Takhtajan Armen. Flowering Plants

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Superorder ASTERANAE 493

(Donatiaceae) or 5-lobed, lobes mostly unequal.

Stamens two or three (Donatiaceae) or only two

(Stylidiaceae); ﬁ laments free (Donatia) or completely

connate with the style into a column; anthers basiﬁ xed,

extrorse, opening longitudinally. Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains 2-celled

or 3-celled, 3–8-colpate (Stylidiaceae) or 3(4)- colporate

(Donatiaceae). Nectary disc or a pair of nectary glands

often present. Gynoecium of 2–3 (Donatiaceae) or two

(Stylidiaceae) united carpels; stylodia free (Donatiaceae)

or connate between themselves and with the ﬁ laments

(Stylidiaceae); ovary inferior, 2- or 3-locular, but in

Stylidiaceae the posterior carpel sometimes reduced or

obsolete and the gynoecium pseudomonomerous with

1-locular ovary. Ovules many in each locule, anatro-

pous, unitegmic, tenuinucellate, with endothelium.

Tapetal cells binucleate. Female gametophyte of

Polygonum-type. Endosperm cellular, with terminal

haustoria. Fruits septicidal capsules or sometimes inde-

hiscent. Seeds with minute embryo and copious ﬂ eshy

endosperm. Storing carbohydrate as inulin, producing

carbocyclic iridoids and caffeic acid. Alkaloids absent.

Iridoids (Stylidiaceae) present; n = 5–16, 18, 24, 26,

28, and 30.

Related to the Asterales.

Key to Families

1 Stamens and stylodia free. Very densely branched

perennial herbs with erect crowded stems forming

compact, often broad, cushions. Roots ﬂ eshy, clit-

like. Vessels with oblique scalariform perforation

plates. Secondary thickening absent or developing

from a conventional cambial ring. Leaves alternate,

imbricate, non-sheating, simple, entire, more or less

hairy at base, without a persistent basal meristem;

mucilage cells abundant in the mesophyll. Stomata

paracytic (Donatia magellanica) or anomocytic (D.

novae-zelandiae), Flowers solitary, sessile, termi-

nal, bisexual, essentially actinomorphic. Calyx 5–7-

lobed, lobes equal or unequal, open, with obconical

tube adnate to the ovary, imbricate. Petals 5–10,

free, oblong-linear or ovate, imbricate, more or less

ﬂ eshy. Stamens two or three, inserted on top of the

ovary within the nectary disc; ﬁ laments free, short

or ﬁ liform; anthers broadly ellipsoidal, extrorse.

Pollen grains 3(4)-colporate. Epigynous disc pres-

ent. Gynoecium of two or three carpels; stylodia

subulate, recurved, with capitate stigma; ovary infe-

rior, 2–3-locular, with numerous (15–100) ovules

on axile placentas near to top of the locule. Fruits

turbinate, indehiscent, 2–3-locular capsules. Seeds

few, with minute embryo and copious, oily

endosperm. Present inulin, iridoids not detected,

n = 24. . . . . . . . . . . . . . . . . . . . . . 1. donatiaceae.

1 Stamens and stylodia not free. Small, rhizomatous

or tuberous, mostly erect or rarely climbing herbs or

rarely subshrubs, more or less xerophytic, occasion-

ally rush-like, often bearing stalked glands with two

to multicellular head; some of glandular cells secrete

mucilage. Vessels with simple perforations.

Secondary thickening anomalous. Leaves alternate

or whorled (Stylidium scandens), imbricate, simple,

entire, linear, often in basal rosettes with ﬂ owers on

a scape (successive rosettes may be separated by a

slightly leafy piece of stem but sometimes crowded

on creeping or shortly erect stems); rosettes some-

times almost bulbous, with aerial roots. Stomata

anomocytic. Flowers in terminal racemes or cymes

or solitary in the upper axils, usually zygomorphic,

bisexual or sometimes unisexual (monoecious,

polygamomonoecious, or dioecious ?). Calyx (2-)5(-

7)-lobed, often covered with stalked viscid glands,

imbricate, persistent. Corolla 5-lobed, lobes imbri-

cate, mostly unequal, the odd lobe originates in a

ventral position and in the zygomorphic genera takes

the form of small or more or less reduced (Stylidium)

or large, hooded (Levenhookia) labellum, but the

ﬂ ower commonly semiresupinate, so that the label-

lum is lateral; the remaining four lobes are more or

less similar in size and shape or heteromorphic; the

corolla throat is often marked by a series of various

glandular appendages. Stamens 2(-3); ﬁ laments free

from the corolla but united with the style to form a

column, often irritable; anthers extrorse. Pollen

grains 2-celled or 3-celled, 3–8-colpate. Epigynous

nectary disc or a pair of nectary glands often pres-

ent. Gynoecium of two carpels; stigmas papillate,

diverging above the anthers; ovary inferior, more or

less completely 2- locular or almost 1-locular, with

axile to free central placentation, or the posterior

locule reduced or suppressed, so that the ovary is

pseudomonomerous; ovules numerous in each loc-

ule, pendulous or horizontal, or ascending. Fruits

ﬂ eshy to non-ﬂ eshy capsules opening from the sum-

mit in two valves, or rarely indehiscent. Seeds

mostly numerous, minute; seed coat thin, 2-layered,

exo-endotestal (Plisko 2005) with small, dicotyle-

donous or sometimes monocotyledonous, straight

494 Subclass VII. ASTERIDAE

embryo and copious, ﬂ eshy oily endosperm. Iridoids

detected; storing inulin, proanthocyanidins, ﬂ a-

vonols (kaempferol and quercetin) present or absent,

n = 5–16, 18. . . . . . . . . . . . . . . . . . 2. stylidiaceae.

1. DONATIACEAE

Chandler 1911. 1/2, New Zealand, Tasmania, sub-

antarctic South America.

Donatia

In spite of some important morphological differ-

ences, including free stamens and 3(4)-colporate pol-

len grains, which are slightly similar to those of Cyphia

(Erdtman 1952; Avetisian 1973), Donatiaceae are

rather closely related to the Stylidiaceae, an afﬁ nity

that is conﬁ rmed by many embryological similarities.

