Takhtajan Armen. Flowering Plants

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A. Takhtajan, Flowering Plants, 293

Trees, shrubs, subshrubs, or perennial or annual herbs.

Vessels with simple or less often scalariform perfora-

tions (sometimes with many bars). Sieve-element

plastids mostly of S-type. Leaves alternate, opposite or

verticillate, simple or often compound, stipulate or

estipulate. Stomata of various types. Flowers in vari-

ous types of inﬂ orescences or solitary, bisexual or less

often unisexual, actinomorphic or zygomorphic, cyclic,

usually with double perianth or sometimes the petals

much reduced or wanting. Sepals and petals free or

more or less connate. Stamens from numerous to few,

rarely solitary. Tapetum secretory, rarely amoeboid.

Microsporogenesis simultaneous or very rarely suc-

cessive. Pollen grains 2-celled or less often 3-celled,

mostly 3-colporate. Nectaries of various type, often of

staminodial origin, often forming an intrastaminal or

extrastaminal disc. Gynoecium apocarpous or more

often of united carpels; stylodia free or connate into a

style; ovary superior, or sometimes semi-inferior, often

with only one or two ovules per carpel or per locule,

but sometimes with several or many. Ovules usually

anatropous, apotropous or epitropous, bitegmic or

sometimes unitegmic or ategmic, crassinucellate or

sometimes tenuinucellate. Endosperm nuclear or cel-

lular (not produced in Podostemales). Fruits of various

types. Seeds with small to large, straight, or curved

embryo, with copious or scanty endosperm or

endosperm wanting. Frequently with alkaloids.

The subclass Rosidae demonstrates ambivalent

afﬁ nities to both the Hamamelididae and Dilleniidae.

Bibliography

Dickison WC. 1989. Comparisons of primitive Rosidae and

Hamamelidae. In: P Crane and S Blackmore, eds. Evolution,

systematics, and fossil history of Hamamelidae, vol. 1,

pp. 47–73. Clarendon Press, Oxford.

Endress PK and ML Matthews. 2006. First steps towards a ﬂ oral

characterization of the major rosid subclades. Plant Syst.

Evol. 260: 223–251.

Endress PE and S Stumpf. 1991. The diversity of stamen

structures in “Lower” Rosidae (Rosales, Fabales, Proteales,

Sapindales). Bot. J. Linn. Soc. 107: 217–293.

Fehrenbach S and W Barthlott. 1988. Mikromorphologie der

Epicuticular-Wachse der Rosales s.l. und deren system-

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Hallier H. 1903. Über Verwandtschaftsverhältnisse bei Engler’s

Rosalen, Parietalen, Myrtiﬂ oren, und in an-deren Ordnungen

der Dikotylen. Abh. Naturw. Ver. Hamburg 18: 1–98.

Hallier H. 1908. Über Juliania, eine Terebinthaceen-Gattung

mit Cupula, und die wahren Stammeltern der Kätzchenblütler:

Neue Beiträge zur Stammesgeschichte nebst einer Übersicht

über das natürliche System der Dicotyledonen. Dresden.

Heimsch CH. 1942. Comparative anatomy of the secondary

xylem in the “Gruinales” and “Terebinthales” of Wettstein

with reference to taxonomic grouping. Lilloa 8: 83–198.

Hijwegen T. 1981. Fungi as plant taxonomists: II. Afﬁ nities of

the Rosiﬂ orae. Acta Bot. Neerl. 30: 479–491.

Huber H. 1963. Die Verwandtschaftsverhältnisse der Rosiﬂ oren.

Mitt. Bot. Staatssamml. München 5: 1–48.

Hufford L. 1992. Rosidae and their relationships to other

nonmarginal dicotyledons: A phylogenetic analysis using

morphological and chemical data. Ann. Missouri Bot. Gard.

79: 218–248.

Mauritzon J. 1939. Contributions to the embryology of the orders

Rosales and Myrtales. Acta Univ. Lund. 2 (35): 1–121.

Superorder ROSANAE

Order 78. CUNONIALES

Trees and shrubs. Vessels with scalariform perforation

(sometimes with many bars), mixed scalariform and

simple perforations, or rarely only with simple

perforations; lateral pitting from scalariform to alternate.

Subclass VI. ROSIDAE

294 Subclass VI. ROSIDAE

Fibers with bordered or simple pits. Rays heterogeneous.

Axial parenchyma apotracheal or (Brunelliaceae) want-

ing. Sieve-element plastids of S- or Pc-type. Nodes

trilacunar or multilacunar, rarely (Bauera) unilacunar.

Leaves alternate, opposite or verticillate, simple or

compound, stipulate. Stomata paracytic, anomocytic,

anisocytic or encyclocytic. Flowers small or rarely

large, in axillary or terminal inﬂ orescences, rarely soli-

tary in the axils, mostly bisexual, actinomorphic, with

double perianth or less often apetalous (some

Cunoniaceae and Brunelliaceae). Stamens numerous to

few, free; ﬁ laments slender; anthers tetrasporangiate,

opening longitudinally or (Davidsonia) by apical pores

that elongate into longitudinal slits. Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains

2-celled, small, mostly 3-colporate, 2-colporate, or

2-colpate. Nectary disk mostly present. Gynoecium

apocarpous or mostly of more or less united carpels;

stylodia free, with capitate or (Brunelliaceae) decurrent

stigma. Ovary superior, semi-inferior or inferior, with

two to numerous ovules per carpel or per locule. Ovules

anatropous or less often campylotropous, apotropous,

or epitropous, bitegmic, crassinucellate. Female game-

tophyte of Polygonum-type. Endosperm nuclear. Fruits

mostly capsular. Seeds exotegmic, with small or large,

straight embryo; endosperm more or less copious or

(Davidsonia) wanting.

