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 megaphyllous [determinancy evolved first, then ad/abaxial symmetry], 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, pollen germinating in less than 3 hours, siphonogamy, 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 , 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, nucellus at apex of ovule 1-3 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, cavity not lined by distinct epidermal layer, stigma ± decurrent, dry [not 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, 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... For other features such a 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), where on the tree a thicker nucellus and a stylar epidermal layer are acquired has not yet been indicated.

NYMPHAEALES [AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]: vessels + [one position], elements with elongated 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 +; inflorescence?; C tube initiation early; G [2-3], inferior.

Evolution. Magallón and Castillo (2009) offer estimates of ca 94 and 9.6 million years for relaxed and constrained penalized likelihood datings for the crown group of this clade, i.e. divergence of Asteraceae, the stem group being ca 99 and 99.7 million years old (relaxed and constrained estimates) - but note topology.

Chemistry, Morphology, etc. Polyacetylenes are sporadic, occuring in Asterales, Dipsacales and Apiales, but always in much embedded clades. The position of early initiation of the corolla tube on the tree is quite uncertain. Although a number of families in this clade have members with such initiation, not only is sampling within the larger orders poor and the smaller orders largely non-existent, while in the asterid I group both Oleaceae and Rubiaceae, "basal" or almost so in their orders, have early initiation. Corolla initiation in Garryales and associated families is unknown (Leins & Erbar 2003b for a summary), and there may anyway be a connection between earely corllla tube formation and an inferior ovary (Ronse Decraene & Smets 2000); although the character "ovary inferior" can be placed at this level on the tree, it then must show frequent reversion to "ovary superior", and whether corolla development changes accordingly is of some interest. The I copy of the RPB2 gene is lost in this clade (Oxelman et al. 2004; Luo et al. 2007), but it occurs both in Escalloniaceae and Apiales. Several unplaced taxa, largely erstwhile Saxifragaceae s.l., now in Escalloniales, Paracryphiales and Bruniales (Lundberg 2001e) have not been sampled for this gene (but see Escallonia).

Previous Relationships. There are some similarities between [Pittosporaceae [Araliaceae [Myodocarpaceae + Apiaceae]]] and some Asterales (esp. Campanulaceae s.l. and Asteraceae) (e.g. Erbar & Leins 2004; Leins & Erbar 2004b). However, given the rather strongly supported set of relationships at the base of Apiales, the other clades that are in this part of the asterid II phylogeny, and relationships within Asterales (Campanulaceae s.l. and Asteraceae are not immediately related), these similarities will probably be plesiomorphies or more likely parallelisms.

ASTERALES Lindley  Main Tree, Synapomorphies.

(Route I secoiridoids, oligo- or polyfructosans, inc. inulin, with isokestose linkages [starch generally 0] +); apotracheal parenchyma 0; nodes 3:3; leaves spiral; flower size?, C valvate, petal apiculi inflexed; A (basifixed), free from C, pollen grains trinucleate, nectary disc +, many ovules/carpel, integument <7 cells thick, endothelium +, antipodal cells ephemeral, style long; micropylar and chalazal endosperm haustoria +; mitochondrial rpl2 gene lost. - 11 families, 1649 genera, 25790 species.

Evolution. Asterales contain ca 13.6% eudicot diversity (Magallón et al. 1999). Fossils assignable to the order are known from the Oligocene, ca 29 million years before present (but these are only from the Menyanthaceae-Asteraceae clade); Wikström et al. (2003) suggest an age of 96-93 million years before present for the crown group, Bremer et al. (2004) 90-82 million years before present, Janssens et al. (2009) date stem group Asterales to 104±12.1 million years ago and the crown group to 94±11.2 million years, and Magallón and Castillo (2009) offer estimates of ca 201 and 128 and 177 and 125 million years for relaxed and constrained penalized likelihood datings for stem and crown group Canellales respectively, the stem group being 123 to 111 million years old (relaxed and constrained estimates) - but note topology.

Chemistry, Morphology, etc. Unfortunately, the condition of corolla and endosperm development, and endothelium presence, not to mention chemistry (for a partial summary, see Grayer et al. 1999), etc., is unknown in some critical families, so understanding character evolution is particularly difficult. Absence of apotracheal parenchyma and x = 9 may also be features of Asterales (Lundberg & Bremer 2001; Bremer et al. 2001). For a study of petal vasculature, which shows interesting variation, see Gustafsson (1995). Tobe and Morin (1996) summarize embryological knowledge of the order (they also include Sphenocleaceae, for which, see Solanales here). Monosymmetry is often associated with a slit the length of the corolla. Several families have forms of secondary pollen presentation (Erbar & Leins 1995a; Leins 2000; Leins & Erbar 1997, Erbar 2003b: Yeo 1993 for a general summary), and Leins (2000) and Leins and Erbar (2006) in particular discuss in considerable detail the evolution of secondary pollination presentation mechanisms. Variation of ovary position in Asterales is considerable. For integument thickness, see Inoue and Tobe (1999) and for pollen, see Polevova (2006). For a general discussion of variation in the order, see J. Kadereit (2006).

Phylogeny. There is quite a lot of phylogenetic structure in Asterales, but with rather weak support, as was suggested by D. Soltis et al. (2000; see also Olmstead et al. (2000). More recent studies have improved support for many clades, although there is still a basal polychotomy (Kårehed et al. 2000; Lundberg 2001a, b; Kårehed 2002a; especially Bremer et al. 2001 and Lundberg & Bremer 2001, 2003), as is shown in the tree here. Note that Olmstead et al. (2000) and B. Bremer et al. (2002) suggest a sister group relationship between Campanulaceae and Stylidiaceae (but not Donatia, which goes elsewhere!), and the latter authors suggest that Pentaphragmataceae are also linked to this clade, although Donatia (also Stylidiaceae - see below) is there very weakly linked with Alseuosmiaceae et al. Stylidiaceae and Donatiaceae are associated as weakly supported sister taxa (D. Soltis et al. 2000) or quite strongly linked (Kårehed et al. 2000; Lundberg 2001); in some studies Donatiaceae are sister to Abrophyllum (Carpodetaceae: Gustafsson et al. 1997), not to Stylidiaceae. There are suggestions that Rousseaceae, Pentaphragmataceae and Campanulaceae are together sister to the other Asterales (Lundberg & Bremer 2003), although the support for this is not very strong, while Soltis et al. (2007a) found Campanulaceae to be sister to rest of Asterales (1.0 p.p.). However, Rousseaceae seem to be sister to Campanulaceae (Kårehed 2002a; Tank et al. 2009). Relationships within some of the clades, e.g. Asteraceae and its immediate relatives, also vary somewhat according to the gene studied (A.P.G. 2003 for references).

Previous Relationships. Asterales are basically Takhtajan's (1997) Asteridae, with the addition of sundry Hydrangeales. Cronquist (1981) also included many families here included in Asterales in the orders placed towards the end of his Asteridae, although some families were also included in Cornales (Rosidae), etc.

Includes Alseuosmiaceae, Argophyllaceae, Asteraceae, Calyceraceae, Campanulaceae, Goodeniaceae, Menyanthaceae, Pentaphragmataceae, Phellinaceae, Rousseaceae, Stylidiaceae.

Synonymy: Alseuosmiales Doweld, Ambrosiales Dumortier, Anthemidales Link, Boopidales Berchtold & J. Presl, Brunoniales Lindley, Calendulales Link, Calycerales Reveal, Campanulales H. G. L. Reichenbach, Carduales Small, Cichoriales Link, Echinopales Link, Goodeniales Lindley, Menyanthales Takhtajan, Pentaphragmatales Doweld, Phellinales Doweld, Rousseales Doweld, Scaevolales R. Brown, Stylidiales Reveal - Asteranae Takhtajan, Campanulanae Reveal, Phellinanae Doweld - Asteridae Takhtajan - Asteropsida Brongniart, Campanulopsida Bartling

Rousseaceae + Campanulaceae: flowers large; A free.

ROUSSEACEAE Candolle   Back to Asterales

Young stem with separate bundles; leaf margins gland-toothed; G [5], mostly superior, opposite petals, integument 5-8 cells across.

4[list]/13 - two subfamilies below. Mauritius, scattered from New Guinea to New Zealand.

1. Rousseaoideae

Evergreen climber to small tree; chemistry?, tannins 0; cork?; resin canals +; petiole bundle cylindrical; bud scales +; hairs tufted-stellate and glandular-peltate; leaves opposite, base broad; flowers single, terminal, (4-merous), K valvate, C connate, anthers basifixed, attached their entire length to stout connective, sagittate, extrorse, pollen 6- or 8-porate, tectum complete; G [5-7], style expanding apically, stigmatic lobes narrower, erect; fruit a berry, K persistent; seed coat with thick-walled exotesta, the rest crushed; micropylar haustorium +, embryo long; n = ?

1/1: Roussea simplex. Mauritius (see Map above: green). [Photo - Flower © D. Lorence]

2. CarpodetoideaeJ. Lundberg

Trees; chemistry?; petiole bundles arcuate or annular plus accessories; hairs unicellular, thick-walled, strongly curved, warty; inflorescence paniculate, flowers small, 4-6-merous, C becoming free; A attachment?, (pollen in tetrads - Carpodetus); G [3-6], style short (0), stigma capitate (± divided - Cuttsia); fruit dry, baccate, or a loculicidal and septicidal capsule, K deciduous; seeds many, funicle elongated, exotestal cells massively thickened on anticlinal and inner periclinal walls (all around - Carpodetus); endosperm hemicellulosic [Carpodetus], ?haustoria; embryo?; n = 14, 15.

3/12. New Zealand, E. Australia, Papuasia (see Map above: red). [Photo - Inflorescence]

Synonymy: Abrophyllaceae Nakai, Carpodetaceae Fenzl

• Rousseaoideae may be recognised by their whorled, serrate leaves, bud scales, and their single, terminal flower with large calyx that persists reflexed on the fruit, long sagittate anthers, and apparently superior ovary tapering gradually into a stout, persistent style.

