EXTANT SEED PLANTS

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

MAGNOLIOPHYTA

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

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

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

ERICALES + ASTERID I + II: ovules tenuinucellate.

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

ASTERID II: Myricetin 0; vessel elements with scalariform perforation plates; flowers rather small, style short; endosperm copious, embryo short/very short.

ASTERALES [ESCALLONIALES [BRUNIALES [APIALES [PARACRYPHIALES + DIPSACALES]]]]: iridoids +; C tube initiation early; G [2-3], inferior.

ESCALLONIALES [BRUNIALES [APIALES [PARACRYPHIALES + DIPSACALES]]]: ?

[BRUNIALES [APIALES [PARACRYPHIALES + DIPSACALES]]: ?

APIALES [PARACRYPHIALES + DIPSACALES]: ?

PARACRYPHIALES + DIPSACALES]]: nodes 3:3; true tracheids +; leaves serrate; inflorescence terminal.

DIPSACALES Dumortier  Main Tree, Synapomorphies.

Route I secoiridoids +; cork cambium deep-seated; pericyclic fibers 0; petiole bundles arcuate; buds with scales; leaves opposite, bases ± confluent, margins gland-toothed; inflorescence cymose; pollen grains trinucleate; G ± inferior; K persistent in fruit; testa vascularised, exotestal cells enlarged (palisade), variously thickened and lignified. - 2 or 7 families, 45 genera, 1090 species.

Evolution. The first split within crown Dipsacales may have occured in the mid-Cretaceous, some 111-93 million years before present, 20-30 million years older than some estimates (Bell & Donoghue 2005a and references). However, major diversification within Dipsacaceae and Valerianaceae - and hence a major part of the diversification of the whole order - is relatively recent, occuring within perhaps ca 10 million years (Bell & Donoghue 2005a). For a very detailed discussion of morphological evolution, see Donoghue et al. (2003). Howarth and Donoghue (2004, esp. 2005, 2008, 2009) noted intriguing correlations between gene duplications with possible neofunctionalisation of e.g. CYC-like genes and changes in floral form in the order, although direct connections between changes both in morphology and in diversification rates remain to be established. Indeed, diversification has often seemed to increase as plants moved into new geographical areas, especially if these were mountainous (Moore & Donoghue 2007).

Rates of molecular evolution vary considerably in Dipsacales, and this is usually associated with chages in the life style, woody plants usually having lower rates of change than herbaceous (Smith & Donoghue 2008).

Chemistry, Morphology, etc. Anthocyanin variation does not allow a sharp distinction between Caprifoliaceae and Adoxaceae (Jordheim et al. (2006). Involute ptyxis is more common in Adoxaceae than in Caprifoliaceae (Cullen 1978). Weberling (1957) discussed the morphology of the stipule-like structures and amplexicaul leaves in Caprifoliaceae and Adoxaceae; amplexicaul leaves in particular occur in several families in the order. Weberling (1989) also described the inflorescence structure, emphasizing the woody Dipsacales, in considerable detail. The morphology of the basic nectary type for the order remains unclear. In Viburnum (and Sambucus?) it is a disc-like structure on top of the inferior ovary, in Adoxa it is associated with multicellular hairs on the corolla, in Caprifoliaceae, unicellular hairs (Takhtajan 1997). For variation in floral anatomy, which clearly separates two groups of genera in the old Caprifoliaceae, see Wilkinson (1949 and references). There is considerable variation in the presence of the mitochondrial coxII.i3 intron.

Phylogeny. The position of Dipsacales in early phylogenetic analyses was unclear. Downie and Palmer (1992) associate Adoxaceae with Asterales, while they are sister to Apiales in some studies (Backlund & Bremer 1997).

The order has strong support in D. Soltis et al. (2000: see also Wagenitz 1997; Bremer et al. 2001; cf. A.P.G. 1998). The phylogeny of the expanded order is based especially on work by Donoghue et al. (2001b, 2003), Zhang et al. (2003), Davis et al. (2001), Bell (2003) and Moore and Donoghue (2007). There are some (not major) disagreements with the phylogeny in Pyck and Smets (2001), while Judd et al. (1994) provide an early morphological analysis which gives a similar topology. Note that the position of Heptacodium was initially somewhat uncertain (Pyck & Smets 2000, 2001), although it is probably to be included in Caprifoliaceae s. str (e.g. Donoghue et al. 2001a, 2003). Valerianaceae are sister to the rest of the Caprifoliaceae s. str. part of the clade in some analyses (Soltis et al. 2007a), while analyses of mitochondrial data are perhaps particularly challenging (Winkworth et al. 2008b).