According to Philipson and Philipson (1973: 458), the

embryological details of Donatia are so similar to

those of Forstera and the other members of the

Stylidiaceae described in the literature that the position

of this genus close to the Stylidiaceae is conﬁ rmed.

2. STYLIDIACEAE

R. Brown 1810. 5/170. Mainly extratropical regions of

Australia, also Tasmania, New Zealand, southern and

especially Southeast Asia, and the southernmost part

of South America.

phyllachneae: Phyllachne, Forstera; stylidieae:

Oreostylidium, Levenhookia, Stylidium.

Related to Campanulaceae, but differs in the pres-

ence of carbocyclic iridoids, glandular hairs with two-

to multicellular stalks (Stylidiaceae), anomalous

secondary growth, reduced number of stamens that in

Donatiaceae are free from the style, and extrorse anthers.

However, there are many similarities between the

Stylidiaceae and the Campanulaceae, including pollen

morphology (Erdtman 1952; Bronckers and Stainer

1972; Avetisian 1973), ﬂ oral development (Erbar 1992),

the presence of inulin, and some embryological features.

According to Crete (1951), the haustoria in the

Stylidiaceae arise from endospermal cells by the same

sequence of divisions as in Codonopsis in Campanulaceae.

The embryological similarities between Stylidiaceae

and some members of Campanulales (especially the

Lobelioideae) have been further substantiated by Rosen

(1935, 1949), Subramanyam (1950a, b, 1951a, b, 1953,

1970), and Philipson and Philipson (1973).

Bibliography

Bronckers F and F Stainer. 1972. A l’étude morphologique du

pollen de la famille des Stylidiaceae. Grana 12: 1–22.

Burns GP. 1900. Beiträge zur Kenntnis der Stylidiaceae. Flora

87: 313–354.

Carlquist S. 1969. Studies in Stylidiaceae: new taxa, ﬁ eld obser-

vations, evolutionary tendencies. Aliso 7: 13–64.

Carlquist S. 1981a. Studies in Stylidiaceae: monocotyly in the

family; nomenclatural change. Aliso 10: 35–38.

Carlquist S. 1981b. Types of cambial activity and wood anatomy

of Stylidium (Stylidiaceae). Am. J. Bot. 68: 778–785.

Carolin RC. 1960a. Floral structure and anatomy in the family

Stylidiaceae Swartz. Proc. Linn. Soc. N. S. W. 85: 189–196.

Carolin RC. 1960b. The structures involved in the presentation

of pollen to visiting insects in the order Campanales. Proc.

Linn. Soc. N. S. W. 85: 197–207.

Carolin RC. 1967. The concept of the inﬂ orescence in the order

Campanulales. Proc. Linn. Soc. N. S. W. 92: 7–26.

Carolin RC. 2007. Stylidiaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 8, 614–619. Springer,

Berlin/Heidelberg/New York.

Chandler B. 1911. Note on Donatia novae-zelandiae Hook. f.

Notes Roy. Bot. Gard. Edinb. 6: 43–48.

Darnowski DW, DM Carroll, B Plachno, E Kabanoff, and

E Cinnamon. 2006. Evidence of protocarnivory in trigger-

plants (Stylidium spp.; Stylidiaceae). Plant Biol. 8:

805–812.

Dunbar A. 1975b. On pollen of Campanulaceae and related fam-

ilies with special reference to the surface ultrastructure:

II. Campanulaceae subfam. Cyphioideae and subfam.

Lobelioideae; Goodeniaceae; Sphenocleaceae. Bot. Not.

128: 102–118.

Erbar C. 1992. Floral development of two species of Stylidium

(Stylidiaceae) and some remarks on the systematic position

of the family Stylidiaceae. Canad. J. Bot. 70: 258–271.

Findley N and GP Findley 1989. The structure of the column in

Stylidium. Aust. J. Bot. 37: 81–101.

Gustafsson MHG. 1996. Phylogenetic studies in the Asterales

sensu lato. Acta Universitatus Upsaliensis, pp. 1–36.

Uppsala.

James SH. 1979. Chromosome numbers and genetic systems

in the triggerplants of western Australia (Stylidium;

Stylidiaceae). Aust. J. Bot. 27: 17–25.

Laurent NB, B Bremer, and K Bremer. 1999. Phylogeny and

generic interrelationships of the Stylidiaceae (Asterales),

with a possible extreme case of ﬂ oral paedomorphosis. Syst.

Bot. 23: 289–304.

Leins P and C Erbar. 1990. On the mechanisms of secondary

pollen presentation in the Campanulales-Asterales complex.

Bot. Acta 103: 87–92.

Philipson WR and MN Philipson. 1973. A comparison of the

embryology of Forstera L. and Donatia J. R. et G. Forst.

N. Z. J. Bot. 11: 449–460.

Plisko MA. 2005. Fruit and seed structure of Levenhookia stipi-

tata and Stylidium species (Stylidiaceae). Bot. Zhurn. 90(4):

575–582 (in Russian with English summary).

Rapson LJ. 1953. Vegetative anatomy of Donatia, Phyllachne,

Forstera, and Oreostylidium and its taxonomic signiﬁ cance.

Trans. Proc. Roy. Soc. N. Z. 80: 399–402.

Superorder ASTERANAE 495

Rauling EJ and PY Ladiges. 2001. Morphological variation and

speciation in Stylidium graminifolium (Stylidiaceae):

description of S. montanum and reinstatement of S. armeria.

Aust. Syst. Bot. 14: 901–935.

Rosen W. 1935. Beiträge zur Embryologie der Stylidiaceen. Bot.

Not. 1935: 273–278.

Subramanyam K. 1950. Development of embryo sac and

endosperm in Stylidium lenellum. Curr. Sci. 19: 294.

Subramanyam K. 1951. A morphological study of Stylidium

graminifolium Swartz. Lloydia 14: 65–81.

Subramanyam K. 1953. The nutritional mechanism of embryo

sac and embryo in the families Campanulaceae, Lobeliaceae,

and Stylidiaceae. J. Mysore Univ. B. 13: 1–4.