Evolution. Roussea is pollinated by a gecko (the pollen is embedded in a slimy substance), which may also disperse its seeds (Hansen & Müller 2009.

Chemistry, Morphology, etc. Mauritzon (1933) suggested that Roussea might have bitegmic ovules. For a summary of what little is known about the plant, see Koontz et al. (2006).

Abrophyllum and Cuttsia both have clusters of small, unlignified cells in the mesophyll that look like little white raphide bundles (Hils 1985). For a useful summary, see Gustafsson (2006).

Details of vegetative anatomy of Carpodetoideae are taken from Hils (1985), of indumentum from Al-Shammary and Gornall (1994), of floral morphology from Tobe and Raven (1999), and of seed anatomy from Takhtajan (2000). For anatomy of Roussea, see Watari (1939) and Ramamonjiarisoa (1980). For general anatomy, see Gornall et al. (1998), for some general information, see Gustaffson and Bremer (1997).

Lundberg (2001a) suggested little in the way of similarities between Roussea and Carpodetoideae other than carpel number (which is variable) and similar-looking petals, however, Roussea in particular is poorly known. It has an endodermis in its petiole, and its seed is drawn as if it were carunculate (Engler 1930a).

Phylogeny. For the circumscription of the family, see Lundberg (2000a).

Previous Relationships. Relationships of Rousseaceae were previously of uncertain position. Thus Takhtajan (1997) placed them in Rosidae-Celastranae-Brexiales, and they have often been associated with Saxifragaceae s.l.; the history of the classification of Roussea is summarized by Lundberg (2001a). The anthers in particular are quite unlike those of ex-Saxifragaceae, especially Escalloniaceae (see Escalloniales here) that are otherwise somewhat similar to Rousseaceae. Carpodetaceae, another group often thought of as "woody saxifrages", were included in Hydrangeales by Takhtajan (1997). None of these genera was mentioned by Cronquist (1981).

CAMPANULACEAE Jussieu, nom. cons.   Back to Asterales

Annual or perennial herbs to shrubs and pachycaul rosette plants; inulin +, iridoids and tannins 0, little oxalate accumulation; cork also inner cortical; vascular cylinder +; (medullary vascular bundles +); vessel elements with simple perforation plates; nodes 1:1; articulated laticifers +; crystals acicular; petiole bundles incurved-arcuate; leaves (opposite), ptyxis variable, margins entire to toothed (lobed), hydathodes common; inflorescence terminal, racemose; flowers monosymmetric, (3-)5(-10)-merous, median K abaxial, C with early tube formation, connate, secondary pollen presentation + [flowers protandrous, A basifixed, hairs at the tip of the style, anthers at least initially close to stigma and connivent when dehiscing, style elongating subsequent to anther dehiscence], pollen also binucleate; G [2(-5)],(± superior; placentation parietal), style elongating after A dehiscence, stigma lobed; K persistent; seeds many, exotesta of lignified cuboid or fibriform cells, (endotestal cells, esp. inner walls, thickened); (endosperm starchy); expansion of chloroplast inverted repeat into small single copy region, 5bp ndhF deletion, chloroplast infA and accD genes lost [but see Haberle et al. 2008a], biparental plastid transmission [?levels], mitochondrial coxII.i3 intron 0.

84[list]/2380. World-wide - five subfamilies below.

Nemocladoideae + Campanuloideae: pollen spheroid to oblate-spheroid, verrucate or with spicules, style with long hairy part; fibrillar protein bodies in nuclei.

1. Nemacladoideae M. H. G. Gustafsson

Annuals (perennials); C polysymmetrical, 3:2 [inverted], or not inverted, A (adnate to C), filaments forming tube, basally and also anthers free, latter flip back after pollen release, (fimbriate scales abaxial to some filaments where they join the C), pollen tricolporate or 6-colpate; G also [3], half inferior, style head bending towards or away from median K; (fruit circumscissile - Parishella); n = 9.

3/15: Nemacladus (13). S.W. U. S. A., Mexico (map: from Wimmer 1968).

Synonymy: Nemacladaceae Nuttall

2. Campanuloideae Burnett

Perennials (annuals), roots often thick; caffeic acid, p-coumaric acid, polyacetylenes [14-C aliphatic tetrahydropyran derivatives] +; (vessel elements with scalariform perforation plates); inflorescence often ± cymose; flowers polysymmetric at maturity, median K adaxial, stamens often sprawling at bottom of corolla tube after pollen is shed although bases conceal nectar, echinate, G (1 [2) 3-5(-10)], opposite sepals (C), or median member adaxial, style hairs with bulbous bases, retractile, stigma dry; chalazal and micropylar endosperm haustoria; (fibrillar protein intranuclear inclusions); extensive rearrangements in the chloroplast inverted repeat.

50/1050. More or less world-wide, but very few in the Australia-New Zealand area (map: from Hultén 1971; Thulin 1975; Shulkina 1978).

1. Cyanantheae Meisner

Leaves often opposite; pollen colpate/colporate, ((A 3), [G 5] - Cyananthus); n = 7-9 (17).

4/60: Codonopsis (30), Cyananthus (23). Central Asia to West Malesia, few in Canaries and Africa, often (sub)tropical, not Europe or northern Asia.

2. Campanula, Wahlenbergia, etc.

Leaves often spiral; pollen porate; fruit also dehiscing through sides (apically), with pores or slits [caused by activity of axicorn on drying - Campanuleae, inc. Edraianthus]; n = 6+ [17 common]; epicotyl and hypocotyl usually not elongated.

46[notional]/975: Campanula (420), Wahlenbergia (260), Adenophora (65). Especially N. temperate Old World, very few in the Australia-New Zealand area (map: from Hultén 1971; Thulin 1975; Shulkina 1978).

Synonymy: Cyananthaceae J. Agardh, Jasionaceae Dumortier

Lobelioideae + Cyphocarpoideae + Cyphioideae: pollen prolate, stylar tip at base of opening anthers, hairs only at tip of style.

3. Lobelioideae Schönland

(Annuals) perennials, herbs to small trees; chelidonic acid, pyrrolizidine alkaloids +, p-coumaric acid, caffeic acid 0; leaves supervolute [Lobelia]; flowers resupinate by pedicel (hypanthium) torsion, (not); C (3:2), 2:3, 0:5 (split-monosymmetric), (spurred - Heterotoma); filaments connate at least apically, anthers connate; pollen reticulate-striate; style with brush hairs and pollen in pollen box, pump mechanism [Nudelspritze], stigma wet; (fruit a berry; a circumscissile capsule; dehiscing through sides,); n = (6-)7(-13).

29/1200: Lobelia (400+), Siphocampylus (230+), Centropogon (215), Burmeistera (100+), Cyanea (80). Almost world-wide, not Arctic and absent from the Near East and central Asia, largely tropical, especially common in the New World (map: see Wimmer 1943; Meusel & Jäger 1992; FloraBase 2007). [Photo - Flower] [Photo - Fruit]

Synonymy: Dortmannaceae Ruprecht, Lobeliaceae Bonpland, nom. cons.

4. Cyphocarpoideae Gustafsson

Annual to perennial herbs; leaves deeply lobed; C induplicate-valvate, 1:4, adaxial petal cucullate, with apical appendage; A epipetalous, free, ovary notably elongated; fruit dehiscing through sides; n = 9; nuclear inclusions fibrillar.

1/3. Chile.

Synonymy: Cyphocarpaceae Reveal & Hoogland

5. Cyphioideae Schönland

Perennial (twining) herbs with tuberous roots; C subpolysymmetrical, also 3:2; A basally connate, (anthers slightly coherent), pollen psilate; G semi-inferior, style bends away from median K but does not elongate after A dehiscence, style hairs with bulbous bases, pollen deposited in pollen box, stigma with a fluid-filled terminal cavity with a lateral (terminal) pore; fruit septi- and loculicidal [valves bifid]; n = 9.

1/65. Africa, esp. the south (map: from Thulin 1978, N. B., not known from Cape Verde Islands).

Sympetaly in Cyphioideae is early. For information on secondary pollen presentation, see Leins and Erbar (2003a, 2005).

Synonymy: Cyphiaceae A. de Candolle

• Campanulaceae are usually herbs that can be recognised by their white latex, often rather soft, spirally inserted leaves, and flowers with an inferior ovary and secondary pollen presentation.

• Polysymmetric campanulate flowers with sprawling dehisced anthers at the base inside distinguish Campanuloideae; pollen is held among hairs along the style, and these hairs later retract. There are often three carpels.

• Lobelioideae have monosymmetric flowers in which the corolla may be split to the base down one side; the anthers (and filaments) are connate and the former at least initially enclose the stigma. The style pushes up within the anther tube and exposes the pollen. There are two carpels.

• Nemacladoideae are often small herbs that have monosymmetric flowers in which the filaments are more or less connate and the anthers are free, flipping backwards after they have dehisced. There are two carpels.

• Cyphocarpoideae have monosymmetric flowers with a single adaxial lobe that has an apical appendage, the stamens are free from each other but adnate to the corolla, and the inferior ovary, with two carpels, is much elongated; dehiscence of the fruit is through the sides.

• Cyphioideae are herbs with tuberous roots that also have monosymmetric flowers in which the stamens are more or less completely free. There are two carpels.

