Takhtajan Armen. Flowering Plants

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232 Subclass V. DILLENIIDAE

opposite petals present in Tremandra. Stamens numer-

ous or very 12–8, free, originating centrifugally, often

weakly grouped into ﬁ ve antesepalous clusters, inserted

on a lobed or annular disc (large and 5-lobed in

Sericolea) or enlarged receptacle that is sometimes

more or less elongated to form an androgynophore;

anthers introrse, except Sloanea, which has lartorse,

extrorse and introrse anthers in the same ﬂ ower

(Matthews and Endress 2002), basiﬁ xed, tetrasporan-

giate, generally relatively large and linear, often with a

conspicuously prolonged connective, opening only

apically, by two short slits or pores in Aristotelia,

Sloanea and Vallea, or by a single pore in Elaeocarpus,

Crinodendron, and Tremandreae (Matthews and

Endress 2002). Tapetum secretory. Microsporogenesis

simultaneous. Pollen grains 2-celled, small, globose,

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

stylodia united into a simple or shortly lobed style.

Ovary 2–5-loculed (in Peripentadenia two of three

loculi becoming abortive), with 2-many ovules per

locule, rarely (Platytheca) with solitary ovule; placen-

tation secondarily axile. Ovules anatropous, bitegmic,

crassinucellate, with zig-zag micropyle and endothe-

lium; unicellular hairs on ovules are present in

Elaeocarpus and Platytheca. Endosperm nuclear.

Fruits loculicidal, sometimes also septicidal capsules,

spiny or nor, or less often drupaceous or baccate. Seeds

massive to very small, sometimes (Peripentadenia,

Echinocarpus, Sloanes spp) with an aril arising from

the raphe and chalazal end of the seed or from the cha-

laza only, the appendage differentiates as an elaiosome

in Tetratheca and Tremandra (Boesewinkel 1999);

exotegmen ﬁ brous; embryo straight or strongly curved;

endosperm more or less copious, oily or proteinaceous.

Often producing pyrrolizidine and tropane alkaloids,

ellagic acid, n = 12, 14, 15.

The Elaeocarpaceae are usually included in the

Malvales near the Tiliaceae. However, from the

Tiliaceae and related families they differ markedly in

the absence of stellate and peltate trichomes, the

absence of mucilage cavities or canals, in usually

septate ﬁ bers with bordered pits, usually opposite lat-

eral pitting of the vessels, paratracheal axial paren-

chyma, unstratiﬁ ed phloem, the absence of nectar

hairs at the calyx base, the presence of a nectariferous

disc at the base of the stamens and the presence of

endothelium, the absence of fatty acids with a cyclo-

propenyl ring, and important differences in seed anat-

omy (according to Corner 1976, the ﬁ brous exotegmen

denies the immediate afﬁ nity with the Tiliaceae,

which are characterized by the palisade exotegmen).

On the other hand, the Elaeocarpaceae share many

features with the Flacourtiaceae, including gross

morphology, pollen morphology, and wood anatomy.

Fibrous exotegmen is typical both of the Flacourtiaceae

and the Elaeocarpaceae (Corner 1976). Both these

families have so many external similarities, that

“members of the Flacourtiaceae are often confused

with Sloanea” (Cronquist 1981: 349). Elaeocarpaceae

are especially allied to the Prockieae (particularly to

Prockia and Hasseltia according to Schumann

[1895]). According to recent molecular and morpho-

logical studies Tremandraceae are nested within

Elaeocarpaceae (Savolainen et al. 2000; Doweld

2001; APG II 2003). In my opinion, Platytheca,

Tetratheca, Tremandra constitute a separate tribe in

the Elaeocarpaceae.

1. ELAEOCARPACEAE

A.L. de Jussieu 1824 (including Aristoteliaceae

Dumortier 1829, Tetrathecaceae R. Brown 1814,

Tremandraceae R. Brown ex A.P. de Candolle 1824).

12/625. Madagascar, Mauritius, Socotra, eastern

Himalayas, India, eastern Asia and Southeast Asia,

Moluccas, New Guinea, New Caledonia, New

Hebrides, Fiji, Samoa, Tonga, and some other Paciﬁ c

Islands, Australia, Tasmania, New Zealand, America

from Mexico to Chile, West Indies.

elaeocarpeae: Peripentadenia, Dubouzetia, Ela-

eocarpus, Aceratium, Sericolea, Crinodendron; aristo-

telieae: Aristotelia; sloaneae: Vallea, Sloanea;

tremandreae: Platytheca, Tetratheca, Tremandra.

Bibliography

Bawa KS and CJ Webb. 1983. Floral variation and sexual dif-

ferentiation in Muntingia calabura (Elaeocarpaceae), a spe-

cies with hermaphrodite ﬂ owers. Evolution 37:

1271–1282.

Biddle JA and DC Christophel. 1978. Intergynoecial develop-

ment in Tremandraceae. Phytomorphology 28: 411–418.

Boesewinkel FD. 1999. Ovules and seeds of Tremandraceae.

Aust. J. Bot. 47: 769–781.

Brizicky GK. 1965. The genera of Tiliaceae and Elaeocarpaceae

in the southeastern United States. J. Arnold Arbor. 46:

286–307.

Superorder VIOLANAE 233

Carlquist 1977. Wood anatomy of the Tremandraceae:

phylogenetic and ecological implications. Am. J. Bot. 64:

704–713.

Coode MJE. 1978. A conspectus of Elaeocarpaceae in Papuasia.

Brunonia 1: 131–302.

Coode MJE 2004. Elaeocarpaceae. In: K Kubitzki, ed. The fami-

lies and genera of vascular plants, vol. 6, pp. 135–144.

Springer, Berlin/Heidelberg/New York.

Dehay Ch. 1961. Remarques sur 1∋ανατομιε comparee des

Elaeocarpacees. Bull. Soc. Bot. France 14(4): 89–96.

Gasson P. 1996. Wood anatomy of the Elaeocarpaceae. In: LA

Donaldson, AP Singh, BG Butterﬁ eld, and LJ Whitehous,

eds. Resent advances in wood anatomy, pp. 47–71. New

Zealand Forest Institute, Rotorua.

Lodkina MM. 1992. Elaeocarpaceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol. 4, pp. 249–254. Nauka, St.

Petersburg (in Russian).

Matthews ML and PK Endress. 2002. Comparative ﬂ oral

structure and systematics in Oxalidales (Oxalidaceae,

Connaraceae, Brunelliaceae, Cephalotaceae, Cunoniaceae,

Elaeocarpaceae, Tremandraceae). Bot. J. Linn. Soc. 140:

321–381.

Mauritzon J. 1934. Zur Embryologie der Elaeocarpaceae. Ark.

Bot. 26A(10): 1–8.

Plisko MA. 2000. Malpighiaceae, Trigoniaceae, Vochysiaceae,

Krameriaceae, Tremandraceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol. 6, pp. 49–11. Nauka, St.

Petersburg (in Russian).

Sandhya Rani R. 1995. Floral anatomy of Tremandraceae.

J. Indian Bot. Soc. 74(1–4): 69–74.

Schumann KM. 1895. Elaeocarpaceae. In: A Engler and K

Pranti, eds. Die natürlichen Pﬂ anzenfamilien, 3(6): 1–8.

Smith CE, Jr. 1954. The New World species of Sloanea

(Elaeocarpaceae). Contr. Gray Herb. 175: 1–114.

Tang Y and Z-Y Wu. 1990. Study on the pollen morphology of

Chinese Elaeocarpaceae. Acta Bot. Yunn. 12: 397–403 (in

Chinese with English summary).

Thompson J. 1976. A revision of the genus Tetratheca

(Tremandraceae). Telopea 1: 139–215.

Venkata Rao C. 1953. Floral anatomy and embryology of two

species of Elaeocarpus. J. Indian Bot. Soc. 32: 21–33.

Weibel R. 1968. Morphologic de 1∋εμβρψον et de la graine des

Elaeocarpus. Candollea 23: 101–108.

West WC, JE Gunkel, and MA Johnson. 1970. Morphology of

the shoot apex in Elaeocarpaceae. Phytomorphology 20:

58–67.

Order 68. CUCURBITALES

Perennial or annual herbs, rarely soft-wooded lianas,

or less often shrubs and trees. Vessels with scalariform

or simple perforations. Rays heterogeneous. Axial

parenchyma paratracheal. Sieve-element plastids of

S-type. Nodes trilacunar. Leaves alternate, usually pal-

mately lobed, sometimes palmately compound with

three or more leaﬂ ets, stipulate or less often estipulate.

Stomata mostly anomocytic. Flowers in axillary or less

often terminal inﬂ orescences, sometimes reduced to a

solitary, nearly always unisexual (monoecious or

dioecious), rarely bisexual usually actinomorphic.

Calyx (3-)5(6–8)-lobed, imbricate or open. Corolla

more or less deeply (3-)5(6–8)-lobed, valvate or indu-

plicate-valvate. Stamens three, less often 5(1–5), or

numerous, free or sometimes more or less connate;

anthers tetrasporangiate or disporangiate, extrorse,

opening longitudinally or rarely by pores. Tapetum

secretory. Microsporogenesis simultaneous. Pollen

grains 2-celled, of various types, the most primitive

ones are 3-colpate or 3-colporate. Nectaries in both

the male and female ﬂ owers commonly present.

Gynoecium of (3-)4–5(-8) united carpels or rarely

secondary monomerous; stylodia usually more or less

united into a style. Ovary inferior or very rarely semi-

inferior, basically 1-locular; ovules numerous or rarely

(Sicyoeae) solitary, anatropous, bitegmic, crassinucellate.

Female gametophyte of Polygonum-type or of Allium-

type. Endosperm nuclear. Fruits berries or pepos, less

often dry or ﬂ eshy capsules. Seeds usually numerous,

rarely solitary, mostly with thick, woody exotesta and

papery endotesta, commonly compressed and some-

times even winged; embryo straight, with two large ﬂ at

cotyledons; endosperm wanting or vestigial. Bitter

substances known as cucurbitacins are widespread.

Closely related to the Violales, especially to the

Passiﬂ oraceae.

Key to Families

1 Fruits usually baccate, rarely (Zanona) capsules.

Perennial herbs with swollen, tuberous rootstock

formed by swelling of hypocotyl, or less often

annuals, mostly climbing or prostrate, rarely soft-

wooded lianas, very rarely (monotypic Socotran

genus Dendrosicyos) small trees with soft and juicy

stem. Herbaceous species usually with spirally

coiled (often branched) tendrils at the side of each

leaf base, which represent modiﬁ ed shoots with

lower part corres ponding to the stem and upper

twining portion cor responding to highly modiﬁ ed

foliage; the tendrils sometimes modiﬁ ed into spines

or wanting. Glandular hairs are very characteristic,

but simple unicellular or uniseriate hairs as well as

wartlike or spiny trichomes also occur. Calcareous

cystoliths and similar bodies occur, usually at the

bases of the hairs or in nearby cells. Vascular bun-

dles of the stem are predominantly bicollateral and

234 Subclass V. DILLENIIDAE

frequently arranged in two rings. Anomalous struc-

ture is fairly frequent in thick stems and roots.

Leaves alternate, usually palmately lobed, some-

times palmately compound with three or more leaf-

lets, estipulate. Petioles usually with a crescent or

ring of unequal vascular bundles, the larger ones

bicollateral. Flowers in axillary inﬂ orescences that

are sometimes reduced to a solitary, nearly always

uni sexual (monoecious or dioecious), very rarely

bisexual (sometimes in the eastern asiatic genera

Actinostemma and Schizopepon), usually actino-

morphic, mostly 5-merous. The perianth together

with the bases of stamens forms a ﬂ oral tube (hypan-

thium) that is shortly or strongly prolonged beyond

the ovary. Calyx (3-)5(6)-lobed, imbricate or open.

Corolla more or less deeply (3-)5(6)-lobed, valvate

or induplicate-valvate. Stamens three or less often

5(1–5), inserted on the lower part of the hypanthium

and alternating with the corolla lobes; ﬁ laments

more or less free, as in Fevillea, or more often vari-

ously united, as also the anthers; anthers straight or

arcuate to triplicate or contorted, when three in

number usually two tetrasporangiate and disporan-

giate, when ﬁ ve in number all disporangiate,

extrorse, opening longitudinally. Pollen grains

2-celled. Nectaries in both the male and female

ﬂ owers commonly present. Gynoecium of 3 (4–5)

united carpels or rarely secondary monomerous

(Cyclanthera and related genera); stylodia usu-

ally more or less united into a style; stigmas as many

as carpels, usually forked, commissural. Ovary

inferior or very rarely semi- inferior, basically 1-loc-

ular, mostly almost completely ﬁ lled with much

enlarged placentas, which are sometimes joined in

the center making the ovary plurilocular, usually

with numerous ovules or rarely (Sicyoeae) ovule

solitary. Female gametophyte of Polygonum-type.

Chalazal end of the female gametophyte usually

developing into a tubular aggressive haustorium of

various lengths. Fruits berries or pepos (ﬁ rm-

walled berries with hardened pericarp), less often

dry or ﬂ eshy capsules, or leathery and indehiscent.

Seeds usually numerous, rarely solitary, mostly

with thick, woody exotesta and papery endotesta,

commonly compressed and sometimes even

winged; in some genera with a false scarlet aril

(placental tissue – Corner 1976); embryo straight,

with two large ﬂ at cotyledons; endosperm wanting

or vestigial, n = 7(8)-14. . . . . . 1. cucurbitaceae.

1 Fruits capsules, dehiscing apically, rarely berry-like.

2 Leaves simple, entire, dentate or deeply lobed, or

pinnate, often asymmetrical, estipulate. Stomata

anomocytic. Vessels with simple perforations; lat-

eral pitting alternate. Fibers with minute simple

pits. Rays heterogeneous. Axial parenchyma para-

tracheal, scanty. Flowers more or less actinomor-

phic, dioecious or in Datisca glomerata functionally

androdioecious. Sepals 3–8, mostly unequal. Petals

only in male ﬂ owers of Octomeles, 6–8, small,

free, valvate, inserted on the calyx, in the other

genera wanting in both the male and female ﬂ ow-

ers. Staminodia generally wanting, but pistillodia

sometimes present. Stamens of the same number

as the sepals and opposite them, or more numerous

(up to about 25). Anthers 2-locular, opening longi-

tudinally. Gynoecium of 3–8 carpels; ovary infe-

rior, 1-locular. Operculum without collar cells.

Fruits capsules. Perennial herbs or very large trees.

3 Perennial herbs engaging in a nitrogen-ﬁ xing

symbiosis with the actinomycetes Frankia.

Leaves pinnate or pinnatiﬁ d, with long nar-

row caudate-acuminate pinnae. Inﬂ orescences

greatly shortened with ﬂ owers appearing axil-

lary. Stamens sessile, indeﬁ nite in number,

without deﬁ nite arrangement in regard to the

sepals; ﬁ laments very short, slender; anthers

large, straight, oblong. Styles elongate-subu-

late, densely papillate from base to apex, with

slender bifurcating stigmas. Capsules opening

apically between the persistent stylodia. Seeds

with lid, exotegmic cells large, cuboid;

endosperm slight; Producing unusual B-ring

unsubstituted and 2'-hydroxylated ﬂ avonols.

n = 11.. . . . . . . . . . . . . . . . . .1. datiscaceae.

3 Large or very large trees, often buttressed,

with soft wood; hairs glandular or lepidote.

Nodes with two traces from the lateral gaps.

Leaves simple, cordate-ovate (sometimes

obliquely), entire or dentate, pubescent or

lepidote, long-petiolate, palmately 3–5-veined.

Flowers in long, pendant axillary spikes or

panicles. Stamens isomerous with and oppo-

site to the petals; ﬁ laments elongate, some-

what ﬂ attened-subulate; anthers relatively

short and ovoid, or, if larger and more or less

oblong, then conspicuously recurved. Styles

shorter or very short, stout, with large, capitate

or obliquely oblong-decurrent stigmas. Fruits

Superorder VIOLANAE 235

septicidal capsules, opening down the sides.

Seeds small, numerous, winged (Tetrameles).

Producing simple ﬂ avonols, kaempferol, and

quercetin. n = ca. 23. . . .2. tetramelaceae.

2 Leaves simple, often palmately lobed, with

mostly large stipules, usually asymmetrical,

mostly distichous. Nodes swollen. Stomata sur-

rounded by 3–6 subsidiary cells, these often

arranged in two rings. Vessels with simple or

(and) scalariform perforations with many bars;

lateral pitting scalariform. Flowers often some-

what zygomorphic, monoecious. Sepals in male

ﬂ owers two, rarely ﬁ ve, free or connate, valvate,

in female ﬂ owers usually 2–5, rarely 6–8, imbri-

cate; sepals (calyx lobes in female ﬂ owers) free,

but in Begoniella and in female ﬂ owers of

Symbegonia united into a tube. Petals absent or

(Hillebrandia and Begoniella) very small and

rudimentary. Stamens four (Begoniella) or numer-

ous, in many cycles, free or more or less connate;

anthers basiﬁ xed, opening by longitudinal slits or

rarely by apical pores. Gynoecium of (2)3(6) car-

pels; stylodia free or sometimes basally connate,

usually deeply biﬁ d; stigmas often twisted; ovary

inferior or (Hillebrandia) semi-inferior, 1- locular

(Hillebrandia and Begonia sect. Meziera) or sec-

ondary 2–4-locular. Ovules with zig-zag micro-

pyle. Fruits capsules (loculicidal or in Hillebrandia

opening at apex between the stylodia) or rarely

berries. With collar cells arranged in a transverse

ring around the operculum. Rather succulent

herbs or subshrubs with often jointed stems,

usually from thick rhizomes or tubers; present

sclereids and uncalciﬁ ed cystoliths, some climb

with the aid of aerial roots. Plants tanniniferous,

n = 10–21 +. . . . . . . . . . . . . . . . 3. begoniaceae.

1. CUCURBITACEAE

A.L. de Jussieu 1789 (including Bryoniaceae G. Meyer

1836, Cyclantheraceae Lilja 1870, Fevilleaceae

L. Pfeiffer 1879, Nhandirobaceae Lestibudois 1826,

Zanoniaceae Dumortier 1829). 123/800–900. Widely

distributed in tropics and subtropics, especially in the

rain forests of South America and the drier parts of

Africa, with relatively few species in Australasia and

the temperate regions.

Classiﬁ cation after C. Jeffrey (2005).

1.1 NHANDIROBOIDEAE

Tendrils two-branched from near the apex, spiraling

above and below the point of branching. Perennial,

dioecious herbs. Gynoecium with two or three stylodia.

Pollen grains small, striate, and uniform. Ovules

pendulous. Seeds often winged, n = 8. – zanonieae:

Fevillea, Alsomitra, Bayabusua, Zanonia, Siolmatra,

Gerrardanthus, Zygosicyos, Xerosicyos, Neoalsomitra,

Hemsleya, Gomphogyne, Gynostemma, Bolbostemma,

Actinostemma, Sicydium, Chalema, Pteropepon,

Pseudosicydium Cyclantheropsis.

1.2 CUCURBITOIDEAE

Tendrils unbranched or two-to seven-branched from

the lower part, spiraling only above the point of branch-

ing. Often annual, monoecious herbs. Stylodia united

into a style. Pollen grains colpate and (panto)porate.

Seeds unwinged, n = (7)8–14. – joliffieae: Baijiania,

Indofevillea, Sinobaijiania, Siraitia, Microlagenaria,

Thladiantha, Momordica, Telfairia, Odosicyos;

bryonieae: Bryonia, Ecballium; trichosantheae:

Hodgsonia, Ampelosicyos, Tricyclandra, Gymnopetalum,

Trichosanthes; herpetospermeae: Biswarea,

Herpetospermum, Edgaria; schizopeponeae:

Schizopepon; luffeae: Luffa; sicyeae: Hanburia,

Echinopepon, Apatzingania, Marah, Echinocystis,

Vaseyanthus, Brandegea, Elateriopsis, Pseudocyclanthera,

Cyclanthera (including Cremastopus

), Rytidostylis,

Sicyos, Sicyosperma, Parasicyos, Microsechium,

Sechium, Sechiopsis (including Pterosicyos); conian-

dreae: Kedrostis, Dendrosicyos, Corallocarpus,

Ibervillea, Tumamoca, Halosicyos, Ceratosanthes,

Doyerea, Trochomeriopsis, Seyrigia, Dieterlea,

Cucurbitella, Apodanthera, Guraniopsis, Melothrianthus,

Wilbrandia, Helmontia, Psiguria, Gurania; benin-

caseae: Solena, Borneosicyos, Papuasicyos, Trochomeria,

Dactyliandra, Ctenolepis, Peponium, Cephalopentandra,

Cogniauxia, Ruthalicia, Lagenaria, Benincasa,

Praecitrullus, Citrullus, Acanthosicyos, Eureiandra,

Bambekea, Nothoalsomitra, Coccinia, Diplocyclos,

Raphidiocystis, Lemurosicyos, Zombitsia, Melancium,

Cucumeropsis, Posadaea, Melothria, Muellerargia,

Zehneria, Cucumella, Cucumis, Oreosyce, Myrmecosicyos,

Mukia, Dicoelospermum; cucurbiteae: Cucurbita,

Sicana, Tecunumania, Calycophysum, Peponopsis,

Anacaona, Polyclathra, Schizocarpum, Penelopeia,

Cionosicyos, Cayaponis, Selysia, Abobra.

The Cucurbitaceae are more or less closely related

to Datiscaceae and Begoniaceae.

236 Subclass V. DILLENIIDAE

2. DATISCACEAE

Berchtold et J. Presl 1820. 1/2. Crete to western

Himalayas and from northern Baja California to north-

ern California.

Datisca.

According to Boeswinkel (1984) Datiscaceae (s. l.)

are less specialized than Begoniaceae because they

have a larger, less reduced nucellus without strikingly

swollen epidermal cells and because there is no special

rupture layer aiding the lifting of the operculum. In

addition, the vegetative characters of the Datiscaceae

are less specialized than those of the Begoniaceae.

3. TETRAMELACEAE

Airy Shaw 1965. 2/2. Malay Archipelago, New Guinea,

New Britain, and Solomon Islands (Octomeles) and

Sri Lanka, Andaman Islands, Indochina, and Malesia

(Tetrameles).

Tetrameles, Octomeles.

Close to the Datiscaceae, but sufﬁ ciently well dif-

ferentiated in morphological characters (Airy Shaw

1965; Davidson 1973, 1976), ﬂ avonoid proﬁ le

(Bohm 1988) and nucleotide sequence data (Swensen

et al. 1994) to deserve familial rank.

4. BEGONIACEAE

Berchtold et J. Presl 1820. 4/950. Widespread in tropics

and subtropics, especially in northern parts of South

America and in tropical Asia, absent in Australia;

Begonia (about 900 or more) is widespread; very closely

related to Begonia is South American Semibegoniella

with two species in Ecuador; the other two genera are

Paciﬁ c (the monotypic Hillebrandia in Hawaii and

Symbegonia with 13 species in New Guinea).

Begonia, Semibegoniella, Symbegonia, Hillebrandia.

Begoniaceae are rather closely related to the

Datiscaceae but are more advanced.

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Superorder CAPPARANAE

Order 69. ACANIALES

Evergreen or deciduous trees. Idioblastic and stomatal

myrosin cells present. Vessels mostly with simple perfo-

rations, but in mature wood occasionally with scalari-

form perforations and with numerous bars, often in

aberrant arrangements. Fibers with bordered pits. Rays

heterogeneous. Axial parenchyma scanty and vasicen-

tric. Sieve-element plastids of S-type (Bretschneidera).

Leaves alternate, imparipinnate, leaﬂ ets entire or ser-

rate; stipules small or reduced. Inﬂ orescences axillary or

terminal, simple racemes of paniculate, bracts minute or

lacking. Flowers actinomorphic or somewhat zygomor-

phic, bisexual. Sepals ﬁ ve, imbricate; petals ﬁ ve, longer

than sepals, clawed or slightly constricted at base,

contorted or imbricate. Stamens 8–10, free, inserted at

the base of receptacular cup; ﬁ laments pubescent at

base. Anthers dorsiﬁ xed or sub-basiﬁ xed, versatile,

tetrasporangiate, opening longitudinally. Tapetum

secretory. Microsporogenesis simultaneous. Pollen grains

2–3-colpate. Gynoecium of 3(5) carpels; style simple,

slender. Ovary superior, sessile, surrounded by copular

tube, 3-locular, pubescent, placentation axile; style

simple, slender; ovules 2–3 in each locule, pendulous,

anatropous or campylolotropous bitegmic, crassinucellate.

Female gametophyte of Polygonum-type (Akaniaceae),

or Allium-type (Bretschneideraceae). Endosperm nuclear.

Fruits loculicidal, leathery, pubescent capsules. Seed

broadly ellipsoid; seed-coat exo-mesotestal with

long palisade cells; testa vascularised, mesotesta and

endoteasta thickened; embryo straight, cotyledons thick;

endosperm copious or absent, n = 9.

Superorder CAPPARANAE 239

Morphological date support a relationship Akaniales

with Tropaeolaceae (Ronse Decraene and Smets 2001;

Ronse Decraene et al. 2002; Ronse Decraene and

Haston 2006).

Key to Families

1 Flowers actinomorphic; nectary disc absent.

Evergreen or deciduous small trees; the freshly cut

wood and bark with a turniplike odour. Rays very

wide, multiseriate. Leaves large, imparipinnate,

coriaceous; petiolules swollen both proximally and

distally, leaﬂ ets serrate, stipules small, subulate, or

reduced. Inﬂ orescences axillary, paniculate, bracts

minute or lacking. Flowers fragrang, white to pink-

ish. Sepals ﬁ ve, imbricate. Petals ﬁ ve, contorted.

Stamens usually eight, rarely 9–10, the ﬁ ve outer

ones opposite the sepals; ﬁ laments free; anthers

dorsi- to sub-basiﬁ xed, latrorse. Gynoecium of

three united carpels, with simple style and a small

3-lobed stigma; ovary 3-locular, with two axile,

superposed, pendulous ovules in each locule.

Ovules epitropous, anatropous, or hemitropous.

Fruits 3–6-seeded capsules. Seeds yellow, exaril-

late, with a straight, massive embryo and copious,

ﬂ eshy endosperm smelling of bitter almonds.

Present calcium oxalate druses and solitary crystals,

mustard-oils, proanthocyanidins (cyanidin and del-

phinidin), ