported set of groupings [[Fontanesieae + Jasminieae] [Oleeae + Forsythieae]]; unfortunately, they did not sample other members of the family, nevertheless, the relationships suggested above may well need to be changed. Franzyk et al. (2001) noted that Myxopyrum and Nyctanthes, both in Myxopyreae, had similar iridoids. Besnard et al. (2009a) and Guo et al. (2011) examined relationships in some Oleeae.

Classification. The tribes recognised above are those of Wallander and Albert (2000). Generic limits in Oleeae in particular need much attention; Olea itself, Osmanthus, and Chionanthus are all polyphyletic (Besnard et al. 2009a; Guo et al. 2011).

Previous Relationships. The position of Nyctanthes has been uncertain, and it has often been included in Verbenaceae in the old sense; Filonenko et al. (2010) consider the genus distinct from both families. Nyctanthes has a rough lamina surface, transverse carpels with 1 ascending ovule/carpel, and seeds with exotesta little developed, mesotesta persistent, and the endotestal cells tangentially elongated and sclerotised.

[Tetrachondraceae [Peltanthera [Calceolariaceae + Gesneriaceae], [Plantaginaceae [Scrophulariaceae [Byblidaceae, Linderniaceae, [Stilbaceae [[Lamiaceae [Mazus etc. [Phrymaceae [Paulowniaceae + Orobanchaceae]]]] [Thomandersiaceae, Verbenaceae, Pedaliaceae, [Schlegeliaceae + Martyniaceae] Bignoniaceae, Acanthaceae, Lentibulariaceae]]]]]]]]: 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 free, thecae ?; pollen in tetrads, 6-sulcate, psilate; nectary 0; G 4 partite, slightly inferior and placentation basal [Tetrachondra], placentae peltate [Polypremum], style gynobasic [Tetrachondra], stigma small, subglobose; ovules 2/carpel, or many [Polypremum], integument 3-4 cells across [Polypremum]; fruit with persistent green K, either a schizocarp, or a loculicidal (+ septicidal) capsule; testa thin, endothelial cells with persistent thickened inner walls; 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.

[Peltanthera, [Calceolariaceae + Gesneriaceae], [Plantaginaceae [Scrophulariaceae [Byblidaceae, Linderniaceae, [Stilbaceae [[Lamiaceae [Mazus etc. [Phrymaceae [Paulowniaceae + Orobanchaceae]]]] [Thomandersiaceae, Verbenaceae, Pedaliaceae, [Schlegeliaceae + Martyniaceae] Bignoniaceae, Acanthaceae, Lentibulariaceae]]]]]]]: shikimic-acid derived anthraquinones [see also Rubiaceae], 6- and/or 8- hydroxylated flavone glycosides + [? Tetrachondraceae], storage substances stachyose and other oligosaccarides; flowers vertically monosymmetric; K ± asymmetrical, C bilabiate, 2-lobed upper lip, 3-lobed lower lip [= (2:3)], adaxial lobes outside the others in bud [ascending cochleate], tube formation late; A 4, didynamous, anthers connivent; pollen tubes lacking callose; ovules many/carpel; endosperm also with chalazal haustoria [?position in tree].

Evolution. Divergence & Distribution. This clade includes the bulk of the diversity within Lamiales; many members are herbaceous or shrubby and have often quite large monosymmetric, bilabiate flowers, and fruits with many small seeds. A didynamous androecium with stamens in two pairs of unequal lengths, is common, and the fusion, or at least close attachment, of the paired anthers may improve pollen removal from the flower (Ren & Tang 2010). The evolution of monosymmetric flowers may be a key innovation in Lamiales (Endress 2011) and would be pegged to this level, but monosymmetry is not a simple character. It is clear (e.g. Damerval & Manuel 2003; Gübitz et al. 2003; Preston et al. 2011b) that CYC genes have duplicated independently within the clade and become separately involved in the development of the strongly monosymmetric flowers of Antirrhinum, Mimulus and Gesneriaceae.

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 occurrence 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 vascularized by branches from the main carpellary vascular traces - examples are Schlegeliaceae, some Pedaliaceae, Verbenaceae. On the other hand, the nectary is vascularized 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 callose, see Prospéri and Cocucci (1979: Oleaceae, etc., not sampled), and 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).

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

Chemistry, Morphology, etc. In paired-flower cymes the two flowers of the flower-pair have the same orientation. 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". 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.

Phylogeny. The exact position of Peltanthera remains to be determined.

PELTANTHERA   Back to Lamiales

Small tree; verbascosides, cornoside derivatives +; nodes 3:3; petiole bundle flattened-annular, with (medullary and) rib bundles; hairs branched-moniliform; lamina vernation involute, margins serrate; inflorescence axillary, thyrsoid; flower slightly monosymmetric; K largely free, C valvate; A 5, thecae confluent; seeds dust-like; n = ?

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

Chemistry, Morphology, etc. For information on general morphology, see Hunziker & Di Fulvio (1957), and for chemistry, see. However, the plant is very poorly known.

Phylogeny. Placed with Gesneriaceae (e.g. Oxelman et al. 1999a), and perhaps to be included in Gesneriaceae, but cf. Soltis et al. (2011).

CALCEOLARIACEAE Olmstead   Back to Lamiales

Herbaceous to shrubs; cork?; lamina margins toothed (entire); flowers 4-merous; K orthogonal, 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. Divergence & Distribution. Crown group diversification began (27-)15(-4) million years ago (Renner & Schaefer 2010). The bulk of the diversity in the family is included in Calceolaria.

Floral Biology & Seed Dispersal. Sternotribic flowers pollinated by Centris bees seem to be plesiomorphous in Calceolaria; species with this and some other morphological features are diploid and have a geographic range that is basically Chilean (Cosacov et al. 2009). 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 or specialised hairs, or pollen if there are no oil glands; specialised food bodies are the reward for species that are pollinated by non-nectarivorous birds (Vogel 1974; Sérsic & Cocucci 1996; Rasmussen & Olesen 2000). All told, about 4/5 of the genus have oil flowers (Sérsic 2004), and the ability to produce oil has been lost several times (Renner & Schaefer 2010). Calceolaria crown group age is only (6-)5(-1) million years, but oil glands may have been acquired ant time after the split of Calceolaria from Jovellana (i.e., the crown group age for the whole family, see above: Renner & Schaefer 2010).

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 Richard & Jussieu, nom. cons.   Back to Lamiales

Usu. herbs or weak-stemmed trees (trees), often epiphytes [ca. 1/5 spp.]; hairs of stalked glands, or with thickened terminal cells; (cambium storied); (vessel elements with scalariform perforation plates); nodes 1(-3 or more):1(-3 or more), (+ split laterals); petiole bundle(s) arcuate; stomata anisocytic; leaves joined at the base, (anisophyllous; two-ranked; spiral), vernation involute, margins toothed or entire; inflorescence axillary (terminal); K connate, abaxial C lobe outside others in bud [aestivation descending cochleate], (C spurred); A (5, 2), anthers didynamous, connivent in pairs, (thecae apically confluent), staminode 1 (0); nectary vascularized; placentation intrusive parietal, placentae ± triangular, usu. covered by ovules, stigma broadly bilobed to trumpet-shaped, wet or dry; integument 3-5 cells across; fruit a septicidal capsule; exotestal cells variously elongated and thickened, endotestal cells at most simply persisting; GCyc duplication.

147(+)[list]/3311 - 3 main groups below. Largely tropical. [Photo - Flower]

Didymocarpoideae + Epithematoideae: 3-desoxyanthocyanins 0, chalcones, aurones +; ?stomata; ovary wall not richly vascularized, 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]); placentae lamelliform-recurved, ovules restricted to distal end, ovary gradually narrowed into the style; (fruit with septicidal and loculicidal dehiscence; ± elongated, twisted; circumscissile; a berry); testa cells little elongated, (ornamented: with [extremely long] hairs); n = (4, 8) 9-11 (12, 13) 14-17, etc., polyploidy not uncommon.

82/2150: Cyrtandra (600), Aeschynanthus (185), Chirita (180), Henckelia (155), Streptocarpus (155), Paraboea (120), Agalmyla (100), Didymocarpus (100), Oreocharis (85). S. Europe (scattered), Africa and Madagascar, mostly Sri Lanka to Malesia (especially southern China) 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); (placentation axile), ovary short, abruptly narrowed into the style; endosperm ?0; n = (8-)10(-12); (seedling primary root not developed).

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.

63/1086. Predominantly Neotropical, a few S.W. Pacific, East Asia.

3a. Coronanthereae

Trees to ± shrubby-herbaceous (rooting from the nodes); stomata anomocytic (paracytic); (flowers polysymmetric); (C fringed); (A 2 [adaxial pair]; 5); nectary embedded in G wall, vascularized from A traces; capsules septicidal (and loculicidal; fruit a berry, placentae fleshy); 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, nectary vascularized from numerous vascular bundles in wall; (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 quite fleshy and softly hairy and have arching, ascending secondary veins that do not 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. Divergence & Distribution. Gesneriaceae may have diverged from other Lamiales 74-71 million years ago, crown group diversification beginning very soon afterwards (Wikström et al. 2001; Bremer et al 2004).