Evolution. The pachycaul giant lobelias are derived from herbaceous ancestors (Knox et al. 1993), and giant lobelias from widely separated parts of the globe may be in the same immediate clade (Antonelli 2008), indeed, some South American taxa may be derived from within the African giant lobelia clade, although the relationships of the giant lobelias and the biogeographic implications of these relationships need more detailed study. Knox et al. (2006, no Nemocladoideae or Cyphocarpoideae included) suggested that [Cyphia + Lobelioideae] originated in southern Africa, dispersing quite widely, and with at least two returns to Africa. Givnish et al. (2006a, 2008b) note that the some 120 species of Hawaiian Lobelioideae appear to have evolved from a single woody ancestor a mere ca 13 million years ago; fleshy fruits have evolved more than once in this clade. This clade represents a major plant radiation on the islands, the species having a variety of growth habits, pollinators, fruit dispersers, etc. Within Campanuloideae there are also remarkable distribution patterms, although sampling needs to be improved; are Wahlenbergia linifolia (St Helena) and W. berteroi (Juan Fernandez sister taxa (Haberle et al. 2009)? Interestingly, Campanuloideae on Crete seem to be largely remnants of a flora that was on the island when it was originally isolated (Cellinese et al. 2009).

Diversification within Campanuloideae may have begun 37.4-23.5 million years ago (Roquet et al. 2009), a possible age for stem Campanuliodeae being ca 41 million years (Wikström et al. 2001). Subsequent diversification within the clade began substantially later, 26.3-15.8 million years ago, and its biogeographic history is complex and involves much movement, the area from the Balkans to western Asia being particularly critical in its diversification (Roquet et al. 2009 for details); there are over100 species of Campanula in Turkey alone.

For the evolution of the secondary pollen presentation devices in the family, see Leins and Erbar (especially 2003b, 2006); sampling (inc. Nemocladoideae, some Lobelioideae) is still incomplete. Associated with the secondary pollen presentation that occurs throughout the family, the flowers are protandrous. Leins and Erbar (2003b) note that the flowers in Campanulaceae s.l. are initially polysymmetrical in bud. Stein (1992) discusses extrafloral nectaries of Andean Centropogon that are found on the outside of the inferior ovary, although mostly in species growing at lower altitudes, where ants are also found; in species at higher altitudes such nectaries were rare and pollination was by sickle-bill hummingbirds (Eutoxeres: Heliconia [Heliconiaceae] is the nectar resource for Eutoxeres at lower altitudes). In high-altitude species of Burmeistera there is both bird and bat pollination (Muchhala 2006), while in Centropogon nigricans there seems to have been coevolution with a remarkably long-tongued bat, Anoura fistulata (Muchhala & Thomson 2009: cf. Angraecum - Orchidaceae).

The nectar of some Campanuloideae may be brightly colored and then the filament bases are not persistent; normally they are, and then they enclose the nectar.

Chemistry, Morphology, etc. Since the pedicel of Lobelia and its relatives is twisted (resupinate), the flowers appears to have a "normal" orientation with the median petal abaxial, however, this does not usually occur in Lysiopoma (= pseudo-resupinate - Ayers 1997). Nemacladus (Nemacladoideae) has groups of remarkable reflexed pseudonectaries at the bases of two filaments. It is not known if the style hairs there are retractile. In Ostrowskia (Campanuloideae) the anther has a placentoid and the integument is massive (Kamelina & Zhinkina 1998). Phyteuma has coherent corolla lobes although the corolla is open laterally; the style hairs are only partly retractile. Some species of Wahlenbergia have an almost superior ovary. The chloroplast gene accD (= ORF512, zpfA) has been lost (Doyle et al. 1995 and references) in at least some Campanulaceae.

For node-stem anatomy of Campanuloideae, see Col (1904), for fruit morphology, see Kolakovsky (1985), for rearrangements in the chloroplast inverted repeat, see Cosner et al. (1997), for morphology, see Shulkina et al. (2003). Further general information on the subfamilies was taken from Schönland (1889), Wimmer (1968) and Lammers (1998 and especially 2006); for pollen see Dunbar (1975a, b), for embryology, etc., see Subramanyam (1970: possible taxonomically interesting differences in cell number of the haustoria), for the protein bodies in the nuclei, see Bigazzi (1986) and Haberle (1998), for the inverted repeat and chloroplast genome rearrangement, see Knox and Palmer (1999: Cyphocarpus, Nemacladus, etc., were not studied, Cyphia was) and Haberle et al. (2008a), and for a world checklist and bibliography, see Lammers (2007).

Other variation that needs to be incorporated into this schema includes inflorescence type, whether basically determinate (Campanuloideae) or indeterminate (Lobelioideae). Also, details of the major variation in secondary metabolites within the clade need to be established.

Phylogeny. The major groupings above are only tentative. Both the monophyly and the relationships of the poorly known Cyphiaceae (there are three subgroups) have been unclear (Lammers 1992), although they are to be included within a monophyletic Campanulaceae s. l. (see Cosner et al. 1994; Gustafsson & Bremer 1995; Gustafsson 1996b; Gustafsson et al. 1996). ITS sequence data suggest that Cyphocarpus is a member of Lobelioideae (Haberle 1998, Ayers & Haberle 1999) - see also pollen (Dunbar 1975b), but if so, it has several characters in common (parallelisms?) with the Campanuloideae + Nemacladoideae clade. Furthermore, the tree presented by Haberle (1998) suggests the groupings [[Nemacladoideae + Campanuloideae] [Cyphioideae + Lobelioideae]], which, if supported by other data, may in turn suggest that the polysymmetric flower of Campanuloideae with the median sepal adaxial (the "normal" condition) is a reversal from a disymmetric flower with the median sepal abaxial. Lundberg and Bremer (2003) found what was basically a trichotomy of Cyphia, Lobelioideae, and Campanuloideae. Finally, Tank and Donoghue (2009) suggest that Nemacladoideae may be basal and paraphyletic, being sister to the rest of Campanulaceae...

General relationships within Campanuloideae are discussed by Eddie et al. (2002, 2003) and Cosner et al. (2004). The Campanula clade can be divided into two main clades centered on Campanula s. str. and Rapunculus, with Wahlenbergia and other genera interspersed in other clades; Platycodon and relatives are strongly supported as being sister to the whole lot. Clearly, neither Campanula nor Wahlenbergia are monophyletic (Haberle et al. 2008b, 2009; Cellinese et al. 2009; Roquet et al. 2008, 2009; Borsch et al. 2009).

Molecular data suggest that Lobelia is wildly paraphyletic (Knox & Muasya 2001; Antonelli 2008; Knox et al. 2008). Within South American lobelioids, Centropogon, for example, is paraphyletic, but there is little support for relationships in general, while the pachycaul giant lobelias are derived from herbaceous ancestors (Knox et al. 1993).

Taxonomy. A.P.G. II (2003) suggests as an option keeping Lobeliaceae separate from Campanulaceae, but the two are best combined in view of their substantial similarities (see A.P.G. III 2009) - and as a practical matter for the time being, given that details of the main pattern of relationships in the clade are unclear. Generic limits in general need much attention, with a much more broadly delimited Campanula being a reasonable solution to its extensive paraphyly; the segregate genera are based on floral features which are unreliable guides to broad relationships (Haberle et al. 2008b; Roquet et al. 2008). Current taxonomy is no better a reflection of relationships in Lobelioideae.

Previous Relationships. Takhtajan (1997) divided Campanulaceae s.l. into four families; these, plus Pentaphragmataceae and Sphenocleaceae (for the latter, see Solanales here), made up his Campanulanae.

PENTAPHRAGMATACEAE J. Agardh, nom. cons.   Back to Asterales

Rather fleshy herbs, rooting at base of stem; chemistry?; cork?; wood rayless; nodes ?; hairs with uniseriate branches; leaves usu. asymmetrical, two-ranked, margins ± serrate (entire); inflorescences cymose, usu. scorpioid; K petaloid, 2 large + 3 small, C ± deeply lobed (free), nectariferous cavities between septae joining hypanthium to G, C with marginal wings, stamens adnate to corolla, extrorse, basifixed; G [2-3], integument 3 cells across, embryo sac protruding from micropyle, style short, stigma capitate; fruit baccate, K and C persisting; seeds minute, exotestal cells cubes, inner walls lignified; micropylar haustorium only, endosperm starchy; n = 54-56.

1[list]/30. South East Asia to Malesia, esp. W. Malesia (map: from Airy Shaw 1954). [Photo - Flower]

• Pentaphragmataceae are distinctive plants. They are rather fleshy herbs with two-ranked, asymmetrical, usually serrate leaf blades and scorpioid cymes with sessile flowers. The flowers are radially symmetrical and have relatively large, conspicuous sepals, extrorse anthers and an inferior ovary; there are nectariferous pockets between the septae that join the hypanthium to the ovary. The fruit is baccate.

Chemistry, Morphology, etc. The family is very poorly known. The micropylar haustorium is single-celled and the embryo is about a third the length of the seed (Kapil & Vijayaraghavan 1965).

For wood anatomy, see Carlquist (1997b), for the flower, see Vogel (1998b), and for general information, see Lammers (2006).

[Alseuosmiaceae [Phellinaceae + Argophyllaceae]] Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae + Asteraceae: ?

Alseuosmiaceae [Phellinaceae + Argophyllaceae]: lamina serrate, gland-toothed; x = 8.

Phylogeny. There is a possible grouping [Alseuosmiaceae [Phellinaceae + Argophyllaceae]] or [Alseuosmiaceae [Stylidiaceae [Phellinaceae + Argophyllaceae]]] (e.g. Kårehed et al. 2000; Lundberg & Bremer 2001 respectively), and although jacknife support for the position of Alseuosmiaceae is not very strong, the posterior probability for the first grouping is 1.0 (Kårehed 2002a).

ALSEUOSMIACEAE Airy Shaw   Back to Asterales

± Woody; condensed and ellagitannins +, inulin?, iridoids 0; young stem with separate bundles; true tracheids +; starch-storing living fibers +; pericyclic fibers weakly developed; petiole bundle(s) arcuate; endodermis in both stem and leaf; hairs axillary; leaves conduplicate; flowers (4-)5(-7)-merous; (hypanthium present), K free, valvate, C margins with (fringed; erose) wings (hardly - Platyspermation), stamens adnate to corolla, anthers ± basifixed, pollen (in tetrads); G [2, 3], 2 or more ovules/carpel, stigma barely expanded; fruit baccate, calyx usually persistent; exotesta little thickened, lignified, mesotesta persistent; ?haustoria; n = 9 [Alseuosmia].

