Takhtajan Armen. Flowering Plants

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A. Takhtajan, Flowering Plants, 11

Woody or herbaceous plants. Secretory cells with oily

contents ordinarily present in the parenchymatous

tissues. Vessels with scalariform or simple perfora-

tions or vessels wanting. Sieve-element plastids usually

containing protein crystalloid (Pc-type) or ﬁ laments

(Pf-type), often also starch, in some families only

starch (S-type). Stomata commonly paracytic. Flowers

bisexual or less often unisexual, frequently spiral or

spirocyclic, actinomorphic. Stamens mostly numer-

ous. Tapetum usually secretory. Microsporo genesis

successive or simultaneous. Pollen grains 2-celled or

less often 3-celled, 1-colpate, 2-colpate, 3–6-colpate,

rugate, porate, or often inaperturate. Gynoecium

mostly apocarpous. Ovules bitegmic or much less

often unitegmic, usually crassinucellate. Endosperm

cellular or nuclear. Seeds mostly with small or minute

embryo and copious endosperm, sometimes accompa-

nied or largely replaced by perisperm. Cotyledons

typically 2, but occasionally 3 or 4 (Degeneriaceae,

Idiospermaceae). Commonly producing neolignans

and/or benzyl isoquinoline alkaloids, but without

ellagic acid and iridoid compounds.

The subclass Magnoliidae includes a number of

relatively very archaic orders and families of ﬂ owering

plants. All of them are extremely heterobathmic, that

is, they have a very disharmonious combination of

both primitive and derived characters.

Different families of the magnoliids developed in

different directions. Although all of them most proba-

bly evolved from a common ancestral stock.

The basal group of ﬂ owering plants are superorder

Nymphaeanae, which include the most archaic fami-

lies, beginning with Amborellaceae and ending with

Ceratophyllaceae.

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Subclass I. MAGNOLIIDAE

12 Subclass I. MAGNOLIIDAE

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Superorder NYMPHAEANAE 13

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Superorder NYMPHAEANAE

Order 1. AMBORELLALES

Evergreen shrubs with several stems from common

base up to 8 m; young parts tomentose or glabrous.

Nodes unilacunar with a single trace. Xylem ves-

selless. Axial parenchyma scanty diffuse; pericycle

with hippocrepiform sclereids. Rays 1-2-(5)-seriate;

ray cells have ligniﬁ ed secondary walls and contains

droplets or massive yellowish to brownish deposits.

Piths homogeneous, composed of thick-walled, pitted

parenchyma, but devoid of sclerenchymatous cells.

Sieve-element plastids of Ss-type (Behnke 2002). Oil

cells and mucilage cells absent. Leaves alternate, at

ﬁ rst spirally arranged, later distichous, simple, entire,

or repand and obscurely dentate. Stomata paracytic to

anomocytic. Flowers in small, botryoids or poorly

ramiﬁ ed panicles (Endress and Igersheim 2000), uni-

sexual, dioecious, with a slightly convex receptacle

bearing bracteoles that grade into the larger perianth

segments; ﬂ oral members spirally arranged. Male

ﬂ owers with 9–11, inconspicuous perianth segments;

approximately 5 perianth segments are the largest

ones. They are reﬂ exed in the open ﬂ owers. Toward the

base of the ﬂ ower these organs are gradually smaller

and bractlike, but there is no clear separation into a

zone of bracts and a zone of perianth segments (Endress

and Igersheim 2000). Stamens 12–21 with short ﬁ la-

ment, inserted spirally on the inner surface of the

receptacle; anthers triangular, tetrasporangiate, two

pollen sacs somewhat separated, opening longitudi-

nally; connective slightly prolonged. Pollen grains

oblate to spheroidal and monoaperturate (ulcerate) at

the distal pole, with a poorly deﬁ ned aperture margin;

the exine consists in the nonapertural region of ektex-

inous unique tecta elements (“cupules”), a smooth foot

layer, and a thin, nonlamellate endexine (Hesse 2001).

Microsporogenesis successive. Female ﬂ owers with

7–8 perianth segments, 1–2 staminodia (rarely 3 or

none); gynoecium of 5, obovoid, shortly stipitate, free

carpels in a single cycle at the center of the receptacle;

commonly only 3 carpels develop into a fruitlets

(Endress and Igersheim 2000), carpels unsealed at the

tip, with single ovule attached to the lower ventral

wall; stigma large, cup-like, sessile, with 2 conspicu-

ous ﬂ anges, shortly decurrent. Ovule pendant, typi-

cally orthotropous (the chalaza and micropyle are in a

line although the funiculus is curved – Yamada et al.