Möller and Cronk (2002) discussed biogeographic relationships within the large African genus Streptocarpus. Within Coronanthereae there seems to have been one (Smith et al. 2006) or two (Woo et al. 2011) E to W dispersal events across the Pacific. Cyrtandra, with its baccate fruits, is a very diverse genus found throughout Malesia, being particularly speciose in places like New Guinea. It is also 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). Fiji may have been the first area in the Pacific colonized somewhat over 20 million years ago; the Hawaiian colonization, also from the west, was independent of that of the other Pacific islands and happened very soon afterwards; there are now ca 60 endemic species on the islands. Note, however, that diversification within the genus may have begun much earlier, ca 48 million years ago in Malesia. (Clark et al. 2009). Within Gloxinieae (Gesnerioideae) diversification occurred some 30-20 million years ago (Roalson et al. 2008b: see also biogeographic relationships).

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

Floral Biology & Seed Dispersal. 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; for instance, Saintpaulia-type flowers have the buzz pollination syndrome (Clark et al. 2011). Wiehler (1978) estimated that perhaps 60% of neotropical Gesnerioideae - some 600 species - were hummingbird pollinated, indeed, the centers of diversity of both neotropical Gesneriaceae and of humminbirds are in the Colombia-Ecuador region (Weber 2011; see also Ericaceae). Wiehler (1978) divided the floral morphologies involved into three common and one less common "types". He thought that another ca 30% of Gesnerioideae were pollinated by euglossine bees of both sexes (cf. e.g. Orchidaceae where it is only male bees seeking scents that are involved); divergence of these bees occurred only 42-27 million years ago (Ramírez et al. 2010). Pollination by birds is relatively less common in Old World Gesneriaceae. More or less polysymmetric flowers - the corolla may be rotate, but the androecium is often technically monosymmetric - 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 here compared with some other families of the strongly monosymmetric Lamiales (Endress 1997a). Relatively little is known about the pollination of such flowers, although as might be expected buzz-pollination has sometimes been recorded (Clark et al. 2011 and references). Martén-Rodriguez et al. (2010) discuss the variety of pollinators of Caribbean Gesnerieae.

Flowers with inverted orientation are known from some Episcieae (Clark & Zimmer 2003); they seem to have evolved ca 3 times. 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 by some definitions.

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.

Ecology & Physiology. 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). Hardly surprisingly, epiphytes are common, with well over 400 epiphytic species being known from neotropical Episcieae alone (Madison 1977; Weber 1978; Gentry & Dodoson 1987); the evolution of epiphytism within Coronanthereae is described by Salinas et al. (2010).

Vegetative Variation. 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. Harrison et al. (2005a) had found that genes involved in shoot development were expressed on the peiole in rosulate species of the genus, plants with this growth form producing leaves, etc., from the "petiole". These genes were not expressed in strictly unifoliate species (Harrison et al. 2005a). Plant architecture is also 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. The developmental pathways controlling meristem development become relocalised (Mantegazza et al. 2009). Thus a meristem develops at the base of the cotyledon and inflorescences also develop at the base of the lamina, however, in some species the flowers arise along the midrib of the blade rather than from separate inflorescences (Imaichi et al. 2001; see also Tsukaya 2005); the "petiole" (petiolode) itself of Streptocarpus, at least, is unifacial, although not at the seedling stage, when it is bifacial (Tononi et al. 2010). The cotyledon that keeps on growing is the one that is exposed to more light (Saueregger & Weber 2005). The plant body of some Monophyllaea can become more complex by repetition of the cotyledonary unit. For anisocotyly, although more accurately, one cotyledon is accrescent, and its development in [Didymocarpoideae + Epithematoideae], see Burtt (1970) and Saueregger and Weber (2004). The seedling radicle may not develop, although a failure to develop has been noted in other Epithematoideae (Imaichi et al. 2001).

Monophylly in general seems to be an adaptation for life on rocks. The plant grows in cracks of the rock and the single leaf hangs down and covers the rock surface quite efficiently; that the better lighted cotyledon of Monophyllaea develops make sense in this context. Note that plants whose stems consist of sprays of alternating leaves - if they are opposite, then there is often strong anisophylly - are also common in such situations; again, a photosynthetic surface covering the rock is deployed.

Chemistry, Morphology, 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 fibres 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. Nodal anatomy is quite variable (see also Howard 1970), but this has never been systematically surveyed. In addition to its distinctive anatomy, Gesneria also 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), Wang et al. (2010), and especially Mayer at al. (2003).

Within Didymocarpoideae, Haberlea and Ramonda, temperate, European, and with polysymmetric flowers and five stamens, may be sister to the rest (e.g. Mayer et al. 2003) or more likely near basal (Möller et al. 2009; Wei et al. 2010; Wang et al. 2010); they have dihydrocaffeoyl ester found nowhere else in flowering plants (Jensen 1996). Some studies (Möller et al. 2009) placed a number of small Asian and European clades all with four or five, rarely two, 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). Wang et al. (2010: Jerdonia not included) found that Corallodiscus, the Ramonda clade, and Streptocarpus are successive branches in the phylogeny; taxa with actinomorphic flowers are scattered through the tree (see above).

Although Didymocarpus itself has been dismembered (Weber & Burtt 1998) and many species placed in Henckelia, the circumscription and relationships of that genus are still unclear (Möller et al. 2009). For the phylogeny of the didymocarpoid Gesneriaceae, see also Möller et al. (2011a). 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; Clark et al. 2009). For relationships in Streptocarpus, to include Saintpaulia, see Möller and Cronk (2001), for a study of Aeschynanthus linking seed morphology and geography, see Denduangboripant et al. (2001), for a phylogeny of Chirita and relatives, see Wang et al. (2011) and in particular Weber et al. (2011), and for relationships in an expanded Oreocharis, see Möller et al. (2011b).

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) and Woo et al. (2011). 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. Relationships along the base of the spine of Gesnerioideae were only weakly supported in Woo et al. (2011). For other relationships in Gesnerioideae, 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 above) go is unclear; the latter is very similar in wood anatomy to Buddleja (but both genera are woody!), 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 in particular (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 current tribal classifications of Didymocarpoideae are decidedly unsatisfactory (Möller et al. 2009), and some genera, perhaps most notably Chirita, are polyphyletic. Indeed, Möller et al. (2011a) found that only 12/29 genera with more than one species they sampled were monophyletic, and there are also many monotypic genera - thus Oreocharis has recently been expanded to include eleven mostly very small Chineae genera (Möller et al. 2011b).

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. Primulina, originally monotypic, has been greatly expanded in the course of understanding the limits of Chirita (Weber et al. 2011), the lmits of Paraboea have been adjusted (Puglisi et al. 2011), and so it goes on.

[Plantaginaceae [Scrophulariaceae [Byblidaceae, Linderniaceae, [Stilbaceae [[Lamiaceae [Mazus etc. [Phrymaceae [Paulowniaceae + Orobanchaceae]]]] [Thomandersiaceae, Verbenaceae, Pedaliaceae, [Schlegeliaceae + Martyniaceae] Bignoniaceae, Acanthaceae, Lentibulariaceae]]]]]]: route II decarboxylated iridoid glucoides [aucubin, catalpol widespread], 6- or 8-hydroxyflavones or 6 methoxyflavones +, cornosides 0; (embryo sac haustoria +).

Evolution. Plant-Animal Interactions. 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 this shift may have been followed by an increased diversificatioon rate (Fordyce 2010). 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 here (Lamiaceae, Pedaliaceae, Plantaginaceae: Noguchi et al. 2009); iridioid acquisition seems best placed here, with an independent origin in Oleaceae. 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, sagittate, or on connective arms, confluent [Penstemon] or not, (staminode + [esp. Cheloneae, Antirrhineae]/0); pollen exine tectate and reticulate; (placentation intrusive parietal); ovules (-1/carpel), campylotropous?, 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 +, mannose-rich polysaccharides + [?distribution], (embryo green; short; curved); n = 6-10 +; protein bodies in nucleus amorphous [excluding Angelonieae and Gratioleae].

Ca 90[list]/1900: 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]

• Plantaginaceae s.l. are very difficult to identify and distinguish from Scrophulariaceae s. str., Gesneriaceae, Stilbaceae, etc., all having largish, monosymmetric flowers, furthermore, Hippuris and Callitriche and genera like the more or less wind-pollinated Plantago are also in this clade, and morphological diversification has been very extensive 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 common in the family.

Evolution. Plant-Animal Interactions. 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).

Floral Biology & Seed Dispersal. Floral morphology is very variable (see Reeves & Olmstead 1998), but Plantaginaceae are predominantly pollinated by large insects and birds; Kampny (1995: as Scrophulariaceae) discussed pollination in the family as a whole. 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 (Kampny 1995; see Baldwin et al. 2011 for floral evolution in the genus). 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 single well-developed abaxial spur, but its 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 the distinctive morpgology of the mutant that he proposed to place it in a genus of its own, Peloria. Of course, Antirrhinum majus is a model organism used for understanding the development of monosymmetric flowers and the involvement of the CYC gene in this (e.g. Rosin & Kramer 2009; Preston et al. 2011 for references); duplication of the gene is evident in Antirrhineae, but not in Digitalis (Gübitz et al. 2003).

