Takhtajan Armen. Flowering Plants

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Superorder ASTERANAE

Order 115. ROUSSEALES

Small trees or shrubs, or (Rousseaceae) climbing shrubs.

Vessels with scalariform perforation; lateral pitting

scalariform to alternate, or occasionally transitional.

Fibers with bordered pits. Rays mostly heterogeneous.

Axial parenchyma scanty paratracheal, or diffuse-in-

aggregates. Sieve-elements plastids of S-type. Nodes

trilacunar. Leaves alternate, opposite or verticillate, estip-

ulate. Stomata anomocytic. Flowers small, in terminal

and axillary panicles, or ﬂ owers solitary, actinomorphic,

mostly bisexual, rarely unisexual, polygamo-dioecious.

Sepals 4–5, or 5–6, valvate, persistent or deciduous.

Petals 4–6, valvate, persistent or deciduous. Stamens

free or connate; anthers tetrasporangiate, opening longi-

tudinally. Tapetum secretory. Microsporogenesis simul-

taneous. Pollen grains 3-celled, mostly 3-colporate,

rarely (Roussea) pantoporate. Nectary disc present.

Gynoecium of 5–7 or 3–5 (Carpodetaceae) united car-

pels; style thick or slender with capitate or lobed stigma.

Ovary superior or semi-inferior to inferior, (4-)5–7-loc-

ular. Ovules numerous, anatropous, unitegmic, tenu-

inucellate. Female gametophyte of Polygonum-type.

Endosperm cellular. Fruits baccate. Seeds numerous,

small; embryo small; endosperm copious.

Rousseales belong to Asteranae, and occupied the

basal position there (see Soltis et al. 1997a, b; Savolainen

et al. 2000; Bremer et al. 2002; Soltis et al. 2006).

Key to Families

1 Climbing shrubs, sometimes strangler, up to 4 m in

height. Trichomes glandular-peltate with multicel-

lular heads, or unicellular, borne either solitary or in

clusters; resin canals present. Vessels with very

oblique end walls and exclusively with scalariform

perforations that have an average of 20 (up to 64)

bars; lateral pitting scalariform or occasionally

transitional. Fibers short, with bordered pits. Rays

tri- to multiseriate, hetero geneous and comprise

procumbent, upright, and square cells. Axial paren-

chyma scanty paratracheal. Leaves opposite and

verticillate, simple, petiolate, serrate-glandular,

with unicellular conic hairs and wartlike cuticle

protrusions on both surfaces and peltate, multicel-

lular, and stalked glands on the lower surface and

on the petiole, estipulate; petiole with radially elon-

gated, schizogenous resin canals. Flowers solitary

or few in the leaf axils, arising from bracteate buds,

nodding; pedicels thick, arcuate. Copious amounts

of nectaries present in ﬂ owers (Koontz et al. 2007).

Calyx 4–5, united at the base, calyx lobes rather

large, thick, valvate, persistent. Petals thick, hairy,

connate into a 4–5-lobed tube, revolute at the apex,

villous outside, persistent, valvate in bud. Stamens

alternating with petals, inserted within the lobes of

the nectary disc; anthers large, oblong-sagittate,

basiﬁ xed, extrorse. Pollen grains in monads, 5–6-

or 8-porate, with smooth and complete tectum.

Style thick, persistent, expanding apically; stigma

capitate, 4–5-lobed, stigmatic lobes narrower, erect,

with revolute margins and like an indusium. Ovary

superior, piramidal, (4-)5–7- locular, with numerous

ovules many-seriate on thick, axile placentas.

Superorder ASTERANAE 485

Micropylar haustoria and a ﬁ lamentous suspensor

are developed. Fruits large, pale green, ﬂ eshy ber-

ries, acutely 5–7-angled, angles dilated at the base,

(4-)5–7 locular. Seeds numerous, ﬂ attened, ovoid,

exotestal. Testa with elongated epidermis cells with

a strongly undulating outline and thickened and lig-

niﬁ ed cells walls (Lundberg 2001). Embryo minute,

straight; endosperm copious.. . . . 1. rousseaceae.

