EXTANT SEED PLANTS

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

MAGNOLIOPHYTA

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

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

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

Relationships between the lineages immediately above the basal pectinations in the main tree, the ANITA grade (Amborellales, Nymphaeales and Austrobaileyales here), remain somewhat a matter of conjecture. The topology of the main tree in this area is thus definitely provisional; it differs somewhat from that in A.P.G. (2003). For further information, see the discussion immediately preceding the Magnoliales, i.e. the magnoliid clade; Ceratophyllales, Chloranthales, and monocots are the other clades involved.

MONOCOTS + EUDICOTS: (stamens opposite [two whorls of] P).   Back to Main Tree

EUDICOTS  Back to Main Tree

Myricetin, delphinidin scattered, asarone 0 [unknown in some groups, + in some asterids]; root epidermis derived from root cap [?Buxaceae, etc.]; nodes 3:3; stomata anomocytic; flowers (dimerous), cyclic, K/outer P members with three traces, "C" with a single trace, few, (polyandry widespread), filaments fairly slender, anthers basifixed, pollen with endexine, tricolpate, G with complete postgenital fusion, style solid [?here]; seed coat?

Tricolpate pollen from the Late Barremian-Early Aptian of the Cretaceous some 120-125 million years before present is known, and an age of some 125 my for the eudicots seems reasonable (e.g. Magallón et al. 1999; Sanderson & Doyle 2001), and this age is similar to that of the oldest monocot fossils; Moore et al. (2007) suggest an age of 131.1-118.5 million years for diversification of the eudicot crown group. Subsequent divergence of eudicot clades like Proteales, Buxales, etc., was rapid, occuring 120-116 million years before present (Anderson et al. 2005), Note, however, that Wikström et al. (2003) suggest an age of 131-147 million years before present for the clade. For the possible functional significance of the evolution of triaperturate - or multiaperturate - pollen, see e.g. Dajoz et al. (1991), Furness and Rudall (2004); thus many apertures may increase the speed of germination of the pollen but decrease its viability.

Cuticle waxes as clustered tubules, nonacosan-10-ol the main wax could be optimised to this position, being lost in Sabiaceae, Platanaceae, Buxales, and perhaps also in the internode below Gunnerales... (but they are present in a few Santalales, also in woody Saxifragales: see Barthlott et al. 2003). Doyle (2007) scored chloranthoid teeth as plesiomorphous for eudicots; given current ideas of phylogeny, they may be an apomorphy here. he also considers plmate-crowded veins to be an apomorphy for all eudicots, but Sabiaceae are not mentioned, the interpretation of the venation of Euptelea is debatable, as he notes (Doyle 2007), and the palmate venation of aquatics like Nelumbo confuses the situation. Interestingly, dimerous flowers are scattered through the basal eudicot grade, but are at least very uncommon in the core eudicots and in monocots (Drinnan et al. 1994; Soltis et al. 2003; Wanntorp & Soltis et al. 2005; Ronse De Craene 2005; Doust & Stevens 2005; Kramer & Zimmer 2006; Moody & Les 2007 [this with examples of variation in the core eudicot Haloragaceae]). Stamens are also quite often inserted opposite the tepals in the basal eudicot grade (e.g. Endress 1995 for illustrations of these in Ranunculales; Doust & Stevens 2005); this feature has been placed at the [monocots + eudicots] node here. Taxa with androecia that are initiated as antesepalous triplets are scattered (but really rather uncommon) throughout the group (Hufford 2001a).

Within the eudicots, most recent studies place Gunnerales sister to core eudicots (or sister to other core eudicots: Senters et al. 2000; Hilu et al. 2001; S. Kim et al. 2004; especially D. Soltis et al. 2003a); there was a weakly supported [Gunnera + Dillenia clade in Zhu et al. (2007). Relationships basal to this group have, however, been uncertain. Ranunculales are generally been considered to be sister to all other eudicots, however, recent work suggests that Ceratophyllaceae may occupy this position (e.g. Moore et al. 2007: see the discussion immediately preceding the Magnoliales). An unresolved Proteales-Sabiaceae group is often sister to eudicots minus Ranunculales (e.g. S. Kim et al. 2004b); the relationships of Buxales and Trochodendrales were also unclear. Savolainen et al. (2000a) suggested that there was weak evidence for an association of Sabiaceae with Proteales. A three-gene analysis by Soltis et al. (2003) found that that Sabiaceae were near Proteales, Buxales, etc., while a four gene analysis (Kim et al. 2004a) had a weakly supported clade [Trochodendrales [Sabiaceae + Buxales]]. Hilu et al. (2003) suggested that Buxales were sister to core eudicots, although other details of their phylogeny are congruent with those presented here. Other permutations, none strongly supported, were found by Zhu et al. (2007). Worberg et al. (2006, 2007) present a 3-gene (chloroplast) phylogenetic analysis focusing on the eudicots; the relationships they suggest are followed here. They found most of the relationships along the eudicot spine to be strongly (>90% jacknife) supported, although the position of Sabiales had only 83% support, of which most came from the matK gene (the other genes examined were petD and trnL-F). Although Buxales are sister to Gunnerales plus core eudicots in most analyses, relationships in this general area were scrambled using the PetD marker alone (Worberg et al. 2007). Qiu et al. (2006b) found a scrambled set of relationships in a three-gene analysis of mitochondrial data, but with eight genes a topology similar to that used here was found, although Sabiaceae were weakly supported as sister to Proteales.

See Hegnauer (1990) for a discussion of the chemistry of the Polycarpicae, which also includes the magnoliids and Austrobaileyales. The Eudicot Evolutionary Research website should also be consulted.

RANUNCULALES Dumortier  Main Tree, Synapomorphies.

(O-methyl)flavonols, flavonols +; vessel elements?; young stem with separate bundles, vessels only in central part, true tracheids +; rays exclusively wide multiseriate [and in wood, where present]; (wood fluorescence +); cambium storied; sieve tube plastids large S-type, dispersive P-protein +; petiole bundles annular; leaf cuticle waxes as clustered tubules with nonacosan-10-ol the main wax; leaf ?teeth; G opposite P; P deciduous in fruit; seed exotestal; endosperm development?, embryo size? - 7 families, 199 genera, 4445 species.

Ranunculales contain ca 1.6% of eudicot diversity. Magallön et al. (1999) suggest an age of ca 70 million years before present for them, based on the earliest fossil occurence assignable to it, however, this is a member of the decidely non-basal Menispermaceae. Anderson et al. (2005) date stem group Ranunculales at 122-120 million years before present, divergence within it beginning 121-114 million years before present. All families diverge before 105 million years before present except Ranunculaceae/Berberidaceae (104-90 million years before present). See Anderson et al. (2005) for further details of the timing of diversification within the clade. Ranunculales - perhaps especially Menispermaceae and Ranunculaceae - is not much used as food plants of butterfly caterpillars (Ehrlich & Raven 1964). There are a number of reports of delayed fertilisation (up to some two months or more after pollination), including Eupteleaceae, Circeasteraceae, Lardizabalaceae, and Ranunculaceae (Sogo & Tobe 2006d for references).