In many respects Cunoniales are relatively the most

archaic member of the subclass Rosidae, and they are

probably the nearest to the hamamelids. The similarities

between the cunonialean families, such as Cunoniaceae,

and the archaic members of the Hamamelididae are

discussed in detail by Dickison (1989).

Key to Families

1 Axial parenchyma present. Trees and erect shrubs.

Indumentum generally of simple hairs, or occasion-

ally stellate and peltate trichomes, or rarely stinging

hairs (Davidsonia). Vessels with scalariform

(Acsmithia, Gill beea, Pullea, Spiraeanthemum, and

Weinmannia p.p.) and mixed simple and scalariform

perforations. Vessel-ray pits coarse and scalariform

to opposite. Axial parenchyma mainly diffuse,

diffuse-in- aggregates, and scanty paratracheal.

Nodes mostly trilacunar with three traces, some-

times (Codia and Pulled) multilacunar or (Bauera)

unilacunar. Sieve-element plastids of S-type, or of

Pc-type with single polygonal protein crystal loid

(Eucryphia). Leaves opposite or sometimes

verticillate, rarely alternate (Davidsonia), imparip-

innately compound or trifoliolate, seldom unifoli-

olate; the leaﬂ ets vary from entire or serrate to

dentate. Stipules sometimes large and conspicuous,

usually united in pairs to form the characteristic

interpetiolar stipular condition, commonly with

small colleters ( multicellular, glandular secretory

structures). Flowers small, bisexual or less often

unisexual (plants dioecious or polygamous), actino-

morphic, mostly both sepals and petals. Sepals (3)-

6(-10), valvate or imbricate, in some genera enlarged

in fruit, in others persistent or caduceus. Petals as

many as sepals and alternate with them, or more

numerous than the sepals (Bauera). Stamens 8–10

or less often 4–5, sometimes numerous; ﬁ laments

usually long and thin; anthers introrse, more or less

dorsiﬁ xed, versatile (except Bauera), opening

longitudinally. Pollen grains mostly 3-colporate,

rarely (Bauera) syncolpate, smooth or nearly so.

A saucershaped, annular nectary disc usually

developed around the gynoecium. Gynoecium of

two or less often 3–5 carpels, syncarpous or some-

times (Acsmithia, Acrophyllum, Pancheria, and

Spiraeanthemum) apocarpous; carpellary margins

mostly inrolled; stylodia free or more or less con-

nate, usually with stigmas ranging from relatively

undifferentiated to capitate or subcapitate, but in

some species of Cunonia extending downward and

in Vesselowskya decurrent on the ventral surface of

the conspicuously elongate stylodium. Ovary supe-

rior or sometimes semi-inferior or inferior. Ovules

(1)2-many per locule or per carpel, apotropous or

sometimes (Acsmithia and Spiraeanthemum)

epitropous. Fruits dehiscent or indehiscent, mostly

capsular, sometimes follicular (Acsmithia and

Spiraeanthemum), drupaceous (Aistopetalum and

Schizomeria), or samara (Gillbeea), or pseudosa-

mara (Ceratopetalum). Seeds small to relatively

large, in dehiscent fruits usually with winges at

one or both ends, or hairy without wings; embryo

embedded in abundant starchy endosperm (except

Davidsonia). Contain much tannin, glycosides of

kaempferol and quercetin, ﬂ avonoids prodel-

phinidins and myricetin; Bauera rubioides con-

tain proanthocyanidins n = 16, 15 (Weinmannia,

Euchriphia), 12 (Pancheria).. . . . 1. cunoniaceae.

1 Axial parenchyma absent. Evergreen, tall trees up to

40 m tall, usually tomentose throughout; hairs

unicellular. Twigs angular, with rather large pits.

Superorder ROSANAE 295

Vessel elements elongate, thin-walled, some of

them with scalariform perforations that have up to

35 bars, others with simple and scalariform perfo-

rations. Fibers with small, simple, or obscurely

bordered pits, often septate. Nodes trilacunar or

pentalacunar. Leaves opposite or ternate, pinnately

compound, trifoliolate to unifoliolate or simple,

the leaﬂ ets opposite, pinnately veined, entire to

double-dentate; stipules small, caducous, often

more than two. Stomata anomocytic. Flowers

small, in axillary or terminal cymes, bisexual to

more or less unisexual (commonly dioecious or

gynodioecious), apetalous; male ﬂ owers with a

vestigial gynoecium, and female ﬂ owers with a ves-

tigial androecium. Sepals (4)5–6(-8), shortly con-

nate below, valvate, persistent in the fruits. Nectary

disc intrastaminal, adnate to the calyx, cupular,

8–10-lobed. Stamens 8-10(-14), in two cycles; ﬁ la-

ments slender, hairy, inserted in the notches of the

nectary disc; anthers dorsiﬁ xed, versatile, introrse,

opening longitudinally. Pollen grains 3-colporate.