Subramanyam K. 1970. Stylidiaceae. Bull. Indian Natl. Sci.

Acad. 41: 317–320.

Wagstaff SJ and J Wege. 2002. Patterns of diversiﬁ cation in New

Zealand Stylidiaceae. Am. J. Bot. 89: 865–874.

Wege JA. 2001a. Corolla venation in Stylidiaceae. J. Roy. Soc.

West. Aust. 84: 97–101.

Wege JA. 2001b. Scape anatomy in Stylidium (Stylidiaceae).

Kew Bull. 56: 955–963.

Order 118. PHELLINALES

(ALSEUOSMIALES)

Small trees or shrubs. Vessels with scalariform or sca-

lariform and simple perforation, with numerous bars;

lateral pitting opposite to alternate, sometimes scalari-

form. Fibers with bordered or simple pits, often sep-

tate. Rays heterogeneous and homogeneous, rarely

(Alseuosmiaceae) absent. Axial parenchyma scanty

paratracheal, or even absent. Sieve-element plastids of

S-type. Nodes mostly trilacunar with three traces,

rarely unilacunar or pentalacunar. Leaves alternate to

subopposte, entire or dentate; lamina serrate, gland-

toothed; estipulate. Stomata anomocytic. Flowers in

axillary cymes, paniculate or corymbs, or solitary,

bisexual or rarely unisexual, usually actinomorphic,

4–6-merous. Sepals free or basally connate, valvate.

Petals free or basally connate, valvate. Stamens as

many and alternate with petals; ﬁ laments free or basally

connate or adnate to the corolla tube; anthers opening

longitudinally. Tapetum secretory. Microsporogenesis

simultaneous. Pollen grains 2-celled or 3-celled, mostly

3-colporate. Gynoecium of 2–5 united carpels; stylodia

free or connate into a style; style slender or stout, with

capitate or clavate, 2–5-lobed stigma, or stigma sessile.

Ovary semi-inferior or inferior, rarely (Phellinaceae)

superior, 2–6-locular; ovules one to many per locule,

mostly anatropous, unitegmic, tenuinucellate or crassi-

nucellate (Phellinaceae). Female gametophyte of

Polygonum-type. Endosperm cellular. Fruits loculi-

cidal capsules, drupes or berries. Seeds with small

straight or slightly curved embryo and with copious

endosperm. Contain gallic and ellagic acids, ellagita-

nins, and alkaloids (Phellinaceae).

The monophyly of the order Phellinales has very

high support values in both the molecular and com-

bined analyses (see also Soltis et al. 2006).

Key to Families

1 Ovary superior. Evergreen dioecious small trees or

shrubs with uniseriate hairs; cuticle waxes as plate-

lets and rodlets. Vessels with scalariform perfora-

tions and with numerous bars; lateral pitting

scalariform or opposite. Rays heterogeneous with

short ends. Axial parenchyma scanty paratracheal.

Leaves alternate or subopposite, simple, usually

papyraceous, more or less crowded towards tips of

branched, estipulate. Stomata anomocytic. Flowers

in axillary spicate or paniculate inﬂ orescences, uni-

sexual, 4–6-merous. Sepals small, free, more or less

persistent. Petals free, ﬂ eshy, valvate with a small

inﬂ exed apiculus. Stamens as many and alternate

with petals, free; anthers onlong and introrse. Pollen

grains 3-colporate. Nectary disc absent. Staminodia

present in female ﬂ owers. Gynoecium of 2–5 car-

pels, with sessile lobed stigma. Ovary with solitary

(rarely two – Doweld 2000), pendulous, hemitrop-

ous or slightly campylotropous, ovule in each loc-

ule. Pistilodia present in female ﬂ owers. Fruits

drupes with 2–5 pyrenes. Seeds with small embryo

and copious endosperm. Contain benzylisoquino-

line alkaloids, irdoid ?; n = 17. . . . 1. pellinaceae

1 Ovary inferior or semi-inferior.

2 Small trees or shrubs with characteristic two-

armed trichomes consisting of a very elongate

terminal «T-cell» that tapers to a ﬁ ne point at

each end and is attached at its middle to multicel-

lular, uniseriate stalk. Vessels with scalariform

perforations; lateral pitting commonly alternate.

Fibers with bordered and bordered and simple

pits, septate. Rays heterogeneous and homoge-

neous. Axial parenchyma absent or very scarce.

Nodes trilacunar, unilacunar (Corokia virgata) or

pentalacunar (Argophyllum laxum). Leaves alter-

nate; petioles subvaginate. Stomata anomocytic.

Flowers small. Petals mostly 4–6, basally con-

nate, valvate, silky outside. Petals usually (except

Corokia macrocarpa) with ligulate, nonvascularized

496 Subclass VII. ASTERIDAE

appendages on their inner side near the base of

their free parts just above the tube. Pollen grains

3-colporate.

3 Fruits loculicidal capsules. Shrubs, often

silky pubescent. Leaves often snow-white

below, entire or coarsely dentate, ovate.

Flowers 4–5(-8)-merous, whitish or green-

ish, in terminal and axillary panicles or

corymbs. Corolla basally connate. Anther

disporangiate. Style short, with capitate,

5–2-lobed stigma. Ovary semisuperior,

6–2-locular; ovules many, many-seriate on

tumid placentas in the axils of the locules.

Seeds very small, with large, elongate, nearly

straight embryo in the middle of the copious,

ﬂ eshy endosperm; exotestal cells with inner

walls massively thickened and ligniﬁ ed.

Ellagic acid present. . .2. argophyllaceae.

3 Fruits drupes crowned by the persistent calyx

teeth and style and with 2–1-locular and

2–1-seeded bony endocarp with characteristic

germination plug. Small trees or shrubs with

branchlets, petioles, leaves below, peduncles

and pedicels densely white, tomentose. Wood

diffuse-porous. Lateral pitting opposite-

alternate. Perforations scalariform with 16–23

bars. Rays heterogeneous. Vascular tracheids

present, except in C. collenettei (Noshiro et

Baas 1998). Crystals absent. Leaves entire.