Floral evolution in the Veronica/Plantago clade is becoming better understood. Veronica has an open, 4-lobed corolla, but only two stamens; in some taxa there are two main veins in the adaxial corolla lobe, perhaps suggesting that it is formed by the fusion of the two adaxial lobes so common in monosymmetric members of the family. Wulfenia, sister to Veronica, has tubular but rather weakly lipped (2 + 3) flowers. Aragoa has 4-merous, polysymmetric flowers (cf. Oleaceae and Tetrachondraceae!), but with five sepals. Plantago, sister to Aragoa and also polysymmetric, has four sepals, petals and stamens, and wind pollination occurs there; the evolution of such flowers is connected with the degeneration of some floral symmetry genes (Preston et al. 2011a). The Plantago clade is 5-17 million years old (Cho et al. 2004; Rønsted et al. 2002b); for relationships within the clade, see Rønsted et al. (2002b, also Rahn 1996). Bello et al. (2004: see Bello et al. 2002 for a phylogeny) discuss floral evolution in this part of the family, also emphasizing the evolution of polysymmetry. Muñoz-Centeno et al. (2006) describe seed morphology and phylogeny; the seeds are mucilaginous, which may have facilitated the three dispersals in the genus from Australia to New Zealand (Tay et al. 2010).

Ecology & Physiology. Philcoxia, a quite recently described white sand endemic from Brasil, has been suspected of being carnivorous (Fritsch et al. 2007). This has recently been confirmed by Pereira et al. (2012). The plants lack mycorrhizae, as is common when there is carnivory. Nematodes become stuck to the glandular secretions covering the leaves, which are underground, and are digested by the plant - phosphatase activity has been recorded in the hairs (Pereira et al. 2012).

Genes & Genomes. 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 (but not in the chloroplast genome) that are 3,000-4,000 times those of nearly all other angiosperm clades (Cho et al. 2004; Mower et al. 2007). In an interesting development, at least three mitochondrial genes have recently been transferred from Cuscuta to species of the Plantago coronopus group (Mower et al. 2010).

In Veronica s.l. no correlation was found between speciation rates and rate of molecular evolution (Müller & Albach 2010).

Chemistry, Morphology, etc. Details of characters like the distribution of hair morpholgies and timing of androecium initiation remain to be clarified, but 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).

Penstemon is reported () to have a storied cambium. Veronica lyallii has successive subhypodermal phellogens (Gray 1937), while Besseya and Plantago have a foliar endodermis. Hair morphology may be of interest, although 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 have similar hairs (Raman 1991 and references).

Penstemon may have paired-flower cymes (elsewhere in Lamiales to be found in Gesneriaceae and Calceolariaceae). In a number of taxa in this clade the androecium is initiated before the corolla, but other patterns also occur, so androecium initiation is perhaps unlikely to be a synapomorphy for Plantaginaceae (Bello et al. 2004, cf. Judd et al. 2002). Bello et al. (2004) note that the Veronica/Plantago, as well as Digitalis, are members of a clade that has descending-cochleate aestivation, i.e. in bud the abaxial corolla lobes are outide the others. Petals have sometimes been lost in Synthyris (Hufford 1992b). Illustrations in Chatin (1874) suggest that the ovules of Veronica may be crassinucellate...

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 Schmid (1906: ovules, Scrophulariaceae s.l.), Junell (1961: gynoecium), Elisens and Tomb (1983: considerable seed variation even in Antirrhineae), 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 Schrock and Palser (1967), Leins and Erbar (1988, 2010), and Endress (1999).

Phylogeny. For the circumscription of Plantaginaceae, 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), also Olmstead and Reeves (1995) and Reeves and Olmstead (1998). 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 included in it, although it was not sampled. However, the limits and correct placement of this clade need confirmation. Estes and Small (2008) found that a clade including Angeloneae were sister to Gratioleae, with Limnophila being part of Gratioleae (1.0 posterior probabilities: Limosella [see Linderniaceae] was not sampled). 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 Gratiolaceae in Plantaginaceae, although Limosella itself is again in Scrophulariaceae. Although Gratiolaceae may form a major clade in Plantaginaceae (see also Schäferhoff et al. 2010), there seems no point in putting them in a separate family.

For the phylogeny of Antirrhineae, see Ghebrehiwit et al. (2003), and for that of Veroniceae, see Albach et al. (2004a, c, 2005c), Taskova et al. (2004, 2006), and Albach and Meudt (2010); the "new" molecular relationships are at least sometimes supported by other data such as chromosome number and iridoid type (Albach et al. 2004b, 2005c; Albach & Meudt 2010). Pedersen et al. (2007 and references), Jensen et al. (2008a) and Maggi et al. (2009) 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) and the genus is polyphyletic. Albach (2008) discusses the limits of Veronica s.l., which includes Hebe, and Wolfe et al. (2006) outline phylogenetic relationships in Penstemon. For the phylogeny of Collinsia and the related Tonella, see Baldwin et al. (2011).

• Philcoxia - a white sand endemic from Brasil (see also above). It has small, peltate leaves with circinate vernation borne at about ground level or underground and coming from underground stems; glandular hairs are conspicuous. The anthers are paired and monothecous (Fritsch et al. 2007; see also Taylor et al. 2000: basic morphology; McPherson 2010: photographs, etc.). The genus is on a long branch in the analysis of Schäferhoff et al. (2010). Pollination is probably by insects.

• Callitrichaceae - Water plant; hairs stellate; plant monoecious; flowers very small; staminate flowers; P 0; A 1(-3), pollen trinucleate, (exine, apertures 0 - see Cooper et al. 2000); carpellate flowers: G [2], transverse, subdivided, styles ± separate, slender; ovules 2/carpel, integument "thin"; fruit a schizocarp, 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. Pollination is by insects.

• Hippuridaceae - Water plant; flavones 0; hairs peltate, glandular; leaves whorled, linear; plant monoecious; flowers very small, monosymmetric by reduction, K a 2-4-lobed or entire rim, C 0; staminate flower: A 1; pollen trinucleate; carpellate flower: G 1, inferior, style single, stigmatic its entire length, stigma dry; ovules 2/carpel, pendulous, apotropous; chalazogamy occurs; fruit achenial or drupaceous; testa structureless, endosperm thin, starchy; n = 16. Pollination is by wind.

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

• Trapellaceae - Rhizomatous/stoloniferous aquatic herb; flower quite large, slightly monosymmetric, A 2 [the adaxial pair], connective apically peltate, staminodes 2; ovary inferior, the abaxial loculus much reduced, stigma unequally bilabiate; ovules 2/carpel, apical; fruit indehiscent, K persistent, with an appendage from below the apex of the sepals; testa with thin-walled cells; n = ca 25. Pollination is by insects.

Classification. The circumscription of Plantaginaceae adopted here is broad on the one hand (it incorporates several highly divergent but small clades) but narrow on the other (it is but a part of the old Scrophulariaceae). Molecular studies suggest that these small but florally very distinctive families are to be included in a clade with relatively large but undistinguished monosymmetric flowers. Maintaining these morphologically distinct families as separate would entail the recognition of a number of other families. Nevertheless, since aquatic taxa like Callitriche and wind-pollinated taxa like some species of Plantago have very distinctive morphologies partly associated with their particular pollination syndromes (and that caused them to be segregated as separate families in the first place), they are briefly characterized below. This allows us to get a better understandiong of the patterns and extent of variation in Plantaginaceae.

Sutton (1988) monographed Antirrhineae.

Previous Relationships. Both Cronquist (1891) and Takhtajan (1997) recognise several of the smaller families just mentioned, but they are in the same general part of their sequences; note that 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 to those of Pedaliaceae.

Thanks. I thank Dirk Albach for comments.

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

[Scrophulariaceae [Byblidaceae, Linderniaceae, [Stilbaceae [[Lamiaceae [Mazus etc. [Phrymaceae [Paulowniaceae + Orobanchaceae]]]] [Thomandersiaceae, Verbenaceae, Pedaliaceae, [Schlegeliaceae + Martyniaceae], Bignoniaceae, Acanthaceae, Lentibulariaceae]]]]]: ?

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), or spiral, lamina (punctate), vernation flat, (± foliaceous, stipuliform structures +); (inflorescence racemose); (flowers polysymmetric; 4-merous; bracts recaulescent); 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; stigma capitate (lingulate), dry or wet; ovules ³1 /carpel, apo/epi/pleurotropous, integument 5-11(-12) cells thick; 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]/1800: Verbascum (360), Eremophila (215), Scrophularia (200), Selago (190), Buddleja (125: inc. Nicodemia, Emorya, Gomphostigma), Jamesbrittenia (85), Manulea (75), Diascia (70), Nemesia (65), Zaluzianskya (55), Chaenostoma (46). 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. Plant-Animal Interactions. 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).

Floral Biology & Seed Dispersal. There are quite a few taxa that have oil-flowers with oil-secreting hairs (Vogel 1974; Vogel & Cocucci 1995 for a list; Renner & Schaefer 2010). Details of the pollination of the remarkable two-spurred oil-flowers of 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; Rasmussen & Olesen 2000; Steiner & Whitehead 1991); flowers of some Orchidaceae from the same area are rather similar. The South American Monttea and Angelonia also have weakly bisaccate oil-producing flowers; in the latter genus the visiting bees have either their front (Centris) or middle (Tapinotaspis) legs elongated. (Sérsic & Cocucci 1999; Machado et al. 2002). Scrophularia is a well-known wasp-pollinated genus (Kampny 1995 for general literature on pollination in the family).

