Takhtajan Armen. Flowering Plants

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534 Subclass VIII. LAMIIDAE

endosperm nuclear; embryo curved. Present unique

tropane alkaloids with senecioic, mesaconic and

itaconic acids, n = 10. – Schizanthus.

1.6 ANTHOCERCIDOIDEAE

Woody shrubs or trees. Flowers with non-accrescent

calyx, inﬂ exo-valvate aestivation of the corolla lobes,

a short relatively broad actinomorphic corolla tube,

extrorsely dehiscing stamens inserted low in the

corolla tube, and an oblong to ellipsoid, slightly curved

seed with reticulate testa, embryo slightly curved.

Fruits capsules or rarely (Duboisia) berries. Endosperm

copious, sometimes including a small oily sector

(Hunziker 2001) Produce nicotinic and tropane alka-

loids, n = 9, 10. – Anthocercis, Anthotroche,

Cyphanthera, Crenidium, Duboisia, Gramnosolen,

Symonanthus.

2. NOLANACEAE

Dumortier 1829. 1–2/60. According to Mesa (1981)

Nolana comprises 18 species of which 15 are in tropi-

cal America, other modern authors have suggested that

there may be as many as 83 American species: Southern

Peru, northern Chile, Galspogos Is.

Nolana (? including Alona)

3. SCLEROPHYLACACEAE

Miers 1848. 1/15. Argentina, Paraguay, Uruguay.

Sclerophylax

Close to Solanaceae.

4. DUCKEODENDRACEAE

Kuhlmann 1950. 1/1. Amazon Basin in Brazil.

Duckeodendron

Related to the Solanaceae (see Kuhlmann 1950;

Carlquist 1988).

5. GOETZEACEAE

Miers ex Airy Shaw 1965. 4/5. Greater Antilles (Cuba,

Hispaniola, Puerto Rico).

Coeloneurum, Goetzea, Henoonia, Espadaea

Close to the Solanaceae (Radlkofer 1888; Hunziker

1979; Carlquist 1988; Zona 1989), but differ from

them markedly in many respects, including pollen

morphology (see Gentry 1986).

6. HYDROLEACEAE

Berchtold et J. Presl 1820. 1/12. Tropical.

Hydrolea

Related to the Solanaceae (APG II 2003; Erbar et al.

2005).

7. CONVOLVULACEAE

A.L. de Jussieu 1789 (including Cressaceae Raﬁ nesque

1821, Dichondraceae Dumortier 1829, Erycibaceae

Endlicher ex Meisner 1840, Poranaceae J.G. Agardh

1858). 56/1700. Cosmopolitan, but mainly in subtropi-

cal regions of Asia and America.

ipomoeeae: Argyreia, Astripomoea, Blinkworthia,

Ipomoea (including Pentacrostigma), Lepistemon,

Lepistemonopsis, Paralepistemon, Stictocardia, Rivea,

Turbina, merremieae: Merremia, Hewittia,

Hyalocystis, Decalobanthus, Xenostegia, Remirema,

Operculina; convolvuleae: Convolvulus, Calyste gia,

Polymeria; aniseieae: Aniseia, Hewettia, Iseia,

Odonellia, Tetralocularia; jacquemontieae:

Jacquemontia; cresseae: Cressa, Bonamia,

Cladostigma, Evolvulus, Seddera, Hildebrandtia,

Sabaudiella, Stylisma, Itzaea, Neuropeltis,

Neuropeltopsis; wilsonieae: Wilsonia; maripeae:

Dicranostyles, Maripa, Lysiostyles, dichondreae:

Dichondra, Falkia, Nephrophyllum, Petrogenia,

Porana, Metaporana, Calycobolus, Dipteropeltis,

Rapona; erycibeae: Erycibe; cardiochamyeae:

Cordisepalum, Poranopsis, Cardiochlamys,

Tridynamia, Dinetus.

8. CUSCUTACEAE

Berchtold and J. Presl. 1820. 1/170. Nearly cosmo-

politan, but best represented in America, especially in

warmer regions.

Cuscuta

Very close to the Convolvulaceae, but differing from

them markedly in parasitic habit, corolla morphology,

Superorder LAMIANAE 535

seeds with more copious endosperm, the absence of

intraxylary phloem, and embryological features (Tiagi

1951; Johri and Tiagi 1952).