1 Evergreen trees to 20 m tall or shrubs with unicel-

lular haris. Young branchlets, petioles, peduncles,

and pedicels pubescent. Vessels with long scalari-

form perforation plates; lateral pitting scalariform

to alternate. Fibers with bordered pits. Rays broad,

heterogeneous, with prismatic crystals in cells.

Axial parenchyma sparse and predominantly dif-

fuse or diffuse-in- aggregates. Leaves alternate,

sharply glandular-serrate, with domatia on the

abaxial surface in the axils of the secondary veins.

Flowers small, white, in terminal and axillary pani-

cles. Calyx tube turbinate, adnate to ovary, lobes

5–6, short, deciduous. Petals 5–6, inserted under

margin of epigynous nectary disc, deciduous, val-

vate. Stamens (4-)5(-7), alternipetalous; ﬁ laments

free, short, sometimes hairy; anthers oblong,

introrse. Pollen grains in tetrahedral tetrads, 3-col-

porate. Gynoecium of 3–5 carpels; style slender,

with capitate stigma; ovary semi-inferior or infe-

rior; ovules numerous, many-seriate on placentas

projecting into the loculi. Fruits 3–5-locular, berry-

like, with more or less leathery pericarp, surrounded

in the middle by the scar of calyx. Seeds numer-

ous, small, pendulous, ovoid, with elongated

funiculus; testa coriaceous, foveolate; embryo

small, terete, at the base of copious, ﬂ eshy

endosperm. Contain leucodelphinidin, quercetin

and kaempferol; n = 14, 15. . . 2. carpodetaceae.

1. ROUSSEACEAE

A.P. de Candolle 1839. 1/1. Mauritius.

Roussea

Systematic position of Roussea is debatable.

Hutchinson (1967) put it in the Escalloniaceae.

According to Lundberg (2001), Roussea is close related

to Carpodetus and they even combine their Carpodetaceae

sensu lato (which includes also Abrophyllum and

Cuttsia) with Rousseaceae. However, I prefer to keep

Rousseaceae as a separate family.

2. CARPODETACEAE

Fenzl 1841. 1/10. New Guinea, Solomon Islands, New

Zealand, and Stewart Island.

Carpodetus

Bibliography

Brook JP. 1951. Vegetative anatomy of Carpodetus serratus

Forst. Trans. and Proc. Roy. Soc. N. Z. 79: 276–285.

Gustafsson MHG. 2007. Carpodetaceae. In: K Kubitzki, ed. The

families and genera of vascular plants, vol. 8, pp. 57–60.

Springer, Berlin/Heidelberg/New York.

Gustafsson MHG and B Bremer. 1997. The circumscription and

systematic position of Carpodetaceae. Aus. Syst. Bot. 10:

855–862.

Koontz JA, J Lundberg, and DE Soltis. 2007. Rousseaceae. In:

K Kubitzki, ed. The families and genera of vascular plants,

vol. 8, pp. 611–613. Springer, Berlin/Heidelberg/New York.

Lundberg J. 2001a. The asteralean afﬁ nity of the Mauritian

Roussea (Rousseaceae). Bot. J. Linn. Soc. 137: 267–276.

Nemirovich-Danchenko EN. 2000a. Carpodetaceae. In: A

Takhtajan, ed. Comparative seed anatomy, vol. 6,

pp. 253–255. Nauka, St. Petersburg (in Russian).

Praglowski J and E Grafstrom. 1985. The genus Carpodetus

(Escalloniaceae). Grana 24: 11–21.

Order 116. CAMPANULALES

Mostly herbs (perennial or rarely annual), but some-

times secondarily woody (shrubs or less often small

trees). Hairs when present mostly unicellular. In leaves

and stems (Campanulaceae) articulated, anastomosing

laticifers forming a well-developed system in the

phloem and commonly extending into the surround-

ing tissues. Vessels mostly with simple or less often

scalariform perforations, or scalariform and simple;

lateral pitting scalariform to alternate. Wood paren-

chyma absent or indistinct. Fibers with simple or indis-

tinctly bordered pits. Rays commonly 4–8 cells wide.