Flowers bright yellow, in axillary or terminal

panicles, racemes, fascicles, or solitary. Anthers

tetrasporangiate. Nectary disc bright orange,

epigynous, pulvinate. Style short or subelon-

gate, with 2-lobed stigma. Ovary inferior,

1(-2)-locular; ovules solitary in the locule;

micropylar and chalazal haustoria present. Seeds

linear-oblong, with large, elongate, slightly

curved embryo in ﬂ eshy endosperm. Contain

gallic and ellagic acids, n = 9. 3. corokiaceae.

2 Shrubs (sometimes up to 6 m tall – Periomphale),

or creeping or epiphytic subshrubs, with unicel-

lular, bicellular, or multicellular, unbranched

trichomes. Stems and usually also petioles have

a morphologically differentiated uniseriate

endodermis of cells with casparian strips.

Secondary thickening developing from a con-

ventional cambial ring. Vessels with scalariform,

or scalariform and simple perforations in almost

vertical wall and composed of 20–45 bars; lat-

eral pitting opposite to alternate, sometimes sca-

lariform. Fibers with indistinctly bordered or

simple pits, often septate, mostly living and with

stored starch at maturity. Rays absent. Axial

parenchyma scanty or even absent. Nodes trila-

cunar with three traces. Leaves alternate or sub-

opposite, entire or dentate, simple, pinnately

veined, with multicellular and uniseriate hairs in

the axils containing dark, reddish pigment,

estipulate. Stomata anomocytic. Flowers in axil-

lary cymes or in terminal umbel-like inﬂ ores-

cences, or ﬂ owers solitary in the axils or on old

wood, bisexual, often including cleistogamous

ones, heavily scented, actinomorphic, perianth

and androecium (4)5(-7)-merous. Sepals free,

more or less triangular, valvate, persistent or

caduceus (Alseuosmia). Petals connate, with

valvate lobes; corolla margins with wings.

Stamens alternipetalous, attached to the throat

of the corolla tube or free; anthers dorsiﬁ xed,

introrse; in Platyspermation the anthers sessile

and extrorse, brown hairy below, and with a

large ﬂ at connective appendage at the apex (van

Steenis 1982). Pollen grains 3-colporate.

Gynoecium of two carpels; style slender or

stout, with more or less 2-lobed, clavate or capi-

tate stigma; ovary inferior or sometimes semi-

inferior, 2-, rarely (Wittsteinia vacciniaceae)

3-locular, crowned by a more or less developed

nectary disc, with 1-many ovules in each locule.

Fruits 2-locular berries, 2–3-locular (Wittsteinia),

or 2-locular (Platyspermation) capsules, with

persistent calyx and one to many seeds. Seeds

with small, straight embryo and copious

endosperm; exotesta little thickened, ligniﬁ ed,

mesotesta persistent. Contain quercetin and kae-

mpferol, caffeic and p-coumaric, stearic acids,

ellagitanins and triterpenoid saponins; n = 9

(Alseuosmia).. . . . . . . . . . .4. alseuosmiaceae.

1. PHELLINACEAE

Takhtajan 1967. 1/12. New Caledonia.

Phelline

Wood structure of Phelline is very primitive and as

Baas (1975: 339) points out, “Amongst the dicotyle-

dons with vessels, the wood anatomy of Phelline,

belongs to the most primitive.”

Superorder ASTERANAE 497

2. ARGOPHYLLACEAE

Takhtajan 1987. 1/11. Tropical Australia and New

Caledonia.

Argophyllum

3. COROKIACEAE

Kapil ex Takhtajan 1997. 1/6. Australia (1, NSW),

New Zealand (2), Lord Howe Island (1), Chatham Is.

(1), and Rapa I. (1)

Corokia

Corokia is usually placed in the Cornaceae s. 1.

However Hallier (1908, 1923) came to the conclusion

that Corokia is closely related to Argophyllum

(“Corokia ist durch Reduktion aus Argophyllum

enstanden und bildet mit ihm und Cuttsia die Sippe

der Argophylleen» (1908: 258). Engler (1930)

included Corokia in his Escallionioideae-

Argophylleae. The afﬁ nity of Argophyllum and

Corokia was convincingly shown by Eyde (1966).

Both Hallier and Eyde emphasized the remarkable

similarity between the peculiar multicellular

T-shaped trichomes of these genera, which are

known only in few groups of ﬂ owering plants. They

also emphasized the similarity of corolline append-

ages in the two genera. Moreover, Corokia and

Argophyllum are similar in gross vegetative

characters, wood anatomy (including septate trache-

ids, which are absent in Escalloniaceae [see Patel

1973; Hils 1985]) and pollen grain structure. Also,

both Corokia and Argophyllum have a lamellate

endexine in the region of the endoaperture, which

resembles Cornaceae (Ferguson 1977; Ferguson and

Hideux 1978). Thus, palynologically Corokia and

Argophyllum occupy a somewhat intermediate posi-

tion between Escalloniaceae and Cornaceae. There

are also important differences between Argophyllum

and Corokia, including placentation, the number of

ovules, and the structure of fruits and seeds. It is

appropriate to consider both these related genera

members of two separate families. According to

Kapil (1992) embryologically Corokia is distinctive

enough to deserve family status. However, in his

opinion the Corokiaceae resemble the Stylidiaceae

more rather than the Cornaceae and the Escalloniceae.

According to Gustafsson and Bremer (1995), Corokia

is the most likely sister group of the Asterales s. 1.

4. ALSEUOSMIACEAE

Airy Shaw 1965 (including Platyspermatiaceae Doweld

2001). 5/10. Southwestern Paciﬁ c. Wittsteinia (2), New

Guinae and southeastern Australia; Periomphale (1)

New Caledonia; Platyspermation (1) New Caledonia;

Alseuosmia (5) is endemic to New Zealand, and

Crispiloba (1) is endemic to Queensland.

Wittsteinia, Periomphale, Alseuosmia, Crispiloba,

Platyspermation

Bibliography

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498 Subclass VII. ASTERIDAE

Paliwal GS and LM Srivastava. 1969 (1970). The cambium of

Alseuosmia. Phytomorphology 19: 5–8.