9. HUMBERTIACEAE

Pichon 1947. 1/1. Madagascar

Humbertia

Closely related to the Convolvulaceae.

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Superorder LAMIANAE 539

Order 122. BORAGINALES

Herbs, shrubs, or less often trees, sometimes lianas,

rarely herbaceous parasites. Stems and leaves of the

herbaceous forms and less frequent in the woody gen-

era provided with characteristic harsh unicellular hairs

that usually have a basal cystolith and often calciﬁ ed

or siliciﬁ ed walls; less often also with gland-tipped

hairs or (Lennoaceae) provided only with stalked,

glandular hairs. Vessels with simple or very rarely

reticulate perforations; lateral pitting alternate. Fibers

with bordered or more often simple pits, often septate.

Rays heterogeneous to homogeneous. Axial paren-

chyma of various types, but mostly apotracheal.

Vestured pits present or absent. Sieve-element plastids

of S-type. Nodes unilacunar, very rarely (Cordia) trila-

cunar with three traces. Leaves alternate or sometimes

partly or wholly opposite, simple, usually entire, some-

times lobed, pinnatisect or pinnate, rarely palmate,

estipulate. Stomata usually anomocytic. Flowers in

scorpioid cymose inﬂ orescences (Buys and Hilger

2003), rarely solitary, mostly bisexual, actinomorphic

or slightly zygomorphic, perianth and androecium

usually 5-merous. Sepals more or less connate, imbri-

cate or rarely valvate. Corolla sympetalous, imbricate

or contorted, rarely valvate. Stamens as many as and

alternate with the corolla lobes, attached to the tube;

anthers tetrasporangiate, introrse, or rarely

(Hoplestigma) latrorse, opening longitudinally. Pollen

grains (2)3-celled, 3-colpate, 3-colporate, 6-colpate,

pericolpate, pericolporate or 3-porate. Nectary disc

present around the base of the ovary or wanting.

Gynoecium of two median carpels, rarely of 4–5 (in

Zoelleria, Boraginaceae), 6–14 (Lennoaceae) carpels;

stylodia usually more or less united into terminal or

gynobasic style; ovary superior or rarely semi-inferior

(as in Nama, Hydrophyllaceae), with two to many

ovules on each placenta or in each locule. Ovules ana-

tropous, rarely hemitropous (some Boraginaceae) or

campylotropous (Lennoaceae), unitegmic, tenuinucel-

late or rarely (Ehretioideae) pseudocrassinucellate

(crassinucellate according to Gottschling 2004),

sometimes with endothelium (Hydrophyllaceae,

Heliotropium in Boraginaceae). Female gametophyte

usually of Polygonum-type, rarely (some Boraginaceae)

of Allium-type. Endosperm cellular or less often

( species of Phacelia in Hydrophyllaceae and some

Boraginoideae) nuclear, rarely (species of Echium and

Pulmonaria) helobial. Often micropylar and/or chala-

zal haustoria present. Fruits of various types. Seeds

exotestal, with straight or nearly straight embryo and

copious (many Hydrophyllaceae) or scanty endosperm

or often (most of Boraginaceae) without endosperm; in

seed coat, funicule and placenta of most member or

the order (except Boraginoideae) present specialized

transfer cells (Diane, Hilger and Gottschling 2002).

Boraginales are connected with the Solanales and

most probably share a common origin.

Key to Families

1 Autotrophic plants.

2 Petals usually ﬁ ve.

3 Ovary basically 1-locular, with two, often more

or less intruded, parietal placentas, which are

sometimes meeting and joined, dividing the

ovary into two secondary locules (locelli).