Chemistry, Morphology, etc. Harpagide and harpagioside, occurring 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), and for that of Verbascum, see Geogiev et al. (2011). 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 fibres 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 Manuleeae 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.

Additional information is taken from Hartl (1959: seed coat/rumination), Jnsen (1999) and Harborne and Williams (1971), both chemistry, and Rogers (1986: as Loganiaceae, but including some genera belonging here, general), 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) and Endress (1999).

Phylogeny. The old Selaginaceae/Selagineae with a single apical ovule per loculus link with Scrophulariaceae-Manuleeae, although the latter have more ovules; Manuleeae 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, a deeply lobed calyx, nectary to one side of the ovary, lingulate stigma, etc. (Kornhall et al. 2001, which see for phylogeny and optimisation of characters). Also included here are Scrophulariaceae - Hemimerideae (see also 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, ex Loganiaceae, is very much paraphyletic and includes Nicodemia, Emorya, and Gomphostigma; several lines of evidence place it in Scrophulariaceae (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), while Kornhall et al. (2001) find most of Scrophulariaceae to be sister to Buddleja and immediate relatives. Other genera such as Androya that used to be in Loganiaceae also belong in Scrophulariaceae, Androya being sister to Myopyrum (Kornhall et al. 2001).

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 having 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 in the past, 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 (Gesneriaceae) in his Buddlejaceae.

Thanks. To F. Zapata, for useful comments on the family.

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

[Byblidaceae, Linderniaceae, [Stilbaceae [[Lamiaceae [Mazus etc. [Phrymaceae [Paulowniaceae + Orobanchaceae]]]] [Thomandersiaceae, Verbenaceae, Pedaliaceae, [Schlegeliaceae + Martyniaceae], Bignoniaceae, Acanthaceae, Lentibulariaceae]]]]: ?

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, lamina linear, abaxially curved or straight, veins parallel; flowers single, axillary; flowers subpolysymmetric, bracteoles 0; K connate only basally, C contorted, connate only basally, margins fimbriate; A ± monosymmetric, stamens = and opposite sepals, shortly epipetalous, anthers dehiscing by short slits or pores, epidermal cells ephemeral; nectary 0; stigma punctate to capitate (slightly bilobed); ovules 2-several/carpel, ± apical; exotestal cells tangentially somewhat elongated, anticlinal walls not uniformly 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 often 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 open by pores and whose apices form a cone, and a long style.

Evolution. Divergence & Distribution. A single seed, now destroyed, from the Middle Eocene of South Australia may be assignable to this family (Conran & Christophel 2004).

Ecology & Physiology. Although there is no evidence that the plant absorbs nutrients from the insects that often 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.

Floral Biology & Seed Dispersal. 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 vascularized 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, 2010: 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 as Byblidaceae and then included in Rosales, as by Cronquist (1981), or the two split, and both placed in Byblidales, a member of Aralianae, as by Takhtajan (1997).

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

Ephemerals to suffruticose perennials; iridoids 0; cork?; nodes 1:3; leaves opposite (basally connate), lamina venation also pamate, margins entire or serrate; inflorescence racemose or flowers from the axils of leaves; bracteoles 0; C with glandular hairs on the inside; 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, surface alveolate or furrowed (smooth); 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).

• Linderniaceae are often 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 very much reduced.

Evolution. Ecology & Physiology. Although many Linderniaceae seem to be rather delicate little herbs, a number are dessication tolerant (poikilohydric). These include the remarkable Chamaegigas intrepidus, which grows in transient pools - it is an aquatic resurrection plant - on inselbergs and probably uses glycine and serine as nitrogen sources (Heilmeier & Hartung 2011).

Floral Biology & Seed Dispersal. 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 in the couple of species that I have looked at. 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).

[Stilbaceae [[Lamiaceae [Mazus etc. [Phrymaceae [Paulowniaceae + Orobanchaceae]]]] [Thomandersiaceae, Verbenaceae, Pedaliaceae, [Schlegeliaceae + Martyniaceae], Bignoniaceae, Acanthaceae, Lentibulariaceae]]]: ?

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; lamina vernation revolute or not, (margins minutely toothed); 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, or unilocular, [1 G infertile, or septum 0], or bilocular, stigma slightly bifid or punctate; ovules 1-2/carpel, ascending and/or descending, apo/epitropous, or many; 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.]

Evolution. Floral Biology & Seed Dispersal. Oil flowers are quite common in the family (Renner & Schaefer 2010).

Chemistry, Morphology, etc. The C-8 iridoid glucosides common in Stilbaceae 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). By and large, the gynoecium is reminiscent of that of Scrophulariaceae - Manueleae. Thesmophora appears to have two collateral carpels, each with one descending ovule (Rourke 1993). This perhaps perhaps represents an abaxial carpel that is divided by a false septum.

For anatomy and morphology, see Carlquist (1986) and Dahgren et al. (1979), for some ovules, see Junell (1934), 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 Choisy

[[Lamiaceae [Paulowniaceae, Phrymaceae, Orobanchaceae]] [Thomandersiaceae, Verbenaceae, Pedaliaceae, [Schlegeliaceae + Martyniaceae], Bignoniaceae, Acanthaceae, Lentibulariaceae]]: ?

[Lamiaceae [Mazus etc. [Phrymaceae [Paulowniaceae + 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 unicellular; stellate); leaves simple or palmately compound, lamina vernation variable, margins toothed; A (2 [e.g. Salvia]), staminode 0 (+); tapetal cells multinucleate; pollen 3-colpate, 2-nucleate, exine not thickened near apertures; G [2(-5)], style (unequally) bifid, stigma inconspicuous, not expanded, dry (wet); ovules 2/carpel, ± erect, borne on inner side of carpel margin, apotropous, integument 5-9 cells across; fruit a schizocarp, berry or drupe, K persistent or accrescent; testa usu. thin, 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 of 3-7-flowered capitate cymes each with an involucre of bracts; flowers polysymmetrical; K 5-8, C 5-16; A 4-18; nectary 0; G imperfectly 2-locular; ovules apical, straight, funicle 0; embryo sac ± on surface of ovule; ?endosperm; n = 12, 14, 17, 18.

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

Synonymy: Symphoremataceae Wight

2. Viticoideae Briquet

Often woody; (hairs branched); (leaves compound); nectary 0 or poorly developed; fruit 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]; pollen exine with branched (simple, granular, etc.) columellae; nectray slight-0 (+); style often ± terminal; (antipodal cells numerous); nutlets reticulate; endosperm several-layered/0, cotyledons investing embryo [?common]; n = 7, 10, 13, 14, 16+.

24/1115: Clerodendrum (150), Teucrium (250), Aegiphila (120), Rotheca (50-60), Ajuga (40-50). Cosmopolitan, but many temperate, and esp. South East Asia to Australia.

Synonymy: Aegiphilaceae Rafinesque, Ajugaceae Döll, 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. 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 +; calyx with epidermal prismatic calcium oxalate crystals; pollen trinucleate, hexacolpate; style gynobasic; exocarp with mucilaginous cells producing hygroscopic spiral fibrils; endosperm development highly asymmetrical, the two haustoria lying close to each other, 1-layered [0?], cotyledons investing embryo; n = 6+; frequently attacked by Puccinia menthae.

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, Monardaceae Döll, Nepetaceae Berchtold & J. Presl, Salviaceae Berchtold & J. Presl, Saturejaceae Döll

[Scutellarioideae + Lamioideae]: stem endodermis +.

6. Scutellarioideae Caruel

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

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

Synonymy: Salazariaceae F. A. Barkley, Scutellariaceae Döll

7. Lamioideae Harley

(Aromatic), laballenic fatty acid [CH₃CH₂)₁₀CH=C=CH(CH₂)₃COOH] +; (calyx mesophyll with narrow prismatic calcium oxalate crystals); (stamens 4, about the same length - Pogostemon and relatives); style gynobasic; (ovule with glandular hairs - Leonurus, Teucrium); embryo sac with micropylar lobe longer and broader than chalazal lobe; nutlets hardly reticulate; endosperm several-layered, embryo spatulate; n = 6+.

63/1260: Stachys (300), Sideritis (140), Leucas (100), Phlomis (100), Pogostemon (80), Eremostachys (5-60). Esp. Europe and Africa to Asia, some cosmopolitan, but v. few Antipodean.

Synonymy: Melissaceae Berchtold & J. Presl, Stachydaceae Döll

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. Divergence & Distribution. The ca 60 species of mints endemic to Hawaii represent a major diversification there. Although they are currently placed in three separate genera, they are all polypoids and are derived from west North American Stachys (Lamioideae); they probably represent but a single introduction to the islands (Lindqvist & Albert 2002; cf. also the silversword alliance - Asteraceae).

Plant-Animal Interactions. 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 midges of the Tephretidae-Tephrellini are found here (and on Acanthaceae and Verbenaceae: Korneyev 2005), as are gall-forming wasps of the Cynipidae-Cynipinae (Redfern 2011) and agromyzid dipteran leaf miners (Winkler et al. 2009).

