EXTANT SEED PLANTS

Plant woody, evergreen; nicotinic acid metabolised to trigonelline; primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins rich in guaiacyl units; true roots present, xylem exarch, branching endogenous; arbuscular mycorrhizae +; shoot apical meristem complex; stem with ectophloic eustele, endodermis 0, xylem endarch, branching exogenous; vascular tissue in t.s. discontinuous by interfascicular regions; vascular cambium + [xylem ("wood") differentiating internally, phloem externally]; wood homoxylous, tracheids +; tracheid/tracheid pits circular, bordered; sieve tube/cell plastids with starch grains; phloem fibers +; stem cork cambium superficial, root cork cambium deep seated; nodes ?; stomata ?; leaf vascular bundles collateral; leaves spiral, simple, axillary buds?, prophylls [including bracteoles] two, lateral, veins -5(-8) mm/mm²; plant heterosporous, sporangia eusporangiate, on sporophylls, sporophylls aggregated in indeterminate cones/strobili; true pollen [microspores] +, grains mono[ana]sulcate, exine and intine homogeneous, ovules unitegmic, crassinucellate, megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development first endo- then exosporic, tube developing from distal end of grain, to ca 2 mm from receptive surface to egg, gametes two, with cell walls, with many flagellae; female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large", first cell wall of zygote transverse, embryo straight, endoscopic [suspensor +], short-minute, with morphological dormancy, white, cotyledons 2; plastid transmission maternal; two copies of LEAFY gene, PHY gene duplication, mitochondrial nad1 intron 2 and coxIIi3 intron present.

MAGNOLIOPHYTA

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

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

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

ERICALES [ASTERID I + II]: (corolla tube in flower), ovules tenuinucellate.

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

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

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

Polyacetylenes are sporadic, occuring in all three orders, but always in much embedded clades. Although in the past groups with them have been considered thereby related, the argument has little force now. The position of early initiation of the corolla tube on the tree is quite uncertain. Although a number of families in the next three orders have members with such initiation, not only is sampling within them poor, but corolla initiation in the unassigned families - which are probably basal to the orders themselves - is largely unknown, while in the asterid I group both Oleaceae and Rubiaceae, "basal" or almost so in their orders, have early initiation, while Garryales and associated families are unkown (Leins & Erbar 2003b for a summary). 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., that are at the base of the Apiales + Dipsacales (Lundberg 2001e: see Escalloniaceae, paracryphiaceae, ), but most of these have not been sampled for this gene (but see Escallonia).

Unplaced:

Bruniaceae, Columelliaceae.   Back to Main Tree

Bruniaceae [Columelliaceae + Desfontainiaceae]: plant woody; nodes 1:1; flowers polysymmetric, anthers basifixed.

Backlund (1996) included Bruniaceae and Columelliaceae in Dipsacales, but other relationships have been suggested (Gustafsson et al. 1996; Backlund & Bremer 1997; Pyck & Smets 2000; Bell et al. 2001). Indeed, a position of Columelliaceae s.l. (inc. Desfontainiaceae, but not Bruniaceae, in or near Dipsacales was suggested by Bremer et al. (2001) and especially by Lundberg (2001e), although the support was at best moderate. In the first eight versions of this site it was thought appropriate to place Columelliaceae s.l. immediately before the ordinal characterisation of Dipsacales as it made morphological "sense". For instance, they all have opposite leaves, and Columellia has amoeboid tapetum (cf. Bremer et al. 2001) although Desfontainia does not (Maldonado de Magnano 1986a). If the pair are not immediately related to Dipsacales, the substantial similarity that they have with Dipsacales may indicate either substantial homoplasy or a suite of lower-level synapomorphies in the asterid II group of which there is currently no indication. Indeed, Bruniaceae are sister (1.0 Bayesian p.p.) to [Columelliaceae + Desfontainiaceae] in many analyses in Whitworth et al. (2008), although not when coding coding chloroplast sequences were used alone, and so this where they are placed now.

BRUNIACEAE Berchtold & J. Presl, nom. cons.   Back to Unplaced

Ericoid shrubs; myricetin +, iridoids?; stomata cyclocytic, orientation transverse; leaves spiral, margins entire, apex glandular, stipules colleter-like or 0; inflorescence spicate or (involucrate and) capitate (flowers single); flowers often small, (4-merous), (K connate), C often clawed (connate), A (adnate to C), anthers sagittate (not Audouinia), pollen 3-10 colporate, disc +/0, G 1 or [2 (3)], (superior), ?oblique, placentation apical-axile, 1-4(-12) pendulous weakly crassinucellate ovules/carpel, micropyle long, endothelium 0, hypostase +, (styles +, style 0), stigma capitate; fruit a schizocarp, achenial or nut-like, single-seeded, or capsule, also splitting adaxially, K persistent; seed (arillate), coat ?; endosperm type?, haustoria?; n = 10-11 (21, 23).