4[list]/10: Alseuosmia (5). New Guinea, E. Australia, New Zealand, New Caledonia (map: from van Balgooy 1993).

• Alseuosmiaceae may be recognised even vegetatively by their spiral, serrate leaves with axillary tufts of uniseriate hairs. Their flowers have an inferior ovary and usually a tubular corolla, and the corolla lobes have fringed or erose winged margins. The fruit is baccate and contains rather small seeds.

Evolution. Both Platyspermation and other Alseuosmiaceae on New Caledonia seem commonly to have galled fruits or flowers.

Chemistry, Morphology, etc. Ellagitannins are reported from Alseuosmia (Kårehed 2006 for references); this should be checked. Most Alseuosmiaceae have rayless wood, living mature fibres with stored starch, and stem with an endodermis. The pollen of Alseuosmia linariifolia is described as being tricolpate, with an ectexine made up of a thick, tubercular tectum and massive, spherical columellae (Polevova 2006); whether this can be generalised to the family is unclear (see also Kårehed 2006).

Some details of vegetative anatomy are taken from Dickison (1989a) and Gornall et al. (1998), and of testa anatomy from Nemirovich-Danchencko and Lobova (1998) and Takhtajan (2000). The embryology is poorly known. See Kårehed (2006) for general information.

Phylogeny. Platyspermation is strongly associated with other Alseuosmiaceae (Lundberg & Bremer 2001), and although its corolla is only shortly tubular, the lobes being rather spreading (buzz pollination?), in other respects it agrees well with the rest of the family. Uniseriate hairs in Platyspermation are not restricted to the leaf axils, although they are particularly dense there, rather, they cover the whole plant. Their persistent, reddish bases look rather like glands, hence perhaps the past inclusion of the genus in Rutaceae... The margins of the corolla lobes have narrow flanges and papillae. Generic limits are in some dispute (Tirel 1996).

Previous Relationships. Genera now included in Alseuosmiaceae have previously been placed in Caprifoliaceae, Rubiaceae, Rutaceae, Ericaceae, Epacridaceae, etc. Although the family is separated by Takhtajan (1997), it is included in Hydrangeales.

Synonmy: Platyspermataceae Doweld

Phellinaceae + Argophyllaceae: cork subepidermal; pollen (spiny) with rugulose exine, ovules apotropous, style short.

PHELLINACEAE Takhtajan   Back to Asterales

Trees; benzylisoquinoline [homoerythrina] alkaloids +, inulin, iridoids?; true tracheids +; rays very broad; sclerenchyma surrounding leaf veins; petiole bundles annular; cuticle waxes as platelets and rodlets; (lamina margin entire); plant dioecious, flowers small, 4-6-merous; K connate basally, ± open, C free, nectary 0; staminate flowers: pistillode +; carpellate flowers: staminodes +; G [2-5], 1 apical ovule/carpel, stigmas quite large; fruit a drupe, stones separate; testa ?; endosperm haustoria?; n = 17.

1[list]/12. New Caledonia.

Chemistry, Morphology, etc. The guard cells are huge, with inner and outer stomatal ledges. The ovules are reported as being hemitropous to campylotropous, but Phelline is embryologically poorly known.

See Baas (1975) for wood anatomy (it appeared to be extremely primitive), and Kårehed et al. (2000) and Barriera et al. (2006) for much additional information.

Previous Relationships. Takhtajan (1997) placed the family in Icacinales, describing the leaves as being mostly estipulate, while Cronquist (1981) placed it in his Aquifoliaceae (adjacent to Icacinaceae - both in Celastrales).

ARGOPHYLLACEAE Takhtajan   Back to Asterales

Shrubs; gallic acid +, inulin?; (nodes 1:1, 5:5); petiole bundles arcuate; hairs T-shaped, multicellular, with slits over the stalk cell; lamina supervolute-curved [Corokia macrocarpa], margins entire; flowers 4-5(-8)-merous, K valvate (always?), C basally connate, with adaxial fringed ligule (and marginal wings), G (1) [2, 3(-5)], (semisuperior), disc + [Corokia], 1 apical apotropous or many ovules/carpel, integument ca 6 cells thick, stigma punctate or lobed, wet; fruit a capsule [Argophyllum] or drupe [Corokia]; exotestal cells with inner walls massively thickened and lignified [Argophyllum] or all walls somewhat thickened [Corokia]; endosperm hemicellulosic, (embryo medium); n = 9.

2[list]/21: Argophyllum (15). S.W. Pacific, including Rapa (map: from van Steenis & van Balgooy 1966). [Photo - Corokia Flower © Gardenweek.org, Argophyllum.]

Chemistry, Morphology, etc. Septate fibers and vascular tracheids are present (Patel 1973; Noshiro & Baas 1998), but the significance of this is unclear. The guard cells in Argophyllaceae are raised above the epidermis (Kårehed et al. 2000). There are tannin cells in the flower both here and in Alseuosmiaceae. The pollen is like that of Cornaceae, with complex H-shaped endoapertures (Ferguson 1977).

See also Eyde (1966) and Kårehed (2006) for general information, Lobova (1997) for testa, and Gornall et al. (1998) for vegetative anatomy.

Previous Relationships. Along with Cornaceae, Argophyllaceae were placed in Hydrangeales by Takhtajan (1997), while Cronquist (1981) included them in his very heterogeneous Grossulariaceae.

Synonymy: Corokiaceae Takhtajan

STYLIDIACEAE R. Brown, nom. cons.   Back to Asterales

Inulin +; young stem with separate bundles; nodes 1:1; leaves entire, sessile; C imbricate, nectary +; A fewer than C, anthers extrorse, pollen colpate; micropylar and chalazal endosperm haustoria +.

6[list]/245 - 2 subfamilies below. Scattered in South East Asia to New Zealand, S. South America, mostly Australia.

1. Donatioideae B. Chandler

Dwarf cushion herbs; iridoids?, tanniniferous; cork cortical?; mucilage cells +; stomata also paracytic; hairs uniseriate, axillary; flowers solitary, terminal; K 3-7, free, C 5-10, free; A 2-3, free, pollen nuclei?; G [2-3], style with recurved branches, stigmas capitate; fruit indehiscent; seed coat?; n = 24.

1/2. New Zealand, Tasmania, S. South America. [Photo - Habit, Flower © Univ. of Tasmania.]

Synonymy: Donatiaceae B. Chandler, nom. cons.

2. Stylidioideae

Herbs (climbers), cushion plants; cork also outer cortical; vascular bundles closed, scattered or in a single ring; 2ndary thickening anomalous; cambium storied; vessel elements with simple perforation plates; hairs glandular; leaves pseudoverticillate or in rosettes, with axillary hairs; flowers with median K abaxial, but often semi-resupinate, split-monosymmetric, (polysymmetric, to 8-merous); K connate, C connate, 4 in two pairs, or 4 + labellum, early tube formation, often with coronal appendages, (spurred); nectary also paired glands; A 2, completely adnate to style [= gynostemium], anther thecae set end to end; pollen 2 or 3 nuclear, 3-8-colpate; G [2] (adaxial much reduced), stigma small, dry; (placentation free-basal), synergid cells elongated; fruit capsular, septicidal (indehiscent); seed exo-endotestal, exotestal cells sclerosed; embryo often with single cotyledon; n = 5-16, 18, protein bodies in nucleus.

5/240: Stylidium (220). Mostly Australia, but scattered in the central part, also South East Asia, Malesia, New Zealand, and S. South America (map: see Erickson 1958; FloraBase 2009). [Photo - Flower.]

• Stylidiaceae are usually rosette or tussock herbs with sessile or obscurely petiolate leaves. Stylidioideae have very distinctive semi-resupinate, split-monosymmetric flowers. The two stamens are usually adnate to the style, and the whole complex is often sensitive, moving when brushed by pollinator (hence the name, trigger plants). Donatioideae are small-leaved tussock plants that can be recognised by their solitary, polysymmetric, terminal flowers that have a variable number of free sepals and petals and two to three stamens with extrorse anthers.

Evolution. In Stylidioideae the two stamens are adnate to the style, the extrorse anthers being borne near the stigma, and the whole complex (a gynostemium) is often sensitive, moving when brushed by pollinator; for the literature on stylar movement, see Findlay and Findlay (1989). In Levenhookia, however, the gynostemium is held under tension in the hooded labellum, flipping only when the latter is disturbed. There are suggestions that Stylidium may be carnivorous, insects being trapped by the glandular hairs, which also show yeast-extract stimulated protease acivity; the plants grow in acid, nutrient-poor soil like other carnivores. However, uptake of nutrients by the plant from the insects has yet to be demonstrated (Darnowski et al. 2006).

Chemistry, Morphology, etc. In Stylidioideae the cambium may develop beneath the endodermis; xylem, and sometimes also phloem, is produced towards the inside, and at most cork to the outside (Carlquist 1981a). In older stems of Donatia the cortex is very thick, and the vascular tissue forms a narrow cylinder in the center (Chandler 1911). The fertile stamens are the adaxial pair; the pollen of at least some Stylidiaceae has a very distinctive inner ectexine that lacks columellae but is permeated by numerous sinuous channels (Polevova 2006). Monocotyly is reported to be quite common in Stylidioideae, at least (Carlquist 1981b).

For anatomical differences between Donatioideae and Stylidioideae, see Repson (1953). The proembryo in Donatia is ovate, the suspensor being short (probably a derived feature), but in Stylidioideae it is much longer than broad (Philipson & Philipson 1973), the suspensor being long, as in other Asterales (Tobe & Morin 1997). The leaves of Donatia are very small, and their venation is acrodromous.

Some anatomical details can be found in Thouvenin (1890); for general information, see Carolin (2006: Stylidiaceae and Donatiaceae), Carlquist and Lowrie (1989: Stylidioideae) and Glenny (2009: Forstera, for protein bodies, see Thaler (1966), for the testa anatomy of Stylidium, see Tobe and Morin (1996), and for floral development, see Erbar (1992).