Gynoecium apocarpous, of 2–3 carpels or more

often up to same number as sepals, more or less

adnate to the disc; each carpel gradually attenuates

into long, slender, elongate, curved, or hooked sty-

lodium with a linear, decurrent, sutural stigma that

extends along its entire length and is composed of

bands of papillae. Ovules two per carpel, collateral,

pendulous, epitropous. Fruits of 1–2-seeded fol-

licles, usually densely short reddish to yellowish

tomentose and with long, pointed trichomes; endo-

carp more or less ligniﬁ ed, separating from the

exocarp at maturity. Seeds with thick, hard, shiny

testa and corky, subarillate raphe, attached by a

funicle in dehisced follicle; embryo large, straight,

surrounded by copious, carnose, mealy, white

endosperm, n = 14. . . . . . . . . . 2. brunelliaceae.

1. CUNONIACEAE

R. Brown 1814 (including Baueraceae Lindley 1830,

Belangeraceae J. Agardh 1858, Callicomaceae

J. Agardh 1858, Davidsoniaceae Bange 1952,

Eucryphiaceae Endlicher 1841, Spiraeanthemaceae

Doweld 2001). 27/300. Almost exclusively conﬁ ned to

the Southern Hemisphere between 13° and 35°

S, mainly Australia, New Caledonia, and New Guinea;

a few (Weinmannia spp.) north to the Philippines and

southern Mexico; there are a few genera in tropical

America and South Africa. The largest genus

Weinmannia (190) distributed through Madagascar,

Mascarenes, Malesia, the Paciﬁ c, New Zealand, Chile,

Mexico, and the West Indies. Cunonia has a discon-

tinuous distribution is South Africa (1) and New

Caledonia (16). Bauera (3) and Davidsonia are

endemic to Australia, Eucryphia (5) Eastern Australia,

Tasmania, Chile, Argentina.

1.1 CUNONIOIDEAE

Erect trees and shrubs. Nodes trilacunar or sometimes

(Codia and Pullea) multilacunar. Leaves with inter-

petiolar stipules. Petals mostly present. Anthers

dorsiﬁ xed, versatile. Carpels mostly 2, rarely 5–3. –

Spraeanthemum, Acsmithia, Aistopetalum, Hooglandia,

Schizomeria, Ceratopetalum, Anodopetalum, Platy-

lophus, Gillbeea, Acrophyllum, Lamanonia, Pseudo-

weinmannia, Geissois, Caldcluvia, Opocunonia,

Ackama, Spiraeopsis, Pullea, Codia, Callicoma,

Vesselowskya, Pancheria, Cunonia, Weinmania.

1.2 BAUEROIDEAE

Erect shrubs. Nodes unilacunar with three traces.

Leaves trifoliate and estipulate, or simple with well-

developed leaﬂ ike stipules. Petals present. Anthers not

versatile. Carpels 2. – Bauera.

1.3 EUCRYPHIOIDEAE

Trees and shrubs with simple, unicellular hairs,

producing gum or mucilage and sticky terminal buds.

Sieve-element plastids with protein crystals. Nodes

trilacunar. Leaves simple or pinnately compound;

stipules interpetiolar, small, caducous, with large

colleters. Stomata paracytic. Petals 4(5), imbricate.

Athers versatile. Carpels 4–14(-18). Fruits leathery or

woody septicidal capsules. n = 15, 16. – Eucryphia.

1.4 DAVIDSONIOIDEAE

Small trees; hairs urticating. Leaves alternate, to 1 m

long. Stomata paracytic. Flowers in large, axillary or

supra-axillary panicles with spikelike branches,

bisexual, apetalous. Anthers dorsiﬁ xed, introrse.

Carpels 2, rarely 3; ovules 5–7, pendulous. Fruits large

2-pyrened drupes, red-velvety when young, pruinose

and glaucous at maturity, with ﬂ eshy mesocarp and

ﬂ attened, ﬁ mbriate-laciniate pyrenes. Seeds strongly

compressed, glabrous, broad-ovate to circular, without

endosperm. – Davidsonia.

296 Subclass VI. ROSIDAE

Dickison (1975a, 1980b, 1984) has shown, the gen-

era Ascmithia (17, Moluccas to western Paciﬁ c) and

Spiraeanthemum (6, western Paciﬁ c) are evidently the

most archaic members of the family (primitive wood

anatomy, paracytic stomata, and apocarpous gynoe-

cium.). They differ from the other members also in the

carpels, which are vascularized by ﬁ ve or four major

vascular bundles, and the carpel margins, which are

not inrolled (Dickison 1975b). These differences show

their rather isolated position.

2. BRUNELLIACEAE

Engler 1897. 1/c.60. America from Mexico, Costa

Rica, and West Indies to Venezuela and the Andes of

Colombia, Bolivia, and Peru.

Brunellia.

Close to the Cunoniaceae (Hallier 1903a, 1912;

Engler 1930; Cuatrecasas 1970; Dickison 1989; Thorne

2000, 2006). According to Dickison (1989: 50), strong

similarities exist between the two families in vegetative

morphology, similar ﬂ oral diagram, ovular type (partic-

ularly Acsmithia and Spiraeanthemum), and fruit anat-

omy. As Cuatrecasas (1970) and Dickison (1989) point

out, Brunellia is uniquely characterized by a linear,

decurrent, sutural stigma and a highly specialized carpel

maturation and mature fruit. The Brunelliaceae are a

highly heterobathmic family with a mixture of both ple-

siomorphic and apomorphic characters.