Axial parenchyma scanty paratracheal or often want-

ing. Nodes usually unilacunar. Sieve-element plastids

of S-type. Leaves alternate or sometimes opposite,

rarely (Ostrowskia) verticillate, simple, entire, estipu-

late. Stomata commonly anomocytic. Flowers in vari-

ous cymose or more often racemose inﬂ orescences,

sometimes solitary, usually bracteate and often brac-

teolate, bisexual or rarely unisexual, actinomorphic to

zygomorphic, mostly with 5-merous perianth and

486 Subclass VII. ASTERIDAE

androecium. Calyx (2-)5(-10)-lobed, lobes imbricate or

valvate. Corolla usually 5-lobed, lobes valvate (some-

times induplicate) or less often imbricate (Sphenocleaceae).

Stamens as many as petals and alternate with them, or

2–3, seemingly free from the corolla and attached to the

annular, epigynous nectary disc or variously attached to

the corolla tube; ﬁ laments free, or variously coherent or

connate, the ﬁ lament bases often expanded, forming a

dome-shaped chamber over the nectary disc (the latter

enlarged and glandlike in Adenophora), anthers bas-

iﬁ xed, introrse or extrorse, tetrasporangiate, opening

longitudinally. Tapetum secretory. Microsporogenesis

simultaneous. Pollen grains 2-celled or less often

3-celled, colpate, colporate, or porate. Gynoecium of

2–10 carpels; style simple or lobed; ovary inferior or

seldom semi- inferior, rarely superior, 1–3-locular with

two parietal placentas (as in some Lobelioideae), or

often with as many locules as carpels, commonly

crowned by nectary disc. Ovules usually numerous,

anatropous, unitegmic, tenuinucellate, with well-devel-

oped endothelium. Female gametophyte of Polygonum-

type. Endosperm cellular, with typical terminal haustoria

(but only in Pentaphragmataceae with micropylar haus-

torium). Fruits usually capsules, loculicidal, septicidal,

or less often poricidal, in Sphenocleaceae and Parischella

circumscissile (pyxidia), but sometimes indehiscent.

Seeds with small, winged, or wingless, usually straight

embryo and usually copious, oily or rarely starchy

endosperm. Iridoids wanting, alkaloids usually present,

cyanogenic; storing carbohydrate as inulin, n = 5–20,

rarely up to 56.

Via the Pentaphragmataceae and archaic genus

Cyananthus the order Campanulales is connected with

the Desfontainiales, especially with Escalloniaceae

(Takhtajan 1987).

Key to Families

1 Plants without laticifers. Flowers lacking any spe-

cialized mechanism for secondary pollen presenta-

tion. Inulin and alkaloids absent.

2 Vessels with scalariform perforations with

numerous bars. Fiber-tracheids with prominently

bordered pits. Perennial coarse herbs often with

ﬂ eshy or succulent stems. Leaves alternate, often

relatively large, petiolate, simple, entire or den-

tate, usually strongly asymmetric at base, often

with branching hairs. Stomata anomocytic, sur-

rounded by three or four cells somewhat differ-

entiated from the ordinary epidermal cells.

Flowers in dense, axillary or extra-axillary, sym-

podial helicoid cymes with usually conspicuous

membranous bracts, bisexual or rarely unisexual

(plant dioecious). Sepals ﬁ ve, unequal, imbri-

cate, persistent. Corolla usually ﬂ eshy or carti-

laginous, rarely delicate, persistent, seemingly

choripetalous in some species, with ﬁ ve, rarely

(Pentaphragma tetrapetalum) four, valvate lobes

or petals. Stamens 5(4), alternating with petals,

inserted in the throat of the corolla tube, shortly

below sinuses or on the margins of the top of

ovary in choripetalous species; ﬁ laments free,

ﬂ attened, or anthers sessile (rarely ﬁ laments

almost wanting); anthers 2-thecal. Pollen grains

2-celled, 3-colpate or 3-colporate, trilobate,

smooth. Floral nectaries present and nectar secre-

tion from between the hypanthium and the gynoe-

cium, creating 4–5 nectariferous channels.