Floral Biology & Seed Dispersal. Species in the very big, New World-Mediterranean Salvia often have 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 Salvia, and Walker and Sytsma (2007) suggested that the distinctive stamen with its lever arm might have evolved three times via a bithecate condition with the thecae at opposite ends of the connective; the common ancestor of all unithecate clades had two "ordinary" stamens. Taxa like the 4-stamened Melissa and Lepechinia are at the base of the tree that includes Salvia, and there are other 2-stamened taxa immediately below the clades containing Salvia (Walker & Sytsma 2007). Reith et al. (2007) describe details of pollination in Salvia pratensis, and Wester and Claßen-Bockhoff (2007, 2011) focus on pollination by birds. There are over 300 bird-pollinated species of Salvia, and these are largely restricted to the New World; flowers with an ornithophilous syndrome in fact encompass a remarkable amount of variation (Wester & Claßen-Bockhoff 2011).

The calyx is conspicuous in fruit and is an integral part of the dispersal mechanism of the disseminule od most species, whether being brightly coloured and helping to attract frugivores, as in Clerodendrum, having hooked hairs or being itself hooked (Priva and some species of Salvia respectively), or forming a kind of catapult mechanism (Scutellaria) or a wing. Various kinds of calcium oxalate crystals are found in the sepals, perhaps protecting the nutlets against insect predators (Ryding 2010b). Myxocarpy, the nutlets producing mucilage and so adhering to their disperser, is common in Nepetoideae (Pammel 1892; Ryding 1992), while large genera like Lamium (Lamioideae) and Teucrium (Ajugoideae) have myrmecochorous nutlets (Lengyel et al. 2010).

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 occurrence of such nodes needs to be clarified.

The pollen grains of at least some Lamiaceae become very much flattened as they dry out (Halbritter & Hesse 2004). The ovules are described as being attached (just) to the false septae. 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). 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. Wunderlich (1967b) suggests that there is no endosperm in mature fruits of Nepetoideae.

For ovules, see Guignard (1893 - ovules vascularized), 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. (2009a), for nutlet micromorphology there, see Moon et al. (2009b) and especially Ryding (2010a and references), for nutlets in Stachys, see Salmaki et al. (2009), for calyx anatomy, see Ryding (2010b), and for a comprehensive general treatment, see Harley et al. (2004). Moon et al. (2008) surveyed pollen morphology especially of Salviinae. For some 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. Thus Bendiksby et al. (2011) recovered a set of relationships [Callicarpa [Prostantheroideae [[Symphorematoideae + Viticoideae] [[Cornutia, Gmelina, Tectona, Premna] [Nepetoideae [Garrettia [Scutellarioideae + Lamioideae]]]]]]], mostly with strong support, although sampling was mediocre and analyses of markers individually apparently yielded different topologies. These relationships differ from those in the classification above mainly in the position of Prostantheroideae and in the paraphyly of Viticoideae (cf. Bramley 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).

For relationships in Viticoideae, see Bramley et al. (2009); Vitex is paraphyletic. For the phylogeny of the Australian-centered Chloantheae (Prostantheroideae), see Conn et al. (2009). In Ajugoideae, Clerodendrum is being divided (Steane et al. 1999, 2004).

Within Nepetoideae, the large New World-centred Salvia, with over 900 species, is probably polyphyletic, Rosmarinus, Melissa and several other mostly quite small genera also being involved (Walker et al. 2004; Walker & Sytsma 2007; Moon et al. 2010) - alas for "Scarborough fair". Nepetoideae include the large tribe Ocimeae with synthecous, dorsifixed anthers (Paton et al. 2004). Another large genus is Hyptis, and the other genera of the Hyptinae are embedded in a paraphyletic Hyptis (Pastore et al. 2011); however, there is little resolution along the backbone of the tree, so major clade limits are unclear. Moon et al. (2010) circumscribe major clades within Mentheae, and Drew and Sytsma (2011) explore the limits and relationships of Lepechinia. Relationships within Menthineae inferred from chloroplast and nuclear data suggest major changes in our ideas of relationships there and in the limits of the subtribe, with Clinopodium in particular being polyphyletic (Bräuchler et al. 2010; Drew & Sytsma 2011).

For phylogenetic relationships in Lamioideae, see Wagstaff et al. (1995), Scheen et al. (2010), who found that Cymaria might be sister to the subfamily, and Bendiksby et al. (2011) who added Acrymia to Cymaria, although support for the sister group position of the combined clade was low. For relationships of the ca 60 species of lamioid 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 (see also Scheen et al. 2010; Bendiksby et al. 2011). Leucas is also highly paraphyletic (Scheen & Albert 2009; Scheen et al. 2010; Bendiksby et al. 2011).

Classification. As their distinctive alternative name Labiatae implies, Lamiaceae have always been considered as an "eminently natural" family, being immediately recognisable because of their herbaceous habit, 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); see Cantino and Sanders (1986) for the distinctions between the two biggest subfamilies, Lamioideae and Nepetoideae. 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. For tribal limits in Lamioideae, see Scheen et al. (2010) and Bendiksby et al. (2011). In general, generic limits will need some attention. Thus in both Stachys and Leucas characters associated with pollination prove unreliable indicators of clades (Scheen et al. 2010), and Clerodendrum has been dismembered (Steane & Mabberley 1998; Yuan et al. 2010). How Salvia is to be treated presents a challenge (Walker et al. 2006; Walker & Sytsma 2007). There are many other places where generic/clade rearrangements are to be expected, as in Chloantheae (Prostantheroideae: Conn et al. 2009). All the floral characters used to distinguish genera in Menthineae turn out to be homoplastic, and species of Clinopodium are scattered through much of the tree (Bräuchler et al. 2010).

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.

[Mazus etc. [Phrymaceae [Paulowniaceae + Orobanchaceae]]] : inflorescence racemose; (protein crystal stacks in nucleus).

Chemistry, Morphology, etc. Note that there may be chemical differences within Phrymaceae s.l., thus Mazus has iridoids while Mimulus does not (Hegnauer & Kooiman 1978). However, sampling is poor - and needs to be expanded. For nuclear protein crystals, see Albach et al. (2009).

Phylogeny. There is still some doubt as to basal relationsips in this clade, in particular, whether genera like Mazus are to be included in a broadly-circumscribed Phrymaceae, or not (Oxelman et al. 2005; Tank et al. 2006). Xia et al. (2009), Albach et al. (2009) and Schäferhoff et al. 2010) have all found 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 (see also Nie et al. 2006), so dismemberment is likely to be in order. The clades are kept separate here.

Mazus, etc.   Back to Lamiales

Annual or perennial herbs; iridoids +; cork ?; vessel elements ?; leaves opposite (spiral), margins toothed; corolla 2:3; anther thecae divergent, staminode 0; stigma sensitive; integument 5-6 cells across; (fruit indehiscent); n = 19.

2/32: Mazus (30). India and North China to the Antipodes, rather scattered, but esp. China (map: from Barker 1991 - India, Malesia inaccurate).

Chemistry, Morphology, etc. Mazus has 1:1 nodes and lacks a pericyclic sheath. For more information, see under Phrymaceae.

Classification. Species limits in Mazus need attention.

[Phrymaceae [Paulowniaceae + Orobanchaceae]]: ?

PHRYMACEAE Schauer, nom. cons.   Back to Lamiales

Annual or perennial herbs (woody); iridoids 0 (?+); cork ?; vessel elements ?; lamina margins toothed; K tubular, toothed, subplicate-ribbed (4-, 3-lobed), C 2 + 3 (polysymmetric; 2 + 0 - Mimulus douglasii); A (2) 4, anthers subreniform, thecae confluent; (pollen trinculeate; <10-colpate; each colpus with 2 orae; spiraperturate, etc.); nectary +/0; (fertile carpel 1), placentation various, axile (parietal), (near-basal), etc., stigma broadly 2-lobed (1-lobed; shortly 2-fid), sensitive or not; ovules (1<)many/carpel, straight, integument 3-7 cells across; (fruit indehiscent), K persistent; endosperm +/almost 0, cotyledons convolute; n = 7-10, 14, 22, etc.

Ca 19, but probably fewer[list]/234: Mimulus (150-170). ± 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 - India - Southeast Asia - Antipodes very inaccurate.). [Photos - Collection, Mimulus Flower.]

Evolution. Divergence & Distribution. 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, a mere ca 6-2 million years before present.

Other. The mostly 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. Whipple (1972) described the nodes of Phryma 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 Phrymaceae s. str., see Whipple (1972), Venkata 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).

• Phryma 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 (only one carpel is fertile) - a very derived morphology.

Classification. For a monograph of Mimulus sensu stricto, species of which are subjects of many evolutionary sudies, see Grant (1924) and Thompson (2005).

Previous Relationships. Phrymaceae have previously been placed in a separate monotypic family on account of the distinctive morphology of Phryma, or allied with Verbenaceae. Mimulus was included in Scrophulariaceae s.l.

[Paulowniaceae + Orobanchaceae]: ?

PAULOWNIACEAE Nakai   Back to Lamiales

Trees; cork cambium outer cortical; nodes 1:1; hairs uniseriate-branched; petiolar bundle annular; lamina margins entire; inflorescence branched; flowers large; K ± valvate, deeply lobed, space between K and C [water calyx], C hairs uniseriate with tapering terminal cell; anther thecae head-to-head, endothecium massive, extending across connective, placentoid +, staminode 0; nectary vascularized; placentae protruding, style hollow, head expanded or not, stigma punctate, hollow; seeds with several sinuous wings; exotesta cells broad, with complex reticulate thickenings; endosperm +; n = 19, 20.