12[list]/75. South Africa, almost entirely the Cape Province, also KwaZulu-Natal (Map: from Claßen-Bockhoff 2000). [Photo - Flowers.]

• Bruniaceae are ericoid shrubs from the Cape region of South Africa that can be recognised by an often black projection at the apex of the entire leaf blade; the leaves are spirally arranged. The often small flowers are frequently aggregated into more or less capitate inflorescences, the adaxial surfaces of the petals showing a diversity of swellings or ridges. The ovary is more or less inferior.

There is a striking similarity between the flowers of Actinocalyx, from the Upper Cretaceous (Santomasian/Campanian) of Sweden, and those of Bruniaceae (Hall 1987).

Bruniaceae are poorly known. The leaf apex has suberised cells produced by a localised cork cambium. The corolla "tube" is at least sometimes formed by adnation of the filaments to adjacent free petals, but the petals are developmentally initally free (Quint & Claßssen-Bockhoff 2006b). The androecium is often weakly monosymmetric, the abaxial pair of stamens being larger than the others. Gynoecial variation is considerable; in genera like Berzelia where there is only a single carpel (?pseudomonomery), there is only a single locule, ovule and style. The ovules may be pleurotropous (van Tieghem 1898). There are only one or two cells between the megaspore mother cells and the nucellar epidermis.

For the possible association of Bruniaceae with Asterales, see Lundberg (2001e). Bruniaceae have often been placed with the South African Grubbiaceae, here in Cornales (see Hall 1987 for some references), but the similarities between the two reflect the fact that they are found in similar habitats.

There is strongly supported phylogenetic stucture within the family: [Linconieae (Linconia only, anther thecae apically connate and sterilised) [Audoinieae (anther thecae adnate to connective their entire length) + Brunieae]] (Quint & Claßssen-Bockhoff 2006a).

Some information is taken from from Saxton (1910 - no endothelium is shown), Jay (1968: chemistry), Dahlgren and van Wyk (1988), Gregory (1998: anatomy), Endress and Stumpf (1991), Claßssen-Bockhoff (2000: detailed discussion of inflorescences), and Quint and Claßssen-Bockhoff (2006b: floral ontogeny). For a somewhat dated monograph, see Pillans (1947), for wood anatomy, see Carlquist (1978b).

Synonymy: Berzeliaceae Nakai - Bruniales Dumortier - Brunianae Doweld

COLUMELLIACEAE D. Don, nom. cons.   Back to Bruniales

Cork cambium deep-seated; pericyclic fibers 0; petiole bundles arcuate; bud without scales; leaves opposite, margins gland-toothed, bases meeting; inflorescence terminal, cymose; flowers medium-sized, anther connective well-developed, style long, many ovules/carpel; K persistent in fruit; seeds many, anticlinal exotestal cells much thickened, with plasmodesmata.

2/5. Costa Rica to Chile, montane in the tropical part of this area.

Columellia Ruíz & Pavón

Shrubs or trees; plant bitter tasting, iridoids 0; central petiole bundle much the largest; cuticle waxes as tubules; leaves (with glands on lower surface), teeth glandular (0); flowers (4-8-merous), K ± valvate, A 2, latrorse, thecae semicircular, attached their length to expanded connective, tapetum amoeboid, G [2], largely inferior, placentation intrusive parietal or lobed-axile, endothelium poorly developed, stigma with two broad lobes, almost cup-like; fruit septicidal and part loculicidal down the sides; exotestal cells elongated; endosperm type?; n = ?

1/4. S. Colombia to Bolivia, in the Andes (Map: from Brizicky 1961). [Photo - Flower]

Desfontainia Ruíz & Pavón

Shrubs; route I secoiridoids +, tannin 0; scattered sclereids in the pericyclic position; leaves conduplicate, teeth spiny, colleters +; flowers usu. single; K connate at base, C contorted, A adnate in throat, filaments stout, thecae ± embedded in connective, G [5(-7)], opposite petals, placentation axile but apically parietal, ovules crassinucellate, stigma only slightly expanded; fruit a berry; exotestal cells polygonal, outer walls pectic, other walls lignified; endosperm haustoria small; n = 7.