Berberin, common in the order, is synthesised via the tyrosine pathway. Gleissberg and Kadereit (1999) discuss the evolution of leaf form in the order, with polyternate/acropetal/basipetal-pedate leaves perhaps being plesiomorphic. The glandular leaf teeth have a clear, persistent, swollen cap into which higher order lateral veins also run. What is the distribution of colleters? Drinnan et al. (1994) suggest that petals have been derived from stamens several times in the order, examples would include Lardizabalaceae. Tamura (1965) summarises evidence for the staminal nature of the petals in Ranunculaceae: petals have a single trace, are in the same parastiches as androecial members, are similar to stamens in early development, and are often peltate. Tepalline spurs have evolved four to six times in the order; they may be on members of the outer (Myosurus) or inner (Aquilegia) perianth whorl, and be equal in number as the basic merism of the flower (Dicentra) or single (Delphinium) (Damerval & Nadot 2007). Is the pollen endexine ever lamellate? Antipodal cells are commonly other than simply persistent; data are summarised in Williams and Friedman (2004).

Relationships in the order are fairly well understood - see Hoot and Crane (1995), Kadereit et al. (1995), Oxelman and Lidén (1995), and Hoot et al. (1999: three genes). Soltis et al. (2003a), a four-gene analyses (Kim et al. 2004a), and a study of non-coding chloroplast DNA (Worberg et al. 2006, 2007), all suggest that Eupteleaceae may be sister to the whole of the rest of the order (in some of these studies the support for this position is only moderate), and this is the topology adopted here, although some earlier studies suggested other topologies, such as Ranunculaceae sister to other members of the order (Soltis et al. 2000). A.P.G. II allows as an option the possibility of including Papaveraceae, Fumariaceae, and Pteridophyllaceae in an expanded Papaveraceae, which I follow here (see also Judd et al. 2002). Papaverales are commonly recognised as a separate order next to Ranunculales (Cronquist 1981; Dahlgren 1989), but there is no point in recognising them, especially given the developing support for the position of Eupteleaceae as sister to the rest of the clade.

For information, see Ernst (1964: general), Hennig et al. (1994: cuticle waxes), Drinnan et al. (1994: floral evolution), Barthlott and Theisen (1995: cuticle waxes), Behnke (1995b: sieve element plastids and phloem proteins), Carlquist (1995b: wood anatomy), Endress (1995: floral morphology), Blackmore et al. (1995: pollen, very variable), Brückner (1995: summary of seed anatomy), Loconte et al. (1995: morphological phylogeny, Ranunculales paraphyletic...), Floyd et al. (1999: embryology), Floyd and Friedman (2000: endosperm) and Damerval and Nadot (2007: floral evolution).

Includes Berberidaceae, Eupteleaceae, Circaeasteraceae, Lardizabalaceae, Menispermaceae, Papaveraceae (inc. Fumarioideae Papaveroideae, Pteridophylloideae), Ranunculaceae.

Synonymy: Papaverineae Thorne & Reveal, Ranunculinae Bessey - Berberidales Dumortier, Eupteleales Reveal, Glaucidiales Reveal, Helleborales Nakai, Hydrastidales Takhtajan, Lardizabalales Loconte, Menispermales Bromhead, Nandinales Doweld, Papaverales Dumortier, Podophyllales Dumortier - Berberidanae Doweld, Papaveranae Doweld, Ranunculanae Reveal - Ranunculidae Reveal - Berberidopsida Brogniart, Papaveropsida Brongniart, Ranunculopsida Brongniart

EUPTELEACEAE K. Wilhelm   Back to Ranunculales

Deciduous trees; (dihydro)chalcones +; cork deep in cortex; vessel elements with scalariform-reticulate perforations; rays ca 10-seriate; nodes 1:many; idioblasts?; cuticle wax crystalloids 0; bud scales +; leaves subplicate-conduplicate, margins gland-toothed, 2ndary veins pinnate; inflorescence axillary, fasciculate or umbellate; flowers (staminate), disymmetric; P 0, A 6-20, anthers valvate, latrorse, connective prolonged, G 6-31, stipitate, "intermediate ascidiate", 1-2(-4) epitropous ovules/carpel, outer integument 4-5 cells across, micropyle bistomal, stigma at most weakly secretory, brush-like; fruit a samara; seed with ± enlarged exotestal cells (sclerotic mesotesta), endotesta lignified subpalisade; endosperm cellular; n = 14; germination epigeal.

1[list]/2. Temperate South East Asia (Map: from Fu & Hong 2000). [Photo - Collection]

• Eupteleaceae can be recognised by their spiral, dentate leaves with strong, ascending, pinnate veins going into the teeth, the excavated petiole bases, and the small flowers that lack a perianth and have anthers with a long connective and separate stipitate carpels. These latter develop into small, disc-shaped samaras down one side of which are the remains of the brush-like stigma.

Lateral veins only approach the glandular teeth; the gland itself has an apical cavity. Is the wood storied, what about fluorescence, separate bundles?

Eupteleaceae were placed next to Cercidiphyllaceae in Hamamelidales by Cronquist (1981) or Hamamelididae by Takhtajan (1997). They have been often been linked with Eucommiaceae, for which see Garryales (asterid I!) - coincidentally, the latter also have delayed fertilization.

See Endress (1969, 1993) for some general information, Hegnauer (1973, 1989, 1990) for chemistry, and Li and Ren (2005) for wood anatomy.

Papaveraceae [[Circaeasteraceae + Lardizabalaceae] [Menispermaceae [Berberidaceae + Ranunculaceae]]]: vessel elements with simple perforation plates; leaves (ternately compound or palmately lobed, articulated), venation palmate; stigma wet [optimization?].

PAPAVERACEAE Jussieu, nom. cons.   Back to Ranunculales

Plant herbaceous, non-mycorrhizal [Fumarioideae, too?]; roots diarch [lateral roots 4-ranked]; cork?; numerous alkaloids [inc. berberin, known from Pteridophylloideae?; protopine] +; leaves usu. spiny (or gland?) toothed, quite often glaucous, base broad; inflorescence terminal; flower parts whorled, dimerous, P = K + C, K 2, median, deciduous, C 4, A extrorse, G [2], collateral, occluded by secretion, placentation (protruding-diffuse) parietal, ovules often amphitropous to campylotropous, (stigmatic lobes commissural); capsule septicidal [= placenticidal], (with persistent placental strands); endotesta well developed, with coarse fibrillar network; seed curved.