2001), crassinucellate, bitegmic, outer integument 4–5

cells across, micropyle endostomal. Female gameto-

phyte of Polygonum-type. Endosperm formation is if

ab initio cellular (Tobe et al. 2000). Fruits of oblique,

ﬂ eshy drupaceous, its wall is formed by both endocarp

and mesocarp. Seeds with weakly differentiated mem-

branous testa; seed exsotestal-endotegmic; embryo

basal, minute, endosperm copious, ﬂ eshy, contains

protein. The Amborellaceae are weak accumulators of

aluminium (Metcalfe 1987). n = 13.

1. AMBORELLACEAE

Pichon 1948. 1/1. New Caledonia.

Amborella.

The shrubby Amborella trichopoda is very hetero-

bathmic and combines both very primitive and a highly

specialized features. According to Hesse (2001), “the

unique tectal elements (cupules) and the thick homo-

geneous endexine, which form an odd operculum in

the apertural area, are unusual angiosperm pollen char-

acters”. According to Carlquist and Schneider (2001)

“Amborella shares stomatal conﬁ gurations nodal type

14 Subclass I. MAGNOLIIDAE

(in part), ray types, and porose pit membranes in

tracheary elements with Illiciales s.l., but differs from

that order in lacking oil cells, vessels and grouped

axial parenchyma cells.” However it is very isolated

genus and one of the most archaic taxon within the

Magnoliidae. It differs from all other Magnolliids in its

many respects, including structure of unique type of

pericarp, and deserve the ordered status (Melikian

et al. 1999). Amborella is one of the most remarkable

living fossils which most probably originated directly

from the earlier angiosperm.

Bibliography

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Roman. 2004. Gene sampling versus taxon sampling: Is

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14 Symp. Biodiversität und Evolutionsbiol. Jena.

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Order 2. NYMPHAEALES

Herbaceous aquatic plants with generally submerged

or ﬂ oating leaves; 4-celled uniseriate secretory

trichomes with a large terminal cell. Parenchymatous

tissues without ethereal oil cells and with conspicuous

air-passages and with laticifers. Vascular bundles

closed, scattered, or less commonly in one or more

rings. Vessels absent. Sieve-element plastids of S-type.

Leaves spiral, involute, peltate (divided), simple, entire

Superorder NYMPHAEANAE 15

or margin toothed, stipulate (the stipules median-

axillary), or estipulate. Stomata anomocytic. Flowers

solitary, often large, bisexual, actinomorphic, mostly

spirocyclic. Perianth basically 3-merous. Stamens usu-

ally numerous; ﬁ laments free; anthers extrorse or

latrorse, tetrasporangiate, opening longitudinally.

Tapetum secretory. Microsporogenesis mostly simul-

taneous. Pollen grains 3-celled or (Cabomba) 2-celled,

1-colpate, zonocolpate, or inaperturate, mostly in

monads. Gynoecium apocarpous or syncarpous.

Carpels 1–50, large to very large. Ovules 3–400, usu-

ally anatropous with cup-shaped outer integument,

bitegmic, crassinucellate. Endosperm usually cellular.

Female gametophyte of Polygonum-type or rarely also

both Polygonum- and Oenothera-type (Orban and

Bouharmont 1998). Fruits of various types. Seeds

operculate, exotesta palisade, mostly with small

embryo; endosperm scanty, perisperm present or

absent, with or without arillus. Without benzyl isoqui-

noline alkaloids, but with pseudoalkaloids based on

sesquiterpenes and ellagitanninses.

There is every reason for considering the superorder

Nymphaeanae as a hydrophilous derivative of some

ancient vesselless arborescent or subherbaceous mag-

noliid, most likely of inhabitants of moist or marshy

places, whose vegetative organs became modiﬁ ed in

an aquatic environment. According to Arber (1920:

309), the nymphaeids “descended from a stock closely

related to that which gave rise to the monocotyledons.”

The nymphaeids and some liliopsids, particularly the

Alismatidae, do indeed have much in common. Besides

the apocarpous gynoecium with laminar-diffuse pla-

centation of the ovules, pollen grain morphology, atac-

tostelic vascular cylinder, more or less reduced primary

root, and similar arrangement of the prophylls, there

are also striking similarities in the “liorhizous” type of

root development (because of the presence of second-

ary dermatogen, their root-cap is sharply cut off from

the rhizodermis) (van Tieghem 1886; Voronin 1966;

Voronkina 1974), and similar formation of the plumule

(Meyer 1960). It is therefore not surprising that some

botanists regarded Nymphaeaceae s. 1. as monocots

(Trecul 1845; Seidel 1869; Schaffner 1904, 1934;

Cook 1906; Guttenberg and Müller-Schroeder 1958).

But in spite of many similarities with the monocots,

nymphaeids are dicotyledonous, and their afﬁ nity with

the basal Magnoliidaes, especially with Amborellales,

is supported also by molecular data (see Qiu et al.

1993, 1999; D. Soltis et al. 2000; Zanis et al. 2002,

and others).

Key to Families

1 Plants with leafy stems coated with a mucilaginous

secretion. Sclerenchymatous idioblasts lacking.

Flowers relatively small. Microsporogenesis simul-

taneous or successive. Female gametophyte of

Schisandra-subtype of Oenothera-type (at least in

Cabombaceae). Endosperm helobial with chalazal

haustorium. Fruits coriaceous, follicle like or

achene like. Seeds exarillate.

2 Submerged parts with thick, transparent, gelati-

nous sheath; stems encased in a thick layer of

mucilage; there are paired, glandular patches at

the nodes. Leaves monomorphic, all alternate,

estipulate, with ﬂ oating entire and peltate laminas.

Flowers without nectariferous spurs, anemophil-

ous. Perianth only slightly differentiated into

sepals and petals, dull- purple. Sepals and petals 3

or 4 each. Stamens 18–36. Anthers latrorse.

Microsporogenesis simultaneous. Pollen grains

3-celled, the exine scabrate. Carpels 4–22, large;

stylodium short, with linear, decurrent, and

extremely papillate stigma. Ovules 1 or 2 per car-

pel, large to very large, dorsal or near dorsal.

Antipodals ephemeral. Fruits achenelike, contain-

ing 1–2 seeds, n = 40.. . . . l. hydropeltidaceae.

2 Plants rhizomatous, without a gelatinous sheath.

Leaves dimorphic, submerged opposite or verticil-

late, palmately dissected into linear segments,

estipulate, ﬂ oating ones few, peltate, usually alter-

nate. Flowers with nectariferous spurs at the adaxial

base of petals, myophilous. Perianth clearly differ-

entiated into sepals and petals, white to yellow or

purple. Sepals and petals (2)3(-4) each. Stamens 3

or 6. Microsporogenesis successive. Anthers

extrorse to latrorse. Pollen grains 2-celled, with stri-

ate ornamentation composed of parallel rods.

Carpels (1-)2-18, small to medium-sized, whorled,

occasionally connate at the base; stylodium long,

with a spherical papillate stigma. Ovules (1)2-3 per

carpel, the upper ovule ventral, the two lower ones

lateral. micropyle endostomal. Antipodals lacking.

Fruits follicle like or achenes, dehiscing along the

dorsal side, containing 2–4 seeds. Alkaloids absent,

n = 40, 48, 52. . . . . . . . . . . . . . . .2. cabomba ceae.

1 Acaulescent perennial or rarely annual plants with

leaves arising directly from the horizontal or verti-

cal rhizomes. Branched sclerenchymatous idio-

blasts common. Nonbranched articulated laticifers

occur throughout the ground tissue and usually

associated with the vascular tissue. Flowers large,

16 Subclass I. MAGNOLIIDAE

generally emergent. Fruits spongy berries. Seeds

arillate or exarillate.

3 Petals distinct, very rarely absent. Stamens free.

Pollen grains 3-celled, I-colpate or more often

zonocolpate. Ovules anatropous. Endosperm cel-

lular. Seeds arillate or exarillate, not spiny. Aquatic,

rhizomatous herbs. Leaves medium-sized to large,

alternate, simple, petiolate, usually more or less

peltate, stipulate or estipulate. Flowers solitary,

large. Calyx imbricate; corolla 0–70, imbricate.

Stamens 14–200, petaloid, or laminar, of ﬁ lanther-

ous; anthers adnate, non-versatile, introrse.

Gynoecium of 5–35 carpels; ovary 5–35- locular,

superior to partly inferior. Ovules 10–100 per loc-

ule, orthotropous or anatropous, arillate or non-

arillate. Fruit ﬂ eshy, dehiscent or indehiscent, or a

schizocarp. Seed numerous; embryo green or

white, straight; endosperm and perisperm present.

Proantho cyanidins (cyanidin or delphinidin) pres-

ent, ﬂ avonols (kaempferol, quercetin, or myrice-

tin) present or (Nuphar) absent, n = 10–12,

14–18.. . . . . . . . . . . . . . . . . . 3. nymphaeaceae.

3 Petals connate into a lobed tube. Stamens 20–30,

attached to the corolla tube, free of one another.

Pollen grains 2-celled, inaperturate, with a

reduced exine. Ovary inferior. Ovules orthotro-

pous. Seeds exarillate, spiny. Stemless, laticiferous

herbs. Laticifers in leaves, stems, roots, in ﬂ owers,

and in the fruits. Leaves ﬂ oating, alternate, mem-

branous, simple, long petiolate, estipulate. Flowers

solitary, long-pedunculate, but often not reaching

the surface of the water, at least sometimes cleis-

togamous. Corolla 12–50 lobes; Staminodes

15–20, external to the fertile stamens. Filaments

short; anthers basiﬁ xed, non-versatile, 2-locular.

Gynoecium of 8–14 carpels; ovary 8–14-locular,

ovules 30–50 per locule. Fruit a berry; seeds

numerous, densely rather softly spinulose; embryo

minute, well differentiated; perisperm and

endosperm present, n = 17, 18. 4. barclayaceae

1. HYDROPELTIDACEAE

Dumortier 1822. 1/1. Eastern Himalayas (Bhutan),

Assam, eastern Asia, eastern Australia, tropical Africa,

North America (from Alaska to Florida), Central

America, West Indies.

Brasenia (Hydropeltis).

2. CABOMBACEAE

Richard ex A. Richard 1822. 1/5. Warm-temperate and

tropical regions of America.

Cabomba.

Closely related to the Hydropeltidaceae, which are

usually included in the Cabombaceae. However, Moseley

et al. (1984), comparing anatomical and morphological

features of both vegetative organs and ﬂ owers in Brasenia

and Cabomba) came to the conclusion that the two gen-

era are more distant taxonomically than commonly

assumed. This view is also shared by Collinson (1980).

According to Osborn et al. (1991: 1367), “there is little

argument that Brasenia and Cabomba are closely linked.”

The differences are so many that I ﬁ nd it necessary to

accept Dumortier’s family Hydropeltidaceae.

3. NYMPHAEACEAE

R. A. Salisbury 1805 (including Euryalaceae J. Agardh

1858; Nupharaceae Kerner 1891). 5/75. Almost cos-

mopolitan, in fresh water habitats. Nuphar (20) tem-

perate and cold regions of the Northern Hemisphere,

Victoria (2) in tropical South America, Euryale (1) in

northern India, China, and Japan, Ondinea (1) in north-

ern western Australia (sandstone streams of the north-

ern Kimberley region).

3.1 NUPHAROIDEAE

Gynoecium superior, its distal part forming a ﬂ attened

to somewhat concave disc with rays of stigmatic tissue.

The stigmatic area is displayed on a ﬂ at plate. Pollen

spiny. Carpels 5-23(-30), glabrous. Sepals 5–17. Petals

small, each with an abaxial nectary. Nucellar cap pres-

ent. Seeds exarillate. Aquatic herbs with stout rhizomes.

Leaves large, entire, with basal sinus, submersed, ﬂ oat-

ing or standing erect above water. n = 17. –Nuphar.

3.2 NYMPHAEOIDEAE

Gynoecium inferior to semi-inferior, its distal part

cup-shaped and covered with stigmatic tissue. Leaf

veins, peduncles, sepals, and petals not aculeate. The

stigmatic area is displayed on a concave groove. Pollen

grains in monads. Carpels 3–47, covered by

pluricellular-uniseriate hairs. Sepals 4, rarely 5. Petals

(0)4-5 to numerous, not small, without nectaries (?).

Seeds arillate. n = 10–12, 14–18. – nymphaeeae:

Nymphaea; ondineae: Ondinea.

Superorder NYMPHAEANAE 17

3.3 EURYALOIDEAE

Leaf veins, peduncles, sepals, and petals aculeate.

Pollen grains in tetrads. – Euryale, Victoria.

4. BARCLAYACEAE

H. L. Li 1955. 1/4. Thailand and Burma to East Malesia.

Barclaya (Hydrostemma).

Close to the Nymphaeaceae, but differ from them in

some important features, including congenital carpel

fusion encompasses a considerable part of these “ stylar

processes”, and are more advanced.

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