1/1: Desfontainia spinosa. Costa Rica to Chile. [Photo - Flower]

The intervascular pits of Desfontainia are scalariform or circular and bordered, not vestured. All the cells in even the young stem are slightly lignified. Although both Desfontainia and Viburnum have similar, smooth pollen orbicules, these are quite widely distributed in Gentianales, at least (Vinckier & Smets 2002). Maldonado de Magnano (1986a) suggests that the ovules are weakly crassinucellate, with a single layer of nucellar cells between the megaspore and the nucellar epithelium. An endothelium is at most poorly developed, and there is a large haustorial suspensor. Myricetin? The two stamens of Columellia are the adaxial pair and have latrorse, semicircular thecae attached their length to an expanded connective; the anthers are described as being extrorse by Backlund and Donoghue (1996).

The relationships of Columelliaceae have long been uncertain; the family was placed in Rosales by Cronquist (1981) and in Hydrangeales by Takhtajan (1997). Although Desfontainiaceae were usually included in Loganiaceae, palynology and wood anatomy suggested a relationship to Columelliaceae s. str. (Mennenga, in Leeuwenberg 1980), and this was strongly supported by rbcL and other data (see B. Bremer et al. 1994; Backlund & Bremer 1997; Bell et al. 2001).

For Columellia information was taken from Hasselberg (1937), Stern et al. (1969: anatomy), Backlund and Donoghue (1996: general), and Gregory (1998: anatomy), for waxes, see Theisen and Barthlott (1994) and Fehrenbach and Barthlott (1988: cuticle platelets as ribbons and rodlets); see also Zak & Jaramillo 3266. For Desfontainia information was taken from Backlund and Donoghue (1996) and Hasselberg (1937); see also Qin 710 and Zarucchi et al. 5195.

ASTERALES Lindley  Main Tree, Synapomorphies.

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

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.

Absence of apotracheal parenchyma and x = 9 may also be features of Asterales (Lundberg & Bremer 2001; Bremer et al. 2001). For a study of petal vasculature within the order, which shows interesting variation, see Gustafsson (1995). Tobe and Morin (1996) summarise embryological knowledge of the order (they include Sphenocleaceae, for which, see Solanales here). 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. Monosymmetry is often associated with a slit the length of the corolla. Several families have forms of secondary pollen presentation (Erbar & Leins 1995a; Leins 2000; Leins & Erbar 1997, Erbar 2003b: Yeo 1993 for a general summary). Leins (2000) and Leins and Erbar (2006) in particular discuss in considerable detail the evolution of secondary pollination presentation mechanisms. Variation of ovary position in the order is considerable. For integument thickness, see Inoue and Tobe (1999) and for pollen, see Polevova (2006). For a general discussion of variation in the order, see J. Kadereit (2006).

Asterales are basically Takhtajan's (1997) Asteridae, with the addition of sundry Hydrangeales. There is quite a lot of phylogenetic structure in the order, but with rather weak support, as was suggested by D. Soltis et al. (2000; see also Olmstead et al. (2000). More recent studies have improved support for many clades, although there is still a basal polychotomy (Kårehed et al. 2000; Lundberg 2001a, b; Kårehed 2002a; especially Bremer et al. 2001 and Lundberg & Bremer 2001, 2003), as is shown in the tree here. Note that Olmstead et al. (2000) and B. Bremer et al. (2002) suggest a sister group relationship between Campanulaceae and Stylidiaceae (but not Donatia, which goes elsewhere!), and the latter authors suggest that Pentaphragmataceae are also linked to this clade. On the other hand, there are suggestions that Rousseaceae, Pentaphragmataceae and Campanulaceae are together sister to the other Asterales (Lundberg & Bremer 2003), although the support for this is not very strong, while Soltis et al. (2007a) found Campanulaceae to be sister to rest of asterales (1.0 p.p.). 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).

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

Synonymy: Alseuosmiales Doweld, Ambrosiales Dumortier, Brunoniales Lindley, Calycerales Reveal, Campanulales H. G. L. Reichenbach, Carduales Small, Goodeniales Lindley, Menyanthales Takhtajan, Pentaphragmatales Doweld, Phellinales Doweld, Rousseales Doweld, Stylidiales Reveal - Asteranae Takhtajan, Campanulanae Reveal, Phellinanae Doweld - Asteridae Takhtajan - Asteropsida Brongniart, Campanulopsida Bartling

ROUSSEACEAE Candolle   Back to Asterales

Leaf margins gland-toothed; A free, 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 +; young stem with separate bundles; petiole bundle cylindrical; bud scales +; hairs tufted-stellate and glandular-peltate; leaves opposite, base broad; flowers large, (4-merous), K valvate, C connate, anthers basifixed, attached their entire length to stout connective, sagittate, extrorse, pollen 6- or 8-porate, tectum complete, G [5-7], style expanding apically, stigmatic lobes narrower, erect; fruit a berry, K persistent; seed coat with thick-walled exotesta, the rest crushed; micropylar haustorium +, embryo long; n = ?