44/760 - 3 subfamilies and six tribes below. Largely N. Temperate, also S. Africa, scattered in South America, etc.

1. Pteridophylloideae [undescribed]   Back to Ranunculales

Chemistry ?; vessel elements?; petiole bundles?; cuticle waxes?; leaves pinnate, fern-like; inflorescences indeterminate, flowers disymmetric; K small, petaloid, caducous, A 4, alternating with C, diagonal, 1(2) ovules/carpel, integument thickness and micropyle?, style long, stigma surface?; fruit a capsule, placentae persistent; exotesta ± collapsing, tegmen thin; embryo?; endosperm development?; n = 9.

1/1: Pteridophyllum racemosum. Japan.

Pteridophyllum was included in Fumariaceae by Cronquist (1981). General information is taken from Lidén (1993). Pteridophyllum is poorly known embryologically, although there is a cellulose network in the endotesta like that of some Papaveroideae.

Synonymy: Pteridophyllaceae (Murbeck) Reveal & Hoogland

Papaveroideae + Fumarioideae: isoquinoline protopine +; little oxalate accumulation; laticifers, articulated or not, anastomosing or not; petiole bundles arcuate; leaves soft, ± fleshy; inflorescences determinate; (carpels gaping apically), micropyle bistomal, antipodals endopolypoid; (seeds arillate); endotesta with calcium oxalate crystals, exotegmen fibrous or not, endotegmen walls thickened; endosperm nuclear.

One species in each subfamily has distinctive UV fluorescence of unlignified cell walls (Hartley & Harris 1981).

2. Papaveroideae Eaton   Back to Ranunculales

(Small trees); latex + milky; nodes also unilacunar; leaf ptyxis variable, colleters +; flowers large, K enclosing the bud, lobed [usu. on left], caducous, C also 6, usu. fugacious, crumpled (0), A (4-)many, in multiples of two or three, (placentation ± axile), many ± anatropous ovules/carpel, antipodals also multinucleate, style usu 0, stigmas often confluent, dry; capsule also with transverse dehiscence (indehiscent, schizocarp); exotegmen often with thickened outer walls, unlignified (endotegmen not persistent); n = 5-10 (14, 19).

23[list]/230 - four tribes below. Largely N. temperate; Argemone mexicana commonly introduced in the tropics (Map: from Ownbey 1958, 1961; Hultén & Fries 1986; Fl. N. Am. III 1997; Fu & Hong 2000; Malyschev & Peschkova 2004). [Photos - Collection (except Dicentra and Corydalis]

2A. Papavereae Dumortier

Hairs multicellular and multiseriate; G [3-24]; fruits opening by pores.

8/95-125: Papaver (50-80: paraphyletic), Meconopsis (50). N. (warm) temperate, also South America (Argemone) and S. Africa and Cape Verde Islands (1 sp. in each - Papaver).

Papaver is paraphyletic and Meconopsis polyphyletic; generic limits need adjustment (Kadereit & Sytsma 1992; Kadereit et al. 1997; Carolan et al. 2006).

2B. Chelidonieae Dumortier

Latex orange, yellow or red; hairs multicellular and terminally uniseriate; pollen also polyporate, G [(3)] (as few as 1 ovule/carpel); fruit elongated, seeds often arillate [ant dispersed].

9/48. East Asia and E. North America (also Europe, C. and S. America, West Indies).

Bocconia has a gynophore and basal placentation, Sanguinaria a 2-merous perianth.

Synonymy: Chelidoniaceae Martynov

2C. Eschscholtzieae Baillon

Hairs unicellular; subepidermal collenchyma in stem, pollen polycolpate; capsule dehiscing explosively from base.

3/16. W. North America.

The flowers of Eschscholtzia have a short hypanthium.

Synonymy: Eschscholziaceae Seringe

2D. Platystemoneae Spach

Hairs multicellular and multiseriate; G [3(-25 - Platystemon)], styles free.

3/5. W. North America.

Platystemon has more or less free carpels.

Synonymy: Platystemonaceae (Spach) Lilja

Vascularization of the petals varies, but even if there is more than a single trace, the traces seem to have a single point of origin (Dickson 1935). As in Ranunculaceae, the numerous stamens in Papaver, etc., may be derived from a paucistemonous condition. The outer integument of Dendromecon is seven cells across. The gametophytic self-incompatability system of Papaver is associated with a stigma that is dry (Wheeler et al. 2001).

Some information is taken from Dickson (1935: floral vascularization) and Kadereit (1993); see Gleissberg and Kadereit (1999) for the complexities of leaf development interpreted in a phylogenetic context.

3. Fumarioideae Eaton   Back to Ranunculales

Latex 0; acetylornithine, (berberin) +; nodes uni(-multi)-lacunar; watery sap in often non-articulated sacs; subepidermal collechyma in stem; nodes 1:1+; flowers transversely disymmetric, K and C in 2's, K small, 4, A 6, free to connate, secondary pollen presentation common, (pollen 3-colporoidate, etc.), nectaries at base of stamens, 1-many ovules/carpel, (placentation axile); (fruit a lomentum, nutlet); seeds curved, exotesta palisade or not, endotesta lacking fibrillar network; (embryo long); n = (6-)8(+).

19/530 - two tribes below. Mostly N. temperate, also S. Africa (Map: from Hultén & Lidén 1986; Fries 1986; Hong 1993; Fl. N. Am. III 1997; Fu & Hong 2000; Malyschev & Peschkova 2004; Serban Procheŝ, pers. comm. S. Africa). [Photo - Corydalis Flower, another, Dicentra Flower.]

Acetylornithine is involved in nitrogen transport (Jensen 1995).

3A. Hypecoeae Dumortier

Inner petals 3-lobed, A 4, two with two vascular strands [each 2 A connate], pollen dicolpate, deposited on inner petals; styles 2; seeds covered with rectangular crystals.

1/18. Mediterranean to W. China.

Synonymy: Hypecoaceae (Dumortier) Willkomm & Lange

3B. Fumarieae Dumortier

(Inflorescences racemose); flowers monosymmetric, single spur adaxial), K minute, two (one) outer C spurred, inner C apically coherent, with median joint, A in two groups of three, 2 anthers dithecal and 4 monothecal, style +, white, caducous (green, persistent), (stigma flattened, with marginal lobes), pollen deposited on stigma; (seed with elaiosome); exotesta usu. pigmented, (endotesta palisade, crystalliferous); suspensor cells like a small bunch of grapes.

19/505: Corydalis (400), Fumaria (55). Eurasia, North America, North and South Africa, mountains of E. Africa; ¾ species in Sino-Himalayan region.

Fumaria has only a single ovule and the fruit is nut-like and indehiscent.

Synonymy: Corydalaceae Vest, Fumariaceae Berchtold & Presl, nom. cons.