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

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

Chemistry, Morphology, etc. Erbar and Gülden (2011) noted that the terminal flowers of an inflorescence might be 5-merous and have five stamens. The ad->abaxial direction of development of members of the calyx and the corolla whorls is unusual in Lamiales (Erbar & Gülden 2011), although observations are limited.

For some information, see Schilling et al (1982: verbascoside, etc.), Fischer (2004b: as Scrophulariaceae; he includes Wightia, etc., in the same immediate group - "Paulowniaceae" - but in a separate tribe), and especially Erbar and Gülden (2011: floral morphology and development).

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); on the other hand, Catalpa is definitely to be included in Bignoniaceae.

OROBANCHACEAE Ventenat, nom. cons.   Back to Lamiales

Many turn black on drying; (mannitol +), cork?; head of glandular hairs lacking vertical partitions; lamina margins often toothed to deeply lobed; inflorescence often racemose; C with abaxial lateral lobes outside others in bud [quincuncial]; A 4, didynamous; placentation often parietal; seed with exotestal cells variously thickened on the inner walls.

99[list]/2060. World wide, but especially N. (warm) temperate and Africa-Madagascar.

Rehmannia + Trianeophora

Leaves spiral; bracts ± foliaceous; (staminode +); stigma sensitive.

2/7. China (map: from 2011, in part).

Lindenbergia

Bracts ± leaf-like, bracteoles usu. 0; A thecae on connective arms; testa usu. with hook-shaped thickenings adnate to surface; n = 16.

1/12. N.E. Africa to N. Philipines (map: see Hjertson 1995).

The Rest

Most (hemi)parasitic herbs (shrubs); stomata do not close; orobanchin +, little oxalate accumulation, 6- and/or 8-hydroxylated flavone glycosides 0; leaves spiral to opposite; (K ± free), C (tube development intermediate), (aestivation imbricate); A (free from C - Eremitilla), 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), (tapetum amoeboid); pollen often starchy, commonly colpate with a retipilate surface, (polyporate); ([G 5]), (placentation axile), (placentae [2, bilobed] 4 [-6]), stigma clavate to capitate; ovule (-1/carpel), integument (2-)4-12 cells across; (antipodal cells persistent); capsule loculicidal to septicidal; (seeds with elaiosomes); (cells of seed wings with reticulate thickenings on radial walls), (cells of layers other than the exotesta thickened and lignified); endosperm +, (walls thickened; reserves starch; mannose-rich polysaccharides; 0), (embryo minute, undifferentiated; germination via germination tube); n = 7+.

96/2040: Pedicularis (600-?800), 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 a similar corolla aestivation, but they are autotrophic.

Evolution. Divergence & Distribution. Orobanchaceae may be some 50-40 million years old (Wolfe et al. 2005), and it has been suggested that the evolution of of the holoparasites with minute dust seeds - which may have occured twice - was driven by the expansion of grasslands in the middle of the Tertiary (Eriksson & Kainulainen 2011).

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). Diversification within the large genus Castilleja is becoming better understood. There is a speciose West North American/Central/South American perennial clade - some 120 species - derived apparently quite recently from an annual ancestor; polyploidy is common in the perennials, but not in the annuals (Tank & Olmstead 2008, 2009). Annuals have dispersed more than once to South America (Tank & Olmstead 2009).

Plant-Animal Interactions. There has been diversification of agromyzid dipteran leaf miners on the hemiparasitic members of this clade (Winkler et al. 2009). Larvae of Nymphalidae-Melitaeini butterflies are commonly found here (also on Plantaginaceae, but not on Scrophulariaceae: Wahlberg 2001).

Ecology & Physiology. Holoparasites have evolved from hemiparasites more than once in Orobanchaceae (dePamphilis et al. 1997; Nickrent et al. 1998; Young et al. 1999; Schneeweiss et al. 2004a; Bennett & Mathews 2006, etc.). Interestingly, 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). For the chloroplast genome of the holoparasites - small, but a few genes are still functional - see Wolfe et al. (1992). For the role of strigolactones, exuded from host roots, in stimulating germination of seeds of some species of Orobanchaceae, see Tsuchiya and McCourt (2009) and Akiyama et al. (2010 and references). 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 hemiparasitic plants in general because they increase the transpiration flow in the parasite so faciltating movement of water, etc., from the host to the parasite; the stomata remain open despite the presence of large amounts of abscisic acid, which normally would be expected to result in stomatal closure (Jiang et al. 2010; other papers in Folia Geobotanica 45(4). 2010).

Some Orobanchaceae, particularly the hemiparasitic taxa with chlorophyll, 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). Recently, horizontal nuclear gene transfer between Sorghum or its relatives and Strica sopecies (but not Orobanche) has been demonstrated (Yoshida et al. 2010); the complexity of the relationships between host and parasite is likely to have been under-estimated. For other information on parasitism in Orobanchaceae, see Irving and Cameron (2009 and references).

Floral Biology & Seed Dispersal. Variation in floral morphology in Pedicularis is very great, some species having a corolla tube ca 10 cm long or more, or flowers with an asymmetrical, proboscis-like extension of the upper lip (the galea) that is formed from the two adaxial corolla members. For comments on the floral evolution of the genus, see Ree (2005a); species numbers are very uncertain (Mill 2001). Red-coloured flowers with long tubes of species growing at higher elevations may lack nectar and be pollinated by pollen-collecting bumblebees, which raises the question of the function of these very long tubes... (Huang & Fenster 2007). Pollen morphology - there is quite extensive variation - is linked with corolla morphology and pollinator type (Hong Wang et al. 2009a). See Kampny (1995: as Scrophulariaceae) for earlier literature on pollination in the family.

For myrmecochory of some orobanchaceous seeds (Melampyrum, Pedicularis), see Lengyel et al. (2009, 2010).

Genes & Genomes. 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 primarily on legume and grain crops in warmer and drier areas and especially in sub-Saharan Africa, where they are still spreading. Striga parasitizes monocots, affecting ca 40% of the cereal producing areas there, 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 Ejeta & Gressel 2007; Yoshida & Shirasu 2009; Irving & Cameron 2009). Alectra vogelii can cause the complete loss of legume crops it infects (Morawetz & Wolfe 2009). Orobanche parasitizes eudicot crops in more or less temperate parts of the world.

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 ovules of Cyclocheilon, etc., see Junell (1934), for embryology, see Tiagi (1963) and Arekal (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 floral development, see Canne-Hilliker (1987), 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 was 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). Lindenbergia is autotrophic (Hjertsen 1995) and 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).

More recently it has been found that Rehmannia, a small genus of ca 6 species from China and Korea, and the related Trianeophora, which includes two to three species from China, are together sister to the rest of the family. In a tree found by Oxelman (2005), Rehmannia linked very weakly with Phryma, Paulownia, Mazus and Lancea, as well as with genera of Orobanchaceae. In a more 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 occurring 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 corolla aestivation in which the two abaxial-lateral lobes are outside the others, as is common in Orobanchaceae; its stigma lobes are sensitive. 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 and forming 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.).

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 were then 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, esp. Morawetz et al. 2010); Nesogenes was sister to Graderia and in a clade that includes Strica (Morawetz et al. 2010).

Inclusion of these erstwhile Cyclocheilaceae and Nesogenaceae considerably increase the morphological 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 apotropous ovules/carpel, endothelium?, the funicles are long and the stigma is lingulate. The fruit is a capsule or schizocarp; there is no endosperm (but present in Nesogenes? - Junell 1934). 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).

For a phylogeny of Pedicularis, see Ree (2005), for that of Euphrasia, see Gussarova et al. (2008), of Orobanche, see Park et al. (2008), of Castilleja, see Tank and Olmstead (2008, 2009), and of rhinanthoid Orobanchaceae, see Tesitel et al. (2010 - also other papers in Folia Geobotanica 45(4). 2010). 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). On the other hand, Rehmannia has often been linked with Titanotrichum and included in Gesneriaceae (Xia et al. 2009 for references).

Thanks. To David Tank for useful comments.

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

[[Thomandersiaceae + Verbenaceae], Pedaliaceae, [Schlegeliaceae + Martyniaceae], Bignoniaceae, Acanthaceae, Lentibulariaceae]: ?

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

THOMANDERSIACEAE Sreemadhavan   Back to Lamiales

Shrub or small tree; 2-indolinone alkaloids +; phloem stratified; pericyclic fibres ?short, massively thickened; nodes 1:3; petiole bundles forming a ring or incurved C-shaped; stomata anisocytic; lamina (margins deeply lobed), with flat ?nectary glands abaxially, petiole swollen apically and basally; K with nectaries on the outside; staminode +; pollen 5-6-colpate; nectary vascularized by carpellary traces; gynoecial vasculature 8-shaped; ovules 1-3/carpel, 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. 2007a).

• 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 2007a). 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 that family, however, support for any Thomandersiaceae-Schlegeliaceae association is currently weak (Wortley et al. 2007a).