Gynoecium of 2(3) carpels; style short and thick,

with massive glabrous stigma; ovary 2(3)-locular,

inferior, adnate to the ﬂ oral tube only by means of

ﬁ ve narrow longitudinal septa formed by continu-

ation of ﬁ laments, which leave intervening lacu-

nae or pits in which nectar is produced. Ovules

numerous on biﬁ d axile placenta, pendulous.

Endosperm only with micropylar haustorium.

Fruits berries with the perianth persisting at the

tip. Seeds minute, exotestal cells cuboid, inner

walls ligniﬁ ed; endosperm copious, starchy; n =

54–56. . . . . . . . . . . . . 1. pentaphragmataceae.

2 Vessels with simple perforations. Annual herbs.

Stem somewhat succulent and with large, verti-

cal, cortical air canals, and pericyclic scleren-

chyma. Leaves alternate, entire, papillose; the

mesophyll containing calcium oxalate crystals.

Stomata tetracytic. Flowers in dense terminal

spikes, in the axils of small bracts and bibrac-

teolate, bisexual, actinomorphic, 5-merous.

Sepals ﬁ ve, with imbricate lobes, persistent.

Corolla small, caducous, urceolate-campanulate,

with ﬁ ve imbricate lobes. Stamens as many as

and alternate with petals, attached to the base of

corolla tube; ﬁ laments short, free; anthers dorsi-

ﬁ xed, introrse. Pollen grains 3-celled, 3-colporate

to colporoidate, reticulate. Nectary disc wanting.

Gynoecium of two carpels; stigma sessile or on

very short style, capitate; ovary 2-locular, inferior

or semi-inferior. Ovules numerous on the large,

spongy, pendulous placenta. Endosperm with ter-

Superorder ASTERANAE 487

minal haustoria. Embryogeny of Onagrad-type.

Fruits membranous pyxidia. Seeds small, with

straight embryo and very scanty or no endosperm;

exotestal cells with inner walls thickened.

Producing fructose with isokestose linkages, n =

12, 16, 20, + . . . . . . . . . . . .2. sphenocleaceae.

1 Plants with network of articulated laticifers. Flowers

usually with specialized mechanisms for pollen pre-

sentation. Anthers free or imperfectly connate. Inulin

present. Alkaloids present or absent. Perennial or

less often biennial or annual herbs, or less often

shrubs, woody lianas (sometimes twining), treelets

or trees to 15 m tall, typically terrestrial, rarely aquatic

or epiphytic; trichomes typically unicellular or some-

times uniseriate. Vessels commonly with simple per-

forations, but some genera (Cyananthus, Platycodon,

Canarina, Musschia) have scalariform perforations

with 1–8 bars; or scalariform and simple, lateral pit-

ting scalariform to alternate. Rays wide, consisting

mostly of upright cells. Axial parenchyma wanting

or indistinct. Nodes unilacunar with one trace. Leaves

alternate, sometimes opposite, entire to pinnately

divided, petiolate or subsessile; leaves of hydro-

phytes submerged and emergent. Stomata anomo-

cytic. Flowers mostly in cymose or racemose

inﬂ orescences, solitary and axillary, usually bisexual,

actinomorphic, or more or less zygomorphic; bracts

foliose or reduced, rarely absent; pedicels often brac-

teolate. Calyx (3-)5(-10)-lobed, the lobes imbricate

or valvate; sometimes basally appendaged, the odd

lobe is dorsal (posterior). Corolla campanulate to

tubular, the lobes valvate or rarely induplicate.

Stamens as many as the corolla lobes and alternate,

inserted near the base of corolla or on the nectary

disc or rarely (Siphocodon and Rhigiophyllum) partly

epipetalous; ﬁ laments usually free; anthers basiﬁ xed,

rarely (Berenice) dorsiﬁ xed, introrse, usually con-

nivent, but commonly separating after anthesis.