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species from Australia and South America. Biochem. Syst.

Ecol. 28: 111–118.

Order 79. ANISOPHYLLEALES

Large trees or shrubs of wet primary forests. Vessels

with simple perforations; lateral pitting alternate, with

coalescent apertures. Fibers with distinctly bordered

pits. Rays heterogeneous, very broad, multiseriate.

Axial parenchyma apotracheal and often banded,

varying to paratracheal and irregular. Sieve-element

plastids of S-type. Lysigenous secretory cavities are

present in the parenchymatous tissues of Poga. Nodes

unilacunar with one trace. Leaves alternate, simple,

entire, often 2-ranked, base asymmetrical, sometimes

pellucid-punctate, estipulate or (Polygonanthus) with

2–4 minute stipules, very base of the petiole. Stomata

mostly paracytic. Flowers small (except for the female

ﬂ owers of Polygonanthus), in axillary spikes, racemes,

or panicles of catkin-like spikes, rarely solitary,

bisexual (Combretocarpus and a few species of

Anisophyllea) or more often unisexual and monoecious

298 Subclass VI. ROSIDAE

or dioecious, sometimes polygamous, actinomorphic,

mostly 4- merous, but in Combretocarpus usually

3-merous and in other genera sometimes 5-merous.

Calyx and petals valvate. Petals commonly deeply

incised, generally with three, ﬁ ve, or seven lobes,

which in taxa of Anisophyllea and Poga may have an

enlarged, glandular tip; in Poga and Polygonanthus

petals entire, but sometimes ﬁ nely ﬁ mbriate, in

Combretocarpus they are lacking or reduced. Stamens

(6-)8 (-10), in 2 cycles, incurved in bud, free, with

narrow ﬁ laments; anthers tetrasporangiate or

(Polygonanthus) occasionally the pollen sacs divided

by tapetal septa, dorsiﬁ xed, ovoid, introrse, opening

longitudinally. Tapetum secretory. Microspo rogenesis

simultaneous. Pollen grains 2-celled, 3- colporate,

have a rather thin tectum with reticulate-punctate or

rarely (Anisophyllea disticha) striate surface, endoap-

ertures, when present, are circular and poorly deﬁ ned.

A lobed intra- and interstaminal, discontinuous

nectary disc surrounds the top of the ovary. Gynoecium

of (3)4(5) united carpels with free, subulate stylodia.

Ovary inferior, 3–4-locular, with two (Combretocarpus)

or one ovule per locule. Ovules pendulous from the

axile placenta, anatropous, bitegmic or (Anisophyllea

and Combretocarpus) unitegmic, clearly crassinucel-

late; no endothelium is formed. Female gametophyte

of Polygonum-type or (Combretocarpus) of Allium-

type. Endosperm nuclear. Fruits drupaceous

(Anisophyllea), or dry, woody, strongly winged,

crowned by much enlarged and persistent calyx lobes

(Polygonanthus), samara (Combretocarpus), or

3–4-seeded capsules (Poga). Seeds winged or wing-

less; coat testal (consists of the testal epidermis only,

as in Poga, of multilayered testa, as in Polygonanthus,

or of multilayered or ultimately thin single integu-

ment, as in Anisophyllea and Combretocarpus respec-

tively); embryo has a long hypocotyl and small to

rudimentary cotyledons or no cotyledons at all; no

endosperm. Aluminum accumulation is highly char-

acteristic. Clearly tanniniferous plants. Alkaloids are

unknown, n = 7, 8.

According to Dahlgren (1988: 1274), in gross mor-

phology Anisophylleaceae are “largely Saxifragalean-

Rosalean in nature,” and he agrees with Cronquist

(1981), who places Anisophylleaceae in Rosales sensu

latissimo. According to Dahlgren the ﬂ oral morpho-

logical data as well as embryological conditions in

Anisophylleaceae agree completely with those com-

mon in Rosaceae sensu stricto. “It is probable that

Anisophylleaceae comprise a rather isolated family

evolved from ancestors shared between those in

Rosales, Cunoniales, and Saxifragales,” concludes

Dahlgren (1988: 1275). On the other hand, according

to Tobe and Raven (1988: 1427), “embryological

features suggest strongly that Anisophylleaceae, even

though there are some points of similarity to Rosales

sensu stricto, do not belong in that order.” In their

opinion it is more appropriate to regard Anisophylleaceae

as constituting a distinct order. Matthews et al. (2001),

Schönenberger et al. (2001), Matthews and Endress

(2004) showed striking similarities in ﬂ oral structure

of Anisophylleaceae with the Cunoniaceae, which is

supported by greater similarity between in ﬂ owers of

Anisophyllea (Anisophylleaceae) and the genus

Ceratopetalum (Cunoniaceae) (Matthews et al. 2001).

According to molecular date the Anisophylleaceae

belong in the Cucurbitales, which is not acceptable.

1. ANISOPHYLLEACEAE

Ridley 1922. 4/34 (including Polygonanthaceae Croizat

1943). Tropical Africa, tropical Asia (India to Malesia),

South America; Combretocarpus (1) is endemic to

Borneo, Polygonanthus (2) to Brazil (Amazonia).