Tirel C. 1996. Rétablissement de Periomphale Baill. (Alseuos-

miaceae) genre endémique de Nouvelle Caledonie. Bull. Mus.

Natl. Hist. Nat. Paris 4e sér. sect. B Adansonia 18: 155–160.

Zeman M. 1907. Studien zu einer Monographie der Gattung

Argophyllum Forst. Ann. K. K. Naturalist. Hofmuns. 22:

270–292.

Order 119. ASTERALES

Perennial or annual herbs and subshrubs, less often

shrubs, lianas or small or medium-sized trees, usually

with various types of glandular or nonglandular

trichomes. Lactifers mostly present. Vessels with sim-

ple or sometimes scalariform perforations. Fibers with

simple pits, or less often (Goodeniaceae) with bordered

pits. Rays mostly heterogeneous. Axial parenchyma

scanty paratracheal. Sieve-element plastids of S-type.

Nodes trilacunar to multilacunar, rarely (some

Goodeniaceae) unilacunar. Leaves alternate or less

often opposite, rarely verticillate, simple to variously

divided, gland-dotted or not gland-dotted, estipulate or

rarely with stipules; leaf mesophyll cells contain oil

bodies. Stomata usually anomocytic. Inﬂ orescences of

various types, or less often ﬂ owers solitary. Flowers

bisexual or unisexual, actinomorphic to zygomorphic,

mostly 5-merous. Sepals usually connate at the base,

sometimes adnate to the ovary. Petals connate, valvate.

Nectary disc usually present. Stamens usually ﬁ ve,

alternate with the corolla lobes; ﬁ laments free or some-

times connate; anthers basiﬁ xed, introrse, tetras-

porangiate, or occasionally disporangiate, opening

longitudinally. Tapetum secretory or amoeboidal.

Microsporogenesis simultaneous. Pollen grains mostly

3-celled, 3-colporate. Gynoecium of two united car-

pels; stylodia united into terminal style; ovary mostly

inferior, 1-locular; ovule mostly solitary, usually anat-

ropous, unitegmic, tenuinucellate, with endothelium.

Female gametophyte of various types, but mostly of

Polygonum-type. Endosperm cellular or less often

nuclear, without haustoria. Fruits of various types,

but mostly cypselas. Seeds with straight embryo;

endosperm usually copious, rarely wanting. Iridoids

present or absent.

Very close to the Campanulales.

Key to Families

1 Iridoids present.

2 Fruits cylindrical or prismatic achenes. Perennial,

biennial or annual herbs, rarely suffrutescens

(typically forming only 1–2 mm thick of second-

ary xylem at stem base and adjacent root area),

rarely pubescent. Laticifers and secretory cavi-

ties wanting, but in Boopis gracilis present drop-

lets of a resin-like substance (Hellwig 2007).

Vessels with simple perforations. Fibers with

thick walls and simple pits. Leaves alternate, often

rosulate, simple, entire or pinnately lobed, estipu-

late. Stomata anomocytic. Inﬂ orescences terminal

on scapes or aillary, capitulate, these solitary or in

cymose panicles, surrounded by 1–2 series of

bracts. Flowers numerous, opening centripetally

(Acicarpha) or in a more complex manner, on

short peduncles, or peduncles absent (Acicarpha

and some species of Boopis), bisexual or seldom

the central ﬂ owers of a head functionally unisex-

ual, actinomorphic or zygomorphic. Calyx with

4–6 small lobes or teeth, aerenchymatous or

spine-like. Corolla tubular, with 4–6 valvate

lobes, the outer layer of the tubular corolla is

photosynthetic. Stamens as many as and alter-

nate with the corolla lobes, attached near the

summit of the tube; ﬁ laments often basally or

wholly connate; gland-like nectaries generally

situated basally in the ﬁ lament tube extending to

the top of the stamen-corolla tube; anthers free or

basally coherent to connate; secondarily present-

ing pollen by depositing it on extending style

(Heald 2004). Pollen grains 2-celled, 3-colpo-

rate, spinulate to nearly smooth. Gynoecium of

two carpels; style slender, glabrous, growing up

and pushing out the pollen; stigma capitate,

indistinctly papillate; ovary inferior, essentially

1-locular (incompletely 5-locular due to reduced

septa), with a solitary ovule pendulous from the

top of the locule. Ovules anatropous, with (?)

endothelium. Fruits with persistent, spine-like

calyx (this lignescent and spiny in Acicarpha and

Calycera), ribbed and receptacle small in Boopis.

Seeds with straight embryo and oily, copious, or

scanty endosperm. Storing carbohydrate as inu-

lin. Producing iridoid compounds (secologanin);

n = 7–9, 13, 17, 21, 22. . . . . . .4. calyceraceae

2 Fruits capsules, baccate, drupaceous or nutlike.

3 Terrestrial perennial herbs, subshrubs or

shrubs, rarely arborescent with trichomes

of both glan dular and nonglandular types.

Superorder ASTERANAE 499

Laticifers wanting. Sclerenchymatous idio-

blasts, which are sometimes branched, are

common in the mesophyll of the leaf and

sometimes occur in the cortex of the stem as

well. Vessels with simple or sometimes with

scalariform perforations. Fibers with bordered

pits. Rays heterogeneous to homogeneous.