Pollen grains in tetrahedral or isobilateral tetrads, 2-

or 3-celled, 5–10-colpate (Cyananthus, Codonopsis,

Leptocodon), 6-colpate (Parishella), (2)3(4)-colpo-

rate, 5–6-colporate, or more often 3–8-zonoporate,

rarely pantoporate, echinate. Gynoecium mostly of

2–10, mostly three carpels; style terminates in a more

or less lobed stigma and is provided with collecting

hairs below the stigma; the stylar hairs eventually

invaginate, dislodging pollen grains. Ovary inferior

or rarely semi-inferior or even superior, often

crowned by nectary disc, with as many locules as

carpels or rarely 1- locular. Ovules mostly numer-

ous, horizontal on axile placentas, rarely placenta-

tion apical Siphocodon) or (Merciera) basal. Fruits

septicidal or loculicidal, or valvular capsules, which

open by valves, slits or pores, located either at the

apex of the ovary (i.e. above the calyx lobes) or on

the lateral walls of the hypanthium (i.e. below the

calyx lobes), or splitting irregularly, often crowned

by the persistent calyx lobes, rarely baccate

(Canarinoideae and Clermontia in Lobelioideae) or

very rarely (Theodoravia) schizocarp. Seeds numer-

ous, small, winged or wingless; exotesta of ligniﬁ ed

cuboid or ﬁ briform cells, endotestal cells thickened;

embryo straight; endosperm ﬂ eshy, oily, or rarely

starchy endosperm. Alkaloids and ﬂ avonols present

or absent, contain tyrosine-derived (with triglochi-

nin); producing 14-carbon polyacetelenes, ursolic

and caffeic acid; very commonly inulin; iridoids

and tannins wanting, n = 6–21, 23–30, 32, 34–36,

40, 45, 48, 51, 52 (the commonest number is 17).

. . . . . . . . . . . . . . . . . . . . . . . . 3. campanulaceae.

1. PENTAPHRAGMATACEAE

J. Agardh 1858. 1/c.30. Lower Burma, southern China,

Indochina, Malesia to New Guinea.

Pentaphragma

It is a relatively less specialized member of the

order. However, inﬂ orescences, pollen grains, fruits,

and seed coat are characterized by a number of derived

characters. Pentaphragmataceae are probably nearest

to the ancestral stock of the Campanulales. Rather

isolated within the order.

2. SPHENOCLEACEAE

Baskerville 1839. 1/2. One species pantropical, another

one in West Africa.

Sphenoclea

An isolated family with a number of unique features,

but it deﬁ nitely belongs to the Campanulales.

3. CAMPANULACEAE

A.L. de Jussieu 1789 (including Cyananthaceae J.

Agardh 1858; Cyphiaceae A.P. de Candolle 1839;

Cyphocarpaceae Reveal et Hoogland 1996; Dortman-

488 Subclass VII. ASTERIDAE

naceae Ruprecht 1856; Jasionaceae Dumortier 1829;

Lobeliaceae A.L. de Jussieu 1813; Nemacladaceae

Nuttall 1842). 92/2000–2400. Very widely distributed,

especially in the temperate regions of the Northern

Hemisphere but relatively sparsely developed in the

tropics. There are only a few endemic genera in South

Africa and adjacent islands of Indian Ocean (including

monotypic Berenice on Reunion). In Australia,

Tasmania, and New Zealand there are only several

species of Wahlenbergia, a large complex subcos-

mopolitan widespread in the Southern Hemisphere.

3.1 CYANANTHOIDEAE (subfam. nov. basionym:

Cyanantheae Meissner 1840)

Pollen grains 4–10-colpate or 5–6-colporate. Vessels

with both simple and scalariform perforations. Ovary

superior to semi-inferior. Calyx lobes without append-

ages between them. Fruits capsular, dehiscing by apical

valves. – cyanantheae: Cyananthus; codonopsideae:

Codonopsis, Numaeacampa, Leptocodon; platy-

codoneae: Platycodon.

3.2 CANARINOIDEAE

Pollen grains 3(4)-colporate to 3-porate. Vessels with

both simple and scalariform perforations. Ovary inferior,

4–7-locular. Fruits baccate. – Canarina, Campanumoea.