Anisophyllea, Poga, Polygonanthus, Combretocarpus.

Bibliography

Baehni C and P Dansereau. 1939. La position systématique du

genre Polygonanthus. Bull. Soc. Bot. Suisse 49: 415–416.

Behnke H-D. 1988. Sieve-element plastids and systematic rela-

tionships of Rhizophoraceae, Anisophylleaceae, and allied

groups. Ann. Missouri Bot. Gard. 75: 1387–1409.

Dahlgren RMT. 1988. Rhizophoraceae and Anisophylleaceae:

summary statement, relationships. Ann. Missouri Bot. Gard.

75: 1259–1277.

Juncosa AM and RB Tomlinson. 1988a. A historical and taxo-

nomic synopsis of Rhizophoraceae and Anisophylleaceae.

Ann. Missouri Bot. Gard. 75: 1278–1295.

Juncosa AM and RB Tomlinson. 1988b. Systematic comparison

and some biological characteristics of Rhizophoraceae and

Anisophylleaceae. Ann. Missouri Bot. Gard. 75: 1296–1318.

Keating RC and V Randrianosola. 1988. The contribution of leaf

architecture and wood anatomy to classiﬁ cation of the

Rhizophoraceae and Anisophylleaceae. Ann. Missouri Bot.

Gard. 75: 1343–1368.

Matthews ML and PK Endress. 2004. Comparative ﬂ oral struc-

ture and systematics in Cucurbitales (Corynocarpaceae,

Coriariaceae, Tetramelaceae, Datiscaceae, Begoniaceae,

Superorder ROSANAE 299

Cucurbitaceae, Anisophylleaceae). Bot. J. Linn. Soc. 145:

129–185.

Matthews ML, PK Endress, J Schönenberger, and EM Friis.

2001. A comparison of ﬂ oral structures of Anisophylleaceae

and Cunnoniaceae and the problem of their systematic posi-

tion. Ann. Bot. 88: 439–455.

Plisko MA. 1996. Anisophylleaceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol. 5, pp. 151–152. Nauka,

St. Petersburg (in Russian).

Schwarzbach AE and RE Ricklefs. 2000. Systematic afﬁ nities

of Rhizophoraceae and Anisophylleaceae, and intergeneric

relationships within Rhizophoraceae, based on chloroplast

DNA, nuclear ribosomal DNA, and morphology. Am. J. Bot.

87: 547–564.

Tobe H. and R H. Raven. 1987. Systematic embryology of the

Anisophylleaceae. Ann. Missouri Bot. Gard. 74: 1–26.

Tobe H and RH Raven. 1988a. Floral morphology and evolution

in Anisophylleaceae. Bot. J. Linn. Soc. 98: 1–25.

Tobe H and RH Raven. 1988b. Additional notes on the embry-

ology of Polygonanthus (Anisophylleaceae) and relation-

ships of the family. Ann. Missouri Bot. Gard. 75:

1425–1428.

Order 80. CEPHALOTALES

Small, insectivorous, perennial bog-herbs with a short

underground rhizome. Vascular system composed of a

ring of xylem and phloem interrupted by broad, foliar

rays. Vessels with scalariform perforations. Sieve-

element plastids of Ss-type. Leaves alternate, all basal,

of two kinds, the lower (outer) ones of the rosette

modiﬁ ed into ground level pitchers, the upper ones ﬂ at,

entire, estipulate; the pitchers probably represent an

ascidiate petiole dilated at the top into a lid (the lid is an

outgrowth from the upper surface of the petiole below

the pitcher proper while the pitcher has been produced

by a ventral-dorsal invagination of the upper, more dis-

tal region according to Lloyd 1942); the lower surface of

the lid and the distal inner surface of the pitcher slippery,

coated with overlapping, downwardly directed projec-

tions from the epidermal cells; multicellular, embedded

glands present on the pitcher surfaces as well as on the

petiole and lower surface of the vegetative leaves; ﬂ ask-

shaped embedded glands also present in the interior of

the pitcher and especially large on the brightly colored,

cushionlike projections from the surface. The mouth of

the pitcher is surrounded by a corrugated rim, each cor-

rugation forming a clawlike tooth extending inward and

downward. Stomata paracytic. Flowers small, borne on

a leaﬂ ess scape arising from the center of the rosette and

bearing racemously arranged dichasia, bracteate, bisex-

ual, actinomorphic, 6-merous, apetalous. Calyx colored,

synsepalous, 6-lobed; lobes valvate, hooded. Stamens

12, in two alternating cycles, inserted at the top of the

calyx tube on the inner margin of a broad, thick, green,

papillate, glandular nectary disc; anthers small, strongly

introrse and cruciform, dorsiﬁ xed, versatile, tetraspo-

rangiate, opening longitudinally; connective large,

globose, swollen at the top and glandular. Pollen

grains 2-celled, 3-colpate, smooth. Tapetum secretory.

Gynoecium of 6 free carpels each with more or less

circinately recurved, subulate stylodium and simple

stigma, with one (rarely 2) ovule per carpel; ovary supe-

rior, 1-locular. Ovules basal, ascending, anatropous,

with dorsal raphe, bitegmic, crassinucellate, with an

endothelium. Fruits multifollicles, densely hairy. Seeds

with small, straight embryo surrounded by copious,

ﬂ eshy endosperm; testa and tegmen very thin. Present

myricetin and quercetin, ellagic and gallic acids, and

tannin cells; n = 10.