Axial parenchyma vasicentric or diffuse or

both or wanting. Secondary thickening absent

or developing from a conventional cambial

ring. Nodes unilacunar or sometimes trilacu-

nar to pentalacunar. Leaves well developed or

much reduced, alternate or rarely opposite or

verticillate, nearly always spiral, petiolate to

sessile, simple, entire, serrate or dentate,

estipulate. Stomata anomocytic. Flowers

zygomorphic, variously in cymes, racemes, or

heads, or solitary in the axils. Calyx tube

adnate to the ovary or rarely free. Corolla

bilabiate (with two dorsal and three ventral

lobes) or sometimes unilabiate (with ﬁ ve ven-

tral lobes), the lobes valvate, often indupli-

cate, spurred or not (Selliera). Stamens ﬁ ve,

attached to the base of corolla tube or free;

anthers separate from one another (Velleia,

Scaevola), or connivent around the style, or

(Dampiera, Leschenaultia) connate into

a tube. Pollen grains 2-celled, 3-colporate or

(Lechenaultia) porate. Gynoecium of two

carpels; style one, with a ‘pollen cup’ close

beneath to stigma, bearing an ‘indusium’

beneath the stigma. Ovary inferior or semi-

inferior or rarely (Velleia) superior, 2-locular

or seldom 1-locular, or (some other spp. of

Scaevola) with two vestigial lateral locules in

addition to the fertile median ones, rarely

(Scaevola porocarya) 4-locular, with one to

many ovules per locule mostly erect or

ascending on axile or basal-axile placentas

(in Verreauxia the locule is solitary and

basal). Fruits mostly capsular, less often

drupaceous, or nutlike. Seeds usually ﬂ at,

winged or wingless, exotestal cells usually

palisade, all walls thickened; embryo straight,

endosperm copious, oily. Storing carbohy-

drate as inulin and producing 13-carbon

polyacetylenes, caffeic acid, sometimes iri-

doids (secologanin), O-metyl ﬂ avonols, and

alkaloids, n = 7–9. . . . . . .1. goodeniaceae.

3 Aquatic or wetland herbs, mostly perennial

and with tufted rootstocks or horizontal creep-

ing rhizomes, but some species of Nymphoides

and Villarsia are annual. Vascular system is

composed of isolated or scattered bundles. A

well-developed system of intercellular canals

and spaces is also characteristic. Branched,

sclerenchymatous idioblasts, similar to those

of the Nympheaceae, often present in the

parenchyma of the stem. Vessels usually with

simple perforations, but scalariform and sim-

ple plates may occur in Menyanthes. Nodes

trilacunar or pentalacunar. Leaves alternate,

simple or rarely (Menyanthes) 3-foliolate with

sheating petioles, linear to cordate or reni-

form; stipules wanting or represented by the

expanded wing margins of the petiole. Stomata

anomocytic. Flowers solitary or in simple or

branched cymes or racemes or in dense heads

or clusters, bisexual, sometimes functionally

unisexual, actinomorphic, usually 5-merous

(except gynoecium), often heterostylous.

Sepals more or less connate at base, some-

times adnate to the ovary below, persistent.

Corolla sympetalous, appendiculate (with

intrastaminal scales), or not appendiculate,

valvate or induplicate-valvate; the margins or

the inner surface of the lobes often ﬁ mbriate

or crested. Stamens attached to the base of the

corolla tube alternate with the lobes, or mid-

way down the corolla tube, or in the throat of

the corolla tube; anthers 2-locular, dorsiﬁ xed,

mostly sagittate, versatile. Sometimes fringed

scales ( staminodia?) alternate with stamens.

Pollen grains 3-celled, 3- colpate or 3-colpo-

rate, or rugate (exceptionally 6-rugate).

Nectary disc usually present around the base

of the ovary. Gynoecium of two united car-

pels; stylodia connate into a terminal, 2-lobed

style, with papillate stigma; ovary superior to

semiinferior, 1-locular, with numerous ovules

on two parietal and often intruded placentas.

Ovules horizontal, with short funicle and with

endothelium. Fruits septicidal, loculicidal, or

valvular capsules, or rarely (Liparophyllum)

ﬂ eshy berries. Seeds 4–100, glabrous or with

trichomes of different shape, winged or wing-

less, in some species surrounded by a caruncle

which is rich in lipids (Kadereit 2007);

500 Subclass VII. ASTERIDAE

exotestal cells with outer walls thickened,

often with a variety of projections; embryo

straight, linear; endosperm copious, oily.

Producing iridoid compounds (loganin,

secologanin, sweroside), ﬂ avonols, caffeic

acid (except Menyanthes), and usually storing

inulin. n = 9, 17. . . . . . . 3. menyanthaceae.

1 Iridoids absent.

4 Stomata paracytic. Perennial scapose herbs with

characteristic hairs of short, thin-walled, erect

basal cell and two to several-celled arm attached

at right angles to the basal cell and appressed to

the surface. Secondary thickening developing

from a conventional cambial ring. Leaves all

basal or alternate towards the base of the stems,

more or less petiolate, simple, estipulate. Flowers

actinomorphic or very nearly so, bisexual, borne

in involucrate, cymose heads or spikes, each

ﬂ ower with several hyaline bracts. Calyx tube

free from the ovary and calyx lobes markedly

longer than the tube. Corolla with ﬁ ve spreading

the lobes valvate, blue, markedly longer than the

tube. Stamens ﬁ ve, attached to the base of the

corolla tube; anthers connate into a tube around

the style. Gynoecium of two carpels; style one,

bearing an ‘indusium’ beneath the stigma; stigma

surrounded by a collar or indusium; ovary 1-loc-

ular, superior, with septum only at base and soli-

tary, ascending ovule inserted on one side of the

septum. Fruits dry and indehiscent, enclosed by

persistent calyx tube. Seeds ellipsoid, with

straight embryo and without endosperm. Iridoids

wanting, n = 9. . . . . . . . . . . . 2. brunoniaceae.

4 Stomata anomocytic or occasionally anisocytic.

Perennial or annual herbs and subshrubs, less

often shrubs, lianas or small or medium-sized

trees (up to 30 m tall or more), sometimes nearly

succulents, halophytes, marsh plants, lianas, epi-

phytes and aquatics, usually with various types of

glandular or nonglandular trichomes. Often with

a system of articulated laticifers (mostly

Chicorioideae) in the phloem, containing latex

rich in triterpene, or with a more or less well-

developed system of schizogenous resin canals

that are often lined with an epithelium (the two

secretory systems largely alternative rather than

coexistive); scattered latex cells occasionally

present in addition to resin canals. Secondary

thickening well developed even in many herba-

ceous species, or absent, or anomalous. Vessels

commonly with simple perforations, but some-

times with scalariform and simple, or simple and

reticulate perforations; lateral pitting alternate.