3.3 OSTROWSKIOIDEAE

Pollen grains compressed-spheroidal, with strongly

perforate tectum and massive endexine, but without

footlayer, 5–7-aperturate, apertures poroid, elongate-

oval; ornamentation gemmate, with two different types

of gemmae – numerous small and sparsely dispersed

large ones. Vessels only annular and helical. Vascular

bundles widely separated. Pith and primary cortex well

developed. Calyx lobes without appendages. Corolla

very large (up to 10 cm long). Anthers with a placen-

toid. Gynoecium of (5-)7(-9) carpels; ovules with

massive integument. Fruits large capsules with oblong,

longitudinal pores of dehiscence, twice as many as

sepals; each pore is covered by translucent tissue that

later ruptures by vertical ﬁ ssure in the middle. Seeds

winged. High geophytes with thick stem and tuberous

root. – Ostrowskia, Central Asia.

3.4 CAMPANULOIDEAE

Pollen grains zonoporate or sometimes pantoporate.

Roots often thick. Vessels usually with simple perfora-

tions. Ovary inferior or semi-inferior, rarely nearly

superior. Calyx with or without appendages between

the lobes. Fruits capsules, dehiscing in various ways in

different genera. – wahlenbergieae: Wahlenbergia

(including Cephalostigma), Berenice, Theilera,

Gunillaea, Nesocodon, Heterochaenia, Microcodon;

azorineae: Azorina; musschieae: Musschia; echi-

nocodoneae: Echhinocodon; campanuleae: Campan-

ula (including Hyssaria, Annaea, Fedorovia, Mzymtella,

Megalocalyx, Sicyocodon), Gadellia, Symphyandra,

Zeugandra, Brachycodonia, Hanabusaya, Adenophora,

Popoviocodonia, Astrocodon, Trachelium, Feeria;

peracarpeae: Peracarpa, Homocodon; michauxieae:

Michauxia; phyteumateae: Phyteuma, Physoplexis,

Asyneuma, Sergia, Petromarula, Cryptocodon, Cylindro-

carpa, Legouzia, Triodanis, Heterocodon, Githopsis;

edraiantheae: Edraianthus, Muehlbergella; jasion-

eae: Jasione; prismatocarpeae: Prismatocarpus,

Namacodon, Roella, Craterocapsa, Treichelia; siphoc-

odoneae:

Siphocodon, Rhigiophyllum; merciereae:

Merciera.

3.5 NEMACLADOIDEAE

Pollen grains 6-colpate (Parishella) or 3-colporate, with

spinules. Stamens inserted a top the ovary. Nemacladus

has groups of remarkable reﬂ exed pseudonectaries

at the bases of two ﬁ laments. Small, annual or

(Pseudonemacladus) perennial herbs with erect or pros-

trate stems and without tubers. Leaves small, basal.

Flowers small, in racemes or capitate clusters. Calyx

5-lobed. Corolla bilabiate, with two ventral and three

dorsal lobes and lack any sort of palate. Filaments con-

nate in their upper or middle parts; anthers distinct.

Stigma 2-lobed. Ovary inferior, 2-locular, with

numerous ovules. Fruits capsules, 2- or 4-valved from

the tip or (Parishella) its low-conic apex circumscis-

sile just above the calyx lobes. Seeds minute, n = 9.

– Pseudonemacladus, Nemacladus, Parishella.

3.6 CYPHOCARPOIDEAE

Pollen grains 3-colporate, with characteristic reticulate

ornamentation. Stamens epipetalous. Annual herbs.

Leaves spinulose-sinuato-dentate, rigid. Flowers ses-

sile, solitary in the axils of the upper leaves, often

in leafy spikes, with two leaﬂ ike bracteoles. Calyx

5-lobed. Corolla bilabiate, with one-lobed upper (dor-

sal) lip and four-lobed ventral lip with a gibbous pal-

ate; corolla tube curved, long. Stamens attached to the

corolla tube under the limb; ﬁ laments short, distinct;

anthers basiﬁ xed, introrse. Style ﬁ liform, glabrous,

Superorder ASTERANAE 489

apically curved, surrounded at the base by ﬂ eshy annu-

lar nectary disc; stigma divided into two broad lobes

with a small, sticky gland between the lobes; lobes of

the closed stigma are externally covered with setulose

hairs; lobes of the open stigma are glabrous (the exter-

nally hairy stigma functions as a collector of pollen);

ovary 2-locular, with numerous ovules on very thin

septa (fused intrusive placentas), which in very early

stage begins to separate from the ovary wall and trans-

forms into free-central placenta. Fruits membranous

capsules crowned by the calyx lobes. Seeds numerous,

cylindrical, longitudinally costate. – Cyphocarpus.