Previous Relationships. Stylidiaceae have been treated as two families in Stylidiales (Takhtajan 1997) or merged in one family (Philipson & Philipson 1973). A.P.G. II suggests as an option keeping Donatiaceae and Stylidiaceae separate, although the two can reasonably be combined (e.g. Lundberg & Bremer 2003).

Menyanthaceae [Goodeniaceae [Calyceraceae + Asteraceae]]: inulin, caffeic acid +; vessel elements with simple perforation plates; C connate, early tube formation, with strong ± fused marginal [commissural] veins joining the median near the apex, stamens adnate to corolla, tapetal cells multinucleate, pollen spiny [not Barnadesia!]; G [2], integument >10 cells thick, vascular bundle proceeding to the micropyle; endosperm haustoria 0, embryo long.

Chemistry, Morphology, etc.The androecium has spiral initiation in some Menyanthaceae, Asteraceae and Goodeniaceae - and also Araliaceae (Erbar 1997). For other characters common to this clade, see Anderberg et al. (2006, but cf. vessel perforation plates). Vegetatively and florally Menyanthaceae are rather different from many other Asterales, however, both Menyanthaceae and Goodeniaceae have corolla lobes with marginal flanges, here placed as apomorphies for both groups (or they could be a synapomorphy for the whole clade, being lost later).

Phylogeny. Menyanthaceae do not link with the other three families in the four-gene study of Albach et al. (2001b).

MENYANTHACEAE Berchtold & J. Presl, nom. cons.   Back to Asterales

Usu. aquatic herbs; flavonols only +, little oxalate accumulation, tannin 0; cork?; vascular bundles separated; nodes also 5:5; branched sclereids +; petiole bundles arcuate to annular or scattered; leaves also two-ranked, involute, 2ndary veins palmate (palmately compound), margins usu. with hydathodal glands, crenate, colleters +, base broad, (with lateral flanges - Nephrophyllidium); flowers heterostylous (not); K basally connate, C lobes with marginal wings and/or with (marginal) fimbriae; ("staminodes" +, fringed and scale-like), anthers sagittate; tapetal cells with fused nuclei; pollen colpate; G [2], (style 0), stigma bilobed-spathulate, wet; placentation parietal; fruit a (septi- and) loculicidal capsule (berry - Liparophyllum); seeds many, (hairy; carunculate), exotestal cells with outer walls thickened, often with a variety of projections, (meso- and endotestal cells sclerotic); endosperm oily; n = 9 (17), protein bodies in nuclei; rpl2 intron missing.

5[list]/ca 58: Nymphoides (40). World-wide (map: from Hultén 1971; Heywood 1978). [Photo - Flower, Collection, Flower.]

• Menyanthaceae are aquatic plants with more or less orbicular or palmately-compound leaves with broad bases and quite large usually heterostylous polysymmetric flowers. The petals are connate, the corolla lobes have disinctive marginal wings or fringes and the ovary is superior.

Evolution. Has the superior ovary of Menyanthaceae with its parietal placentation been derived from a more of less inferior ovary witrh axile placentation (cf. Pittosporaceae - Apiales)?

Chemistry, Morphology, etc. Kasinathan and Kumari (2001) thought that the leaves of Nymphoides may be opposite. The vascular anatomy of the flower implies a basic monosymmetry - the lateral corolla traces are fused (Wood & Weaver 1982). There are "staminodes" on the corolla tube alternating with the stamens; there are also nectar glands at the base of the ovary. Johri et al. (1992) suggested that the endosperm stores starch.

For seed morphology, see Chuang and Ornduff (1992), for floral development, see Erbar (1997), for the missing rpl2 intron, see Downie et al. (1991b), and for general information, see G. Kadereit (2006).

Phylogeny. Relationships within Menyanthaceae have been clarified by Tippery et al. (2006, 2008) and Tippery and Les (2008); [Menyanthes + Nephrophyllidium] are sister to the rest of the family while Villarsia is very much paraphyletic.

Previous Relationships. The iridoids of Menyanthaceae differ chemically from those of Gentianaceae, in which Menyanthaceae used to be included, although placentation, etc., are similar. Menyanthaceae were placed in Solanales by Cronquist (1981), but inulin storage i.a. is inconsistent with that position. Branched sclereids and air canals are similarities between Menyanthaceae and Nymphaeaceae, but both are aquatics.

Goodeniaceae [Calyceraceae + Asteraceae]: secondary pollen presentation + [protandry, anthers connivent at dehiscence, the style elongates after pollen deposition], pollen with bifurcating columellae, stigma dry, ± papillate; K persistent in fruit; x = 8.

Chemistry, Morphology, etc. There is considerable variation in details of pollen presentation in this clade (Leins & Erbar 2003b, 2006 and references). DeVore et al. (2000) discuss variation in details of pollen wall morphology. Leins and Erbar (2003b) concluded that Goodeniaceae are probably sister to Asteraceae, noting i.a. that Barnadesia polyacantha has a bulge beneath the style branch, perhaps homologous with the stylar cup of Goodeniaceae. Interestingly, study of early capitulum development in Arnaldoa macbrideana (Asteraceae - Barnadesioideae) suggests that the capitulum there is built up of partial inflorescences with cymose branching, so perhaps linking the apparently racemose heads of Asteraceae with the apparently rather different inflorescences of many Calyceraceae and Goodeniaceae (Leins & Erbar 2003b, see their polytelic thyrses). Acicarpha is the only Calyceraceae with the possibly plesiomorphic n = 8, and it is also the only member of that family with an also possibly plesiomorphic condensed spicate inflorescence (DeVore 1994). For a comparison of the pollen of the three families, see DeVore et al. (2007).

Phylogeny. Soltis et al. (2007a) found the relationships [Asteraceae [Calyceraceae + Goodeniaceae]].

GOODENIACEAE R. Brown, nom. cons.   Back to Asterales

Herbs (woody, arborescent); O-methyl flavonols only, alkaloids, polyacetylenes +; cork subepidermal or cortical; (medullary vascular bundles +; cortical bundles = leaf traces); (vessel elements with scalariform perforation plates); nodes 1:1 (3:3, 5:5); branched sclereids +; indumentum variable, hairs often minutely warty; leaf margins entire to toothed; flowers split-monosymmetric (polysymmetric); C induplicate-valvate, (spurred; not slit - Brunonia), lobes with marginal wings (not Brunonia, Selliera); A free or adnate to base of C, basifixed, (anthers connate); pollen binucleate, mesocolpia concave; nectary usu. 0; G also [4] ([2]), (placentation ± basal), 1 or more ovules/carpel, style with apical hairy pollen-collecting indusium and stylar cup, stigma bilobed; ovule with hypostase; fruit dehiscing laterally, septicidal (and loculicidal) (drupe, nut); testa 7-14 cells thick, exotestal cells usu. palisade (crystalliferous), all walls (especially inner) thickened, (hypodermal layers lignified), (endosperm 0); n = 7-9; rpl16 intron missing.

12[list]/440: Goodenia (180), Scaevola (130), Dampiera (66). Largely Australian, Scaevola pantropical, with the coastal S. taccada in the E. and C. Pacific and Indian Oceans and S. plumieri in the W. Indian, E. Pacific and the Atlantic oceans (map: van Balgooy 1975). [Photo - Flower]

• Goodeniaceae are more or less herbaceous plants with spiral leaves and monosymmetric flowers. The corolla is often divided to the base adaxially, the lobes have marginal wings that may make each lobe look trilobed, and the style is curved, with an apical cup and (in older flowers) a bilobed stigma in the middle. The anthers dehisce in bud and the ovary is inferior.

Evolution. Some diversification in Scaevola may be associated with the aridification of the Nullarbor Plain some 14-13 million years ago separating eastern and western clades (Crisp & Cook 2007).

For the diversity of pollen presentation devices in the family, see Leins and Erbar (2003b, inc. Brunoniaceae). Most Goodeniaceae seem to be myrmecochorous (Lengyel et al. 2009).

Chemistry, Morphology, etc. The lateral veins of the corolla of Brunonia unite in the receptacle (Erbar 1997). The pollen is distinctive; endoapertures are bordered (with orae) and lalongate and the spines are rounded (Gustaffson et al. 1997). The mitochondrial genes cox1, atp1 and matR showed massive divergence (Barkman et al. 2007: Scaevola only sampled). See Carolin (1978, 2006) for general variation and morphology. I am grateful to Mats Gustafson for comments.

Phylogeny. Relationships within Goodeniaceae are becoming fairly well understood (Gustafsson 1996a; Gustafsson et al. 1996). There are two main clades, Leschenaultia and allies, and the other Scaevola, a paraphyletic Goodenia, and allies. Within the latter clade Brunonia is sister to the rest, and it differs from all other Goodeniaceae in its polysymmetric flowers, connate anthers, superior ovary, glabrous indusium, and lack of endosperm, and has a condensed inflorescence, two carpels each with a single, basal ovule, and a thin-walled, compressed testa. Some of these features might seem to suggest relationships with Asteraceae (Gustafsson 1996a), however, most are probably derived within Goodeniaceae (and connate anthers and glabrous indusium, at least, occur elsewhere in the family) and the exclusion of Brunonia would make that family paraphyletic.

Classification. For a general account of most of the family, see George (1992).

Synonymy: Brunoniaceae Dumortier, nom. cons., Scaevolaceae Lindley

Calyceraceae + Asteraceae: flowers small, C tubular, commissural veins connate, (median veins 0), filament collar +, 1 ovule/flower; fruit a cypsela, K persistent, modified, involved in dispersal.

Phylogeny These two clades may diverge 49-42 million years before present (K.-J. Kim et al. 2005). Both Calyceraceae and Asteraceae-Barnadesioideae, sister to the rest of Asteraceae, are South American.