Fibers typically with small simple pits, or the pits

with very small borders. Rays mostly heteroge-

neous, mostly 4–10(-18) cells wide. Axial paren-

chyma scanty paratracheal. Nodes trilacunar to

multilacunar. Leaves alternate or less often oppo-

site, rarely verticillate, simple or variously

divided, gland-dotted or not gland-dotted, aro-

matic or foetid, or without marked odour; leaves

estipulate or rarely with stipules. Flowers in

dense racemose heads (capitula) with one to

many sessile ﬂ owers on common receptacle, the

head nearly always subtended by an involucre of

one to several series of bracts; capitula occasion-

ally aggregated into cymose secondary heads

sometimes with a secondary involucre; common

receptacle of the head ﬂ at to conical or cylindri-

cal, sometimes with a bract subtending each

ﬂ ower (mainly in Heliantheae) or densely bristly

(mainly in Cynareae); heads homogamous (all

ﬂ owers alike) or heterogamous; in homogamous

heads all ﬂ owers are usually bisexual; in heterog-

amous heads the peripheral (radiate or ﬁ liform)

ﬂ owers female or neutral and the central (disc)

ﬂ owers bisexual or unisexual (monoecious or

dioecious, or androdioecious (rarely), or gyno-

dioecious (rarely), or polygamomonoecious.

Perianth and androecium 5-merous or rarely

4–3-merous. Calyx tube together with the base of

corolla tube completely fused with the ovary and

its free lobes usually greatly modiﬁ ed and trans-

formed into a pappus, which is often strongly

reduced or obsolete. Corolla of three basic types:

(1) tubular or discoid (actinomorphic, 5-lobed or

5-dentate, with a conspicuous tube and usually a

short limb), ligulate (stap-shaped with ﬁ ve apical

teeth) or bilabiate in bisexual (or functionally

male) ﬂ owers; (2) radiate ( strap-shaped with 0–4,

usually three, apical teeth) in peripheral female

or sterile ﬂ owers; (3) ﬁ liform, narrow, apically

truncate, minutely lobed or minutely radiate, in

peripheral female ﬂ owers; rarely corolla absent

(in peripheral female ﬂ owers). Stamens usually

ﬁ ve, epipetalous, alternate with the corolla lobes;

ﬁ laments usually free, but sometimes connate

into tube (e.g., in Silybum, Galactites, and

Superorder ASTERANAE 501

Dipterocome); anthers usually tetrasporangiate,

or occasionally disporangiate, often with short

apical appendage and often basal tails. Tapetum

amoeboidal or less often secretory. Micro-

sporogenesis simultaneous. Pollen grains

3-celled, mostly 3-colporate, psilate, spinulose or

lophate, often caveate. Gynoecium of two car-

pels; style terminal, usually papillate, biﬁ d

(branches commonly separating after passage

through the anther tube), with annular or short

tubular nectary disc at the base; ovary inferior,

1-locular, with one basal, erect ovule. Female

gametophyte of various types, but mostly of

Polygonum-type. Endosperm cellular or less

often nuclear, without haustoria. Fruits cypse-

las (achenes), usually with persistent pappus,

very rarely drupaceous with ﬂ eshy pericarp

(Chrysanthemoides, tribe Calenduleae, and

Wulfﬁ a, tribe Heliantheae). Seeds with straight,

oily embryo; endosperm wanting or sometimes

present as a very thin vestigial layer; exotestal

cells thickened, palisade or ﬂ attened, or undistin-

guished. Usually storing carbohydrate as inulin,

and producing polyacetylenes, terpenoid essen-

tial oils, alkaloids and ﬂ avonols present or absent;

producing diverse array of polyacetylenes, bitter

sesquiterpene lactones, caffeic and ursolic acids,

and others; n = 2–19 + . . . . . . . . 5. asteraceae.

1. GOODENIACEAE

R. Brown 1810 (including Scaevolaceae Lindley 1830).

12/420. Widely dispersed on tropical shores of the

Atlantic and Indian oceans while the remainder extend

north and east in the Paciﬁ c as far as China, Japan, the

Philippines, the Hawaiian Islands, New Zealand, and

Chile. Over 95% of the species are endemic to Australia

(especially western Australia) and Tasmania.

Velleia, Goodenia (including Calogyne), Selliera,

Scaevola, Leschenaultia, Anthotium, Pentaptilon,

Diaspasis, Verreauxia, Dampiera, Coopernookia.

Morphological and molecular data support the

relationships the Goodeniaceae, Calyceraceae, and

Asteraceae.

2. BRUNONIACEAE

Dumortier 1829. 1/1. Australia and Tasmania.

Brunonia

A rather heterobathmic family that differs from the

Goodeniaccae mainly in actinomorphic or nearly acti-

nomorphic ﬂ owers borne in involucrate heads and the

endospermless seeds.

3. MENYANTHACEAE

Dumortier 1829. 5/60. Subcosmopolitan.

Menyanthes, Villarsia, Nymphoides, Liparophyllum,

Nephrophyllidium

Belong to the Asterales, but occupies somewhat

isolated position. The afﬁ nity with the Asterales is

conﬁ rmed by molecular data (Michaels et al. 1993;

Gustafsson and Bremer 1995).

4. CALYCERACEAE

Richard 1820. 5/60 (including Boopidaceae Cassini

1816). South America, mainly in the southern Andes

and Patagonia.

Boopis, Calycera, Gamocarpha, Moschopsis,

Acicarpha (including Nastanthus).

Calyceraceae have similarities with Goodeniaceae,

but differ from them mainly in the absence of pollen-

collecting cups, epipetalous stamens, in multinucleate

tapetal cells, and fruit-type. Calyceraceae also related

to Asteraceae: they have similar second pollen presen-

tation, and also Calyceraceae and Barnadesioideae

have a similar simple ﬂ avonoid proﬁ le. However they

markedly differ from Asteraceae in having en entire

stigma, glabrous terminal style, and in lacking a pap-

pus, as well as the presence of iridoids.