Previous Relationships. Takhtajan (1997) included Dipsacales, Adoxales and Viburnales in his Dipsacanae. In earlier versions of this site (pre April 2008), [Columelliaceae + Desfontainiaceae] (= Columelliaceae s.l.) were placed sister to Dipsacales, with which they do show considerable morphological similarity. However, recent work suggests they are to be associated with Bruniaceae and are placed in Bruniales, which are in turn sister to [Apiales [Paracryphiales + Dipsacales]] (Winkworth et al. 2008a; Tank & Donoghue 2009).

Includes Adoxaceae, Caprifoliaceae, Diervillaceae, Dipsacaceae, Linnaeaceae, Morinaceae, Valerianaceae.

Synonymy: Adoxales Nakai, Caprifoliales Lindley, Lonicerales T. Leibe, Sambucales Berchtold & J. Presl, Valerianales Berchtold & J. Presl, Viburnales Dumortier - Dipsacanae Takhtajan - Caprifoliopsida Endlicher, Loniceropsida Brongniart

ADOXACEAE Trautvetter, nom. cons.   Back to Dipsacales

0-methylated flavonols only +; true tracheids?; pericyclic sheath poorly developed; (nodes 1:1, 5:5); (stomata paracytic); (petiole also with adaxial inverted bundles; closed C-shaped and with wing bundles); glandular hairs common; (bud scales 0); leaves involute or conduplicate (plicate), 2ndary veins palmate to pinnate, (stipules on leaf base); (flowers 4-merous); K open, usu. very small, with one trace, C rotate; G not entirely inferior, lateral and dorsal carpellary bundles free, ventral bundles move from axis to ovary septae, 1 median crassinucellate ovule/carpel, endothelium +, style short, stigma lobed, dry or wet; fruit drupaceous; seed often ruminate, exotestal cells palisade or not; (embryo large); n = (8-)9.

5[list]/200. North temperate, tropical montane, but not in Africa (map: from Hultén 1958, 1971; Meusel et al. 1992). 2 groups below. [Photos - Collection]

1. Viburnum

Shrubs (trees); hairs stellate and variants; G [3], odd member abaxial, 2 abort, nectary on top of G.

1/175. North temperate, tropical montane, but not in Africa. [Photo - Flower]

For a phylogeny of Viburnum, see Winkworth and Donoghue (2005), for variation in its endocarp and seed, see Jacobs et al. (2008).

Synonymy: Tinaceae Martynov, Viburnaceae Rafinesque

2. Adoxa, etc.

Shrubs or herbs; (cambium storied - Sambucus); vessel elements with simple perforation plates; leaves compound (deeply lobed); anthers extrorse (8, monothecal); G 1, [5], opposite petals, embryo sac tetrasporic, but eight nucleate [Adoxa-type], nectary on top of G or on C associated with multicellular hairs, styles +, separate, stigmas capitate.

4/23. Mainly north temperate, esp. China, also (sub)tropical montane.

Sambucus has crystal sand. Adoxa is distinctive in lacking any obvious corolline ring primordium and in its eight stamens with monothecate anthers, amoeboid tapetum, nectary on the petal lobes, tenuinucellate ovules, weak endothelium, etc. (Erbar 1994). The leaves at the bases of the rosettes and on the rhizomes are spiral and there are no extrafloral nectaries (cf. Sambucus) In Sinadoxa the stamens appear to be divided into two half stamens, each with a separate filament and a monothecal anther; it is molecularly close to Adoxa (Liu et al. 2000). Donoghue et al. (2003) describe the flowers of Sinadoxa as having three corolla lobes and a single carpel. In the curious genus Tetradoxa the half anthers are peltate, there are separate styles, the ovary was described as being superior, and the sepals are persistent (Ying et al. 1993). Adoxa itself has five carpels with five styles (Leins 2000).

For more details of the synapomorphies of this clade, and of clades within it, see Donoghue et al. (2001b, 2003).