Chemistry, Morphology, etc. For other similarities or possible synapomorphies, see DeVore (1994) and Lundberg and Bremer (2001, 2003), these include libriform fibers with simple pits and vasicentric parenchyma. Calyceraceae and Barnadesioideae have a similar simple flavonoid profile. Pesacreta et al. (1994) suggest similarities in the micromorphology of the filaments and connective bases between at least some members of Calyceraceae and Asteraceae. Placement of characters like pollen with intercolpar depressions on the tree is difficult to ascertain (see also DeVore 1994; DeVore et al. 2000); DeVore and Skvarla (2008) suggest that pollen characters thought to suggest a relationship between the two families are different in detail and are therefore not homologous. Although both families have but a single ovule, the position and orientation of the ovule is such that the single ovule condition may well have been derived independently.

Previous Relationships. Both Cronquist (1981) and Takhtajan (1997) placed the two families in separate, if adjacent, orders.

CALYCERACEAE Richard, nom. cons.   Back to Asterales

Herbs; ?flavonols 0; cork?; vascular bundles separated; nodes ?; pericyclic fibers 0; leaf margins entire; inflorescence an involucrate and bracteate strongly condensed capitulum made up of cymose units (Acicarpha, basically racemose with centripetal development of flowers); K connate, aerenchymatous or spine-like, C outer layer separates and photosynthesises, filaments ± connate, anthers basically free, pollen binucleate, with intercolpar depressions, glands [?nectaries] internally alternating with filaments (0), ovule apical, pendulous, apotropous, stigma minutely capitate, pollen deposited on its top; apex of fruit with a conical body [persistent base of C and style]; seed coat undistinguished; endosperm +; n = 8, 12, 13, 15, 17, 18, 20-22.

4[list]/60: Boopis (30). South America (map: from Heywood 1978 [S. part of range]; DeVore 1994). [Photos - Acicarpha Habit, Calycera © H. Wilson., Undetermined Flowers]

• Calyceraceae are herbs that may be recognised by their capitate inflorescences the flowers of which often open centripetally. The flowers are small and polysymmetric, the sepals are spines or are thick and aerenchymatous, the outer layer of the tubular corolla is photosynthetic, the stamens are free, and the ovary is inferior.

<Evolution. There is secondary pollen presentation of the pump mechanism type.

Chemistry, Morphology, etc. Cronquist (1981) described the flowers as being sometimes "slightly irregular"; they are commonly polysymmetric. The integument is described as being "thick" and the outer cell layers contain chloroplasts (Dahlgren 1915). Some general information is taken from Hansen (1992: especially useful), DeVore (1994), DeVore and Stuessy (1995) and Hellwig (2006), some details of morphology come from Pontiroli (1963) and of embryology from Dahlgren (1915: one species!).

Synonymy: Boopidaceae Cassini

ASTERACEAE Martynov, nom. cons.//COMPOSITAE Giseke, nom. cons. et nom. alt.   Back to Asterales

Herbs to trees or vines; iso/chlorogenic acid, isoflavonoids, sesquiterpene lactones, pentacyclic triterpene alcohols, terpenoid essential oils, various alkaloids, acetylenes [cyclic, aromatic, with vinyl end groups], a variety of fatty acids in the seeds +, tannins, iridoids 0; vascular bundles separated (a cylinder - woody taxa); (cork deep seated); (cortical or medullary vascular bundles +); cambium storied or not; (vessel elements with scalariform or reticulate perforations); nodes also 5:5; schizogenous secretory canals +; leaves also opposite, often conduplicate or revolute, margins various; inflorescence capitulate, involucrate, ebracteate; flowers poly- or variously monosymmetric, K reduced, C midveins 0 (+), anthers connate, with conspicuous apical and basal [the latter = calces, hence calcarate] appendages, caudate, endothecial cells elongated parallel to main axis of anther [?level], tapetum plasmodial (secretory), pollen 37-49.1 µm in diam., exine 6.1-6.7 µm across, tectum foraminate, ovule basal, erect, epitropous, embryo sac with elongated synergid cells and persistent multinucleate antipodal cells; (K deciduous); (testa not vascularized), exotestal cells thickened, palisade or flattened, or undistinguished; endosperm (nuclear), scanty to 0; protein bodies in nuclei; sporophytic incompatibility system present.

1620[list]/23,600 - eleven groups below. World-wide (map: Vester 1940; Hultén 1971). [Photo - Flowers, and more Flowers.]

1. Barnadesioideae Bremer & Jansen

Usually woody; notably poor in flavonoids, flavones 0; axillary thorns/spines common; esp. flowers with long tricellular hairs [even on the bristles of the achene]; corolla bilabiate [4 + 1], pollen (lophate, not spiny), with intercolpar depressions, style glabrous or papillate below bifurcation, stigma lobed; achene with spines, pappus?

9/94. South America, esp. Andean (map: see Karis et al. 1992; Ezcurra 2002). [Photo - Flower.]

Stifftioideae + Mutisioideae [Wunderlichioideae [Gochnatioideae [Hecastocleidoideae [Carduoideae [Pertyoideae [Gymnarrhenoideae [Cichorioideae [Corymbioideae + Asteroideae]]]]]]]]: often herbaceous; style with rigid sweeping hairs, style branches often long; achene with twin hairs [unicellular to uniseriate base, apical cell un/equally 2-armed], pappus capillary (scales, etc.); 22.8 kb chloroplast DNA inversion, and 3.3 kb inversion nested within it; n = 2-100+.

Evolution. The age of the stem of this clade is some 42-36 million years before present (K. J. Kim et al. 2005).

Chemistry, Morphology, etc. For the chloroplast DNA inversions, see Jansen and Palmer (1987), K.-J. Kim et al. (2005) and Timme et al. (2005).

2. Mutisioideae Lindley

Corolla bilabiate [2 + 3], (stigma lobes short); n = (6-)9(+).

44/630: Acourtia (65), Chaptalia (60), Mutisia (50), Trixis (50), Gerbera (35). South America.

Synonymy: Mutisiaceae Burnett, Nassauviaceae Burmeister, Perdiciaceae Link

3. Stifftioideae Panero

Ca 10/40. Venzuelan-Guianan (Andes, N.E. South America).

Wunderlichioideae [Gochnatioideae [Hecastocleidoideae [Carduoideae [Pertyoideae [Gymnarrhenoideae [Cichorioideae [Corymbioideae + Asteroideae]]]]]]]: ?

4. Wunderlichioideae Panero & Funk

Style and style arms glabrous; deletion in rpoB gene.

Ca 8/24. Venzuelan-Guianan (E. South America, S.W. China).

Gochnatioideae [Hecastocleidoideae [Carduoideae [Pertyoideae [Gymnarrhenoideae [Cichorioideae [Corymbioideae + Asteroideae]]]]]]: ?

5. Gochnatioideae Panero & Funk

Pollen anthemoid, style branches short (long), glabrous, apices rounded; pappus of bristles; n = 22, 23, 27.

4 (?5)/90: Gochnatia (70). Central and South America, esp. the Caribbean and southern South America.

Hecastocleidoideae [Carduoideae [Pertyoideae [Gymnarrhenoideae [Cichorioideae [Corymbioideae + Asteroideae]]]]]: deletion in rpoB gene.

6. Hecastocleidoideae Panero & Funk

Capitulae 1-flowered; flower polysymmetrical, corolla 5-lobed, style branches short, apices rounded; pappus of scales; n = 8.

1/1: Hecastocleis shockleyi. S.W. U.S.A.

Carduoideae [Pertyoideae [Gymnarrhenoideae [Cichorioideae [Corymbioideae + Asteroideae]]]]: carpels of disc flowers [at least] superposed, sweeping hairs +; deletion and insertion in rpoB gene.

Evolution. The age of this clade is some 38-32 million years (K.-J. Kim et al. 2005).

7. Carduoideae Sweet

Biennial habit common; leaves dissected, teeth spine-tipped; (laticifers +); flower usu. polysymmetric, disc flowers deeply lobed, pollen (psilate), with internal tectum, style branches short, ring of hairs below style branches; (fruits lacking twin hairs); n = 12.

83/2780: Centaurea (695), Cousinia (655), Saussurea (300), Cirsium (250), Jurinea (200), Echinops (120), Carduus (90), Serratula (70), Dicoma (65), Onopordum (60). World-wide, but most N. hemisphere, esp. Eurasia/N. Africa.

Synonymy: Acarnaceae Link, Carduaceae Dumortier, Centaureaceae Martynov, Cnicaceae Vest, Cynaraceae Durande, Echinopaceae Dumortier, Serrulataceae Martynov

Pertyoideae [Gymnarrhenoideae [Cichorioideae [Corymbioideae + Asteroideae]]]: ?

8. Pertyoideae Panero & Funk

Like Mutisieae, but flowers not bilabiate, corolla deeply but unequally divided, style arms short, pilose to papillose abaxially, apices variable; pappus of (plumose) bristles; n = 12-15.

5-6/70: Ainsliaea (50). Afghanistan to East (and Southeast) Asia.

Gymnarrhenoideae [Cichorioideae [Corymbioideae + Asteroideae]]: ?

9. Gymnarrhenoideae Panero & Funk

Plant amphicarpic; heads grouped into synflorescence; style branches long, apices rounded; pappus of bristles; n = 10.

1/1: Gymnarrhena micrantha. North Africa to the Middle East.

Gymnarrhena was excluded from Asteroideae by Anderberg et al. (2005).

Cichorioideae [Corymbioideae + Asteroideae]: style branches medium to long [strict definition?, position on tree?]; deletion in ndhF gene.

10. Cichorioideae Chevalier

(Latex + - Cichorieae); disc flowers deeply lobed, (all flowers ligulate - Lactuceae; true ray florets +), (pollen with cavities separating columellae from foot layer [caveate; ?distribution], lophate, with internal tectum), carpels collateral, apices of style branches acute, hairs usually acute; n = (7-)9-10(-13).