• Papaveraceae are usually herbs with an exudate of some kind whose fleshy leaves have broad bases. The flowers are 2-merous, and the calyx and corolla are clearly distinguishable; the gynoecium has two (or more) carpels, parietal placentation, and usually a short style.

• Pteridophylloideae are herbs with a rosette of fern-like leaves and a scapose inflorescence with rather small, 2-merous flowers which have a distinct style.

• Papaveroideae are usually herbs that may be recognised by their often soft if rather bristly-hairy leaves, copious latex, and flowers with two, large, fugaceous sepals enclosing the bud, crumpled petals, usually numerous stamens, and syncarpous gynoecium with at most a short style and parietal placentation.

• Fumarioideae are often fleshy herbs with watery sap and distinctive 2-merous mono- or disymmetrical flowers that are usually spurred; the calyx is very small or almost invisible.

Several species of Fumaria and its relatives are chasmophytes in the Old World, growing in the apparently most inhospitable habitats despite their delicate and rather succulent habit. Papaveraceae have gametophytic self-incompatability systems which, however, are very different from those in core eudicots (Charlesworth 2005).

For the unusual (transverse) plane of monosymmetry in Fumarioideae, see e.g. Troll (1957), Ronse Decraene and Smets (1992), Endress (1999), etc. Corydalis, etc. have only one outer petal spurred and the flower is monosymmetric; there is a 90° rotation of the flower rather late in development so the spur is in the adaxial position (Ronse Decraene & Smets 1992) and the monosymmetry is functionally vertical. There is a correlation between flowers with monosymmetry and indeterminate inflorescences, and there has been duplication of CYCLOIDEA genes in the Papaveraceae s.l. (Kölsch & Gleissberg 2006). The nature of the androecium of Fumarieae in particular has occasioned much discussion, and it has sometimes been suggested that two anthers have each split into two, monothecal units, so there would be only 4 stamens altogether, however, it is likely that the androecium consists of two dithecal and four monothecal stamens, and in Hypecoum the monothecal stamens have fused in pairs, hence the double vascular supply to two of what appear to be dithecous stamens (see Ronse Decraene & Smets 1992, for literature). The androecium of Pteridophyllum has also been interpreted as being derived from a a flower with six stamens; there the lateral stamens have been lost (Ronse Decraene & Smets 1992). The stigma of Fumaria and relatives, in which the pollen is deposited (secondary pollen presentation), can have a complex morphology; there is also secondary pollen presentation in Hypecoeae, and here the pollen is deposited on the inner petals. I am unsure if all/some Papaveroideae have extrorse anthers, but anthers are clearly extrorse in the other subfamilies (Murbeck 1912). Papaveraceae are described as having hollow styles, although the hollow may become occluded by papillae (Hanf 1935).

For a phylogeny of Fumarioideae, see Lidén et al. (1997); Dicentra is dismembered and the old Corydaleae becomes paraphyletic. Various other studies have been carried out on the Papaveraceae s.l., and these include Nikolic (1995: numerical taxonomy) and Judd et al. (1994: morphology) suggest relationships, the groupings above are taken from Hoot et al. (1997) and Kadereit et al. (1994, 1995). In Fumarioideae in particular morphological studies tend to recover a Fumarieae and Corydaleae.

See also Murbeck (1912) for floral morphology, Bersillon (1955) for nodal anatomy, Hegnauer (1969, 1990) and Preininger (1986) for chemistry, G. Dahlgren (1981) for stigma secretions, and Brückner (1983, 1985) and Fukuhara and Lidén (1995) for testa anatomy. In Fumarioideae, for ovule orientation, see Goebel (1932), for general information, Lidén (1986 [esp. Fumarieae], 1993), and for arils (?originated several times), see Fukuhara (1999).

[Circaeasteraceae + Lardizabalaceae] [Menispermaceae [Berberidaceae + Ranunculaceae]]: rays broad; flowers 3-merous, K, C, and A, or P and A opposite each other, outer P members with three or more vascular traces; AP₃ gene triplicated.

The sepals/perianth usually have three or more traces (e.g. see Hiepko 1965).

For pollen morphology, see Nowicke and Skvarla (1982).

Circaeasteraceae + Lardizabalaceae: A extrorse.

CIRCAEASTERACEAE Hutchinson, nom. cons.   Back to Ranunculales

Herbs; chemistry?; cork ?; true tracheids?; nodes 1:1; petiole bundle ?arcuate; prophyll adaxial; leaves also 2-ranked, palmate or simple, margins toothed, venation largely dichotomous; inflorescence terminal, cymose or thyrsoid, or flowers terminal, perfect or not; parts spirally arranged, P 2-3, 5(-7), with a single trace, (8-13 clavate glands - Kingdonia), A (1-)2-6(-8), (anthers bisporangiate/monothecal - Circaeaster), exine layered-striate, G 1-9, occlusion by ?secretion, 1-2 ± apical atropous [Circeaster] or hemianatropous unitegmic tenuinucellate ovules/carpel, integument 3-5 cells thick, embryo sac tetrasporic, 4- or 8-nucleate; fruit an achene; seed coat degenerating, thin; endosperm cellular; embryo relatively large; n = 9, 15.

2[list]/2. N. India to S.W. and W. China (Map: from Fu & Hong 2000). [Photo - Circaeaster Habit]

• Circaeasteraceae are easily recognised. They are small herbs, their leaves have largely dichotomous venation, and their rather small flowers have separate carpels.

Circaeasteraceae have been placed in the Ranunculaceae-Anemoneae, as by Kosuge et al. (1989)

Kingdonia may have up to four bundles departing from the single trace, endotrophic mycorrhizae and, like Circaeaster, several root hair zones on the roots (Ren & Hu 1998). Xylem perforation plates may also be scalariform. Kingdonia at least appears to have an adaxial prophyll (see s.e.m. of axillary buds in Ren et al. 2004 - no comment is made about this). Circaeasteraceae do not show the regular relationship between the stamens and perianth members of many other Ranunculales. Circaeaster lacks bracteoles. The tepals of Kingdonia have a single trifid vein, indeed, all floral organs are innervated by a single vein, apart from the first tepal, which has two traces (as in Ranunculaceae, see Ren et al. 2004). Its anthers are extrorse, while those of Circaeaster may be latrorse. Kingdonia has 8-13 glistening clavate glands inside the perianth whorl; these are described as petals (perhaps unlikely) by Tamura (1993) and as staminodes by Ren et al. (2004) and may secrete nectar. Mesogamy, i.e. the pollen tube entering the ovule laterally by penetrating the integument, is reported for Circaeaster by Junell (1931); Circaeaster also has endosperm with a chalazal haustorium.