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

Mauritzon (1933) suggested that Roussea might have bitegmic ovules.

For a summary of what is known about the plant, see Koontz et al. (2006).

2. CarpodetoideaeJ. Lundberg

Trees; chemistry?; young stem with separate bundles; 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; n = 14, 15.

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

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).

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.

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).

Rousseaceae may be sister to Campanulaceae (Kårehed 2002a). For the circumscription of the family, see Lundberg (2000a).

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 summarised by Lundberg (2001a). The anthers in particular are quite unlike those of ex-Saxifragaceae (Escalloniaceae in particular - see Apiales relatives here) that are otherwise somewhat similar to Rousseaceae. Carpodetaceae, another group often thought of as woody saxifrages, were included in Hydrangeales by Takhtajan (1997).

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).

CAMPANULACEAE Jussieu, nom. cons.   Back to Asterales

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

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

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

1. Nemacladoideae M. H. G. Gustafsson

Annuals (perennials); C 3 [adaxial]: 2 [abaxial lobes], A (adnate to C), filaments forming tube, anthers free, flip back after pollen release, pollen tricolporate or 6-colpate, G also [3], half inferior, style head bending towards or away from median K; (fruit circumscissile - Parishella); n = 9.

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

Nemacladus has groups of remarkable reflexed pseudonectaries at the bases of two filaments. It is not known if the style hairs are retractile.

Synonymy: Nemacladaceae Nuttall

2. Campanuloideae Burnett

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

50/1050: Campanula (420, 100 in Turkey alone), Wahlenbergia (260), Adenophora (65), Codonopsis (60). Especially N. temperate Old World, very few in the Australia-New Zealand area (Map: from Hultén 1971; Thulin 1975; Shulkina 1978).

Codonopsis, Cyananthus, etc.

Leaves often opposite; pollen colpate/colporate; n = 7-9 (17).

Campanula, Wahlenbergia, etc.

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

The latter clade can be divided into two main clades centered on Campanula s. str. and Rapunculus, however, generic limits are a mess, neither Campanula nor Wahlenbergia being monophyletic (Haberle et al. 2008b). Generic limits in Campanuleae in particular 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 (Roquet et al. 2008). Phyteuma has coherent corolla lobes although the corolla is open laterally; the style hairs are only partly retractile. The nectar of some Campanuloideae may be brightly colored and then the filament bases are not persistent. Some species of Wahlenbergia have an almost superior ovary.

For node-stem anatomy, see Col (1904), for fruit morphology, see Kolakovsky (1985), for rearrangements in the chloroplast inverted repeat, see Cosner et al. (1997), for relationships within Campanuloideae, see Eddie et al. (2002, 2003) and Cosner et al. (2004), for morphology, see Shulkina et al. (2003).

Synonymy: Cyananthaceae J. Agardh, Jasionaceae Dumortier

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

3. Lobelioideae Schönland

(Annuals) perennials, herbs to small trees; chelidonic acid, pyrrolizidine alkaloids +, p-coumaric acid, caffeic acid 0; leaves supervolute [Lobelia]; flowers (split-monosymmetric); resupinate by pedicel (hypanthium) torsion, C 2:3, 0:5, filaments connate at least apically, anthers connate, pollen reticulate-striate, style with brush hairs and pollen in pollen box, pump mechanism [Nudelspritze, stigma wet; (fruit a berry; a circumscissile capsule; dehiscing through sides,); n = (6-)7(-13).

29/1200: Lobelia (400+), Siphocalymus (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]

There is much of interest in the floral biology and biogeography of Lobelioideae. Molecular data suggest that Lobelia is wildly paraphyletic (Knox & Muasya 2001; Antonelli 2008; Knox et al. 2008: within South American lobelioids, Centropogon [e.g.] paraphyletic, but little support for relationships in general), and that the pachycaul giant lobelias are derived from herbaceous ancestors (Knox et al. 1993) Giant lobelias from widely separated parts of the globe may be in the same immediate clade (Antonelli 2008), indeed, some South American taxa may be derived from within the African giant lobelia clade, although the relationships of the giant lobelias and the biogeographic implications of these relationships need more detailed study. Knox et al. (2006, no Nemocladoideae or Cyphocarpoideae included) suggested that [Cyphia + Lobelioideae] originated in southern Africa, dispersing quite widely, and with at least two returns to Africa. Givnish et al. (2006a) note that the some 120 species of Hawaiian Lobelioideae appear to have evolved from a single ancestor a mere ca 13 million years before present.