Synonymy: Sambucaceae Borkhausen

• Adoxaceae may be recognised by their opposite, toothed leaves that are compound or simple. The inflorescence is usually flat-topped, cymose, and has numerous small flowers. The flowers are 5- or sometimes 4-merous, the calyx is open during development, the corolla is rotate, the style is at most short, and the fruit is drupaceous. Some Cornaceae look rather similar to Viburnum, but the former often have 4-merous flowers, they lack stellate indumentum, and their lateral veins ascend towards the apex of the blade.

Evolution. Early divergence within this clade can be dated to ca 91-75 million years before present (Bell & Donoghue 2005a); the evolution of one-seeded fruits in Viburnum seems to have increased speciation rates there (Moore & Donoghue 2009).

Chemistry, Morphology, etc. Tannin-secreting tubes in Sambucus are coenocytic and up to 32.8 cm long, i.e., the length of an internode (Zobel 1986). A fibrous pericyclic sheath in Viburnum may be absent, and that of Sambucus interrupted; that of Caprifoliaceae s.l. is better developed. The cells in the latter are also longer, except those of Linnaea (Cooper 1939). There is the same number of stamens as petals (cf. many Dipsacaceae s.l.). The chromosomes are distinctive. They are large (cf. Caprifoliaceae, etc.), continuously condensed from prophase to metaphase, the proximal regions normally condensing first, but when cold, areas of the mitotic chromosomes are not condensed (Benko-Iseppson & Morawetz 2000).

Synonymy: Sambucaceae Borkhausen, Tinaceae Martynov, Viburnaceae Rafinesque

Diervillaceae [Caprifoliaceae [Linnaeaceae [Morinaceae [Valerianaceae + Dipsacaceae]]]] [= Caprifoliaceae s.l.]: bark papery-flaky; O-methylated flavones, flavonols +; cork cambium deep-seated; wood often fluorescing; (pith diaphragms +); pericyclic fibers moderately developed; flowers rather weakly monosymmetric [not the K], medium-sized, C 2 + 3, nectary on C associated with unicellular hairs, anthers sagittate, tapetum plasmodial, pollen grains spheroid, echinate, lateral and dorsal carpellary bundles 0, ventral bundles in axis, stigma capitate, wet; (testa hypodermis also differentiated, fibrous or not); chromosomes "small"; duplication of dipsCYC2 and dipsCYC3 genes, horizontal transfer of mitochondrial rps11 gene [?Morinaceae and upwards in the tree].

Early divergence within this clade can be dated to ca 81-60 million years before present (Bell & Donoghue 2005a).

The glandular leaf teeth have a main vein plus two accessory veins, or one accessory vein proceeding above the tooth. Transpetal veins, branch veins linking adjacent median veins, occur in the upper part of the corolla in at least Diervilla, Kolkwitzia and some Valerianaceae and Dipsacaceae (Gustafsson 1995). The calyx often persists, characteristically remaining small and perched on top of the fruit, which may be narrowed towards the apex. Biparental plastid transmission occurs in some taxa, but sampling is poor.

The circumscription, etc., of families follows Backlund and Bremer (1997) and Backlund and Pyck (1998). Note that the small families recently recognised in Dipsacales are a consequence of maintaining the well known Dipsacaceae and Valerianaceae in their old circumscriptions - the small clades resulting from the break-up of the old, broadly-circumscribed Caprifoliaceae remained unaccounted for. The whole lot might usefully be combined in a Caprifoliaceae s.l. (see also A.P.G. 2003), since similarities between the families are considerable and differences are mostly slight.

Some information is taken from Magócsy-Dietz (1899: pith diaphragms, woody members only) Carlquist (1982: wood anatomy), Wagenitz and Laing (1984: nectaries), Verlaque (1988: relationships), Ogata (1988, 1991: wood anatomy), Hoffmann and Göttmann (1990: general), Backlund and Donoghue (1996: general), Backlund and Nilsson (1997: pollen), Backlund and Pyck (1998: general), Benko-Iseppson and Morawetz (2000: general), Nilova (2001: bark anatomy) and Donoghue et al. (2001b) and especially Bell et al. (2001: both general).

DIERVILLACEAE Pyck   Back to Dipsacales

Shrubs; leaves involute [Weigela]; nectary at base of C, filaments hairy, pollen pororate, membrane granulose; G [2], many ?marginal ovules/carpel; fruit beaked, septicidal down sides; n = 9.