Evidently related to the Cunoniales, which is

supported by the 18S rDNA and rbcL sequences data

(Sotlis and Soltis 1997; Soltis et al. 2006) and even

more to Brunelliaceae, they have an identical ﬂ oral

diagram (Conran 2004).

1. CEPHALOTACEAE

Dumortier 1829. 1/1. Southwestern Australia.

Cephalotus.

Bibliography

Arber A. 1941. On the morphology of the pitcher-leaves in

Heliamphora, Sarracenia, Cephalotus, and Nepenthes. Ann.

Bot. 5: 563–578.

Conran JG. 2004. Cephalotaceae. In: K Kubitzki, ed. The fami-

lies and genera of vascular plants, vol. 6, pp. 65–68. Springer,

Berlin/Heidelberg/New York.

Conran JG and MD Denton. 1996. Germination in the Western

Australian Pitcher Plant Cephalotus follicularis and its

unusual early seedling development. W.A. Nat. 21: 37–42.

Dickson A. 1878. The structure of the pitcher of Cepha-lotus

follicularis. J. Bot. (London) 16: 1–5.

Dickson A. 1882. On the morphology of the pitcher of

Cephalotus follicularis. Trans. Proc. Bot. Soc. Edinb. 14:

172–181.

Hamelton AC. 1904. Notes on the West Australian pitcher plants

(Cephalotus follicularis La Bill.). Proc. Linn. Soc. N.S.W.

29: 36–53.

300 Subclass VI. ROSIDAE

Nicholls KW, BA Bohm, and R Ornduff. 1985. Flavonoids and

afﬁ nities of Cephalotaceae. Biochem. Syst. Ecol. 13:

261–263.

Peng C-I and P Goldblatt. 1983. Conﬁ rmation of the chromo-

some number in Cephalotaceae and Roridulaceae. Ann.

Missouri Bot. Gard. 70: 197–198.

Order 81. SAXIFRAGALES

Trees and shrubs, or perennial and annual herbs.

Vessels mostly with scalariform or more often simple

perforations. Sieve element plastids of S- or P-types.

Nodes trilacunar, multilacunar, or unilacunar. Leaves

alternate, opposite or verticillate, simple or compound,

stipulate or estipulate. Stomata of various types.

Flowers in various kinds of inﬂ orescences or solitary,

mostly bisexual, actinomorphic, actinomorphic or less

often zygomorphic, usually with double perianth,

sometimes apetalous. Perianth forming a more or less

well-developed ﬂ oral tube, free or more or less adnate

to the base of the ovary. Sepals mostly ﬁ ve, less often

less (3, or 4) or more (up to 10), free or more or less

connate, imbricate or valvate. Stamens few to

numerous, mostly as many or twice as many as the

sepals; anthers basiﬁ xed or less often slightly dorsi-

ﬁ xed, introrse or latrorse, tetrasporangiate, opening

longitudinally. Tapetum secretory. Microsporogenesis

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

3-colporate to pantoporate. Nectary disc mostly present.

Gynoecium of free or more or less united carpels, stylo-

dia free, stigma decurrent to capitate; ovary superior,

semi-inferior or inferior, with 1–2 to numerous ovules

per carpel or per locule. Ovules usually anatropous,

bitegmic or rarely unitegmic (Tetracarpaeaceae,

Darmera, section Micranthes of Saxifraga, Choristylis),

crassinucellate. Female gametophyte of Polygonum- or

Allium-type, or rarely (Gunneraceae) Peperomia-type.

Endosperm cellular or less often helobial or nuclear.

Fruits multifollicles (Penthoraceae, majority of

Crassulaceae), capsular, baccate (Grossulariaceae),

drupaceous (Gunneraceae), or nutlike (Aphanopetalaceae).

Seeds small or minute, exotestal or rarely (Astilbe and

Rodgersia in Saxifragaceae) exotestal-endotegmic;

embryo small or large, mostly straight; endosperm

usually copious. Present ﬂ avones, ﬂ avonoids, proantho-

cyanidins, alkaloids.

The basal member of the Rosidae.

Key to Families

1 Flowers usually 4-merous.

2 Leaves mostly alternate.

3 Ovules bitegmic.

4 Embryo straight, cylindrical. Perennial or

annual aquatic or terrestrial herbs, subshrubs

or shrublets, or seldom (Haloragodendron)

shrubs or small trees up to 3 m tall, glabrous

or scabrous with simple uniseriate hairs;

often calcium oxalate crystals in hair-like

cortical cells. Vessels with simple perfora-

tion. Rays heterogeneous to homogeneous.

Leaves alternate, opposite or verticillate,

simple, entire to more or less deeply

dissected, pinnatiﬁ d or multiﬁ d, with minute

vestigial stipules. Stomata usually anomo-

cytic. Flowers in dichasial, often com-

pounded inﬂ orescences, sometimes reduced

to solitary axillary ﬂ owers, usually small,

bisexual or unisexual (monoecious), actino-

morphic, basically 4-merous, but often

reduced to 3- merous or sometimes 2-merous

condition, 2-bracteolate. Sepals valvate, per-

sistent, absent in the female ﬂ owers of

Myriophyllum. Petals imbricate, keeled,

hooded, or navicular in the apex and more or

less unguiculate in the base, deciduous with

stamens, absent in Proserpinaca and female

ﬂ owers of Myriophyllum and Laurembergia.