5. ASTERACEAE

Martynov 1820 or Compositae Giseke 1792 (nom.

altern.) (including Ambrosiaceae Marynov 1820;

Anthemidaceae Martynov 1820; Aposeridaceae Raﬁ ne-

sque 1838; Arctotidaceae Bessey 1914; Artemisiaceae

Martynov 1820; Athanasiaceae Martynov 1820;

Calendulaceae Link 1829; Carduaceae Dumortier

1822; Cassiniaceae C.H. Schultz 1852; Centauraceae

Martynov 1820; Cichoriaceae A.L. de Jussieu

1789; Cnicaceae Vest 1818; Cynaraceae Durande

1782; Echinopaceae Dimortier 1822; Eupatoriaceae

502 Subclass VII. ASTERIDAE

Martynov 1820; Gnaphaliaceae Link ex F. Rudolphi

1830; Heleniaceae Raﬁ nesque 1824; Helianthaceae

Dumortier 1822; Inulaceae Bessey 1914; Lactucaceae

Drude 1979; Matricariaceae J.Voigt 1845; Mutisiaceae

Burnett 1835; Nassauviaceae Burmeister 1836;

Picridaceae Martynov 1820; Santolinaceae Martynov

1820; Senecionaceae Berchtold et J. Presl 1820;

Serratulaceae Martynov 1820; Syngenetiaceae Horani-

now 1847; Tanacetaceae Vest 1818; Vernoniaceae

Burmeister 1837; Xanthiaceae Vest 1818). C.1600/

23000 or more. Cosmopolitan, but mostly in temperate

and subtropical regions.

Delpino (1871), Small (1919), Rosen (1946,

1949), and many others derive the Asteraceae from

the Lobeliaceae or Lobeliaceae-like ancestor. In the

Englerian system, they occupy a position near and

after the Calyceraceae at the top of Campanulales

sensu lato. Recent morphological and molecular data

suggest that the Asteraceae form a sister group to the

clade Calyceraceae and Goodeniaceae (Jansen et al.

1991b; Harris 1995). The Asteraceae differ from

them mainly in producing various types of sesquiter-

pene lactones, in the absence of iridoid compounds,

in calyx transformed into pappus, and in mostly

amoeboid tapetum. The similarities between

Asteraceae and Calyceraceae include involucrate

capitula, specialized pollen presentation mechanism,

epipetalous stamens, uniovulate ovary, and the pres-

ence of unique intercolpar concavities. Both families

evidently share a common origin from some

Goodeniaceae-like ancestor.

Classiﬁ cation after C. Jeffrey 2007.

5.1 BARNADESIOIDEAE

Shrubs, trees, or perennial or annual herbs, usually

with fascicled nodal spines. Capitula homogamous

or heterogamous, discoid or pseudoradiate or ligu-

late, sessile or pedunculate. Floral parts having

3-cellular hairs with swollen basal cell (barnade-

sioid hairs), often with axillary spines; corolla tubu-

lar, often villous; ﬁ laments free or rarely fused,

inserted at different levels; style shortly bilobed or

biﬁ d, glabrous or papillose below bifurcation;

cypselas densely villous, with straight simple hairs,

rarely glabrous. Pappus uniseriate or rarely absent.

Poor in ﬂ avonoids, ﬂ avonols absent; n = 8, 24, 25,

27. – Schlechtendalia, Doniophyton, Duseniella,

Fulcaldea, Barnadesia, Huarpea, Dasyphyllum,

Arnaldoa, Chuquiraga.

5.2 MUTISIOIDEAE

Shrubs, trees or perennial or rarely annual herbs, rarely

scandent. Leaves usually alternate, denticulate or lobu-

late. Capitula homogamous, bilabiate or discoid, rarely

ligulate. Florets 1-many, 5-merous. Corolla lobes long,

with an apical tuft of minute hairs; style with or without

hairs, without thickening on shaft below branches;

stigma lobes short, pollen spinulose; cypselas usually

with twin hairs. Pappus usually present, commonly of

bristles, often uniseriate; n = (6-)9. – stifftieae:

Stifftia, etc.; mutisieae: Mutisia, Acourtia, Chaptalia,

Anislaea, Trixis, etc.

5.3 CARDUOIDEAE

Perennial, biennial or less often annual herbs, shrubs

or rarely trees, rarely scandent. Leaves alternate,

denticulate or lobulate, especially in herbaceous

members often spiny. Capitula homogamous or

heterogamous, discoid or discoid with marginal ﬂ orets

sterile and radiant, rarely bilabiate-radiate or ligulate.

Florets one to many, 5-merous; corolla lobes long;

anthers calcarate and caudate; pollen spinulose or

spiny; plant often spiny; style usually with a papillose

or hairy thickening on the shaft below the style

branches; cypselas with twin hairs, simple hairs or

glabrous. Pappus usually present, of bristles or scales;

n = 12. – gochnatieae: Ainsliaea, Gochnatia,

Brachylaena; hecastocleideae: Hecastocleis;

tarcho nantheae: Tarchonanthus; dicomeae:

Dicoma; cynareae-carlininae: Atractylodes,

Thevenotia, Carlina, Atractylis, etc.; pertyeae:

Ainsliaea, cynareae-echinopsinae: Echinops, Acan-

tholepis, Amphoricarpos, Xeranthemum, Char dinia,

etc.; cynareae-carduinae: Arctium, Cousinia,

Onopordum, Saussurea, Jurinea, Carduus, Cirsium,

Silybum, Cynara, Ptilostemon, Lamyropsis, Alfredia,

Olgaea, Galactites, Picnomon, etc.; cynareae-

centaureinae: Serratula, Stemma cantha, Leuzea,

Tricholepis, Acroptilon, Callicephalus, Centaurea,

Centaurodendron, Chartolepis, Stizolophus, Zoegea,

Cnicus, Carthamus, Carduncellus, Amberboa,

Oligochaeta, Volutaria, Crupina, etc.

5.4 CICHORIOIDEAE (Lactucoideae)

Perennial, biennial or annual herbs, shrubs or trees, rarely

scandent, very rarely aquatic. Leaves alternate or oppo-

site, entire to deeply lobed, sometimes spiny. Capitula

homogamous, ligulate, radiate or discoid, less often

heterogamous, radiate or discoid. Disc ﬂ orets usually