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 +; (pits vestured); petiole bundles arcuate (also medullary, associated with median bundle); needle crystals common; stomata dia(anomo)cytic; stems often square; eglandular hairs unicellular; lamina margins toothed to deeply lobed; inflorescence racemose; flower weakly bilabiate; space between K and C [water calyx]; (A of two lengths, but free), tapetal cells 2-4-nucleate; pollen (colpate, por[or]ate), exine thickened near apertures; G collateral, placenta on the margin of the carpel, otherwise position various, stigma bilobed, with conspicuous stigmatoid tissue, wet; ovules 2/carpel, apotropous, (1/carpel - Lantana), integument 5-9 cells across, obturator +; (antipodal cells multinuclear); fruit a schizocarp or drupe with 1, 2 or 4 stones, K persistent; testa thin-walled; endosperm ± 0, cotyledons spatulate; n = 5-12+.

31[list]/918. Pantropical (to warm temperate), but mostly New World, esp. S. South America (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?).

1. Petreeae Briquet

Shrubs and vines.

?1/12. Mexico to the Amazona Basin.

Synonymy: Petreaceae J. Agardh

Duranteae [[Casselieae + Citherexyleae] [Priveae [Neospartoneae [Verbeneae + Lantaneae]]]]: ?

2. Duranteae Bentham

Trees to herbs; eglandular hairs multicellular; (A 2 + 2 staminodes); ([G 4]).

6/192: Stachytarpheta (130). S. U.S.A. to Argentina, (Africa to India).

Synonymy: Durantaceae J. Agardh

[Casselieae + Citherexyleae] [Priveae [Neospartoneae [Verbeneae + Lantaneae]]]: ?

Casselieae + Citherexyleae: ?

3. Casselieae Troncoso

G 1 [adaxial carpel].

3/14. Mexico and the Caribbean to Argentina.

4. Citharexyleae Briquet

?

3/135: Citharexylum (130). S. U.S.A. to Argentina. [Photo - Flower.]

Priveae [Neospartoneae [Verbeneae + Lantaneae]]: ?

5. Priveae Briquet

?

?1/21. Pantropical-warm Temperate.

Neospartoneae [Verbeneae + Lantaneae]: ?

6. Neospartoneae Olmstead & O'Leary

(Ephedroid shrubs); glabrous; (staminode +); G 1.

3/6. Argentina, Chile, S. to Patagonia.

Verbeneae + Lantaneae: staminode 0.

7. Verbeneae Dumortier

(A 5 - Verbena).

3/160: Verbena (300), Junellia (48). Mostly American, Eurasia to Africa.

8. Lantaneae Endlicher

Ethereal oils +; stomata anisocytic; inflorescence capitate; G 1 [Coelocarpum, with [G 2], could be sister to rest of clade]; (endosperm + - Lantana).

9/275: Lippia (120), Lantana (100). Mostly New World

Synonymy: Lantanaceae Martynov

• 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. Plant-Animal Interactions. 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 margins of the carpel (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. Marx et al. (2010) present a comprehensive phlogeny of the family, although, as they note, sampling within the big genera needs to be improved. A couple of genera remain unplaced. Dipyrena may be close to Verbeneae while Rhaphithamnus may be close to Priveae, although branches within the latter are rather long; the position within Lantaneae of Coelocarpum, morphologically plesiomorphic, is uncertain (Marx et al. 2010).

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. For a tribal classification, see Marx et al. (2010). The whole Lantana-Lippia complex, speciose although it may be, could perhaps be reduced to a single genus, the larger genera currently recognised being para- or polyphyletic (Lu-Irving et al. 2009), however Marx et al. (2010) disentangle relationships and suggest that nine genera can be recognised in the complex. I provisionally include Glandularia, with about 100 species, within Verbena. The limits of genera around Junellia have been redrawn (O'Leary et al. 2009); for a revision of Junellia in the old sense, see Peralta et al. (2008).

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

Annual to perennial (± succulent stemmed, or swollen roots) herbs to deciduous trees; orobanchin, amyloid +; cambium storied; pericycle also with sclereids (fibers few); petiole bundle interrupted-annular; hairs broadly capitate-stellate, mucilaginous; leaf (spiral - Sesamum), lamina (venation palmate), margins toothed, lobed or entire; flowers usu. axillary (inflorescence dichasial); paired nectaries (modified flowers) at base of pedicel (not - Uncarinia); (C with spur); A (5), thecae ± confluent, at right angles to filaments, staminode + (0); pollen 5-13 stephanoocolpate; G [2-4], ± divided, (8 loculi - Josephinia), stigma with 2 broad lobes, wet, often sensitive; ovules 2-many/carpel, (1 ovule/loculus - Josephinia), integument 7-20 cells across, hypostase +; 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, surface often sculpted, testa multiplicative, exotestal cells palisade or otherwise thickened, (mesotesta with crystals); fat and amyloid [xyloglucans] in cotyledons, endosperm thin; n = 8 (13); protein bodies in nucleus?

14[list]/70: Sesamum (19), Pterodiscus (13). Mostly tropical, in coastal or arid habitats, Old World (map: from Ihlenfeldt & Grabow-Seidensticker 1979; FloraBase 2005; Ihlenfeldt 1994b, 2010).

• 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.]

Evolution. Floral Biology & Seed Dispersal. The diversity of fruit morphology and dispersal "strategies" in this small family is remarkable, as is their variation in growth form (Ihlenfeldt 2010).

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), Ihlenfeldt (1967, 2004, 2010: general), and Jordaan (2011: seed coat of Harpagophytum - complex).

Synonymy: Sesamaceae Berchtold & J. Presl

[Schlgeliaceae + Martyniaceae]: ?

SCHLEGELIACEAE Reveal   Back to Lamiales

Large trees or woody shrubs, vines or epiphytes (many); pericyclic sheath sclereidal; nodes 1:3; petiole bundle solid-(almost)annular, with wing bundles, no pericyclic lignification; sclereids +; stomata variable; laimna margins entire or serrate; flowers quite large; nectaries on outside of K; staminode +/0; nectary vascularized from carpellary bundles/0; 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, most being epiphytes, 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.

MARTYNIACEAE Horaninow, 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, lamina margins toothed; inflorescence racemose; (K free); A (2), connective with apical gland, staminode(s) +; pollen trinucleate, inaperturate, exine dissected into 20-40 platelets; G with parietal placentation, placentae bilobed, stigma bilobed; 2-many ovules/carpel; 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 (for map, see McPherson 2010, vol. 2).

• 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. Ecology & Physiology. Insects may stick to the very viscid indumentum of Martyniaceae, although there is no evidence that the plants are carnivorous (see Rice 2008; 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) and McPherson (2010, vol. 2, esp. photographs) provide 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 (inaperturate and with platelets vs several colpi) and placentation (parietal vs axile) best separate the two.

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, lamina vernation 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), stigma lobes broad, often sensitive, wet; nucellar endothelium +; 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 polysymmetric; A 5); fruits septicidal.

4/6. Indo-Malesian.

[Crescentieae + Coleae]: ?

7. Crescentieae G. Don

Leaves palmate, (unifoliolate; 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 but otherwise there are terminal leaflets. 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. Divergence & Distribution. Based on this fossil [?which], the divergence time of Bignoniaceae and Verbenaceae (sister taxa in the phylogeny used) 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).

Floral Biology & Seed Dispersal. 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 can be associated with the behaviour and type of visitor (Gentry 1974a, b, 1990; Alcantara & Lohmann 2010a, b). Alcantara and Lohmann (2010a, b) found that, in general, ancestral flower size in the lianescent Bignonieae is larger than that of today's species. One of the commonest flower types in the New World is the Anemopaegma type, visited by euglossine bees (along with anthophorids); this may be ancestral in the Bignonieae, and has an infundibular, straight corolla tube, nectar, and is magenta, yellow or white (Alcantara & Lohmann 2010a). The nectarless Cydista type, otherwise rather similar florally, is also visited by euglossines, a group of bees than began diversifying only some 42-27 million years ago (Ramírez et al. 2010).

Dispersal syndromes are also quite diverse (Gentry 1983; 1990) but they are not particularly correlated with pollination syndromes. Wind dispersal is common in the family, and the seeds often 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.

Vegetative Variation. Bignoniaceae are, along with Sapindaceae, the most ecologically important neotropical group of lianes, and Bignoniaceae are the second most speciose family in drier tropical forest types in America (Gentry 1988, 1991). 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). Pace et al. (2011) note that the variant phloem that causes the fluting of the vascular cylinder has large-diameter sieve tubes and numerous fibres, hence contributing substantially to translocation and to stem support; regular phloem has sieve tubes with much smaller diameters.

Palmate leaves may have arisen more than once within Bignoniaceae, but they 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.

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.

ACANTHACEAE Jussieu, nom. cons.   Back to Lamiales

Quaternary methylammonium compounds, amyloid +; (cork cambium deep seated); (intraxylary phloem +); stomata diacytic; lamina margins entire to toothed; inflorescence with main axis indeterminate, 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; embryo sac long, curved, (apex of 4-nuclear sac growing out of the micropyle and eventually into the placenta); (zygote pushed back into the ovule by a long suspensor); capsule dehiscence explosive, walls cartilaginous, K persistent; endosperm development highly asymmetrical, the two haustoria lying 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, lamina margins?; inflorescence a raceme, (branches cymose - Saintpauliopsis); bracts spiral, (bracteoles 0 - Nelsonia); stamens 2 or 4, anthers variable (e.g. thecae ± separate); ovary (with pariteal placentation - Elytraria); stigma broadly (unequally) lobed, (sensitive - Elytraria); ovules many(-4)/carpel, campylotropous, endothelium +; antipodal cells persistent; funicular obturator +; jaculators rudimentary[?]; seeds 2-many, ruminate, testa ± disorganised (± visible - Nelsonia); endosperm +, oily; n = 9.