Kingdonia has been placed in the Ranunculaceae-Anemoneae, e.g. by Kosuge et al. (1989); the dichotomous venation of the leaves and the separate carpels have attracted attention as possibly indicating a very "primitive" group.

Keeping Kingdoniaceae separate from Circaeasteraceae is optional in A.P.G. II (2003).

General information is taken from Tamura (1993: in Ranunculaceae) and Wu and Kubitzki (1993); see also Nowicke and Skarvla (1981) for pollen, Hu et al. (1990), Ren and Hu (1995) and Tian et al. (2006) for information on Circaeaster agrestis, and Ren et al. (1998, 2004) for information on Kingdonia uniflora. See the inside cover of Act. Bot. Bor.-Occid. Sinica 24(1) (2004) for a photograph of Kingdonia uniflora with excellent details of gross morphology.

Synonymy: Kingdoniaceae Airy-Shaw (leaves 2-ranked; 8-13 petaloid staminodes outside ("C" above!), A 3-6, anthers tetrasporangiate, G 3-9; n = 9)

LARDIZABALACEAE R. Brown, nom. cons.   Back to Ranunculales

Lianes; glabrous or with uniseriate hairs; (plant Al-accumulators); petiole bundles arcuate; bud scales +; leaves trifolioliate, conduplicate, margins entire; inflorescence axillary, racemose; parts spiral, K 6, C 6, nectariferous, both with a single trace, staminate flowers: A 6, connective often prolonged into appendage; carpellate flowers: staminodia +, G 3, also spiral, with postgenital fusion and secretion, placentation marginal, micropyle endostomal; placenta fleshy in fruit; endosperm cellular, starchy or with hemicellulose; germination phanerocotylar.

9[list]/36 - two groups below. South East Asia and Chile (Map: see Taylor B. 1967; Ying et al. 1993). [Photos - Lardizibala Staminate flower, Boquila Flowers, Fruit, Fruit close-up]

1. Sargentodoxa

Cork deep-seated; tanniniferous cells +; plant dioecious; G many, 1 pendulous ovule/carpel; surface of testa featureless; n = 11.

1/1: Sargentodoxa cuneata. China.

Synonymy: Sargentodoxaceae Hutchinson

2. Lardizabaloideae Kosteletzky

(Shrubs); (vessel elements with scalariform perforations); (stomata cyclocytic); (leaves palmately or odd-pinnately) compound, with basal tooth or lobe; 2ndary veins pinnate); plant monoecious (dioecious; flowers perfect): (K 3, C 0), staminate flowers: (A 3, 8), filaments ± connate, pollen also colporoidate; carpellate flowers: A staminodial, G (-12), (placentation laminar), (few-)many (hemitropous) ovules/carpel, (stigma peltate); fruit a berrylet or fleshy follicle; exotestal cells lignified, elongated, ± oblong [Descaisnea] or unlignified, fibrous [Akebia, Hoelboellia], hypodermal cells thickened; endosperm cellular, starchy or with hemicellulose; n = 14-16, ?17, 18.

8/35. South East Asia and Chile (Lardizabala, Boquila).

Synonymy: Decaisneaceae (Quin) Loconte (starchy endosperm), Sinofranchetiaceae Doweld

• Lardizabalaceae are usually lianes with compound leaves and broad rays in the wood, the plants being monoecious or dioecious. The medium-sized, three-merous flowers have the perianth members and androecium opposite each other. The carpels are free and the fruits are more or less fleshy, although they are sometimes follicles.

Lardizabalaceae and Menispermaceae are both lianes, sometimes vines, and they have very large sieve tube plastids and unisexual flowers... Wood fluorescence? Smets (1986) suggested that the nectaries are staminal nectaries.

Sargentodoxa is sister to the rest of the family (Hoot et al. 1995b, see also Hoot 1995a), and it has some differences (autapomorphies) from the other genera, but there is no compelling reason to segregate it as a family.

General information is taken from Wu and Kubitzki (1993) and Qin (1997), chemistry from Hegnauer (1966, 1989, also 1973, as Sargentodoxaceae), seed surface from Xia and Peng (1989), and some anatomy from Yong and Su (1993); Zhang et al. (2005) provide a detailed study of Sinofranchetia.

Menispermaceae [Berberidaceae + Ranunculaceae]: (berberin + [a benzylisoquinoline - scraped stems yellow in color]); endosperm nuclear.

MENISPERMACEAE Jussieu, nom. cons.   Back to Ranunculales

Often woody lianes (trees; vines); also/or aporphine alkaloids, sesqui- and diterpenoids +, (plant tanniniferous); secretory cells in files; sclereids common; crystals common; stomata various, often ± cyclocytic; hairs unicellular to uniseriate; leaves simple (compound - Burasia), margins entire (toothed; lobed), often peltate or with the lamina borne on the top of the petiole, petiole often pulvinate at base and apex; plants dioecious, inflorescence axillary; flowers small, parts whorled or spiral, "K" (1-)6(-12), with a single trace, "C" 0-8, often connate, (clasping A), staminate flowers: A 3, 6, 12 (1-40, if many, not all opposite C), (connate and ± extrorse), anthers (bisporangiate/monothecal), thecae superposed, pollen tricolporate, pistillodes +/0; carpellate flowers: staminodes +/0, gynophore common, G (1-)3(-30<), with postgenital fusion and secretion, opposite P [Cissampelos], 1-2 often unitegmic, hemitropous-campylotropous ovules/carpel, antipodals multiplying, multinucleate, micropyle endostomal (zig-zag), stigma ± flaring; fruit a straight drupelet, 1-seeded; seed ruminate (not), with condyle [placentar intrusion], coat undistinguished (exotesta tabular, lignified); endosperm +, embryo long, cotyledons divaricate, ± foliaceous; n = (9-)11-13(+).

70 (many small)[list]/420. Pantropical, usually lowland (Map: see van Balgooy 1993; Fu & Hong 2000: Malyschev & Peschkova 2004; Rosa Ortiz-Gentry, pers. comm. 2004). [Photo - Fruit, Fruit.]

1. Tinomiscieae

Stomata cyclocytic; endaperture longitudinally elongated.

1/1: Tinomiscium petiolare. Southeast Asia, Malesia.

The tangential cell walls of the rays are oblique to the ray axis when viewed in transverse section; this is uncommon in other Menispermaceae, where the walls are at right angles (Jacques & de Franceschi 2007), but I do not know the distribution of this feature in the outgroups.

[Tinosporeae + Coscinieae] + The Rest: serial cambia frequent; (rays narrow); pollen endapertures circular.