Stein (1992) discusses extrafloral nectaries (on the outside of the inferior ovary, mostly lower altitude, where the ants are) and pollination by sickle-bill hummingbirds (Eutoxeres) at higher altitudes in Andean Centropogon (Lobelioideae); Heliconia (Heliconiaceae) is the nectar resource at lower altitudes. Since the pedicel of Lobelia and its relatives is twisted (resupinate), the flowers appears to have a "normal" orientation with the median petal abaxial.

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

4. Cyphocarpoideae Gustafsson

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

1/3. Chile.

Synonymy: Cyphocarpaceae Reveal & Hoogland

5. Cyphioideae Schönland

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

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

Sympetaly 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.

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

For evolution of the secondary pollen presentation devices, see Leins and Erbar (see esp. 2003b); sampling (inc. Nemocladoideae, some Lobelioideae) is still incomplete. Associated with the secondary pollen presentation that occurs throughout the family, the flowers are protandrous. Leins and Erbar (2003b) note that the flowers in Campanulaceae s.l. are initially polysymmetrical in bud.

In Ostrowskia (Campanuloideae) the anther has a placentoid and the integument is massive (Kamelina & Zhinkina 1998). The chloroplast gene accD (= ORF512, zpfA) has been lost (Doyle et al. 1995 and references) in at least some Campanulaceae.

The groupings above are only tentative, and A.P.G. II suggests as an option keeping Lobeliaceae separate from Campanulaceae. 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), 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.

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

Further general information on the subfamilies was taken from Schönland (1889), Wimmer (1968) and Lammers (1998 and especially 2006); for pollen see Dunbar (1975a, b), for embryology, etc., see Subramanyam (1970: possible taxonomically interesting differences in cell number of the haustoria), for the protein bodies in the nuclei, see Bigazzi (1986) and Haberle (1998), for the evolution of secondary pollen presentation, see especially Leins and Erbar (2006), for the inverted repeat and chloroplast genome rearrangement, see Knox and Palmer (1999: Cyphocarpus, Nemacladus, etc., were not studied, Cyphia was) and Haberle et al. (2008a), and for a world checklist and bibliography, see Lammers (2007).

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

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

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

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

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).

There is weak support for the association of Pentaphragmataceae with a Stylidiaceae + Campanulaceae clade in B. Bremer et al. (2002), although Donatia itself is there very weakly linked with Alseuosmiaceae et al.

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

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

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

ALSEUOSMIACEAE Airy Shaw   Back to Asterales

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

5[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.

Ellagitannins are reported from Alseuosmia (Kårehed 2006 for references); this should be checked. Most Alseuosmiaceae have rayless wood, living mature fibres with stored starch, and stem with an endodermis. The pollen of Alseuosmia linariifolia is described as being tricolpate, with an ectexine made up of a thick, tubercular tectum and massive, spherical columellae (Polevova 2006); whether this can be generalised to the family is unclear (see also Kårehed 2006). Generic limits are in some dispute (Tirel 1996).

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

Genera of Alseuosmiaceae have previously been placed in Caprifoliaceae, Rubiaceae, Rutaceae, Ericaceae, Epacridaceae, etc.; the family is included in Hydrangeales by Takhtajan (1997).

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

Synonmy: Platyspermataceae Doweld

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

PHELLINACEAE Takhtajan   Back to Asterales

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

1[list]/12. New Caledonia.

The ovules are hemitropous to campylotropous and the guard cells are huge, with inner and outer stomatal ledges. Phelline is embryologically poorly known.

Takhtajan (1997) placed the family in Icacinales, and says the leaves are mostly estipulate.

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

ARGOPHYLLACEAE Takhtajan   Back to Asterales

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

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

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

Along with Cornaceae, Argophyllaceae were placed in Hydrangeales by Takhtajan (1997).

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

Synonymy: Corokiaceae Takhtajan

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

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

6[list]/245. Scattered in South East Asia to New Zealand, S. South America (Map: from Good 1974).

Donatioideae B. Chandler

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

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

In older stems of Donatia the cortex is very thick, and the vascular tissue forms a narrow cylinder in the center (Chandler 1911).

Synonymy: Donatiaceae B. Chandler, nom. cons.

Stylidioideae

Herbs (climbers), cushion plants; cork also outer cortical; vascular bundles closed, scattered or in a single ring; 2ndary thickening anomalous; cambium storied; vessel elements with simple perforation plates; hairs glandular; leaves pseudoverticillate or in rosettes; flowers with median K abaxial, but often semi-resupinate, split-monosymmetric (polysymmetric); K connate, C connate, early tube formation, often with coronal appendages, nectary also paired glands, A 2, completely adnate to style, anther thecae set end to end, pollen 2 or 3 nuclear, 3-8-colpate, G [2] (adaxial much reduced), (placentation free-basal), synergid cells elongated, stigma small, dry; 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, also outh East Asia, Malesia, and S. South America (Map: see Erickson 1958). [Photo - Flower]

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 1981). The fertile stamens are the adaxial pair.