Stamens typically eight in two cycles, but

sometimes the antesepalous or antepetalous

cycle is absent or rudimentary or (antesepal-

ous stamens in some species Gonocarpus)

transformed into staminodia; ﬁ laments usu-

ally very short, slender; anthers large, oblong

( reniform in Proserpinaca), basiﬁ xed, with

shortly apiculate connective, introrse, the

antesepalous anthers sometimes more or less

longer than antepetalous ones. Pollen grains

3-celled, pertectate, 4–5(6)-colpate or

4–5-porate. Gynoecium usually of four

united carpels, but reduction to three or two

occurs in some genera and species; stylodia

free, clavate, more or less bulbously based,

convergent at tips until anthers shed, with

more or less capitate, ﬁ mbriate stigmas;

ovary inferior, (1–3)4-locular, but sometimes

septa are weakly developed or completely

Superorder ROSANAE 301

lacking (Laurembergia and most species of

Glischrocaryon), with 1–2 pendulous ovules

per locule (if two ovules, then one aborts at

an early stage). Ovules anatropous or

hemitropous, with hypostase and weakly

developed funicular obturator. Haustorial

suspensor present. Endosperm cellular

(Haloragis and some species of

Myriophyllum) or nuclear (Laurembergia

and some species of Myriophyllum). Fruits

small, nutlike or drupelike, in Myriophyllum

splitting septicidally into (2–3)4 mericarps,

but in all other genera indehiscent, 1–4

seeded, variously ornamented with wings,

ribs and tubercles. Seeds with straight, cylin-

drical embryo and usually with more or less

copious, starchy endosperm; exotesta (and

hypodermal payer) persistent, thin-walled.

Present ﬂ avones, tannins, myriophyllin and

ellagic acid; n = 6 (Gonocarpus), 7, 9, 21,

29, mostly 7. . . . . . . . . . 5. haloragaceae.

4 Embryo curved to bent (obcordate). Perennial

herbs with very short upright stems approach-

ing acaulescence, and short or prostrate and

creeping, usually more or less ﬂ eshy rhi-

zomes, and adventitious roots, harboring

symbiotic Nostoc and Chlorococcus colo-

nies in superﬁ cial tissues of stem and petiole

entered through hydathodes or mucilage

glands. The stem has an endodermis. Hairs

unicellular. Vascular system of the stem and

rhizome consists of anastomosing bundles,

each with its own endodermis. Scalariform

perforation plates occur in most vessel ele-

ments of stolons and roots; vessels with

simple perforations are more frequent in the

stems of large leafed-species (Wilkinson

2000). Fibers thick-walled, with simple

pits. Sieve-element plastids of Pcs-type.

Nodes multilacunar. Leaves alternate, all

radical, extremely large (sometimes more

than 3 m across) to small, long- petiolate,

orbicular, reniform or ovate, sometimes

peltate, entire, toothed, or variously lobed;

the margins of larger-leafed species are

crenate-denate with crenations and lobes

having protruding hydathodes, up to 8 mm;

pinnately or subpalmately veined, with

large median axillary scales (squamulae

intravaginales), which sometimes have been

interpreted as stipular. Stomata anomocytic.

Flowers small, in mostly very large, terminal

or upper axillary, often much branched, rac-

emose panicles, usually not bracteolate, the

basal ones often female, upper ones male

and middle ones bisexual, or seldom all the

ﬂ owers bisexual or the ﬂ owers all unisexual

and dioecious, monoecious or gynomonoe-

cious. Sepals two, rarely three, valvate, very

small, often almost absent. Petals two,

miter-shaped, or more often absent. Stamens

one or two, rarely three, with short ﬁ la-

ments; anthers 2-locular, opening longitu-

dinally. Pollen grains 2-celled, with 3(-5)

long, deep colpi (with bulging mesocolpia).

Gynoecium of two united carpels, with free

subulate, long papillate stylodia; ovary infe-

rior, 1-locular, rarely 2-locular (Gunnera

chilensis), with 1(2) pendulous ovules.

Ovules large, hemianatropous, with broad

nucellus; micropyle endostomal, formed by

inner integument. Female gametophyte of

Peperomia-type (tetracyclic, 16-nucleate).

Fruits one-seeded drupes. Seeds with a very

small, cordate embryo and copious, oily

endosperm. Producing caffeic and ellagic

acids, pelargonidin, two ﬂ avonoids: querce-

tin 3–0- galactoside, and quercetin 3–0-glu-

coside and high concentration of an

unidentiﬁ ed ellagitannin, n = 12, 17, 27,

more often 17. . . . . . . . . .6. gunneraceae.

3 Ovules unitegmic. Low, evergreen, glabrous

shrubs. Imperforate tracheary elements are

exclusively very short tracheids. Vessels sca-

lariform, with 5–18 bars per plate; lateral pit-

ting scalariform to opposite. Rays heterogeneous.