3.7 CYPHIOIDEAE

Pollen grains 3-colporate, nanogranulate, almost

smooth. Style in the upper part with a ﬂ uid-ﬁ lled stig-

matic cavity communicating with the air through a small

lateral aperture. Perennial herbs usually with a subter-

ranean root tuber containing both inulin and starch.

Stems erect or twining. Leaves alternate, radical or cau-

line, entire, dentate or divided. Flowers zygomorphic, in

racemes, bracteate and with two bracteoles. Calyx

5-lobed; the odd (unpaired) lobe originates in a ventral

(anterior) position. Corolla either split to the base into

two lips with the upper lip 3-lobed and the lower lip

consisting of two free petals, or all petals united into a

tube with a 5-lobed limb. Stamens ﬁ ve, free from petals;

ﬁ laments free or more or less connate at the base; anthers

free or apically loosely united, glabrous or more or less

pubescent on the back of the connective. Gynoecium of

two carpels; style shorter than the stamens, obliquely

clavate, crowned by hairs; ovary semi-inferior to semi-

superior, 2-locular, with usually many ovules per locule.

Fruits capsules dehiscing loculicidally by two apical,

usually biﬁ d valves. Seeds minute, numerous, with

reticulate testa, n = 9. – Cyphia.

3.8 LOBELIOIDEAE

Flowers resupinate at anthesis (except Downingia laeta)

and with specialized mechanism for protandrous pollen

presentation. Herbs or less often more or less woody,

sometimes (Brighamia) with succulent stem. Vessels

with simple perforations. Fibers with simple pits.

Rays heterogeneous. Axial parenchyma scanty para-

tracheal. Leaves alternate, entire to pinnately divided.

Flowers usually in racemose inﬂ orescences, bisexual

or rarely unisexual, more or less zygomorphic. Calyx

5-lobed. Corolla bilabiate or unilabiate, the odd lobe

originates in a dorsal position. Stamens ﬁ ve, inserted

at the base of the corolla tube or at the annular nectary

disc; ﬁ laments connate for at least part of their length;

anthers connate into a tube around the style; the two

ventral anthers are shorter than the other three. Pollen

grains usually 3-colporate or sometimes 3- colpate,

usually reticulate-striate, never echinate. Gynoecium

of two carpels; style provided with a brush of collect-

ing hairs below the initially appressed stigmas; ovary

inferior or rarely semi-inferior (Diastatea), usually

2-locular, with numerous ovules. Fruits capsular and

variously dehiscent or rarely baccate (Clermontia).

Seeds numerous, small, with small, straight embryo

and copious endosperm. Producing pyridine alkaloids

(which commonly accumulate in the latex), chelidonic

acid and 14-carbon polyacetylenes, but no caffeic acid.

n = 6–7, commonly 14. – lobelieae: Siphocampylus,

Diastatea, Laurentia, Palmerella, Solenopsis, Hippobroma,

Porterella, Isotoma, Lobelia, Trimeris, Wimmerella,

Trematolobelia, Grammatotheca, Monopsis, Heterotoma,

Dialypetalum, Ruthiella, Legenere, Unigenes, Downingia,

Howellia, Brighamia, Sclerotheca, Apetahia, Lysipomia;

delisseeae: Pratia, Hypsela, Burmeistera, Centropogon,

Delissea, Cyanea, Clermontia, Rollandia, Dielsantha.

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Order 117. STYLIDIALES

Small herbs, seldom subshrubs or shrubs. Secondary

growth anomalous (Stylidiaceae). Vessels with simple

or (Donatia) with scalariform perforations. Sieve-

elements plastids of S-type. Leaves alternate, densely

crowded, simple, entire, linear, coriaceous, estipulate.

Stomata anomocytic or less often paracytic. Flowers in

racemes or cymose inﬂ orescences, of less often soli-

tary, bisexual or sometimes unisexual, usually zygo-

morphic. Calyx (2-)5(-7)-lobed. Corolla 5–10-partite