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

Synonymy: Nelsoniaceae Sreemadhavan

Acanthoideae [Thunbergioideae + Avicennioideae]: (inverted vascular bundles in the pith); acicular fibres +; (adaxial C lobes outside others - [descending cochleate]); pollen other than tricolpate or -colporate common; ovules 2/carpel; collateral, endothelium 0; (amyloid [xyloglucans] 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, retinacula]; 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; (ovules 3+/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; (ovules 3+/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: C left-controted; filament bases thickened; pollen sac placentoids +; ovules collateral, embryo sac ± on surface of nucellus; cotyledons folded.

3. Thunbergioideae Kosteletzky

Twining vines (erect); (C-8 iridoid glucosides [unedoside] +); petiole bundles arcuate or annular; lamina vernation 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 (not contorted); anthers with lignified unicellular hairs (multicellular awns), sagittate, (thecae slightly displaced), dehiscing by (elongated) pores/slits, connective elongated, endothecium 0; pollen 8-colpate or spiraperturate; (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); endosperm chalazal haustorium 0, (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 occurring in bands of conjunctive tissue; vessels in radial multiples; nodes 3:3; petiole bundle annular; sclereids +; lamin thick, 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; pollen 3-colporate; nectary small [as tufts of hairs?]; (G with false septae), loculi apically confluent, stigma with 2 blunt lobes; ovules apical on partitions, ± straight, naked, micropyle 0; 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 Miquel, 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. Plant-Animal Interactions. Gall-forming fruit flies of the Tephretidae-Tephrellini are found here (and on Verbenaceae and Lamiaceae: Korneyev 2005). Larvae of Nymphalidae-Melitaeini butterflies commonly feed on Acanthaceae (Wahlberg 2001; Nylin & Wahlberg 2008). Mass defoliation of Avicennia by lepidopteran larvae seems to be not uncommon (Fernandes et al. 2009).

Floral Biology & Seed Dispersal. 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). All told, 500-600 species of Acanthaceae (E. A. Tripp and L. McDade, pers. comm.: also Tripp & Manos 2006) are humming-bird pollinated. Acanthaceae (minus Nelsonioideae) are extremely heterogeneous palynologically, although the functional significance of this variation is unclear.

Full (180^o) or partial resupination has evolved several times independently in Acanthoideae, and this is sometimes 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.

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. "Rudimentary" retinacula are reported from Nelsonioideae (Johri & Singh 1959; Roham Ram & Masand 1963). In a number of taxa the testa is mucilaginous.

In a number of species of Strobilanthes all the individuals of a species flower and fruit in synchrony and then die; this happens in a regular cycle every few years and can affect very large areas (Jansen 1976). Both pollinators and seed dispersers (the seed are rich in oils) are attracted to the plants in large numbers.

Ecology & Physiology. 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). These 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. Robert et al. (2009, 2011) discuss the hydraulic architecture of the wood of Avicennia in which both xylem and phloem are organized in a three-dimensional network.

Chemistry, Morphology, etc. 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), Scotland and Vollesen (2000) and references, and Daniel (2010). Many Isoglossinae (Justicieae) have distinctive "Gürtelpollen" (Kiel et al. 2006) - lenticular biporate pollen with a prominent circumferential band. The Acanthoideae [Thunbergioideae + Avicennioideae] clade 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. For cotyledon folding, see Schwarzbach and McDade (2002).

The distinctive asymmetrical endosperm development found in Acanthaceae is also to be found in Lamiaceae-Nepetoideae (a parallelism). In Acanthoideae other than Acantheae, 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 (Mohan Ram & Wadhi 1964; Johri et al. 1992 and references). In Nelsonioideae this central area is entirely cellular, but other details of endosperm and embryo sac development are similar to those in the rest of the family (Johri & Singh 1959; Moham Ram & Masand 1963). Indeed, embryo sac development in some species is almost unique in flowering plants: the apex of the embryo sac grows through the micropyle and eventually may be lodged in the placenta, and this where the egg apparatus is formed (the movements of the polar nuclei are unclear). As the embryo develops, a very long suspensor is produced and the embryo is pushed back into the endosperm - and so pushed back into the ovule and the developing seed (e.g. Roham Ram & Masand 1963 and references). The ovule of Avicennia is sometimes reported to be straight, but the embryo sac is extra-ovular, the endosperm is asymmetrical, and the micropylar endosperm haustorium at least is extra-ovular, being incredibly branched and reaching the placenta (Junell 1934; Padmanabhan 1964, 1970).

Note that 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 (see above), but even if present, they certainly are not functional.

For general anatomy, capsule dehiscence, see van Tieghem (1908), for embryology, etc., see Mauritzon (1934a), 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) and also Borg and Schönenberger (2011 - also characters uniting Avicennia and Thunbergioideae) and for wood anatomy, see Carlquist (1990b), 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, while Borg and Schönenberger (2011) mention possible floral/cevelopmental apomorphies of Thunbergioideae and Avicennioideae.

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 it is woody, has more or less cymose inflorescence, and a 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.

LENTIBULARIACEAE Richard, nom. cons.   Back to Lamiales

Insectivorous rosette-forming herbs, also other derived growth forms; (verbascoside 0 - Utricularia); 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, lamina margins entire, but 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], (± free - Pinguicula), filaments stout [always?], thecae (superposed), confluent, epidermal cells ephemeral, staminode 0; pollen 4-10-zonocolporate [Pinguicula], trinucleate; nectary 0; G free-central or basal, style hollow, often 0, stigma with broad lobes, wet, (sensitive); ovule with integument 2-6 cells across; (embryo sac protrudes into the micropyle), (antipodal cells persisting); fruit a capsule of various types; exotestal cells variously thickened; endosperm 0, starchy, embryo green, with cotyledons [1 or 2, Pinguicula], or 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. Ecology & Physiology. As befits their carnivory, Lentibulariaceae are notably prominent in acid habitats including those of the ephemeral flora of African inselbergs (Seine et al. 1996). 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, while Peroutka et al. (2008b) discuss aspects of their functional biology. Reifenrath et al. (2006 and references) describe details of trap architecture of Utricularia (see also Merl 1915); many species have sensitive hairs the stimulation of which leads to the rapid opening of the trap, the only suction trap known in land plants. 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 traps. Vincent et al. (2011) distinguish between a slow, energy-dependent phase in which water is pumped out of the trap, the trap becoming deformed and elastic enegy being stored in the trap body, and a fast but passive phase in which the trap door opens and closes in less than a millisecond, water and contained prey rushing inside. Spontaneous opening of traps without stimulation by prey is common (Adamec 2011 and references).

There is a diverse microbial community in the traps - perhaps a mutualistic association - that may aid in the uptake of phosphorous by the plant (Sirová et al. 2009). Recently-fixed carbon may end up in the young traps (Sirová et al. 2010), perhaps for the microbes there. Indeed, 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. 2008a) and nitrogen from ¹⁵N-labelled phytoplankton may be taken up into the plant (Alkhalaf et al. 2009). However, the relationship between plant and algae - and potential animal prey, too - is for the most part unclear (Jobson & Morris 2001; Alkhalaf et al. 2011), and the plant may even nutritionally support the microbial community in prey-free traps (Adamec 2011: see above).

Some species of Utricularia have non-suction traps rather more like those of Genlisea. In that genus animals are passively trapped as they swim up the spiraling branches of an eel-type trap with backwardly-pointing hairs. 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.

Genes & Genomes. 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).

It can be particularly difficult to understand the morphology of some species of Utricularia in particular using conventional morphological terms. The embryo is undifferentiated, and although some exceptions are mentioned by Plachno and Swiatek (2010), even there cotyledons and radicle are not apparent. Small fragments of the "leaves" or the cut peduncle will regenerate the whole plant (Merl 1915), ordinary roots do not occur in Genlisea and Utricularia, and the suction bladder-traps have no parallel in other flowering plants (see e.g. Sattler & Rutishauser 1990 and references; Plachno & Swiatek 2010 and references for development). There are quite commonly leaf-like structures in Utricularia, and U. kuhlmannii is even described as having odd pinnate leaves by Merl (1915).

Chemistry, Morphology, etc. "Nutritive tissue" is described as being derived from the ovule, funicle and placenta, i.e., at both ends of the developing embryo, but it is not know from 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). in Pinguicula a single antipodal cell may persist, enlarge, and divide (Kopczynska 1964). 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), Farooq (1965, 1966) and Farooq and Bilquis (1966 and references) and G. Degtjareva (2004), for some chemistry, see Damtoft et al. (1994), for pollen, see Rodondi et al. (2010: Pinguicula), and for general information, see Goebel (1891: Utricularia), Fischer et al. (2004b), McPherson (2008: Pinguicula, 2010), 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). Cieslak et al. (2005) and Degtjareva et al. (2006) discuss the phylogeny and evolution of Pinguicula and Reut and Jobson (2010) that of Utricularia subgenus Polypompholyx.

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

Synonymy: Pinguiculaceae Dumortier, Utriculariaceae Hoffmannsegg & Link