1-2/16. East Asia, S.E. U.S.A. (map: from Li 1952, approximate).[Photo - Flowers © M. Clayton]

For the early Tertiary fossil history of Weigela, see Manchester et al. (2009). The large nectary at the base of the corolla in Diervilla (perhaps including Weigela) is a swelling covered by nectar-secreting hairs similar to those in Lonicera, etc. Only one carpel may be fertile, and in some species there may be an "epicalyx" (see below) immediately below the ovary.

Caprifoliaceae [Linnaeaceae [Morinaceae [Valerianaceae + Dipsacaceae]]]: fruit indehiscent.

Note that taxa with connate leaf bases, especially of inflorescence bracts, are scattered through this clade.

CAPRIFOLIACEAE Jussieu, nom. cons.   Back to Dipsacales

Shrubs (trees); vessel elements usu. with simple perforation plates; leaves supervolute or conduplicate (involute - Heptacodium), margins entire, 2ndary veins pinnate to palmate; K small; A (4-)5, filaments glabrous; G [3(-4)], all, 3/4 or 2/4 fertile, odd member adaxial (opposite petals - Leycesteria), 1-8 ovules/carpel; fruit baccate [Lonicera] or drupaceous [Symphoricarpus]; n = 9; deletion in the chloroplast gene clpP.

5[list]/220: Lonicera (180). Mostly N. temperate, esp. East Asia and E. North America (map: see Hultén 1971; Hultén & Fries 1986; Meusel et al. 1992). [Photo - Flower]

There is no stem endodermis. Heptacodium has cork forming in the middle of the cortex and well-developed cortical fibers. Caprifoliaceae also include Triosteum and Symphoricarpus. Triosteum seems to lack dorsal carpellary bundles only and has apotropous ovules (Wilkinson 1949), while the drupes in Symphoricarpus have a thick layer of narrow, vertical fibers on the outside, then a layer of horizontal fibers, and the seed coat itself is crushed.

For relationships in Caprifoliaceae - [Heptacodium [Triosteum *[Lonicera [Leycesteria + Symphoricarpos]]]] - are relationships suggested by Theis et al. (2008), although the node with the asterisk has a poor posterior probability, but a good bootstrap value.

Synonymy: Loniceraceae Vest

Linnaeaceae + Morinaceae + Dipsacaceae + Valerianaceae: supernumerary bracts from aborted flowers +; A 4, didynamous; G [3], 2 abort; fruit a cypsela.

For various interpretations of the supernumerary bracts, bracteoles, epicalyx, and involucel in these families, see Weberling (1992), Roels and Smets (1996), Donoghue et al. (2003, and literature cited) and Pyck and Smets (2004). The epicalyx is sometimes described as consisting of four fused bracts and having nothing to do with bracteoles (Hofmann & Göttmann 1990; Mayer 1998); the individual flower that it surrounds may then be the terminal flower of a partial thyrsoid inflorescence (esp. Roels & Smets 1996). In the characterisations below, the particular derived condition for each family is described; what synapomorphies might "be" morphologically is unclear (Bell et al. 2001). In general, where there are only two stamens, the adaxial pair persists. Of the three carpels, it is the adaxial lateral carpel that remains fertile (Donoghue et al. 2003).

LINNAEACEAE Backlund   Back to Dipsacales

Shrubs (± herbaceous); vessel elements usu. with simple perforation plates; leaves supervolute(-curved); nectariferous petal abaxial, filaments hairy; G [(4)], ovule ?apotropous; K and/or bracts often modified in fruit; n = 8, 9.

4-5/36. Temperate South East Asia (N. Temperate, inc. Mexico). Linnaea, Dipelta (subfleshy, 2-seeded fruits), Kolkwitzia, Zabelia, Abelia. [Photo - Habit] [Photo - Flower]

For distinctive Eocene fruits assignable to this group and their implications for biogeography and evolution, see Manchester and Donoghue (1995; see also Manchester et al. 2009 for the fossil distribution of Dipelta).