2. Tinosporeae + Coscinieae: ?

3. The Rest: style lateral to basal; endocarp often sculpted; (endosperm ruminate; 0), embryo curved, cotyledons narrow.

• Menispermaceae are recognisable by their often viney/lianescent habit, broad rays, petioles pulvinate at both ends, and often coriaceous lamina with palmate venation. The lamina is also often peltate, and even if not, the two sides of the lamina nearly meet on top of the petiole. The plants are dioecious, the often 3-merous flowers are small, and the drupelets often have a distinctively sculpted endocarp and can be strongly curved (hence the "moon-seed" family - also Menispermum canadense).

Callicrypta, from the mid-Cretaceous of Siberia, has very small flowers (carpellate) with the parts more or less opposite, or forming spirals, and may be Menispermaceae; it is unclear what a link between Menispermaceae and Amborellaceae - hardly close - might consist of (cf. Krassilov & Goloneva 2004). Jacques and de Franceschi (2005, and references) discuss Lower Eocene endocarps of this family.

Larvae of the large noctuid moths of the subfamily Catocalinae use Menispermaceae as their major food source throughout the topics, although they can also be found on other plants like Erythrina (some Menispermeae have pentacyclic Erythrina-type alkaloids). The adult moths, with their saw-like proboscides, attack ripe or ripening fruits and cause a considerable amount of damage (Fay 1996).

The muscle relaxant D-tubocuranine is obtained from Chondrodendrum tomentosum, a major ingredient of the South American poison curare, put on arrows and darts.

Tinomiscium is strongly supported as sister to all other Menispermaceae, within which the [Tinosporeae + Coscinieae] clade has only moderate to weak bootstrap support. A large clade corresponding to the rest of the family is well supported, within which Menispermum is sister to other taxa, again with strong support, and there are other well supported relationships (Ortiz et al. 2007: cf. Jacques et al. 2007, rather different relationships but morphological data only, variously treated). The old Menispermeae, Fibraureae and Peniantheae are polyphyletic (see also Wang et al. 2007a). There is considerable variation in pollen morphology in the family (Harley & Ferguson 1982 and references) which needs to be integrated with the clades that are becoming evident. Hong et al. (2001) suggest phylogenetic relationships within Menispermeae.

There are few records of cork position. In at least some taxa, the presence of laticifers or sclereids is mutually exclusive (Wilkinson 1986). Cocculus has plagiotropic branches (Keller 1996); does it also have 2-ranked leaves? Flowers can be monosymmetric, as in the carpellate flowers of Stephania dielsiana, where there are 1 + 2 or 1 + 3 sepals and petals and a single carpel (Wang et al. 2006); note that the staminate flowers remain polysymmetric. Tepals in e.g. Menispermum canadense have only a single trace (Smith 1928). The upper of the two ovules is epitropous and fertile, the lower is apotropous (Mauritzon 1936). Does Tinomiscium have a condyle (see Jacques 2006 - no)? There is apparently a period of 6-8 weeks between fertilization and first division of the zygote in Tinospora cordifolia (Sastri 1964).

Some general information is taken from Kessler (1993), Harley (1985 and references) surveyed pollen morphology, Wilkinson (1986) described leaf anatomy, Réaubourg (1906) provided a morphological and anatomical study of the family, Hegnauer (1969, 1990) summarised information on the chemistry, and Jacques and de Franceschi (2007), wood anatomy; see also Jacques (2006: much useful information, inc. phylogeny).

Synonymy: Pseliaceae Rafinesque

Berberidaceae + Ranunculaceae: rhizomatous herbs (woody) [roots or stem bright yellow]; nodes also multilacunar; vascular bundles V-shaped, in herbaceous taxa often closed, scattered or in concentric rings; leaf base broad, (lateral petiolar stipules +); ± petaloid staminodial nectaries +, nucellar cap +, outer integument at least 4 cells thick; seed exotestal; endosperm reserves other than oil or protein.

Nowicke and Skvarla (1981) suggested that aperture columellae might be a synapomorphy for the two families.

BERBERIDACEAE Jussieu, nom. cons.   Back to Ranunculales

Myricetin, isoprenylated flavonoids +, tanniniferous; cork also pericyclic; hairs 0 (unicellular or -seriate), lamina curved or conduplicate (complex in Podophyllum, etc.), margins gland- or spiny-toothed (entire), (2ndary venation pinnate), stipules common; inflorescence terminal (axillary), often racemose; flowers (2-)3(-5)-merous, parts whorled, with cortical vascular system, P = "K" + 6 nectariferous "C", A 6, with flaps, G 1, ascidiate, postgenital occlusion by secretion, outer integument 5-11 cells thick, micropyle bistomal [zig-zag], antipodals endopolypolypoid, stigma broad, dry or wet; exotesta lignified, with oblong-fibrous (cuboid) cells; endosperm with hemicellulose; embryo minute.

14/701 [list] - three clades below. Mostly East Asia and E. North America, also South America, N. Africa, general N. temperate (Map: from Ahrendt 1961; Hong 1993; Fl. N. Am. III 1997; Malyschev & Peschkova 2004).[Photos - Collection]

1. Nandina, Caulophyllum, etc. [Nandinoideae Heintze]

(Woody); petiole concave at the base; lowermost branch of inflorescence from axil of ± expanded leaf; (P many, nectary 0 - Nandina), (A dehiscence by slits), G with 1-2 (four) ovules; fruit a berry or pericarp evanescent or bladder-like; funicle swollen, (seed endotegmic - Nandina); n = 8, 10.

4/15. E. Europe to Japan.

Synonymy: Nandinaceae Horaninow

2. Berberis, Razania. [Berberidoideae Kosteletzky]

(Plant woody;) (vessel elements with scalariform perforations, petiole bundles arcuate - Berberis); leaves odd-pinnate or simple [unifoliolate]; lowermost branch of inflorescence from axil of reduced leaf; K 3-12, "C"/staminodia with paired basal nectaries, sensitive, tapetum plasmodial, pollen 6-12 colpate, or apertures irregular (spiraperturate), wall undifferentiated, 1-many ovules/carpel; fruit a berry; embryo long; n = 7.

2/601: Berberis (600). General N. temperate, also South America, N. Africa.

3. Jeffersonia, Diphylleia, Epimedium, etc.

Lowermost branch of inflorescence from axil of reduced leaf; K (0 - Achlys), 4-18, (C 0; 4, with nectar spurs; 7-9), A (-19, Podophyllum, Achlys; dehiscence by slits), pollen striate (spiny; diads; tetrads), 1-many ovules/carpel; fruit an achene, berry, or a follicle also with transverse dehiscence; n = 6.

8/75: Epimedium (55). Mostly (Europe to) East Asia (some desert xerophytes) and W. or E. North America. [Photo - Podophyllum Flower © R. Kowal, Fruit, Ripe Fruit.]