224/3600: Vernonia (800-1000, but see below), Crepis (200), Scorzonera (175), Lactuca (125), Lepidaploa (115), Tragopogon (110), Lessinganthus (100), Hieracium (90-1000+: apomixis), Vernonanthura (65), Hypochaeris (60), Sonchus (60), Taraxacum (60-500+: apomixis). World-wide.

Synonymy: Aposeridaceae Rafinesque, Arctotidaceae Bessey, Cichoriaceae Jussieu, nom. cons., Lactucaceae Drude, Picridaceae Martynov, Vernoniaceae Burmeister.

11. Corymbioideae Panero & Funk

Leaves with parallel veins; heads with 1 flower enclosed by two innermost involucral bracts; C with broad, patent lobes, apices of style branches variable; pappus of bristles; n = 16.

1/7. South Africa (map: from Weitz 1989).

12. Asteroideae Lindley

Sesquiterpene lactones at biogenetic levels 3 and 4, (6,8-deoxygenation of flavonoids), benzopyrans, benzofurans +; laticifers 0; ray florets [3 lobed] common, female, disc florets with C shallowly lobed, perfect, anthers (free), often ecalcarate and ecaudate, pollen 25.0-34.3 µm in diam., exine 2.3-4.2 µm across, with cavities separating columellae from foot layer [caveate], with a double tectum, style hairs often rounded, only ± at style tip, stigmatic areas in two marginal bands; n = (4-)9-10(-19); rbcL 6bp x 4 inversion.

1135/16360: Senecio (1000), Eupatorium (1200 [s.l.] or 40 [s.s.]), Helichrysum (500-600), Artemisia (550), Mikania (430), Baccharis (400), Verbesina (300), Ageratina (290), Bidens (235), Stevia (235), Anthemis (210), Erigeron (200), Pentacalia (200), Aster (180: Flora of North America, vol. 20, 2006), Viguiera (180), Chromolaena (165), Gnaphalium (150), Solidago (150), Tanacetum (150), Olearia (130), Seriphidium (130), Ligularia (125), Achillea (115), Coreopsis (115), Anaphalis (110), Brickellia (110), Calea (110), Blumea (100), Koanophyllum (110), Euryops (100), Pectis (100), Wedelia (100), Diplostephium (90), Emilia (90), Espeletia (90: secondarily woody), Symphyotrichum (90), Felicia (85), Fleischmannia (80), Pluchea (80), Pseudognaphalium (80), Pulicaria (80), Pteronia (80), Antennaria (70-several hundred), Brachycome (70), Cremanthodium (70), Haplopappus (70), Packera (65), Perityle (65), Celmisia (60), Conyza (60), Gynoxys (60), Leontopodium (60), Monticalia (60), Parasenecio (60), Psiadia (60). World-wide.

Synonymy: Ambrosiaceae Martynov, Anthemidaceae Martynov, Artemisiaceae Martynov, Athanasiaceae Martynov, Calendulaceae Link, Coreopsidaceae Link, Eupatoriaceae Martynov, Gnaphaliaceae Rudolphi, Heleniaceae Rafinesque, Helianthaceae Dumortier, Helichrysaceae Link, Inulaceae Bessey, Madiaceae A. Heller, Matricariaceae J. Voigt, Partheniaceae Link, Santolinaceae Martynov, Senecionaceae Spenner, Tanacetaceae Vest, Xanthiaceae Vest

• Asteraceae are very variable vegetatively, but may be recognised by their capitulate and involucrate inflorescences in which the numerous small centripetally-opening flowers are only sometimes subtended by bracts. The anthers are usually connate and form a tube through which the style extends before the two stigmatic lobes separate and become recurved. The rather small, single-seeded fruits usually have a pappus, a modified calyx, and are frequently dispersed by wind.

Evolution. Diversification of Asteraceae may have occured only some 42-36 million years ago, the stem group perhaps being up to 49 million years old (K.-J. Kim et al. 2005); Bergh and Linder (2009) suggested that diversification of Asteroideae began in the Eocene (56.6-)43.0(-29.6) million years ago. Various clades within the family have been dated and there seems to have been an Oligocene radiation of the subfamilies (Kim et al. 2005; Funk et al. 2005; Barker et al. 2008). Diversification probably began in South America, and many tribes arose in the course of a subsequent radiation from Africa (Panero & Funk 2008; Funk 2009, see also Kim & Jansen 1995). For instance, crown Gnaphalieae (Asteroideae) began diverging (52.3-)34.5(-20.6) million years ago, probably in southern Africa, with various dispersals subsequently; these include one that resulted in the some 550 species of the Australasian part of the tribe (stem age [22.1-]15.6[9.1] million years; crown age [20.6-]14.6[-8.3] million years - see Bergh & Linder 2009). Much diversification of the high-altitude species of Saussurea may have occured in the context of the uplift of the Qinghai-Tibetan plateau within the last 14 million year (Y.-J. Wang et al. 2009).

It is hardly surprising, given the size and ubiquity of the family, that there should be speculation about what has caused its diversification, whether the development of the capitulum itself, the storage of carbohydrates as unbranched-chain fructans, which probably contributes to the ability of Asteraceae to live in the rather dry conditions that many of them prefer (John 1996), or some other reason. There is evidence of a palaeopolyploidy event involving most of the family, and subsequent genome duplications near the base of Asteroideae and in Mutisioideae (Barker et al. 2008). But there are perhaps parallels with Poaceae, which also have large-scale genome duplications and store carbohydrates as fructans - and also in the likelihood that diversification is best explained in the context of the evolution of particular clades within the family... Indeed, although Asteraceae alone contain about 8% of eudicot species, within Asteraceae, Asteroideae alone include over 16,000 species, some two thirds of the family; most of the eleven clades basal to Asteroideae are species poor, with the exception of Cichoroideae, with ca 3,600 species, and Carduoideae, with 2,800 species.

Flaveria (Asteroidae) has some species with C₄ photosynthesis, some with C₃, and some intermediate (Bläsing et al. 2000; McKown & Dengler 2009 and references).

Most Asteraceae can be instantly recognised because of their capitular inflorescence. This is a highly modified raceme, although in Gorteria the outermost (ray) flowers develop centrifugally, and they develop more slowly than the acropetal more central (disc) flowers (Thomas et al. 2009). There are several cases in which the capitulum has independently become reduced to a single flower, and then the single-flowered capitulae reaggregate into a supercapitulum (Claßssen-Bockhoff 1996b for details; Leins & Gemmeke 1979; Katinas et al. 2008a). Whether normal capitulum of a supercapitulum, the inflorescence functions as a flower in terms of attacting pollinators.

The flowers of some Senecioneae and Eupatorieae are visted by male Danainae, Ithomiinae, Arctiidae and Ctenuchidae because the pyrrolizidine alkaloids they contain are used as the basis of the pheromones of these lepidoptera, or are the sources of compounds other organisms find distasteful (see also Crotalaria and some Boraginaceae and Apocynaceae: Edgar et al. 1974; Fiske 1975; Ackery & Vane-Wright 1984; Brown 1987; Weller et al. 1999; Anke et al. 2004); there has been parallel evolution of these alkaloids within Asteroideae. Larvae of Nymphalidae-Melitaeini butterflies are commonly found on Asteraceae, and also on Lamiales, from whence they probably moved (Wahlberg 2001; Nylin & Wahlberg 2008), while caterpillars in a clade of Nymphalidae-Heliconiinae-Acraeini utilise primarily Andean members of this family, probably switching from host plants in the Passifloraceae area (Silva-Brandão et al. 2008). Cecidomyiid gall midges are notably common on the family in North America (Gagné 1989), and galls on Solidago growing in the prairies are conspicuous in the late summer, the Tephritidae-Tephritinae are mostly associated with the family (Korneyev et al. 2005, esp. Urophora), while the probably paraphyletic Cynipidae-Aylacini are also common on Asteraceae (Csoka et al. 2005). There has been a diversification of agromyzid dipteran leaf miners in north temperate Asteraceae; these insects prefer plants with noxious secondary metabolites (Winkler et al. 2009).<

Some Asteraceae have a pump (nüdelspritze) mechanism of secondary pollen presentation (?plesiomorphic), other taxa have a brush mechanism (see Leins & Erbar 2003b for a possibly evolutionary sequence of pollen presentation devices). Within Barnadesioideae, sister to the rest of the family, secondary pollen presentation is by simple deposition on the style/stigma (as in at least some Calyceraceae) or by an unspecialised type of brush mechanism (Leins & Erbar 2006). Pollinators of Asteraceae appear not to be very selective, since it seems that insects simply trample about on top of the capitulum and pollinate indiscriminately as they go, this is not true; pollination is commonly by a wide variety of specialist solitary bees which form complex and partly learned associations with individual species of Asteraceae (Lane 1996). A number of Carduoideae have thigmotropic stamens, the filaments contracting when touched by the pollinator, the pollen then being forced out of the anther tube; this pollination mechanism appears to have arisen more than once, and is associated with short and sticky, not long and dry, stigmas and smooth, not spiny, pollen (López-Vinyallonga et al. 2009).

There is a great diversity of breeding systems in the family (e.g. Burtt 1961, 1977a), and the evolution of different flower types in Inuleae (Asteroideae) has been examined by Torices and Anderberg (2009). The anthers are free in wind-pollinated Asteroideae and the heads have either staminate or carpellate flowers, and since the latter may have only a single flower, the end result is a breeding system very much like that of other wind-pollinated plants like Fagales - pollen-producing units aggregated, and female reproductive units producing a single-seeded fruit (for the phylogeny of Artemisia, see Vallès & Garnatje 2005 and Sanz et al. 2008). Apomixis is quite common, as in Hieracium and Taraxacum (see T. absurdum), both in Cichorioideae, and Antennaria (Asteroideae).