Morinaceae + Dipsacaceae + Valerianaceae: perennial rosette herbs with taproot; monoterpenoids, cathecolic tannins, alkaloids +, ellagic acid 0; libriform fibers +; flank-bridge in stem between lateral bundles; bud scales 0; filaments glabrous, anthers dorsifixed, nectary at base of C; G with sterile loculi much reduced, 1 apical ovule/flower, prominent endothelium with crystal layer, chalazal nuclei variously proliferating; exotesta not thickened; endosperm scanty, embryo large; distinctive expansion of the chloroplast inverted repeat.

For flank-bridges, see Neubauer (1979); it is as if two bundles on each side form split laterals, and one branch from each bundle fuses in the middle, so forming the bridging bundle. Bundles innervating the petiole arise from the bridging bundle, except in Morina.

For a comparison of the three families, see Cannon and Cannon (1984), for embryology, see Johri et al. (1991), and for relationships suggested by some morphological characters, see Peng et al. (1995).

MORINACEAE Rafinesque   Back to Dipsacales

Ethereal oils +, iridoids 0; nodes 5:5; stems hollow; leaves involute, margins spinose to pinnatifid (entire), 2ndary veins palmate, bases sheathing; inflorescence of sessile cymes [verticels], epicalyx 12-ribbed; flowers strongly monosymmetric, K 4, monosymmetric; A pairs inserted at different heights on the C tube (2, + 2 staminodes), extra thickened layer of pollen wall, tapetum ?glandular, pollen binucleate, (turret-like triporate), exine columellae reduced, hypostase +, chalazal cells producing multicellular structures; integument multiplicative, not vascularised, outer layer only persisting; endosperm ruminate; n = 17, chromosomes 0.7-1.5 µm long; duplication of dipsCYC2 and dipsCYC3 genes.

2-3[list]/13. Balkans to China (map: see Cannon & Cannon 1984). [Photo - Morina Inflorescence, Acanthocalyx Inflorescence.]

• Morinaceae are robust, perennial herbs. They may be recognised by their opposite leaves with connate bases and often spiny margins and inflorescences consisting of whorls of flowers, each flower having a 12-ribbed epicalyx. The flowers are sympetalous and strongly monosymmetric.

The calyx is distinctly monosymmetric. Remarkable intine-covered structures like pollen tubes are produced in all taxa before pollen germination (Blackmore & Cannon 1983). The integument is 14-18 cells across when the embryo sac is mature, becoming ca 25 cells across later (Vijayaraghavan & Sarveshwari 1968).

For tables of differences between Morinaceae and Dipsacaceae, see Vijayaraghavan and Sarveshwari (1968), Cronquist (1981), Cannon and Cannon (1984) and Johri et al. (1992). There seems to be total confusion as to the nature of the tapetum of Morina: Vijayaraghavan and Sarveshwari (1968) describe it as being polyploid, multinuclear and secretory, Kamelina (1983) as cellular, binuclear and glandular, and Johri et al. (1992) as being plasmodial (and that of Dipsacaceae as being glandular).

For embryology, see also Vijayaraghavan and Sarveshwari (1968: Morina longifolia only), for a monograph of the family, see Cannon and Cannon (1984), and for its phylogeny and biogeography, see Bell and Donoghue (2003).

Dipsacaceae + Valerianaceae: (annual herbs); saccharose +, quercetin 0; vessel elements with simple (scalariform) perforations; (nodes 5 or more:5 or more); leaves simple to ± pinnate, conduplicate (margins entire); flowers rather small; A ± equal in length, tapetal cells 4 or more nucleate [?level], exine columellae prominent and branched, chalazal cells 2-4-nucleate, stigma dry; embryo green.

DIPSACACEAE Jussieu, nom. cons.   Back to Dipsacales

(Shrubby); starch almost 0; (stomata anisocytic); inflorescence capitate, epicalyx 8-ribbed; lateral abaxial C lobes overlapping adaxial lobes, (C with 4 lobes), (A 2-3), (pollen 3-porate), micropyle very long, hypostase +; G [2, 3] [latter - Triplostegia], micropyle very long, chalazal nuclei polyploid, stigma entire or 2-lobed; fruit with calycine awns or bristles; endosperm + (slight in Triplostegia); n = 5, 7-9(-10), chromosomes 1-5 µm long.

11[list]/290: Scabiosa (80), Cephalaria (65), Knautia (60). Eurasia, Africa, esp. Mediterranean region, to New Guinea (Triplostegia) (map: see Meusel et al. 1976; Hultén & Fries 1986; also Verlaque 1985). [Photo - Habit, Inflorescence, Flower.]