Ovules many to two on each placenta. Annual,

biennial, and perennial herbs, subshrubs,

shrubs, sometimes spiny, rarely small trees (to

5 m in Wigandia). Stems, leaves and ﬂ owers

often variously pubescent, sometimes with

stringing hairs (as in Wigandia). Leaves alter-

nate or sometimes partly or wholly opposite,

basal leaves sometimes in rosettes, petiolate,

simple to pinnately or rarely palmately divided,

entire or lobed. Stomata anomocytic. Flowers

in terminal or axillary scorpioid cymes, rarely

solitary or paired in axils, bisexual, 5-merous

or rarely (South African genus Codon)

10–12-merous. Calyx usually divided some-

times to the base or nearly so, lobes imbricate,

sometimes with appendages between the calyx

lobes. Corolla sympetalous, actinomorphic or

nearly so, with imbricate or less often con-

torted lobes, often with scales inside the tube,

alternating with the stamens. Stamens (4-)5(-

10), inserted near the base of the corolla tube;

ﬁ laments variously, basally appendiculate;

anthers dorsiﬁ xed, usually with small scales at

each side of the base, introrse. Pollen grains

3- colpate, 3-colporate, or 5–6-colporate, with

tectate- perforate to semitectate exine structure,

basically reticulate. Nectaries present at base

of ovary (Hofmann 2004). Gynoecium of two

carpels; stylodia more or less connate, some-

times free essentially to the base, mostly with

capitate stigmas; ovary usually superior, rarely

540 Subclass VIII. LAMIIDAE

semi-inferior, 1–2-locular. Ovules two to many

in each locule, pendulous, anatropous, usually

tenuinucellate, crassinucellate (Nama – see di

Fulvio 1989a), with an endothelium. Endosperm

cellular or less often nuclear. Fruits usually

loculicidal (sometimes also septicidal) cap-

sules or sometimes irregularly dehiscent or

indehiscent. Seeds small, ruminate by inpush-

ings of the exotestal cells or not, sometimes

carunculate; exotestal cells thickened on inner

and radial walls, endotestal cells persistent;

embryo small to large, green or white, straight,

surrounded by copious or scanty, oily

endosperm. Flavonols present ( kaempferol and

quercetin), n = 5–9, 11–13, 19, 38 (Wigandia

kunthii).. . . . . . . . . . . . 1. hydrophyllaceae.

3 Ovary at maturity with twice as many second-

ary locules (locelli) as carpels, each with one

ovule, rarely one carpel suppressed in subfam-

ily Cordioideae and the ovary with only two

locelli. Herbs, subshrubs, less often shrubs or

trees, seldom lianas (Cordia, Tournefortia

spp.); taproots or creeping rhizomes sometimes

present, rarely present tubers (in Lithospermum

and Heliotropium); plant often roughly hairy,

hairs usually strigose to hirsute, with a basal

cystolith or cystolith-like body. Leaves alter-

nate, very rarely opposite (in Tournefortia), or

sometimes the lowest of them or all opposite,

mostly entire, in a very few species serrate.

Inﬂ orescences of one or more terminal or lat-

eral helicoids or scorpioid cymes, or ﬂ owers

contracted into globose or club-shaped heads,

or ﬂ owers solitary; bracts sometimes present.

Flowers mostly bisexual, usually 5-merous,

actinomorphic or rarely zygomorphic. Sepals

free or basally connate, imbricate or rarely val-

vate. Corolla imbricate, rarely valvate, or con-

torted (in Myosotis), with faucal, hairy or

scale-shaped appendages. Stamens of the same

number as corolla lobes, alternating with them;

anthers 2-locular, sessile or on distinct ﬁ la-

ments, sometimes with a sterile pit of connec-

tive tissue or produced to an awn-like structure,

awns of one ﬂ ower twisted of not, basiﬁ xed or

dorsiﬁ xed. Pollen grains from 3-colporate or

3-porate to pericolpate or pericolporate, some-

times with six colpi, the alternate ones with and

without a pore. Annular nectary disc present

around the base of the ovary, often covered by

basal appendages formed by corolla-tube, or

absent. Gynoecium of two or rarely (Zoelleria)

4–5 carpels; stylodia connate into terminal or

gynobasic style; ovary usually 2-locular, undi-

vided, entire or with four deep furrows, break-

ing into two biovulate halves or into four

uniovulate mericarps, sometimes part of the

locules aborted, rarely a greater number of

carpels developed (e.g. in Trigonotis procum-

bens), placentation axillary (Riedl 1997).

Ovules anatropous to hemitropous, erect or

nearly horizontal, rarely pendulous, some-

times (Heliotropium) with an endothelium.

Fruits drupes with 1–4 pyrenes (Cordiodeae,

Ehretroideae, and part of Heliotropi oideae),

separated into four or rarely two (Cerinthe)

1-seeded, nutlike mericarps (Boraginoideae),

or capsular (Wellstedioideae). Seeds with

straight or sometimes more or less curved

embryo; endosperm of several cell layers

(Heliotropoideae) or reduced to a single cell

layer. Producing alkaloids of pyrolizidine group

and ﬂ avonols (kaempferol and quercetin), ﬂ a-

vonol glycosides astragalin and nicotiﬂ orin,

isokestose and higher inulin oligosaccharids as

storage, n = 4–14, 16, 21.. . 2. boraginaceae.

2 Petals 11–14. Deciduous trees; cork superﬁ cial,

secondary thickening developing from a conven-

tional cambial ring. Leaves alternate, very large,

petiolate, chartaceous, simple, entire. Flowers

large, in terminal bractless, brown-hirsute sub-

scropioid cymes, bisexual, actinomorphic. Calyx

globose, persistent, irregularly splitting into 2–4

lobes. Corolla with short tube, lobes imbricate.

Stamens about 20–35, in about three irregular

series attached to the base of the corolla tube; ﬁ la-

ments ﬁ liform; anthers dorsiﬁ xed, latrorse. Pollen

grains 3-colporate with meridional ridges alter-

nating with intercolpar concavities. Gynoecium

of two carpels; ovary superior, 1-locular with two

intruded, forked, parietal placentas; each placenta

with two pendulous, anatropous ovules; stylodia

basally connate, elongate, slender, curved, or bent

in the middle and each with a disclike horseshoe-

shaped stigma. Fruits drupaceous, 4-seeded, sur-

rounded at the base by the calyx, with leathery

pericarp and hard endocarp. Seeds with rather

large, nearly straight embryo with elongate

Superorder LAMIANAE 541

hypocotyl and expanded cotyledons and scanty

endosperm. . . . . . . . . . . 3. hoplestigmataceae.

1 Fleshy perennial (Pholisma) or annual (Lennoa)

parasitic herbs. Stems subterranean or nearly so.

Roots spreading horizontally and initiating new

haustoria by short, lateral branches. Leaves alter-

nate, simple, entire, reduced to mere scales, linear

to broadly deltoid. Flowers in cymose-thyroid,

cymose-paniculate, spicate, or capitulate inﬂ ores-

cences, bisexual, actinomorphic or nearly so,

(4)5–9(10)-merous as to the perianth and androe-

cium. Calyx persistent, tubular or divided nearly to

base into narrow lobes, puberulent to tomentose

with stalked glandular trichomes. Petals united

nearly to apex, limb variously lobed, induplicate

valvate or imbricate, persistent. Stamens 5–10,

inserted in the throat of the corolla tube, 1–2-seri-

ate; ﬁ laments short; anthers introrse. Pollen grains

2-celled, 3–4(5)-colporate or colpate and colporate,

often with alternating aperturate and inaperturate

colpi. Nectary disc wanting. Gynoecium of 5–16

carpels; style terminal, with capitate or slightly

5–9-lobed stigma; ovary superior, entire or shal-

lowly lobed; each primary locule secondarily

divided by false septum into two locelli; ovules two

per locule (solitary in each locellus), horizontal,

anatropous, epitropous. Endosperm cellular. Fruits

slightly depressed capsules, somewhat ﬂ eshy, but

drying at maturity; dehiscence circumscissile by an

irregular ring below the middle of capsule. Seeds

small, ovate to slightly reniform, laterally ﬂ attened,

with reticulate surface; embryo undifferentiated,

globose, embedded in the copious, starchy

endosperm, n = 9. . . . . . . . . . . . . .4. lennoaceae.

1. HYDROPHYLLACEAE

R. Brown 1817 (including Ellisiaceae Berchtold et

J. Presl 1820, Eutocaceae Horaninow 1847, Sagoneaceae

Martynov 1820). 16/300. Mainly America, especially

western parts of the U.S.A., but also Hawaiian Islands

(Nama sandwicensis), and South Africa (Codon).

1.1 HYDROPHYLLOIDEAE

Flowers (4-)5-merous; stigma capitate. – hydrophyl-

leae: Hydrophyllum, Pholistoma, Ellisia, Nemophila;

phacelieae: Eucrypta, Draperia, Phacelia,

Emmenanthe, Eriodictyon, Turricula, Tricardia,

Hesperochiron; nameae: Nama, Wigandia,

Romanzofﬁ a.

1.2 CODONOIDEAE

Flowers 10–12-merous. Calyx deeply linear-lobed.

Corolla campanulate, 10–12-lobed. Style terminal,

stigma dentate. – Codon.

Related to the Boraginaceae.

2. BORAGINACEAE

A.L. de Jussieu 1789 (including Anchusaceae Vest

1818, Buglossaceae Hoffmannsegg et Link 1809,

Cerinthaceae Martynov 1820, Cordiaceae R. Brown

ex Dumortier 1829, Ehretiaceae C. Martius 1827,

Heliotropiaceae Schrader 1820, Onosmataceae

Martynov 1820, Wellstediaceae Novak 1943). Circa

130/2500. Widely distributed in tropical and especially

subtropical and temperate regions; the largest subfam-

ily, the Boraginoideae, is centered in the Mediterranean

and Irano-Turanian regions.

2.1 EHRETIOIDEAE

Trees or shrubs, seldom herbs (Coldenia). Flowers (4-)5-

merous. Corolla imbricate or inwards-folded. Pollen

3-porate. Style terminal, simple or more or less cleft, or

two terminal nearly free stylodia (Rochefortia, Coldenia,

Pteleocarpa). Ovary entire or 4-lobed. Endosperm cel-

lular. Fruits more or less drupaceous or dry, not breaking

at maturity (as in Carmona) or breaking into four one-

seeded or into two two-seeded mericarps. Seeds with

copious to scanty endosperm or endosperm absent.

Cotyledons ﬂ at. Tropical and subtropical regions, n = 5,

7–11, 13, 16 21. – Coldenia, Tiquilia, Pteleocarpa,

Lepidocordia (including Antrophora), Rochefortia,

Rotula, Ehretia, Carmona, Pteleocarpa, Bourreria,

Cortesia, Halgania.

2.2 CORDIOIDEAE

Odoriferous trees, lianes, or shrubs. Corolla contorted

(imbricate). Style terminal, once or twice forked.

Ovary undivided. Endosperm cellular. Fruits variable,

dry with a ﬁ brous or bony pericarp, or thin-ﬂ eshed

drupes with a thin exocarp, juicy to mucilaginous

mesocarp, and bony endocarp, usually 1-locular and

1-seeded. Seeds without endosperm. Cotyledons

plicate, toothed. Present terpenoid-based quinones.

Tropical and subtropical, n = 9, 14–16, 19. – Cordia

542 Subclass VIII. LAMIIDAE

(including Varronia, Gerascanthus, ? Saccellium,

Patagonula, Auxemma).

2.3 HELIOTROPIOIDEAE

Odoriferous trees, shrubs, lianes, or herbs. Corolla imbri-

cate or with involute margins. Stigma receptive only

basi-laterally, discoid, then conical and more or less

2-lobed at sterile apex, or hemispherical, with a ring of

hairs, wet. Ovary entire or 4-lobed. Endosperm cellular.

Fruits ﬂ eshy or dry, lobed or unlobed, at maturity break-

ing into two or four bony, 1- or 2-seeded mericarps.

Seeds exotestal, without endosperm. Cotyledons not pli-

cate. Contain pyrrolizidine alkaloids and salicylic acid

(Tournefortia). Tropical, subtropical, and warm-temper-

ate regions. n = 5, 7–9, 11–14. – Tournefortia, Argusia,

Heliotropium (including Parabouchetia), Valentiniella,

Mallotonia, Beruniella, Ixorhea, Nogalia.

2.4 BORAGINOIDEAE

Mainly herbs. Corolla often rotate. Pollen grains 4–6

colporate. Style gynobasic, entire or lobed, arising

from the base of usually deeply 4-lobed ovary.

Endosperm cellular or nuclear, sometimes helobial.

Fruits of four one-seeded mericarps (8–10 in Zoelleria).

Endosperm scanty or absent. Mostly subtropical and

temperate regions. Containing pyrrolizidine alkaloids,

lakannin, n = 4–13. – trigonotideae: Trigonotis

(including Pedinogyne, Zoelleria), Omphalotrigonotis,

Sinojohnstonia, Brachybotrys, Bothriospermum,

Mertensia, Pseudomertensia, (including Scapiceph-

alus), etc; eritricheae: Plagiobothrys, Allocarya,

Amblynotus, Antiphytum, Echiochilon, Ogastemma,

Sericostoma, Chamissoniophila, Amsinckia, Microula

(including Schistocaryum), Anoplocaryum, Trigono-

caryum, Microcaryum, Chionocharis, Actinocarya,

Hackelia, Asperugo,, Eritrichium, Metaeritrichium,

Tianschaniella, Lepechiniella, Heterocaryum, Cranio-

spermum, Lappula, Lasiocaryum, Stephanocaryum,

Cryptantha; rochelieae: Rochelia; cynogloseae:

Bothrio spermum, Antiotrema, Cynoglossum, Pardo-

glossum, Ivanjohnstonia, Austrocynoglossum, Para-

cynoglossum, Harpagonella, Lindeloﬁ a, Adelocaryum,

Rindera, Trichodesma, Lacaitaea, Omphalodes, Sole-

nanthus, Kuschakewiczia, Trachelanthus, Mattiastrum,

Paracaryum, Thyrocarpus, Suchtelena, Caccinia,

Heliocarya, etc. lithospermeae: Arnebia, Huynhia,

Macrotomia, Lithospermum (including Aegonychon),

Ulugbekia, Neatostema, Buglossoides, Lithodora,

Onosma, Maharanga, Cerinthe, Moltkia, Moltkiopsis,

Stenosolenium, Alkanna, Halacsya, Echium, Lobo-

stemon, Echiostachys, Ancistrocarya, etc.; borag-

ineae: Pulmonaria, Mertensia, Nonea, Nephro carya,

Paraskevia, Elizaldia, Symphytum, Brunnera, Anchusa,

Lycopsis, Anchusella, Phyllocara, Cynoglottis,

Gastrocotyle, Hormuzakia, Pentaglottis, Pectocarya,

Borago, Trachystemon, etc.; myosotideae: Myosotis.

2.5 WELLSTEDIOIDEAE

Low woody herbs or shrublets. Leaves grey-strigose.

Flowers bisexual, solitary, 4-merous. Calyx imbricate

or open in bud; corolla imbricate. Pollen grains

3- colporate. Nectary disc absent. Style terminal,

shortly bind; stigmas two; ovary 2-locular, with soli-

tary, pendulous ovule per locule. Fruits compressed,

broadly obcordate, 1–2-seeded loculicidal capsules.

Seeds comose, without endosperm, embryo curved,

cotyledons accumbent. Southwestern Africa, Somalia,

Socotra. – Wellstedia.

Ehretioideae are the most archaic group within the

family (free or nearly free stylodia and copious

endosperm in some genera). Cordioideae are more

advanced (plicate cotyledons and usually more special-

ized pollen grains). Heliotropioideae are closely con-

nected with the Ehretioideae and probably derived from

them. Boraginoideae, which are by far the largest and the

most diverse subfamily of the Boraginaceae, have proba-

bly a common origin with the Heliotro pioideae.

Wellstedioideae with their 4-numerous ﬂ owers and cap-

sular fruits are the most isolated group within the family.

3. HOPLESTIGMATACEAE

Engler et Gilg 1924. 1/2. Tropical West Africa from

Cameroon to Gabon.

Hoplestigma

Closely related to the Boraginaceae (Hallier [1911,

1912] even included Hoplestigma in Boraginaceae),

especially to the Ehretioideae (Takhtajan 1987), which

is supported by palynological data (Erdtman 1952;

Nowicke and Miller 1989).

4. LENNOACEAE

Solms-Laubach 1870. 3/6–7. California, Arizona,

Mexico, Central America, northernmost Colombia,

and Venezuela.

Superorder LAMIANAE 543

Lennoa, Pholisma, Ammobroma

Related to the Hydrophyllaceae and Boraginaceae

(Hallier 1912, 1923; Suessenguth 1927; Copeland

1935; Wettstein 1935; Avetisian 1956; Drugg 1962;

Takhtajan 1966, 1987; Yatskievych and Mason 1986;

Thorne 1992a, b, 2000), in particular to the Ehretioideae.

However, the Lennoaceae are very advanced and rather

isolated within the order.

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