Synonymy: Diphylleiaceae Schultz-Schultzenstein, Epimediaceae Menge, Leonticaceae Berchtold & J. C. Presl, Podophyllaceae Candolle, nom. cons., Razaniaceae Takhtajan

• Berberidaceae are herbs or shrubs that may be recognised by their often yellow-colored secondary tissue (most obvious in the roots) and their often compound, stipulate leaves with palmate venation and sharply toothed or spiny margins. Their flowers are usually 3-merous, the perianth is often multiseriate, the stamens are opposite the perianth, and anther dehiscence is valvate; there is always a single carpel.

Several genera in Berberidaceae are disjunct and/or occupy only limited areas; Berberis is by far the most widely distributed genus. Many of the taxa involved in these disjunctions may have originated in East Asia and despite the age of the family - probably late Cretaceous - the dijunctions may be relatively recent, within the last 10 million years, although the desert xerophytes Bongardia and Leontice are probably rather older (Donoghue & Smith 2004; Wang et al. 2007b). Berberis is the alternate host for Puccinia graminis, the black stem rust of wheat.

In Berberis s. str. leaves on long shoots are mostly modified as spines; there are usually simple but articulated photosynthetic leaves (i.e. they are unifoliate) on short shoots. Mahonia s. str. (with ca 100 species) has compound leaves, but the two hybridise. Epimedium has four nectar spurs coming from the four inner tepals. In genera like Caulophyllum the carpel walls do not surround the maturing blue seeds, so the plant is a kind of gymnosperm...

In Podophyllum the epidermal waxes are solid rods. Successive microsporogenesis has been reported (Min et al. 1995). The carpels in Berberidaceae vary in their orientation. According to Chapman (1925, cf. e.g. Feng & Lu 1998), the gynoecium is derived from two or three carpels, with the gynoecia of the n = 6 clade alone being derived from two carpels (Kim & Jansen 1998), however, the gynoecium is probably unicarpellate throughout the family. The seeds of several genera (but not in the Berberis/Razania clade) are variously described as having elaisomes or being arillate - not necessarily different conditions.

Nandina is a very distinctive plant, and in the past it has been segregated as a separate family or subfamily (as in versions 8 and earlier of this site). However, Nickol (1995) had suggested on morphological grounds that it was close to Caulophyllum (although sister to the rest of the family in the most parsimonious tree). Early molecular studies (e.g. Adachi 1995) found similar relationships, and these have since been confirmed, as by Kim et al. (2004), although Nandina does tend to wander (e.g. Kim & Jansen 1996, 1998). The three groupings above, which more or less form a polychotomy, come from Kim et al. (2004); the third clade has only moderate support (see also Wang et al. 2007b).

For Nandina: P with ca 3 traces, K many, spiral, nectaries absent, [C and A develop from the splitting of a single primordium], pollen with massive endexine, nectary 0, nucellus early absorbed, endothelium ?+; fruit a berry, seed concave; endotestal cells crystalliferous, but otherwise testa crushed, endotegmic cells enlarged, lignified, thickened esp. internally, crystalliferous. For its floral development, see Feng and Lu (1998); the flower is basically trimerous-whorled, sepals are borne in slightly-spiralling lines up the receptacle, and all perianth parts have similar vasculature.

Fruit dehiscence in at least some Berberidaceae and Papaveraceae is transverse. Although it has been suggested that on this account these families are similar (e.g. Endress 1995), there is little other evidence indicating immediate phylogenetic relationships.

Some general information is taken from Loconte (1993) and chemistry from Hegnauer (1964, 1989); see Nowicke and Skarvla (1981) for pollen. Stearn (2002) provides much information on herbaceous Berberidaceae. For floral development of Caulophyllum (with common stamen-nectary primordia), see Brett and Posluszny (1982).

RANUNCULACEAE Jussieu, nom. cons.   Back to Ranunculales

Tannin 0, little oxalate accumulation; cork deep-seated, rarely developed; when woody with broad primary rays persisting and cambium developing in the primary vascular bundles; vessel elements with simple and/or scalariform perforations; (nodes 1:1, 2:2); (cuticle waxes as platelets); stomata also paracytic; leaves (opposite, 2-ranked), quite often simple, ptyxis variable, margins usu. gland-toothed (stipules - Thalictrum, Caltha, etc.); (plant dioecious), inflorescence terminal, often cymose, or flowers single; flower often large (vertically monosymmetric), parts spiral or whorled, K, "C", and A not opposite each other, K (2-)5(-6), often petaloid, three-trace, A many, spiral, extrose or introrse, (pollen multiporate; inaperturate), G (1-)many, usually with complete postgenital fusion, when 3, orientation variable, several apotropous ovules/carpel, micropyle endostomal, antipodal cells persist, multiply or not, stigma ± dry; fruit a follicle; exotestal cells often thickened, unlignified, or seed ± pachychalazal, coat thin; endosperm starchy (0), embryo minute to long, cotyledons connate or not, cotyledonary tube common; chromosomes small, straight, stout; germination epigeal.

62[list]/2525 - five subfamilies below. ± World-wide, but esp. (N.) temperate, not lowland tropics (Map: from Vester 1940; Hultén 1971; Wilson 2007). [Photo - Flower]

Hydrastidoideae + Glaucidioideae: vessel elements also with scalariform perforation plates; medullary bundles +; vascular bundles flat; petiole bundles annular and medullary; palisade mesophyll 0; leaves 2-ranked; flowers single, terminal, nectaries 0, outer integument 4-13 and inner integument 2-5 cells across, antipodals unmodified, stigma bilobed; follicle dehiscing abaxially as well.

The rhizome is an irregular sympodium. Soltis et al. (2003a) suggest that both Glaucidium and Hydrastis have a bimerous perianth.

1. Hydrastidoideae Martynov

Roots bright yellow; nodes on erect stem swollen, multilacunar; lamina plicate, petiole base on rhizome encircling stem; P (2)3(4), with a single trace, pollen tectum striate-reticulate, G several, 1-4 ovules/carpel, micropyle zig-zag, stigma with multicellular projections; follicle rather fleshy; exotesta strongly palisade, exotegmen lignified, both multiplicative; embryo minute; n = 13.

1/1: Hydrastis canadense. C. and E. North America (Map: from Li 1952, red).

There are cortical bundles in the erect stem, but not in the rhizome.

For much information, see Tobe and Keating (1985).

Synonymy: Hydrastidaceae Martynov

2. Glaucidioideae Loconte

Coumarin +, alkaloids, berberin 0; lamina supervolute-curved and plicate; flowers with cortical vascular system, P 4, A centrifugal, G 2, basally connate, opposite outer P [transverse], plicate, many tenuinucellate ovules/carpel, archesporium multicellular, nucellar cap massive; fruit with stigma on lower abaxial surface; seeds winged, outer integument vascularised; polyembryony common, embryo long, cotyledons foliaceous; n = 10, chromosomes 1.5-2.5 µm long.

1/1: Glaucidium palmatum. Japan (Map: from Li 1952, blue).

The adaxial side of the carpels expands more than the abaxial during the development of the fruit, so the stigma ends up on the "lower" surface. The embryo is shown as being long by Tamura (1972) and Takhtajan (1988), but it is described as being minute by Takhtajan (1997).

See Tamura (1972) for much information, embryological data are taken mostly from Tobe (1981).

Synonymy: Glaucidiaceae Tamura

Coptoideae [Thalictroideae + Ranunculoideae]: xylem surrounding phloem (cf. Takhtajan 1997); petiole bundles with associated lignification; P 5-merous, "C" 0-13, usu. obviously nectariferous, very diverse in form, with a single trace, antipodals modified.

3. Coptoideae Tamura

Woody or perennial herb; nectaries 5-10, petaloid, thick, stalked, carpels stipitate; n = (8) 9, small, rod-like.

3/17. East Asia, E. and W. North America.

Thalictroideae + Ranunculoideae: ovules apotropous [when single]; (fruit an achene).

4. Thalictroideae Rafinesque

(Cyanogenic compounds +); hairs capitate; lamina segments ± curved-involute, papillate; nectaries & petaloid and stalked, integument single, 7-8 cells across; n = 7, small, bean-shaped.

9/450: Thalictrum (330: wind pollinated), Aquilegia (80: little molecular divergence). N. temperate, also South America, Africa and New Guinea.

Tucker and Hodges (2005) discuss floral development in Aquilegia and its immediate relatives, Wang and Chen (2007) discuss phylogenetic relationships and petal evolution in the whole subfamily.

Synonymy: Aquilegiaceae Lilja, Thalictraceae Rafinesque

5. Ranunculoideae Arnott

Lactone-forming glycosides + [ranunculin], benzylisoquinoline alkaloids few or usu. 0, berberin 0; stomata >35 µm long; (petiole bundles arcuate; wing bundles +; medullary bundles +); hairs clavate; lamina segments ± involute (supervolute and/or curved); (1 unitegmic [integument 6-9 cells across] tenuinucellate ovule/carpel); (fruit also berrylet - some Actaea; testa vascularised); endosperm nuclear; n = 8, R-type chromosomes [Ranunculus type - large, long, 2-armed, often curved]; (germination hypogeal - some Clematis).

46/2025: Ranunculus (600: for relationships, see Hörandl et al. 2005 and Paun et al. (2005), perhaps to include Myosurus, etc.), Delphinium s.l. (365), Aconitum (300), Clematis (325), Anemone s.l. (190: Hoot & Palmer 1994; Schuettpelz et al. 2002 for relationships). Actaea includes Cimicifuga (Compton et al. 1998). Worldwide, but not tropical.

For the intimate association between Trollius and its pollinators/seed parasites, the fly Chiastochaeta, see Pellmyr (1992).

Clematis has opposite leaves, sensitive petioles, and secondary woodiness, Myosurus develops a much-elongated receptacle (cf. Saururus!). Laccopetalum (= Ranunculus s.l. - see Hoot et al. 2008) has flowers with about 10,000 carpels. Nigella is weakly syncarpous. In Ranunculus there are small nectaries at base of the "petals", while in Anemone the bracts tend to be calycine.

Synonymy: Actaeaceae Rafinesque, Anemonaceae Vest, Calthaceae Martynov, Cimicifugaceae (Arnott) Bromhead, Clematidaceae Martynov, Delphiniaceae Tamura, Helleboraceae Vest, Nigellaceae J. Agardh

• Ranunculaceae can be recognised by their usually herbaceous habit and spiral, usually exstipulate, lobed leaves with broad bases. The flowers have large and often petaloid sepals, nectariferous "petals", spiral, usually numerous stamens, and separate carpels borne on a well-developed receptacle.

Ranunculaceae are hosts to over 110 species of dipteran agromyzid leaf miners (Spencer 1990), which for the size of the family may be the most diverse assemblage in flowering plants; there are no agromyzids on Paeoniaceae!

Benzylisoquinoline alkaloids are largely absent from Ranunculaceae, although present in Coptis and Isopyyrum (Jensen 1995), which makes placing this feature on the tree difficult (lost and regained versus two losses). Ruijgrok (1966) clarified the distribution of the lactone ranunculin and of cyanogenic compounds. Clematis, secondarily woody, has storied wood (see Carlquist 1995a for wood and bark anatomy). There is extensive variation in petiole anatomy (Tamura 1995). The petaloid/nectariferous structures are usually interpreted as being derived from stamens, with which they have a number of points of similarity (Tamura 1965; Kosuge & Tamura 1989; Erbar et al. 1999; Leins 2000; cf. Kosuge 1994), and the androecium as being secondarily spiral and multistaminate. For an earlier discussion on such matters, and a suggestion that the flower of Ranunculaceae might be fundamentally 3-merous, see Salisbury (1919). Caltha has nectariferous hairs on the carpels. Although the carpels of Nigella are connate, no compitum is developed (Erbar 1998). There are often five traces to each carpel. When there is only one ovule/carpel, it is the basal member of the series (cf. Rosaceae, with which Ranunculaceae share superficial similarity, but where the ovule is the apical member). Uniovulate taxa are usually also unitegmic and with a nucellar cap (Philipson 1974). There is extensive variation in seedling morphology, and the development of a cotyledonary tube is quite common; in a few species, e.g. Ranunculus ficaria, there is only a single cotyledon (Förster 1997).

Ranunculaceae are a classic example of a "famille par enchaînement", nothing in particular holding them together, but recent work suggests that it is largely monophyletic (Hoot 1991, 1995; Jensen et al. 1995). However, Paeonia, frequently associated with Ranunculaceae in the past, is now included in Saxifragales as Paeoniaceae. Glaucidium has quite often been placed in its own family, but this would make Ranunculaceae paraphyletic; Tamura (1996) describes its androecial development as being centrifugal and the androecium as being innervated by branches of staminal trunk bundles.

Hydrastis + Glaucidium are sister to the rest of the family (Hoot et al. 1998). This and other major phylogenetic structure within the family - [Hydrastidoideae + Glaucidioideae] [Coptoideae [Thalictroideae + Ranunculoideae]] - seems quite well established (cf. also in part Ro et al. 1997).

For additional information see Kumazawa (1937: vernation), Tamura (1962: petiole anatomy, 1965: flower, etc., 1993: a summary of the cytology, 1995: general account, including infrageneric groupings), Hegnauer (1969, 1990: chemistry), Tobe and Keating (1985: Hydrastis), Hegnauer (1986: chemistry), Weberling (1989: nectaries), Engell (1995: considerable variation in embryo and suspensor morphology), Endress (1995: floral morphology), Johri et al. (1992: general), and Tobe (2002: Hydrastidaceae).