• Dipsacaceae are herbs with opposite leaves, capitate, involucrate inflorescences and more or less monosymmetric flowers each with a well-developed epicalyx. The dry fruit has a single seed.

Evolution. The well-developed epicalyx suggests that the seeds are often wind-dispersed, but myrmecochory may occur in almost half of the family (Lengyel et al. 2009).

The cork of Knautia may be superficial (Metcalfe & Chalk 1950).

Triplostegia, which has a double epicalyx and valepotriates, seems best assigned to Dipsacaceae (e.g. 100% support in a 30 taxon-5 gene analysis - Davis et al. 2001; see also Bell & Donoghue 2000; Zhang et al. 2001; Bell 2004). However, Pyck and Smets (2004) show that although a two-gene analysis places Triplostegia in this position, morphological data alone and when combined with the molecular data place it sister to Valerianaceae. As they note, [Triplostegia + Valerianaceae] have valpotriate-type iridoids, a pollen aperture with a halo, a granulose colpus, and also endosperm reduction in common, and possibly even epicalyx/bracteole similarities. Avino et al. (2009) found that Triplostegia might be sister to [Valerianoideae + Dipsacoideae], and used this topology in their character state reconstruction. Clearly, where Triplostegia ends up will substantially affect apomorphy positions.

The position of Triplostegia aside, Pterocephalodes, a recent segregate from Pterocephalus (Mayer & Ehrendorfer 2000), has been found to be sister to the rest of the clade (Avino et al. 2009).

For embryology, see Kamelina (1983), for fruit, see Verlaque (1977), for a general study, see Verlaque (1985, 1986), for phylogeny and the evolution of seed dispersal syndromes, see Caputo et al. (2004).

Synonymy: Scabiosaceae Adanson, Triplostegiaceae Airy Shaw

VALERIANACEAE Batsch   Back to Dipsacales

Foetid monoterpenoids and sesquiterpenoids, valpotriates +; true tracheids ?; leaf bases often sheathing; (plant dioecious), flowers small, (polysymmetric), bracteoles +, epicalyx 0; K small, abaxial C spurred (or not); A 1-4 (5), pollen colpate, (G with sterile loculi well developed); K + (a pappus; reduced); testa 1-layered; n = (7-)8(9-12), chromosomes 0.7-4 µm long.

17[list]/315: Valeriana (200: perhaps para- or polyphyletic - Hidalgo et al. 2004), Valerianella (80). N. temperate, especially Mediterranean, and Andean South America (half the family) (map: from Hultén 1958; Meusel et al. 1978). [Photo - Habit, Flower, Flower.]

• Valerianaceae are often foetid herbs with opposite, sometimes connate leaves and rather small flowers. The inflorescence is cymose and bracteolate. The sympetalous corolla is often spurred but other than that may be only weakly monosymmetric; the calyx often develops greatly in fruit.

Evolution. Crown-group Valerianaceae may be 60-55 million years old (Bell & Donoghue 2005a), but diversification in the Andean paramo - ca 1/7 of the family - happened less than 5 million years ago (Bell & Donoghue 2005b).

Chemistry, Morphology, etc. Valepotriates are triesters of route I secoiridoids; for their distribution, see Backlund and Moritz (1998). The calyx in Patrinia and in particular Nardostachys is well-developed. Evident sterile loculi, a well developed calyx, an androecium with either four or five stamens, etc., are likely to be plesiomorphic in the family. The corolla tube of Centranthus is divided vertically into two by a septum. The extra "bracteole" of Patrinia may be an interpolated structure that develops into a fruit wing (Hoffmann & Göttmann 1990). For general morphology, see Eriksen (1989).

Phylogeny. Patrinia may be sister to the rest of the family (Pyck 2002), Nardostachys sister to the remaining taxa (Bell 2004; Bell & Donoghue 2005b: both with very strong support), or the two together form a well-supported clade sister to other Valerianaceae (Hidalgo et al. 2004); the former topology is more likely. Valeriana celtica, Valerianella and Centranthus may be successively sister to Valeriana (Bell & Donoghue 2005b, for which see for further details). For other phylogenies of the family, see Bell (2004) and Hidalgo et al. (2004).