Most fruits are crowned by a plumose pappus, a highly modified calyx (see Yu et al. 1999 for confirmation at the level of gene expression); this is often not the first part of the flower to be initiated (see Mukherjee & Nordenstam 2008 for variation). The hairs that make the pappus up are themselves sometimes hairy, and wind dispersal is very common. However, a number of taxa are dispersed by animals, whether by hooked fruits (Bidens), or hooks on the inflorescence (Arctium), or by myrmecochory, the fruits having some sort of elaiosome, as in Centaurea (Carduoideae) and Osteospermum (Asteroideae: Lengyel et al. 2009).

Within Carduoideae, biennials are ten times commoner than in other Asteraceae, and the stout root stocks and large flower heads in particular are resources for the numerous herbivorous insects that specialize on this clade. More than fifty genera of specialized thistle insects, including representatives of Zygaenidae, Tortricidae, Pterolonchidae (all Lepidoptera), Curculionidae (Coleoptera), Tephritidae (Diptera: see Braändle et al. 2005), Tingitidae (Hemiptera) and Cynipidae (Hymenoptera), are found on Carduoideae of the west Palearctic region, although their numbers are not great considering the diversity of Carduoideae there (Zwölfer 1988; Brändle et al. 2005 and literature). These herbivores are particularly abundant in the Mediterranean region, which is perhaps where the clade evolved (Zwölfer 1988).

Economic Importance. Timme et al. (2007) provide a phylogeny of the important genus Helianthus.

Chemistry, Morphology, etc. Sesquiterpene lactones give the bitter taste of many Asteraceae, e.g. Cnicus benedictus. For fatty acids in the seeds, see Bamai and Patil 91981). There are floral bracts in some Asteroideae-Heliantheae (the tribe includes taxa that were in Eupatorieae), but they seem to have been reacquired more than once and are certainly not plesiomorphic in the family as was once thought. The involucral bracts in Asteroideae-Senecioneae are uniseriate. There is considerable varion in floret morphology in Asteraceae, however, I have seen no study where all this variation has been evaluated in the context of a tree, most work along these lines laying out morphoclines developed from a priori principles (). Many Asteroideae have three-toothed ray florets that give the appearance of being slit-monosymmetric, however, they seem to represent a modification of a 2:3 bilabiate corolla in which the adaxial lobes have been suppressed (Weberling 1989; Gillies et al. 2002)). The corolla hairs of Barnadesioideae are distinctive; of the three cells that make them up, the epidermal cell is undistinguished, the basal cell is short and thick-walled, and the other cell is longer and has thin walls. Some taxa, including Barnadesioideae, have a midvein in the petal... (see Carlquist 1976; Gustafsson 1995). Caveate pollen also is found in Arctoteae and some Lactuceae, and may be more basal on the tree (it is a synapomorphy for Asteroideae at present). Indeed, Blackmore et al. (1984) noted that it is evident early in development in Gerbera (Mutisieae), but not later, and suggested that pollen grains of Asteraceae might all be basically caveate. There is variation in the orientation of the gynoecium and style branches: carpels superposed, style branches arranged radially to the head surface; carpels collateral, style branches tangential to head surface: see Robinson 1984); details of the distribution of this feature are unknown. Buphthalmum has a hollow style (Leins 2000); I do not know how widespread such styles are in Asteraceae.

In general, the very different adult floral morphologies are quite similar early in development (Harris 1995); there seems to be some variation as to whether there is a corolla ring primordium initated first, or whether petals are initiated separately, i.e., between early and late corolla tube development. Asteroideae-Heliantheae have distinctive black fruits that are covered by phytomelan (see Graven et al. 1998 for what is known about this compound); they are also described as being carbonized. Nuclear endosperm is sometimes mentioned as being the only endosperm condition found in the family or as a synapomorphy for it (e.g. Tobe & Morin 1996; Inoue & Tobe 1999), but there is in fact considerable variation in endosperm development (Johri et al. 1992 for references) which I have not attempted to fit to the tree. Syneilesis seems to lack cotyledons entirely (Teppner 2001).

For information on Asteraceae, see e.g. Carlquist (1976: an interesting summary of variation associated with a tribal classification), Skvarla et al. (1977: pollen terminology), Roque and Silvetstre-Capelato (2001: pollen of Gochnatioideae), Wortley et al. (2008: pollen of Arctotidae-Cichoridoideae), Tellería and Katinas (2009: pollen in Mutisia), Osman 2009 (pollen of Cichorioideae-Cardueae), Hong Wang et al. (2009b and references), pollen of Cichorioideae-Cichorieae, K. Bremer (1987, esp. 1994 [a monograph, but the major classification is rather different from that followed here], 1996 [subfamilial groupings]), Seaman (1982: sesquiterpene lactones), Seaman et al. (1990: diterpenes, other references), Jansen et al. (1991: variation in the context of phylogeny), Harris (1995: inflorescence and flower development), Hind et al. (1996: general), Bohm and Stuessy (2001: flavonoid chemistry), Leins and Erbar (2003b: capitulum development), Aniszewski (2007: alkaloids), Watanabe et al. (2007: chromosome numbers), Katinas et al. (2008b: Mutisioideae, but not in the sense used here - general morphology) and Thomas et al. (2009: development of Gorteria flowers). Anderberg et al. (2006) summarize the variation in the family and Funk et al. (2009) discuss biogeography and much more.

Phylogeny. There is much recent phylogenetic work on Asteraceae, and only a few references can be included here. Panero and Funk (2002, especially 2008; see also Funk et al. 2005, a supertree) present the phylogeny reflected in the classification above. Much, but not all, of the uncertainty in relationships around the old Mutisioideae seems to have been resolved, with distinctive gene deletions and insertions characterising a number of the clades (Panero & Funk 2008). Indeed, the large inversion in the chloroplast genome that occurs in most of the family, but not Barnadesioideae and other Asterales, was an exciting discovery in the early days of molecular systematics, in part because it was consistent with a morphological phylogeny that came out at about the same time (cf. Jansen & Palmer 1987; Bremer 1987; see also Y.-D. Kim & Jansen 1995).

For phylogenetic relationships within Barnadesioideae, see Urtubey and Stuessy (2001) and in particular Gustaffson et al. (2001) and Gruenstaeudl et al. (2009); the position of Schlechtendalia is uncertain. For corolla morphology in this subfamily, see Stuessy and Urtubey (2006). For a phylogeny of Carduoideae-Cardueae, see Susanna et al. (2006) and Garcia-Jacas et al. (2002), of Centaurinae, see Garcia-Jacas et al. (2001), of Echinops, see Garnatje et al. (2005: sectional classification), and for that of Cousinia (biphyletic) and relatives, see López-Vinyallonga et al. (2009). In Sonchinae (Cichorioideae), Sonchus is para/polyphyletic (Kim et al. 2007), with woody, island-dwelling forms being independently derived within the clade. The classic studies by Babcock (e.g. 1947) on Crepis that assumed that evolution - in this case of the karyotype in particular - was unidirectional need comprehensive re-evaluation (Enke & Gemeinholzer 2008). Species limits around here are difficult because of apomixis; see Gottschlich (2009) for the complexities of variation in Hieracium in a smallish area of Italy. For a phylogeny of Tragopogon and its relatives, see Mavrodiev et al. (2005), and of the African Goteriinae, see Funk and Chan (2008). For a phylogeny of Vernonia, a genus whose circumscription is problematic, see Keeley et al. (2007).

North American Asteroideae-Astereae are monophyletic and largely herbaceous (Noyes & Rieseberg 1999). For the phylogeny of the helenioid Heliantheae, see Baldwin et al. (2002), of Inuleae (inc. Plucheeae), see Anderberg et al. (2005) and Englund et al. (2009), for circum-Mediterranean Anthemidae, also their biogeography, see Oberprieler (2005), and for that of the Hawaiian silverswords and their immediate relatives, and their relatives in turn, which are now found in west North America (Madiinae), see Baldwin and Wessa (2000) and Carlquist et al. (2004, also Madiinae Showcase). Bidens on Hawaii also shows much variability in growth form, etc., but there is little molecular variation or genetic barriers between the species (Ganders et al. 2000). Within Gnaphalieae, Helichrysum is polyphyletic (Galbany-Casals et al. 2004, 2009; Bergh & Linder 2009). For a phylogeny of Anthemidae, see Himmelreich et al. (2008 and references). Relationships within the huge genus Senecio are beginning to be disemtangled (Pelser et al. 2006, esp. 2007). For the phylogeny of Artemisia, see Vallès & Garnatje (2005) and Sanz et al. (2008). Olearia is likely to be polyphyletic (Cross et al. 2002); for Symphyotrichum and relatives, see Vaezi and Brouillet (2009), for Blumea, whose sections need overhaul, see Pornpongrungrueng et al. (2009), and for Machaerantherinae, see Morgan et al. (2009).and for the Hinterhubera group, see Karaman-Castro and Urbatsch (2009: groupings geographic). Finally, it is worth noting that there may be relatively common and deep hybridisation in Asteroideae in particular which may make life for those interested in phylogeny reconstruction rather interesting (e.g. Fehrer et al. 2007; Pelser et al. 2008; Morgan et al. 2009).

Classification. Panero and Funk (2008) present the subfamilial classification followed above (there are, of course, alternative classifications in the literature - e.g. Jeffrey 2004). Anderberg et al. (2006) have recently enumerated the genera in the family. However, generic limits in many places are in something of a state of flux; Vernonia is a classic case of uncertainty - should it include 800-1000 species, or should these species be placed in 20 subtribes, of which two thirds of the genera are mono- or ditypic? (Keeley et al. 2007 for a phylogeny; Robinson 2006 and references for genera). Should there be lumping or splitting in the even larger genus Senecio (see Pelser et al. 2006, esp. 2007)? Answer: both. Substantial adjustments to generic limits will be needed in Asteroideae-Inuleae-Inulinae (Englund et al. 2009). I am grateful to Jose L. Panero for comments.

Botanical Trivia. Arctium lappa (burdock) infructescences became attached to the dog of a Swiss engineer, George de Mestral, in 1945 and the result was the development of velcro (Wikipedia 2009).