LIGNOPHYTA

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

EXTANT SEED PLANTS/SPERMATOPHYTA

Plant woody, evergreen; nicotinic acid metabolised to trigonelline, (cyanogenesis via tyrosine pathway); primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins derived from (some) sinapyl and particularly coniferyl alcohols, thus containing p-hydroxyphenyl and guaiacyl lignin units, (lignins derived from p-coumaryl alcohol, i.e. S [syringyl] lignin units); true roots present, apex multicellular, xylem exarch, and branching endogenous; arbuscular mycorrhizae +; shoot apical meristem multicellular, interface specific plasmodesmatal network; stem with ectophloic eustele, endodermis 0, xylem endarch, branching exogenous; vascular tissue in t.s. discontinuous by interfascicular regions; vascular cambium + [xylem ("wood") differentiating internally, phloem externally]; wood homoxylous, tracheids and rays alone, tracheid/tracheid pits circular, bordered; mature sieve tube/cell lacking functioning nucleus, plastids with starch grains; phloem fibres +; stem cork cambium superficial, root cork cambium deep seated; leaves with single trace from sympodium ["nodes 1:1"]; stomata ?; leaf vascular bundles collateral; leaves megaphyllous [determinancy evolved first, then ad/abaxial symmetry], spiral, simple, lamina with vein density up to 5 mm/mm² [mean for all non-angiosperms 1.8]; axillary buds associated with at most some leaves; prophylls [including bracteoles] two, lateral; plant heterosporous, sporangia eusporangiate, on sporophylls, sporophylls aggregated in indeterminate cones/strobili; true pollen [microspores, i.e. no distal pore for release of gametes] +, grains mono[ana]sulcate, exine and intine homogeneous; ovules unitegmic, crassinucellate, megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development first endo- then exosporic, tube developing from distal end of grain, to ca 2 mm from receptive surface to egg, gametes two, developing after pollination, with cell walls, with many flagellae; female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large", first cell wall of zygote transverse, embryo straight, endoscopic [suspensor +], short-minute, with morphological dormancy, white, cotyledons 2; plastid transmission maternal; two copies of LEAFY gene, PHY gene duplications [three - [BP [A/N + C/O]] - copies], nrDNA with 5.8S and 5S rDNA in separate clusters; mitochondrial nad1 intron 2 and coxIIi3 intron and trans-spliced introns present.

MAGNOLIOPHYTA

Lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, [cyanogenesis in ANITA grade?], S [syringyl] lignin units common, positive Maüle reaction [syringyl:guaiacyl ratio more than 2-2.5:1], and hemicelluloses as xyloglucans; root apical meristem intermediate-open; root vascular tissue oligarch [di- to pentarch], lateral roots arise opposite or immediately to the side of [when diarch] xylem poles; origin of epidermis with no clear pattern [probably from inner layer of root cap], trichoblasts [differentiated root hair-forming cells] 0; shoot apex with tunica-corpus construction, tunica 2-layered; reaction wood ?, with gelatinous fibres; starch grains simple; primary cell wall mostly with pectic polysaccharides, poor in mannans; tracheid:tracheid [end wall] plates with scalariform pitting, wood parenchyma +; sieve tubes enucleate, sieve plate with pores (0.1-)0.5-10< µm across, cytoplasm with P-proteins, cytoplasm not occluding pores of sieve plate, companion cells from same mother cell that gave rise to the sieve tube; sugar transport in phloem passive; nodes unilacunar [1:?]; stomata with ends of guard cells level with pore, paracytic, outer stomatal ledges producing vestibule; leaves petiolate, lamina [formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, veins (1.7-)4.1(-5.7) mm/mm², endings free; most/all leaves with axillary buds; flowers perfect, pedicellate, polysymmetric, parts spiral [esp. the A], free, numbers unstable, development in general centripetal; P not sharply differentiated, with a single trace, outer members not enclosing the rest of the bud, often smaller than inner members; A many, filament not sharply distinguished from anther, stout, broad, with a single trace, anther introrse, tetrasporangiate, sporangia in two groups of two [dithecal], ± embedded in the filament, with at least outer secondary parietal cells dividing, each theca dehiscing longitudinally by action of hypodermal endothecium, endothecial cells elongated at right angles to long axis of anther; tapetum glandular, binucleate; microspore mother cells in a block, microsporogenesis successive, walls developing by centripetal furrowing; pollen subspherical, tectum continuous or microperforate, ektexine columellar, endexine thin, compact, lamellate only in the apertural regions; nectary 0, G free, several, ascidiate, with postgenital occlusion by secretion, stylulus short, hollow, cavity not lined by distinct epidermal layer, stigma ± decurrent, dry [not secretory]; ovules few [?1]/carpel, marginal, anatropous, bitegmic, micropyle endostomal, outer integument 2-3 cells across, often largely subdermal in origin, inner integument 2-3 cells across, often dermal in origin, parietal tissue 1-3 cells across [crassinucellate], nucellar cap?; megasporocyte single, hypodermal, megaspore tetrad linear, functional megaspore chalazal, lacking sporopollenin and cuticle; female gametophyte four-celled [one module, nucleus of egg cell sister to one of the polar nuclei]; P deciduous in fruit; seed exotestal; pollen binucleate at dispersal, trinucleate eventually, germinating in less than 3 hours, pollination siphonogamous, tube elongated, growing at 80-600 µm/hour, with pectic outer wall, callose inner wall and callose plugs, growing between cells, penetration of ovules via micropyle [porogamous] within ca 18 hours, distance to first ovule 1.1.-2.1 mm, tube moves between nucellar cells; double fertilisation +, endosperm diploid, cellular [micropylar and chalazal domains develop diffently, first division oblique, micropylar end initially with a single large cell, divisions uniseriate, chalazal cell smaller, divisions in several planes], oily and/or proteinaceous, embryo cellular ab initio, minute; germination hypogeal, seedlings/young plants sympodial; Arabidopsis-type telomeres [(TTTAGGG)_n]; whole genome duplication, ndhB gene 21 codons enlarged at the 5' end, single copy of LEAFY and RPB2 gene, knox genes extensively duplicated [A1-A4], AP1/FUL gene, paleo AP3 and PI genes [paralogous B-class genes] +, with "DEAER" motif, SEP3/LOFSEP and three copies of the PHY gene, [PHYB [PHYA + PHYC]].

Evolution. Possible apomorphies for flowering plants are in bold. Note that the actual level to which many of these features, particularly the more cryptic ones, should be assigned is unclear. This is because some taxa basal to the [magnoliid + monocot + eudicot] group have been surprisingly little studied, there is considerable homoplasy as well as variation within and between families of the ANITA grade in particular for several of these characters, and also because details of relationships among gymnosperms will affect the level at which some of these characters are pegged. For example, if reticulate-perforate pollen is optimized to the next node on the tree (see Friis et al. 2009 for a discussion), it effectively makes the pollen morphology of the common ancestor of all angiosperms ambiguous... For other features such as details of sugar transport in the phloem, their placement on the tree is frankly speculative. Finally, for features such as parietal tissue/a nucellus only one (Nymphaeales) to three cells thick above the embryo sac and a stylar canal lacking an epidermal layer, although plesiomorphous for basal grade angiosperms (Williams 2009), I am unsure where on the tree a thicker nucellus and a stylar epidermal layer are acquired.

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

[ERICALES [ASTERID I + II]]: ovules tenuinucellate.

[ASTERID I + II] / CORE ASTERIDS: ellagic acid 0, non-hydrolysable tannins not common; sugar transport in phloem active; inflorescence basically cymose; C forming a distinct tube; A epipetalous, = and opposite sepals or P, polyandry associated with increased numbers of C or G, very uncommon; (pollen with orbicules); duplication of the PI gene.

ASTERID II / CAMPANULIDAE: 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]]]]] / APIIDAE: iridoids +; inflorescence?; C tube initiation early; G [2-3], inferior.

Evolution. Divergence & Distribution. Magallón and Castillo (2009) offer estimates of ca 94 and 96 million years for relaxed and constrained penalized likelihood datings for crown group divergence in this clade (i.e. divergemce of Asterales from the rest), the stem group being ca 99 and 99.7 million years old (relaxed and constrained estimates) - but note topology.

Endress (2011a) suggested that the inferior ovary so common (but there are reversals) here might be a key innovation.

Chemistry, Morphology, etc. Polyacetylenes are sporadic in this clade, occurring in Asterales, Dipsacales and Apiales, but always in much embedded clades. Although usually not co-occurring with iridoids, the two may sometimes be found together, as in Torricellia angulata (Pan et al. 2006; Liang et al. 2009).

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 early corolla 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" (see above), 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 phytochemical similarities between the clade [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 Link  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 +; style long; ovules many/carpel, integument <7 cells thick, endothelium +, hypostase 0; antipodal cells ephemeral; embryo suspensor filamentous, micropylar and chalazal endosperm haustoria +; mitochondrial rpl2 gene lost. - 11 families, 1649 genera, 34153 species.

Evolution. Divergence & Distribution. 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.

Endress (2011a) thought that the character "monosymmetric flower" in Asterales might be a key innovation, although where it is to be optimised on the tree is currently unclear; the speciose Campanulaceae-Lobelioideae will (depending on the phylogeny) almost certainly represent an independent acquisition of this feature, its aquisition near Asteraceae being another. Furthermore, although monosymmetric flowers may occur in most Asteraceae, the pollination unit, the capitulum, is polysymmetric, and major pollinators behave accordingly (see below under Asteraceae). Endress (2011a) also suggested that a key innovation somewhere in Asterales was tenuinucellate ovules.

Floral Biology. Several families, notably Campanulaceae and the Asteraceae area, 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, 2010) in particular discuss in considerable detail the evolution of these pollen presentation mechanisms.

Ecology & Physiology. Fructans may stabilize cell membranes under drought and/or freezing conditions (Livingston III et al. 2009).

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). The corolla lobes quite often appear to consist of a central portion and marginal "wings"; this is cause by the induplicate-valvate aestivation of flowers with such a corolla. For a study of petal vasculature, which shows interesting variation, see Gustafsson (1995). Monosymmetry is often associated with a slit the length of the corolla. Variation of ovary position in Asterales is considerable.

Tobe and Morin (1996) summarize embryological knowledge of many members of the order. For some inflorescence morphology, see Philipson (1953), for fructans/inulins, see Meier and Reid (1982), for integument thickness, see Inoue and Tobe (1999), for some inflorescence development, see Harris (1999), and for pollen, see Polevova (2006). For a general discussion of variation in the order, see J. Kadereit (2006) and Lundberg (2009).

Phylogeny. There is quite a lot of phylogenetic structure in Asterales, but often with rather weak support, as was suggested by D. Soltis et al. (2000; see also Olmstead et al. (2000).Subsequent studies improved support for many clades, although there was still a basal polytomy (Kårehed et al. 2000; Lundberg 2001a, b; Kårehed 2002a; especially Bremer et al. 2001 and Lundberg & Bremer 2001, 2003). 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; Tank & Donoghue 2010); 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 was initially not very strong, while Soltis et al. (2007a) found Campanulaceae to be sister to rest of Asterales (1.0 p.p.). However, Rousseaceae s.l. do seem to be sister to Campanulaceae (Kårehed 2002a; Tank et al. 2007; esp. Tank & Donoghue 2010). 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), while Soltis et al. (2011) found that Alseuosmiaceae were siter to the rest of the clade, but with little support and in any case perhaps because of the pull of some mitochondrial genes. However, relationships along the spine of Asterales were quite well resolved in a ten chloroplast gene analysis of Tank and Donoghue (2010) and are followed here; Soltis et al. (2011) found a largely similar topology, apart from the position of Pentaphragmaceae and a weakly supported set of relationships [Phellinaceae [Alseuosmiaceae + Argophyllaceae]].

Stylidiaceae and Donatiaceae have often been associated, e.g. 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.

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: Alseuosmiineae Shipunov - Alseuosmiales Doweld, Ambrosiales Dumortier, Anthemidales Link, Boopidales Berchtold & J. Presl, Brunoniales Lindley, Calendulales Link, Calycerales Link, Campanulales Berchtold & J. Presl, Carduales Small, Cichoriales Link, Cynarales Rafinesque, Echinopales Link, Goodeniales Berchtold & J. Presl, Lobeliales Link, Menyanthales J. Presl, Pentaphragmatales Doweld, Phellinales Doweld, Rousseales Doweld, Scaevolales Martius, Stylidiales Berchtold & J. Presl

[Rousseaceae + Campanulaceae]: flowers large; A free.

ROUSSEACEAE Candolle   Back to Asterales

Young stem with separate bundles; lamina 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, leaf 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 medium-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. Floral Biology. 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 simplex 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. 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

Herbs, whether annual or perennial, 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), lamina vernation 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 (binucleate), endexine not lamellate by apertures; G [2(-5)],(± superior; placentation parietal), style elongating after A dehiscence, glabrous, stigma lobed; integument ca 6 cells across; K persistent; seeds many, exotesta cells lignified, polygonal or elongated, (endotestal cells, esp. inner walls, thickened); endosperm (starchy), copious; 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]: flowers polysymmetric; pollen spheroid to oblate-spheroid, verrucate or with spicules; fibrillar protein bodies in nuclei.

1. Nemacladoideae M. H. G. Gustafsson

Annuals (perennials); C polysymmetrical, resupinate or monosymmetric, 3:2 not resupinate; A (adnate to C), filaments ± forming tube, anthers flipping back after pollen release, free, (fimbriate scales abaxial to some filaments where they join the C); pollen tricolporate or 6-colpate; large "glands" at base of style; G also [3], half inferior to superior, style head bending towards or away from median K, presenting pollen; ?embryology; (fruit circumscissile - Parishella); n = 9.

2/25: Nemacladus (24). 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] +, latex rich in polysterols; (vessel elements with scalariform perforation plates); inflorescence often ± cymose; median K adaxial; stamens often sprawling at bottom of corolla tube after anthers have dehisced although bases conceal nectar; pollen echinate; G (1 [2) 3-5(-10)], opposite sepals (C), or median member adaxial, (style hollow - how common?), style long-hairy, espeacially in the upper half or so, hairs with bulbous bases, retractile, presenting pollen, stigma dry or wet; (integument ca 8 cells across, vascularized - Azorina); (embryo medium); (fibrillar protein intranuclear inclusions); extensive rearrangements in the chloroplast inverted repeat.

50/1050. More or less world-wide, but few in Australia-New Zealand and South Americar (map: from Hultén 1971; Thulin 1975; Shulkina 1978; FloraBase v.2011).

2A. Cyanantheae Meisner

Leaves often opposite; (A 3); pollen colpate/colporate; G [5], opposite ?; 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.

2B. Campanula, Wahlenbergia, etc.

Leaves often spiral; (A adnate to C); pollen (flattened-triangular), 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, pyr[roliz]idine alkaloids +, p-coumaric acid, caffeic acid 0; lamina vernation supervolute [Lobelia]; flowers resupinate by pedicel 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; synergids hooked, (antipodal cells barely persistent); (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 Jussieu, 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; ?embryology; 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 subpolysymmetric, 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; embryology?; 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. Divergence & Distribution. Ages for diversification within Lobelioideae differ greatly according to the how they are estimated - (59.6-)54.9(-50.9) million years (penalized likelihood) versus (88.2-)72.7(-52.2) milion years (BEAST: see Antonelli 2009). Diversification within Campanuloideae may have begun (41-)37.4-23.5(-3.2) million years ago (Roquet et al. 2009, also ages for other clades, note ages in different analyses varied considerably for deeper nodes), a possible age for stem Campanuloideae being ca 41 million years (Wikström et al. 2001). For dating of crown Wahlenbergia, (45.3-)29.6(-15.2) million years (HPD: Prebble 2011); however, there was little diversification in terms of those taxa in the analysis for ca 10 million years, while stem Wahlenbergia is ca 32 million years old.

The pachycaul giant lobelias are derived from herbaceous ancestors (Knox et al. 1993), and giant lobelias from widely separated parts (Pacific, South America, Africa) of the globe may be in the same immediate clade (Antonelli 2009), 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; Antonelli (2009) also suggested that Lobelioideae originated in Africa, and with much subsequent long distance dispersal of the tiny seeds. Givnish et al. (2006a, 2008b; see also Buss et al. 2001 - seed morphology) note that the some 130 species of Hawaiian Lobelioideae appear to have evolved from a single woody ancestor a mere ca 13 million years ago (ages in Antonelli 2009 are slightly older); these ages are older than that of the oldest island, but presumably there was movement from islands that subsequently have sunk. Fleshy fruits have evolved more than once in Hawaiian Lobelioideae, which represent a major plant radiation on the islands, and species there have a variety of growth habits, pollinators, fruit dispersers, etc.

Subsequent diversification within Campanuloideae began substantially later than their separation from the rest of the family, 26.3-15.8 million years ago (Wikström et al. 2001). The biogeographic history of this subfamily 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 over 100 species of Campanula in Turkey alone. 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).

Floral Biology. For the evolution of the secondary pollen presentation devices in the family, see Erbar and Leins (1988b) and Leins and Erbar (especially 2003b, 2006, 2010); sampling (inc. Nemocladoideae, some Lobelioideae) is still incomplete, and understanding the evolution of these devices awaits a better supported phylogeny (see below). Associated with the secondary pollen presentation that occurs throughout the family, the flowers are are initially polysymmetrical in bud, protandrous and the anthers are introrse and connivent at the time that the anthers dehisce (e.g. Leins & Erbar 2003b, 2010). In Campanuloideae, with their brush pollination devices, the pollen is caught in a brush of hairs on the style, whence they are removed by the pollinator; in the female phase, the hairs retract so any grains present will fall off and selfing is prevented . However, in Pteromarula the stigmatic head is swollen and hairs occur only there (Igersheim 1993), but otherwise pollination is similar. Phyteuma has coherent corolla lobes although the corolla is open laterally; the style hairs are only partly retractile. In Lobelioideae the stigma forces pollen out or the tube formed by the connate anthers before the stigmatic lobes separate, recurve, and become receptive.

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). 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).

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). 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 flowers of Downingia, see Kaplan (1969 and references), 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), for testa morphology and anatomy, see Cupido et al. (2011 and references). 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) and Eddie et al. (2010), for embryology, etc., see Kausik and Subramanyam (1945, 1947), Subramanyam (1949, 1970: possible taxonomically interesting differences in cell number of the haustoria), Murata (1995) and Buss et al. (2001), both seed coat anatomy/morphology, and Shamrov (1998: ovule); 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 Nemacladus, see Morin and Ayers (2011).

Other variation that needs to be incorporated into this schema includes inflorescence type, whether basically determinate (e.g. Campanuloideae) or indeterminate (e.g. 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 (2010) 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 (Cyanantheae) are strongly supported as being sister to the whole lot. Neither Campanula nor Wahlenbergia are monophyletic, and within the latter, W. hederacea is separate from the rest of the genus (Haberle et al. 2008b, 2009; Cellinese et al. 2009; Roquet et al. 2008, 2009; Borsch et al. 2009; Prebble et al. 2010).

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. The pachycaul giant lobelias, with a very scattered distribution, are derived from herbaceous ancestors (Knox et al. 1993; Antonelli 2009).

Classification. 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. For a world checklist and bibliography, see Lammers (2007), while Lammers (2011) provided a sectional classification for Lobelia in its current circumscription.

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 [[Alseuosmiaceae [Phellinaceae + Argophyllaceae]] [Stylidiaceae [Menyanthaceae [Goodeniaceae [Calyceraceae + Asteraceae]]]]]]: corolla lobes with marginal wings [could go here].

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 two-ranked, lamina usu. asymmetrical, margins ± serrate (entire); inflorescences cymose, usu. scorpioid; hypanthium +, nectariferous cavities between septae joining hypanthium to G; K petaloid, 2 large + 3 small, C ± deeply lobed (free), with marginal wings; stamens adnate to corolla, anthers extrorse, basifixed; pollen 2-celled, oblate, 3-lobed, apertures between lobes, ektexine smooth, with surface lamellae, endexine lamellate, esp. by apertures; G [2-3], style short, stigma capitate; integument ca 3 cells across; embryo sac protruding from micropyle; fruit baccate, K and C persisting; seeds minute, exotestal cells cubes, inner walls lignified; chalazal haustorium 0, 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 pollen, see Dunbar (1978), 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]]: plant woody; 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 +; rays 0, narrow or broad; starch-storing living fibres +; axial parenchyma 0 (+); pericyclic fibres weakly developed; petiole bundle(s) arcuate; endodermis in both stem and leaf; hairs axillary; lamina vernation 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], stigma barely expanded; ovules 2 or more/carpel; 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. Plant-Animal Interactions. 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 (Dickison 1986b), 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 Paliwal and Srivastava (1969), 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.

Classification. 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.

Synonymy: Platyspermataceae Doweld

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

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 margins 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], stigmas quite large; ovule 1/carpel, apical; 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), Lobreau-Callen (1977) for pollen, 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 vernation 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), stigma punctate or lobed, wet; nectary + [Corokia]; ovule 1 or several/carpel, apical, apotropous, integument ca 6 cells across, nucellus base massive; 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 fibres 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, Mauritzon (1933) for a little embryology, 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 [Menyanthaceae [Goodeniaceae [Calyceraceae + Asteraceae]]]]: ?

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

Inulin +; young stem with separate bundles; nodes 1:1; lamin margins entire, petiole 0; C imbricate; nectary +; A 2, anthers extrorse; pollen colpate; micropylar and chalazal endosperm haustoria +; ?embryo "minute".

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 also 3, free; pollen nuclei?; G [2-3], styles separate, somewhat recurved, stigmas capitate; hypostase +; fruit indehiscent; seed coat?; embryo suspensor short; 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; cambium from inside the endodermis, storied, xylem with intraxylary phloem developing towards the inside only; vessel elements with simple perforation plates; hairs glandular; leaves pseudoverticillate or in rosettes, with axillary hairs; flowers with the odd sepal adaxial, 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), or 5; nectary also paired glands; A completely adnate to style [= gynostemium], anther thecae set end to end; pollen grains prolate [?level], 2 or 3 nuclear, 3-8-colpate; G [2] (adaxial much reduced), (placentation free-basal), stigma small, dry; integument 4-6 cells across; 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. Divergence & Distribution. Diversification in Stylidioideae has been dated to ca 39 million years ago (Wagstaff & Wege 2002).

Floral Biology. In Stylidioideae the two stamens are adnate to the style, the extrorse anthers being borne near the stigma. 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. Ronse de Craene (2010) describes the flower of Stylidium graminifolium as being obliquely monosymmetric at maturity, and the corolla and parts inside are illustrated as having rotated ca 60^o relative to the calyx, and there is only a single abaxial (in the text described as being adaxial) nectary. See also Erbar (1992) for floral development, which needs more study in the family as a whole.

Ecology & Physiology. 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 carnivorous plants. 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 some embryology, see Subramanyam (1951a), for general information, see Carolin (2006: Stylidiaceae and Donatiaceae), Carlquist and Lowrie (1989: Stylidioideae) and Glenny (2009: Forstera), for protein bodies, see Thaler (1966), and for the testa anatomy of Stylidium, see Tobe and Morin (1996).

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; A.P.G. III 2009).

[Menyanthaceae [Goodeniaceae [Calyceraceae + Asteraceae]]]: inulin, caffeic acid +; vessel elements with simple perforation plates; inflorescence with a terminal flower, signle flowers and then cymes below; 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 more or less psilate; G [2]; integument >10 cells thick, vascular bundle proceeding to the micropyle; endosperm haustoria 0, embryo long; x = 9.

Evolution. Divergence & Distribution. Pollen of all families of this clade - and of some subfamilies of Asteraceae - had differentiated by the Oligocene, and has been found in many places that are fragments of the Gondwanan continent (Barreda et al. 2010a).

Scalariform perforation plates in the vessels, presence of sclereids, binucleate pollen, and multi-nucleate tapetal cells may also be synapomorphies for this clade (Lundberg 2009, which see for other possible synapomorphies).

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 wings, here placed as apomorphies for both groups (or they could be a synapomorphy for the whole clade, being lost later). For inflorescence morphology and evolution in the whole clade, see Pozner et al. (2012).

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

MENYANTHACEAE Dumortier, nom. cons.   Back to Asterales

Aquatic or marsh herbs; flavonols only +, little oxalate accumulation, tannin 0; cork?; vascular bundles separate, often scattered; nodes also 5:5; intercellular canals +; branched sclereids +; petiole bundles arcuate to annular or scattered; leaves also two-ranked, (palmately compound), lamina vernation involute, 2ndary veins palmate, margins usu. with hydathodal glands, crenate, colleters +, leaf base broad, petiole margins ± winged; 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 (two-celled), colpate; placentation parietal, (style 0), stigma bilobed-spathulate, wet; 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 = (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. Divergence & Distribution. For fossil pollen, see Barreda et al. (2010a).

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

Chemistry, Morphology, etc. Kasinathan and Kumari (2001) thought that the leaves of Nymphoides may be opposite, indeed, architecture in that genus is complex (Richards et al. 2010). 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. Many relationships within Nymhoides are as yet poorly supported; whether or not the flowers are heterostylous seems evolutionarily very labile (Tippery & Les 2011).

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.

Evolution. Divergence & Distribution. Kim et al. (2005) date this node to (80-)64.5(-49) million years ago.

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), for chromosome numbers see Semple and Watanabe (2009).

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; (vessel elements with scalariform perforation plates); (medullary vascular bundles +); nodes 1:1 (3:3, 5:5 - cortical bundles = leaf traces); branched sclereids +; indumentum variable, hairs often minutely warty; lamina margins entire to toothed; flowers split-monosymmetric (polysymmetric); C induplicate-valvate, (spurred; not slit - Brunonia), lobes with marginal wings (not Brunonia, Selliera); A basifixed; pollen binucleate, mesocolpia concave; nectary usu. 0; G also [4] ([2]), (placentation ± basal), style curved, with apical hairy pollen-collecting indusium and stylar cup, stigma bilobed; ovules 1 or more/carpel, integument (4 - Levenhookia)6-20 cells across, hypostase +; synergids long, hooked, antipodal cells persistent; 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 - Brunonia); n = 7-9; rpl16 intron missing.

12[list]/440. Very largely Australian, Scaevola alone pantropical (map: van Balgooy 1975). [Photo - Flower]

Anthers connate.

Dampiera (66)

A adnate to base of C.

: Goodenia (180), Scaevola (130). 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 (red in the map: van Balgooy 1975).

• 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. Divergence & Distribution. For fossil pollen, see Barreda et al. (2010a). 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).

Floral Biology & Seed Dispersal. For the diversity of pollen presentation devices in the family, see Erbar and Leins (1988) and Leins and Erbar (2003b, 2010). In nearly all species the pollen is initially enclosed by the indusium, and is pushed out by the elongating style; Brunonia, with its capitate inflorescence and flowers with straight styles, also has pollen caught in a brush formed by stylar hairs.

The seeds of many Goodeniaceae are myrmecochorous (Lengyel et al. 2009, 2010).

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 Subramanyam (1950a) for embryology, Carolin (1978, 2006) for general variation and morphology and Erbar and Leins (1988) and Cave et al. (2010) for the floral development of Brunonia.

Phylogeny. Relationships within Goodeniaceae are becoming fairly well understood (Gustafsson 1996a; Gustafsson et al. 1996; Jabaily et al. 2010). 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 Goodeniaceae paraphyletic.

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

Thanks. I am grateful to Mats Gustafson for comments.

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

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

Evolution. Divergence & Distribution. These two clades may diverge (49-)45.5(-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, and this whole clade may have originated there.

For other similarities or possible synapomorphies, see DeVore (1994) and Lundberg and Bremer (2001, 2003), these include libriform fibres with simple pits and vasicentric parenchyma. 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. Similarly, the inflorescence, although superficially similar in the two families, may be cymose in Calyceraceae and racemose in Asteraceae.

Chemistry, Morphology, etc. Calyceraceae and Barnadesioideae have a similar simple flavonoid profile. For inflorescence morphology, see Pozner et al. (2012).

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 fibres 0; lamina margins entire; inflorescence bracteate; but made up of cymose units (not obvious, but a terminal flower - Acicarpha); K connate, aerenchymatous or spine-like, C outer layer separates and photosynthesises; filaments ± connate, anthers basically free; pollen binucleate, with intercolpar depressions, (spinulate); glands [?nectaries] internally alternating with filaments (0); stigma minutely capitate, pollen deposited on its top; ovule apical, apotropous; 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. Floral Biology. There is secondary pollen presentation of the pump mechanism type (e.g. Erbar 1993).

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), embryology from Dahlgren (1915: one species), floral development from Erbar (1993: one species), and inflorescence development from Harris (1999: two species).

Synonymy: Boopidaceae Cassini

ASTERACEAE Berchtold & J. Presl, 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, lamina vernation often conduplicate or revolute, margins various; inflorescence lacking a terminal flower, involucral bracts in several series; flowers poly- or variously monosymmetric, bracts/bracteoles 0; K reduced, C midveins 0 (+), (3-lobed ray corolla), lobes of disc corollas usu. longer than wide; 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; style branches rounded at apex, stigmatic surface continuous on inner surfaces; ovule basal, epitropous, integument 6-20 cells across; embryo sac with elongated synergid cells, antipodal cells often persistent, proliferating and/or multinucleate; (K deciduous); (testa not vascularized), exotestal cells thickened, palisade or flattened, or undistinguished; endosperm scanty to 0, (nuclear); 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 tricellular hairs, a pedicel cell ["golf-tee cell"], ± globose cell, elongated cell [even on the bristles of the achene]; corolla polysymmetric, (bilabiate [4 + 1 - Arnaldoa]; style strongly curved - Fucaldea stuessyi); pollen (lophate, not spiny), with intercolpar depressions, (± caveate - cavity between the two layers of exine); style glabrous or papillate below bifurcation, style arms short, widened at apex; achene with spines, pappus uniseriate, foxy coloured.

9/94. South America, esp. Andean (map: see Karis et al. 1992; Ezcurra 2002; Funk & Roque 2011). [Photo - Flower.]

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

Evolution. Divergence & Distribution. 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(+); (genome duplication).

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

Synonymy: Mutisiaceae Burnett, Nassauviaceae Burmeister, Perdiciaceae Link, nom. inval.

3. Stifftioideae Panero

(Flowers actinomorphic); (C lobes long, ± coiled).

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]]]]]]: pollen grains spherical; deletion in rpoB gene.

6. Hecastocleidoideae Panero & Funk

Capitulae 1-flowered; flower polysymmetrical, corolla 5-lobed; pollen tricolpate; style branches short; 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 +; x = 10; deletion and insertion in rpoB gene.

Evolution. Divergence & Distribution. The stem 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); exotestal cells palisade; 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 (map: from Hellwig 2004).

Synonymy: Acarnaceae Link, nom. illeg., Carduaceae Berchtold & J. Presl, Carlinaceae Berchtold & J. Presl, Centaureaceae Berchtold & J. Presl, Cnicaceae Vest, Cynaraceae Burnett, Echinopaceae Berchtold & J. Presl, Serrulataceae Martynov, Xeranthemaceae Döll

[Pertyoideae [Gymnarrhenoideae [Cichorioideae [Corymbioideae + Asteroideae]]]]: pollen with unbranched columellae.

8. Pertyoideae Panero & Funk

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

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

[Gymnarrhenoideae [Cichorioideae [Corymbioideae + Asteroideae]]]: pollen echinate.

9. Gymnarrhenoideae Panero & Funk

Plant amphicarpic; heads grouped into synflorescence; style branches long; pappus of bristles.

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 Chevallier

(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), columellae aggregated under spines; 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 Berchtold & J. Presl, Cichoriaceae Jussieu, nom. cons., Lactucaceae Drude, Picridaceae Martynov, Vernoniaceae Burmeister

[Corymbioideae + Asteroideae]: pollen without any columellae spanning space above foot layer [caveate], [?]with internal foramina.

11. Corymbioideae Panero & Funk

Leaves with parallel veins; heads with 1 flower enclosed by two innermost involucral bracts; C with broad, patent lobes; style branches long, hairy outside, apices 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 usu. ecaudate; pollen 25.0-34.3 µm in diam., colpus ends acute, exine 2.3-4.2 µm across, 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 Berchtold & J. Presl, Anthemidaceae Berchtold & J. Presl, Artemisiaceae Martynov, Athanasiaceae Martynov, Calendulaceae Berchtold & J. Presl, Coreopsidaceae Link, nom. inval., Eupatoriaceae Berchtold & J. Presl, Gnaphaliaceae Rudolphi, Grindeliaceae A. Heller, Heleniaceae Rafinesque, Helianthaceae Berchtold & J. Presl, Helichrysaceae Link, nom. inval., Inulaceae Berchtold & J. Presl, Ivaceae Rafinesque, Madiaceae A. Heller, Matricariaceae J. Voigt, Partheniaceae Link, nom. inval., Santolinaceae Martynov, Senecionaceae Berchtold & J. Presl, Tanacetaceae Vest, Tussilagaceae Berchtold & J. Presl, Xanthiaceae Vest

• Asteraceae are very variable vegetatively, but may be recognised by their capitulate and involucrate inflorescences in which the numerous small flowers open first on the outside and are only sometimes subtended by bracts. The anthers are usually fused 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 plumose pappus, a modified calyx, and are frequently dispersed by wind.

Evolution. Divergence & Distribution. Diversification of Asteraceae may have occurred only some 42-36 million years ago, the stem group perhaps being up to 49 million years old (K.-J. Kim et al. 2005); most other suggested ages are similar (Funk et al. 2009c for a summary; see also Torices 2010). For fossil pollen, see Barreda et al. (2010a); depending on the identification of the grains, the first seven subfamilies in the sequence above may all have diverged by the late Eocene about 34 million years ago. A macrofossil, Raiguenrayun, from the Middle Eocene of Patagonia ca 47.5 million years ago also is assignable to this general area, but not to Barnadesioideae (Barreda et al. 2010b, 2012: bird pollination possible!). Bergh and Linder (2009) suggested that diversification of Asteroideae began in the Eocene (56.6-)43.0(-29.6) million years ago; most estimates are younger (Funk et al. 2009c).

Ages for various clades within the family have been suggested (see the papers in Funk et al. 2009a). There seems to have been an (early) Oligocene radiation of the subfamilies (Kim et al. 2005; Funk et al. 2005; Barker et al. 2008; Torices 2010). Diversification probably began in South America, and many tribes arose in the course of a subsequent radiation from Africa (Panero & Funk 2008; Funk 2009; Barreda et al. 2010a, see also Kim & Jansen 1995). For instance, crown Gnaphalieae (Asteroideae) may have begun 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). Within Senecio s. str. there has been intercontinental dispersal between areas with Mediterranean and desert climates (Coleman 2003), while much of the diversification in the Mediterranean-Central Asian Carduoideae-Centaureinae may be Plio-Pleistocene transition and younger (Hellwig 2004).

The iconic giant senecios (Dendrosenecio) of the African mountains are all closely related and not immediately related to Senecio s. str. (Knox & Palmer 1995a, b; Pelser et al. 2007). Much diversification of the high-altitude species of Saussurea may have occurred in the context of the uplift of the Qinghai-Tibetan plateau within the last 14 million years (Y.-J. Wang et al. 2009), and there have been other radiations of Asteraceae in this area (Zhang et al. 2011 and references). Climbing Asteraceae are prominent in montane forests of South America (Gentry 1991).

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 with its high seed set, 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), the diversity of secondary metabolites produced, or some other reason (see also Funk et al. 2009c). More attention should be paid to the significance of pollination of Asteraceae by oligolectic bees (Müller & Kuhlmann 2008: see below). There is also evidence of a palaeopolyploidy event involving most of the family, and subsequent genome duplications near the base of Asteroideae and in Mutisioideae with a distinctive pattern or palaeolog retention - not much in the way of regulatory genes (Barker et al. 2008). However, there are perhaps parallels with Poaceae, which also have large-scale genome duplications in the family 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 rather than treating the family as a unit (see also, perhaps, rate shifts suggested in Smith et al. 2011). 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 have relatively few to very few species, with the exception of Cichorioideae, with ca 3,600 species, and Carduoideae, with ca 2,800 species. Net evolutionary rates within the family are highly heterogeneous.

Floral Biology & Seed Dispersal. Pollinators of Asteraceae might seem not to be very selective, since the frequent and diverse insect visitors to a capitulum apparently trample about on top of it and pollinate indiscriminately as they go, but this is not true. Effective pollination is commonly carried out by a variety of broadly oligolectic solitary bees which form complex and partly learned associations with individual species of Asteraceae (Lane 1996; Müller & Kuhlmann 2008; Kuhlmann & Eardley 2012). For example, within Colletes (plasterer bees) a few species specialize on Asteroideae, but the family is rarely visited by the other species; specialization on flowers of Asteraceae has evolved three or four times there (Müller & Kuhlmann 2008). Pollen of Asteraceae-Asteroideae and -Cichorioideae, at least, may be unsuitable for many potential pollinators. It may lack essential amino acids, have generally lower amino acid and protein concentrations than other pollen, or it may contain harmful secondary metabolites, etc. (Müller & Kuhlmann 2008; Goulson 2010; Sedivy et al. 2011). Indeed, some bees actively avoid collecting pollen from composites, thus female bumblebees may get covered in pollen as they collect nectar, yet they do not transfer that pollen to their corbiculae (Goulson 2010). In general, insect - perhaps especially beee - pollination occurs throughout the family, bird (Cronk & Ojeda 2008) and wind pollination (see below) being uncommon.

Some Asteraceae have a pump (nüdelspritze) mechanism of secondary pollen presentation, 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 (e.g. Erbar & Leins 2000; Leins & Erbar 2006). 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).

The anthers are free in those Asteroideae that are wind-pollinated and the heads have either staminate or carpellate flowers. Since the carpellate heads 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, genome evolution, etc., of the wind-pollinated Artemisia, with its multiple invasions of the Arctic (polyploidy apparently not involved), see Vallès and Garnatje (2005), Sanz et al. (2008) and Trach et al. (2008).

The distinctive capitulate inflorescence of Asteraceae lacks a terminal flower, unlike its immediate relatives (Pozner et al. 2012). In Gorteria the outermost (ray) flowers develop both centrifugally and more slowly than the acropetally-developing more central (disc) flowers (Thomas et al. 2009), and there are several cases where the peripheral flowers develop more or less centrifugally (literature summarized by Pozner et al. 2012). Interestingly, those Calyceraceae which have an inflorescence most similar to that of Asteraceae, and those Asteraceae with some centrifugal development of flowers that can perhaps be linked with the cymose part inflorescences common in the outgroups to Asteraceae, are both derived within their respective families (cf. Pozner et al. 2012). In a number of cases the capitulum has become reduced to a single flower; the single-flowered capitulae may reaggregate into a supercapitulum (this has happened in parallel), and there may occasionally be yet another round of aggregation (Claßssen-Bockhoff 1996b for details; Harris 1994 [tertiary capitulae], 1999; Leins & Gemmeke 1979; Katinas et al. 2008a). Whether a normal capitulum or a supercapitulum, the whole inflorescence functions as a flower in terms of attracting pollinators, and some genes whose expression is normally restricted to individual flowers may be more widely expressed in the capitulum as a whole, as well as in vegetative shoots (Ma et al. 2008).

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). 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 & Harris 1995 and 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).

Ecology & Physiology. Flaveria (Asteroidae) has some species with C₄ photosynthesis, some with C₃, and some intermediate; details of the metabolic shifts involved are quite well understood, and there have been several of these shifts (Bläsing et al. 2000; McKown & Dengler 2009; Ludwig 2011a and references, b, c; Gowik et al. 2011). Christin et al. (2011b) suggest a number of dates for diversification within this clade, all less than 4 million years ago; the repaeeted changes in photosynthetic mechanism may reflect an underlying "predisposition" (McKown et al. 2005).

Plant-Animal Interactions. 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. Pyrrolizidine alkaloids and pentacyclic triterpene saponins obtained from Asteraceae and variously modified are in the secretions of the defensive glands of some Chrysolina and Platyophora beetles (Chrysomelidae, both genera are very speciose: Pasteels et al. 2001; Termonia et al. 2002; Hartmann et al. 2003). Larvae of Nymphalidae-Melitaeini butterflies are commonly found on Asteraceae, and also on Lamiales, from whence they probably moved (Wahlberg 2001; Nylin & Wahlberg 2008), a move perhaps associated with an increase in their diversification rate (Fordyce 2010). 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); here there is no evidence of a shift in diversification rate (Fordyce 2010). 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).

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), 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; Csoka et al. 2005; Brändle et al. 2005 and literature). All these herbivores are particularly abundant in the Mediterranean region, which is perhaps where Carduoideae evolved (Zwölfer 1988). Tephritid flies are particularly noteworthy associates of Carduoideae, either eating fruits, exudates they induce from the plant, or forming galls in the stem or inflorescence (Korneyev et al. 2005, esp. Urophora; Redfern 2011). Introduced insects including a weevil and also species of the tephritid Urophora are often very effective biological control agents of introduced Carduoideae in North America and other parts of the world (Redfern 2011). Tephritid-induced ball galls on Solidago are particularly well known in North America and have been much studied (Abrahamson & Weis 1997); galls on Solidago growing in the prairies are conspicuous in the late summer.

Genes & Genomes. There is accumulating evidence of relatively common and deep hybridisation in Asteroideae in particular (e.g. Fehrer et al. 2007; Pelser et al. 2008, 2010; Soejima et al. 2008; Morgan et al. 2009; Montes-Moreno et al. 2010; Schilling 2011; Smissen et al. 2011). This is going to make genera in some places non-monophyletic.

Economic Importance. Timme et al. (2007) provide a phylogeny of the important genus Helianthus; Simpson (2009) summarised what is known of the otherwise rather slight economic importance of the family.

Chemistry, Morphology, etc. There are tens of thousands of secondary metabolites produced by the family (Calabria et al. 2009 for a convenient summary and entry into the literature), although nothing seems to be known about the secondary chemistry of Hecastocleioideae and Gymnarrhenoideae. See Seaman (1982) for sesquiterpene lactones (Zidara 2008 for those of Cichorieae), Seaman et al. (1990) for diterpenes, Aniszewski (2007) for alkaloids, and Bohm and Stuessy (2001: family) and Sareedenchai and Zidorn (2010: Cichorieae) for flavonoid chemistry; Calabria et al. (2007) produced a phytochemical phylogeny at the tribal level. Sesquiterpene lactones give the bitter taste of many Asteraceae, e.g. Cnicus benedictus, and are sequestered by insects feeding on the family (e.g. Pasteels et al. 2001).

The inflorescence is basically racemose, although ray florets may have their development somewhat delayed relative to the adjacent disc florets (Philipson 1953 and Harris 1995 and references; Leins & Erbar 2003b). 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 (see e.g. Harris 1995 for literature). The involucral bracts in Asteroideae-Senecioneae are uniseriate.

The pappus is often not the first part of the flower to be initiated, often developing well after the corolla (see Mukherjee & Harris 1995, Nordenstam 2008), although otherwise the sequence of initiation is as might be expected. 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, the former perhaps being derived (see observations in Harris 1995). See also Leins and Erbar (2000) and Erbar and Leins (2000) for floral development.

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 (). Koch (1930a, b and references) discusses corolla venation in the family (see also Gustafsson 1995). Many Asteroideae have three-toothed ray florets that give the appearance of being slit-monosymmetric, and 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). 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.

There is much variation in pollen morphology in the family, and it has often been described in terms of pollen "types". However, Blackmore et al. (2009) decompose these types into a number of individually-varying characters, and then discuss the distribution of some of these characters across the Asteraceae tree. Caveate pollen is found in Arctoteae and some Lactuceae, and may be more basal on the tree than it is here (it is a synapomorphy for [Corymboideae + 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. Wortley et al. (2007b) used distinctive pollen characters to help place some genera whose relationships had previously been unclear. See also Skvarla et al. (1977: pollen terminology in Asteraceae), Roque and Silvetstre-Capelato (2001: pollen of Gochnatioideae), Wortley et al. (2008: pollen of Arctotidae-Cichoridoideae), Wortley et al. (2009: a comprehensive bibliography of palynological work in the whole family), Tellería and Katinas (2009: pollen in Mutisia), Osman 2009 (pollen of Cichorioideae-Cardueae), Hong Wang et al. (2009b and references, pollen of Cichorioideae-Cichorieae).

The seed coat is poorly developed, as might be expected; there may be calcium oxalate crystals in the inner layer (Guignard 1893). Guignard (1893) also suggests a), that the ovules may be vascularized, and b), that there is sometimes an antiraphe, as in Centaurea. 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. The embryo of Syneilesis seems to lack cotyledons entirely (Teppner 2001). For fatty acids in the seeds, see Bamai and Patil (1981). For embryo sac development, which I have not thought about, see e.g. Fagerling (1939c): embryo sacs other than the common monosporic 8-nucleate type occur here!

For general information on Asteraceae, see e.g. Carlquist (1976: an interesting summary of variation associated with a tribal classification), K. Bremer (1987, esp. 1994 [a monograph, but the major classification is rather different from that followed here], 1996 [subfamilial groupings]), Hind et al. (1996), and Jansen et al. (1991: variation in the context of phylogeny); Anderberg et al. (2006) summarize the variation in the family. Funk et al. (2009a) discuss biogeography and much more, each tribe in the family is treated in detail, and there is also a glossary. See also Goldflus (1898-9: antipodal cells), Harris (1995: inflorescence and flower development), Watanabe et al. (2007: chromosome numbers), Katinas et al. (2008b: Mutisioideae, but not in the sense used here - general morphology), Thomas et al. (2009: development of Gorteria flowers), Semple and Watanabe (2009: chromosome numbers). For possible plesiomorphic/apomorphic characters in the family, see e.g. Hansen 1991; Leins and Erbar (2000), Erbar anbd Leins (2000).

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, 2009c, a metatree) 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), Gruenstaeudl et al. (2009) and Funk and Roque (2011); 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 Centaureinae, see Garcia-Jacas et al. (2001) and Hellwig (2004), of Echinops, see Garnatje et al. (2005: sectional classification) and Sánchez-Jimenéz et al. (2010), and for that of Cousinia (biphyletic) and relatives, see López-Vinyallonga et al. (2009).

Warionia may be sister to all other Cichorieae (Kilian et al. 2009); although no flavonoids have been reported from them, they are diverse in the rest of the tribe (Sareendenchai & Zidorn 2010 - see Zidorn 2008 for sesquiterpene lactones there). 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).

For extensive analyses of the morphological variation in Gnaphalieae, Inuleae and Plucheae, see Anderberg (1991a, b, and c respectively). 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), for Chrysanthemum and other Anthemidae, see Zhao et al. (2010), and for that of the Hawaiian silverswords and their immediate relatives, and their relatives in turn - west North America tarweeds (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, relationships began to be disentangled by Bayer et al. (2000). Helichrysum is polyphyletic (Galbany-Casals et al. 2004, 2009; Bergh & Linder 2009), and Galbany-Casals et al. (2010) and Montes-Moreno et al. (2010) further clarify relationships in this tribe (see also immediately below). For a phylogeny of Anthemidae in the southern hemisphere, see Himmelreich et al. (2008 and references), and for a delimitation of Anthemis itself, see Lo Presti et al. (2010). Relationships within the huge genus Senecio are beginning to be disemtangled (Pelser et al. 2006, esp. 2007, see also Pelser et al. 2010). For the phylogeny and evolution of Artemisia, originally Eurasian, see Vallès et al. (2003) Vallès and Garnatje (2005), Sanz et al. (2008), Pellicer et al. (2010b: genome size, etc., 2011) and Garcia et al. (2011: North American taxa). 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), for Machaerantherinae, see Morgan et al. (2009), and for the Hinterhubera group, see Karaman-Castro and Urbatsch (2009: groupings geographic). For a phylogeny of Euryops (Senecioneae), see Devos et al. (2010).

Finally, note the relatively common and deep hybridisation in Asteroideae in particular as evidenced by incongruence between topologies based on different genomes, and this is making life for those interested in phylogeny reconstruction rather interesting (e.g. Fehrer et al. 2007; Pelser et al. 2008, 2010; Morgan et al. 2009; Montes-Moreno et al. 2010). Thus there is significant incongruence between relationships suggested by plastid and nuclear sequences in Senecioneae, probably due to ancient hybridisation rather than incomplete lineage sorting (Pelser et al. 2010). Smissen et al. (2011) suggest that complex allopolyploidy may have been involved in the origin of at least four clades in Gnaphalieae, one of which is now globally distributed and that together encompass more than half the ca. species of the tribe. There has also been chromosome reduction there, e.g. from n = 12 to n = 3 in Podolepis (Smissen et al. 2011 and references).

Classification. Panero and Funk (2008) present the subfamilial classification followed above (there are, of course, alternative classifications in the literature - e.g. Jeffrey 2004); it is similar in basic structure to that in Funk et al. (2009b: as 43 tribes, see the accounts there). 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)? In the latter case, Senecio species are found in over eight clades, but even so Senecio s. str., at ca 1,000 species, is paraphyletic (Pelser et al. 2007). Substantial adjustments to generic limits will also be needed in Asteroideae-Inuleae-Inulinae (Englund et al. 2009).

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).

Thanks. I am grateful to Jose L. Panero for comments.