For general information, see Carlquist (1989), for the testa anatomy of Stylidium, see Tobe and Morin (1996), for floral development, see Erbar (1992), and for the literature on stylar movement, see Findlay and Findlay (1989).

• 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 adnate to the style, the extrorse anthers being borne near the stigma, 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.

There are suggestions that Stylidium may be carnivorous, insects being trapped by the glandular hairs, which also show yeast-extract stimulated protease acivity; the plants grow in acid, nutrient-poor soil like other carnivores. However, uptake of nutrients by the plant from the insects has yet to be demonstrated (Darnowski et al. 2006).

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).

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 are very small, and their venation is acrodromous. For protein bodies, see Thaler (1966).

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. They 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).

Some anatomical details can be found in Thouvenin (1890); for general information, see Carolin (2006).

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

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

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

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

Usu. aquatic herbs; flavonols only +, little oxalate accumulation, tannin 0; cork?; vascular bundles separated; nodes also 5:5; branched sclereids +; petiole bundles arcuate to annular or scattered; leaves also two-ranked, involute, venation palmate (palmately compound), margins usu. with hydathodal glands, crenate, 2ndary veins often palmate, colleters +, base broad, (with lateral flanges - Nephrophyllidium); flowers heterostylous (not), K basally connate, C lobes with marginal wings (fringed), (with fringed scale-like staminodes), anthers sagittate, tapetal cells with fused nuclei, pollen colpate, G [2], 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 = 9 (17), protein bodies in nuclei; rpl2 intron missing.

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

• Menyanthaceae are aquatic plants with 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.

Kasinathan and Kumari (2001) thought that the leaves of Nymphoides may be opposite. The vascular anatomy of the flower implies a basic monosymmetry - the lateral corolla traces are fused (Wood & Weaver 1982). Johri et al. (1992) suggest that the endosperm stores starch.

Has the superior ovary of Menyanthaceae been derived from a more of less inferior ovary???

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

The iridoids of Menyanthaceae differ chemically from those of Gentianaceae, in which Menyanthes, etc., 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.

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).

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

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).

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

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

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

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

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

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).

Relationships within Goodeniaceae are becoming fairly well understood (Gustafsson 1996a; Gustafsson et al. 1996). There are two main clades, Lechenaultia and allies and 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, and lack of endosperm, and has a single, basal ovule, etc. (some of these features might seem to suggest relationships with Asteraceae, Gustafsson 1996a). However, these features are derived within Goodeniaceae and the exclusion of Brunonia would make the family paraphyletic.

For the diversity of pollen presentation devices in the family, see Leins and Erbar (2003b, inc. Brunoniaceae); see Carolin (1978, 2006) for general variation and morphology.

Synonymy: Brunoniaceae Dumortier, nom. cons. (inflorescence capitate; flowers polysymmetric, lateral veins of the C unite in the receptacle (Erbar 1997), no hairs on indusium, G [2], 1 basal ovule, testa thin-walled, compressed, endosperm 0), Scaevolaceae Lindley

Calyceraceae + Asteraceae: flowers small, C tubular, commissural veins connate, (median veins 0), filament collar +, G [2], with 1 ovule; fruit a cypsela, K persistent, modified, involved in dispersal.

These two clades may diverge 49-42 million years before present (K.-J. Kim et al. 2005).

For other similarities or possible synapomorphies, see DeVore (1994) and Lundberg and Bremer (2001, 2003), these include libriform fibers with simple pits and vasicentric parenchyma. Calyceraceae and Barnadesioideae have a similar simple flavonoid profile. Pesacreta et al. (1994) suggest similarities in the micromorphology of the filaments and connective bases between at least some members of Calyceraceae and Asteraceae. Placement of characters like pollen with intercolpar depressions on the tree is difficult to ascertain (see also DeVore 1994; DeVore et al. 2000); DeVore and Skvarla (2008) suggest that pollen characters thought to suggest a relationship between the two families are different in detail and are therefore not homologous. Although both families have but a single ovule, the position and orientation of the ovule is such that the single ovule condition may well have been derived independently (e.g. Cronquist 1981 for an extreme taxonomic interpretation of this position).

CALYCERACEAE Richard, nom. cons.   Back to Asterales

Herbs; ?flavonols 0; cork?; vascular bundles separated; nodes ?; pericyclic fibers 0; leaf margins entire; inflorescence an involucrate and bracteate capitulum made up of cymose units (except Acicarpha, basically racemose and centripetal); K connate, aerenchymatous or spine-like, C outer layer separates and photosynthesises, filaments ± connate, glands [?nectaries] internally alternating with filaments (0), anthers basically free, pollen deposited on top of stigma, binucleate, with intercolpar depressions, ovule apical, pendulous, apotropous, stigma minutely capitate; 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 (like Asteraceae-Barnadesioideae) (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.

There is secondary pollen presentation of the pump mechanism type. 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), details of morphology from Pontiroli (1963), and embryology from Dahlgren (1915: one species!).

Synonymy: Boopidaceae Cassini

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

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

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

1. Barnadesioideae (D. Don) Bremer & Jansen

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

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

The corolla hairs 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.

For phylogenetic relationships, see Urtubey and Stuessy (2001) and in particular Gustaffson et al. (2001); for corolla morphology, see Stuessy & Urtubey (2006).

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

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

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

2. Mutisioideae Lindley

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

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

Synonymy: Mutisiaceae Burnett, Nassauviaceae Burmeister, Perdiciaceae Link

3. Stifftioideae Panero

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

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

4. Wunderlichioideae Panero & Funk

Style and style arms glabrous; deletion in rpoB gene.

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

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

5. Gochnatioideae Panero & Funk

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

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

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

6. Hecastocleidoideae Panero & Funk

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

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

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

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

7. Carduoideae Sweet

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

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

For a phylogeny of Cardueae, see Susanna et al. (2006).

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

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

8. Pertyoideae Panero & Funk

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

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

Gymnarrhenoideae [Cichorioideae [Corymbioideae + Asteroideae]]: ?

9. Gymnarrhenoideae Panero & Funk

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

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

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

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

10. Cichorioideae Chevalier

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

224/3600: Vernonia (800-1000, or in 20 subtribes, 2/3 genera mono- or ditypic!: see Keeley et al. 2007 for a phylogeny; Robinson 2006 and references for genera), Crepis (200), Jurinea (200), Scorzonera (175), Lepidaploa (115), Tragopogon (110), Lessinganthus (100), Hieracium (90-1000+: apomixis), Lactuca (75), Vernonanthura (65), Hypochaeris (60), Sonchus (60), Taraxacum (60-500+: apomixis). World-wide.

For a phylogeny of Tragopogon and its relatives, see Mavrodiev et al. (2005), and of the African Goteriinae, see Funk and Chan (2008). In Sonchinae, Sonchus is para/polyphyletic (Kim et al. 2006, 2007), with island-dwelling woody forms 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).

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

11. Corymbioideae Panero & Funk

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

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

12. Asteroideae Lindley

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

1135/16360: Senecio (1000: lump or split? - see Pelser et al. 2006, esp. 2007: both), Eupatorium (1200 [s.l.] or 40 [s.s.]), Helichrysum (600), Artemisia (550: wind pollination, see Vallès & Garnatje 2005 and Sanz et al. 2008 for a phylogeny), 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: polyphyletic, see Cross et al. 2002), 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.

North American Astereae are monophyletic and largely herbaceous (Noyes & Rieseberg 1999). The involucral bracts in Senecioneae are uniseriate. Many Asteroideae have three-toothed ray florets that give the appearance of being slit-monosymmetric. However, they represent a modification of a 2:3 bilabiate corolla in which the adaxial lobes have been suppressed (Weberling 1989). 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. Timme et al. (2007) provide a phylogeny of the important genus Helianthus.

For the phylogeny of the helenioid Heliantheae, see Baldwin et al. (2002), of Inuleae (inc. Plucheeae), see Anderberg et al. (2005), for circum-Mediterranean Anthemidae, also their biogeography, see Oberprieler (2005), and for that of silverswords and their relatives of the Madiinae, see Baldwin and Wessa (2000) and Carlquist et al. (2004, also Madiinae Showcase); Bidens also shows much variability in growth form, etc., on Hawaii, but there is little molecular variation or genetic barriers between the species (Ganders et al. 2000). For a phylogeny of Anthemidae, see Himmelreich et al. (2008 and references). Finally, it is worth noting that there may be relatively common and deep hybridisation in Asteroideae which can make life rather difficult (e.g. Pelser et al. 2008).

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

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

Asteraceae alone contain about 8% of eudicot diversity, and within Asteraceae, Senecionoideae make up the lion's share of species numbers. Given this, there is speculation about what has caused this, whether the development of the capitulum itself, the storage of carbohydrates as unbranched-chain fructans, which contributes to the ability of Asteraceae to live in the rather dry conditions that many of them prefer (John 1996), or some other reason. Diversification of Asteraceae may have been within 42-36 million years, the stem group perhaps being up to 49 million years old (K.-J. Kim et al. 2005, for which see for dates of various other clades within the family; cf. also Poaceae, which also store carbohydrates as fructans... John 1996).

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 (also Crotalaria and some Boraginaceae and Apocynaceae: Edgar et al. 1974; Fiske 1975; Ackery & Vane-Wright 1984; Brown 1987; Weller et al. 1999; Anke et al. 2004); there has been parallel evolution of these alkaloids within Asteroideae. Larvae of Nymphalidae-Melitaeini butterflies are commonly found on Asteraceae, and also on Lamiales, from whence they probably moved (Wahlberg 2001; Nylin & Wahlberg 2008), while caterpillars in a clade of Nymphalidae-Heliconiinae-Acraeini utilise primarily Andean members of this family, probably switching from host plants in the Passifloraceae area (Silva-Brandão et al. 2008). Cecidomyiid gall midges are notably common on the family in North America (Gagné 1989), and galls on Solidago growing in the prairies are conspicuous in the late summer.

Some Asteraceae have a pump (nüdelspritze) mechanism of secondary pollen presentation (?plesiomorphic), other taxa have a brush mechanism (see Leins & Erbar 2003b for a possibly evolutionary sequence of pollen presentation devices). Within Barnadesioideae, sister to the rest of the family, secondary pollen presentation is by simple deposition on the style/stigma (as in at least some Calyceraceae) or by an unspecialised type of brushing mechanism (Leins & Erbar 2006). Although it might seem that pollinators are not very selective, simply trampling about on top of the capitulum and pollinating as they go, this is not true; pollination is commonly by a wide variety of specialist solitary bees which form complex and partly learned associations with individual species of Asteraceae (Lane 1996). Many Carduoideae have thigmotropic stamens, the filaments contracting when touched by the pollinator, the pollen then being forced out of the anther tube. There is a great diversity of breeding systems in the family (e.g. Burtt 1961). The anthers are free in wind-pollinated Asteroideae and the heads have either staminate or carpellate flowers; the latter 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. Apomixis is quite common, as in Hieracium and Taraxacum (see T. absurdum), both Cichorioideae, and Antennaria (Asteroideae).

Most fruits are crowned by the pappus, modified sepals (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. 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.

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 (Lepidoptera), Curculionidae (Colepoptera), Tephritidae (Diptera), Tingitidae (Hemiptera) and Cynipidae (Hymenoptera), are found on Carduoideae of the west Palearctic region, particularly in the Mediterranean region, which is perhaps where the clade evolved (Zwölfer 1988).

Sesquiterpene lactones give the bitter taste of many Asteraceae, e.g. Cnicus benedictus. Caveate pollen also is found in Arctoteae and some Lactuceae, and may be more basal on the tree (it is a synapomorphy for Asteroideae at present). Indeed, Blackmore et al. (1984) noted that it is evident early in development in Gerbera (Mutisieae), but not later, and suggested that pollen grains of Asteraceae might all be basically caveate. There is variation in the orientation of the gynoecium and style branches: carpels superposed, style branches arranged radially to the head surface; carpels collateral, style branches tangential to head surface: see Robinson 1984); details of the distribution of this feature are unknown. 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 capitulum may be reduced to a single flower, and then the single-flowered capitulae reaggregated into supercapitula (Claßssen-Bockhoff 1996b for details; Katinas et al. 2008); for capitulum development, see Leins and Erbar (2003b). Some taxa, including Barnadesioideae, have a midvein in the petal... (see Carlquist 1976; Gustafsson 1995). There are floral bracts in some 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. Buphthalmum has a hollow style (Leins 2000); I do not know how widespread this feature is in Asteraceae.

Syneilesis seems to lack cotyledons entirely (Teppner 2001).

For information on Asteraceae, see e.g. Carlquist (1976: an interesting summary of variation associated with a tribal classification), Skvarla et al. (1977: pollen terminology), K. Bremer (1987, esp. 1994 [a monograph, but the major classification is rather different from that followed here], 1996 [subfamilial groupings]), Seaman (1982: sesquiterpene lactones), Seaman et al. (1990: diterpenes, other references), Jansen et al. (1991: variation in the context of phylogeny), Harris (1995: infloresence and flower development), Hind et al. (1996: general), Y.-D. Kim and Jansen (1995: phylogeny), Bohm and Stuessy (2001: flavonoid chemistry), Panero and Funk (2002, esp. 2008: phylogeny and the subfamilial classification followed here), Jeffrey (2004: an alternative classification), Funk et al. (2005: a supertree), Aniszewski (2007: alkaloids), and Watanabe et al. (2007: chromosome numbers). Anderberg et al. (2006) summarize the variation in the family and enumerate the genera. I am grateful to Jose L. Panero for comments.