Axial parenchyma apotracheal, sparse and

either diffuse or diffuse-in-aggregates. Nodes

unilacunar and one-trace. Leaves small, simple,

serrate, with rounded to acute teeth, pinnately

veined with conspicuous secondary veins ter-

minating near the leaf margin. Stomata anomo-

cytic. Flowers in erect bracteate racemes, small,

bisexual, actinomorphic. Sepals four, small,

essentially free, imbricate, persistent. Petals

four, free, spreading, spatulate, slightly

302 Subclass VI. ROSIDAE

imbricate, caducous. Stamens four or eight,

borne in a single cycle and positioned in both

sepal- and petal-planes although one or more

members of the whorl may be missing; ﬁ la-

ments free, ﬁ liform; anthers elliptic-oblong,

basiﬁ xed, latrorse. Pollen grains very small,

3-colporate, rugulate. Carpels 4(5), fusiform,

supplied by three veins that all reach the

stigma, prominently stipitate, erect, free for

most of their length, but sometimes adjacent

ones are united to the middle of their ventral

surfaces; usually one or more carpels remain

unfused; stigma subsessile, small, lobed.

Ovules numerous, borne on branched submar-

ginal placentas. Fruits multifollicles; follicles

erect, stipitate, coriaceous, many-seeded.

Seeds very small, obovoid- subulate, testa

membranous, slightly prolonged at each end,

with narrow wings extended along their entire

length and parallel ridges on the surface;

embryo minute, at the base of ﬂ eshy copious

endosperm. . . . . . . . . . .1. tetracarpaeaceae.

2 Leaves opposite, simple, ovate-lanceolate or

elliptical, serrate to mostly entire; stipules want-

ing but with minute colleters at each side of the

nodes. Stomata anomocytic. Scrambling or viny,

thin-stemmed, glabrous shrubs, with endodermis

in young stems. Vessels with scalariform perfora-

tions; rays homocellular, uniseriate and hetero-

cellular, multiseriate. Axial parenchyma scarce,

being apotracheal diffuse with occasional para-

tracheal scanty; cells of both ray parenchyma and

axial parenchyma contain abundant starch grains

(Dickison et al. 1994). Nodes unilacunar with one

trace. Flowers solitary or in axillary panicles,

bisexual, 4-merous. Sepals free, imbricate, greatly

enlarged after ﬂ owering and persistent. Petals

very small or absent. Stamens eight, perigynous;

anthers long, almost basiﬁ xed, not versatile, with

connective protrusion, opening longitudinally.

Pollen grains 3-colporate with regulate-stellate

sculpture. Gynoecium of four carpels; style

4-lobed apically, with four canals, stigmas ter-

minal, highly papillate; ovary semi-inferior,

4-locular; ovules one per locule, suspended on

axile placenta, with long, thick funiculus. Fruits

nut-like, involucrate by the enlarged calyx-

lobes. Seed rugose; embryo curved, endosperm

ﬂ eshy. . . . . . . . . . . . . . . 2. aphanopetalaceae.

1 Flowers mostly 5-merous.

5 Gynoecium apocarpous or less often

semiapocarpous (Penthoraceae and some

Crassulaceae).

6 Vessels with scalariform perforations. Vessels

have many to numerous bars (9–57, usually

18–34 per plate); lateral pits bordered.

Imperforate tracheary elements ﬁ ber-tracheids

and vascular tracheids. Axial parenchyma absent.

Nodes unilacunar. Perennial rhizomatous, rather

ﬂ eshy herbs; young stem with pseudosiphonos-

tele. Leaves alternate, simple, serrate, pinnately

veined, nonsucculent. Stomata anomocytic.

Flowers in terminal cymose inﬂ orescences,

bisexual, 6–7(8)-merous. Sepals basally connate,

valvate, persistent. Petals inconspicuous and

devoid of venation or more often wanting.

Stamens twice as many as sepals, in two cycles;

ﬁ laments ﬁ liform; anthers basiﬁ xed, latrorse.

Pollen grains 3-colporate. Gynoecium of 5-6(7-

8) carpels united halfway, recurved, each with

short stylodium and small capitate stigma;

ovules numerous in each carpel, on a thick,

axile placenta. Fruits syncarpous multifollicles;

follicles circumscissile above their union. Seeds

numerous, scobiform, papillate, with an oper-

culum formed by an inner integument in the

micropylar part; seed coat consists of one layer

of exotesta. Present ﬂ avonoids and digluco-

sides, n = 8, 9. . . . . . . . . . . 3. penthoraceae.

6 Vessels with simple perforations. Lateral pits

simple. Imperforate tracheary elements absent,

or libriform ﬁ bers. Nodes unilacunar, trilacu-

nar or multilacunar. Mostly perennial herbs

with ﬂ eshy leaves and stem, sometimes suc-

culent subshrubs and small shrubs, rarely

arborescent. Leaves alternate, opposite, or

sometimes verticillate, simple and usually

entire, often with hydathodes. Stomata aniso-

cytic or diacytic. Flowers in various kinds of

cymose inﬂ orescences or sometimes solitary,

bisexual or rarely (Rhodiola p.p.) unisexual,

usually actinomorphic, mainly 5-merous or

less, free or nearly so, less often more or less

connate, persistent. Sepals usually green, free

or more or less connate, much shorter than the

corolla. Petals usually free or connate only at

the base, rarely corolla sympetalous. Stamens

mostly twice as many as petals, in two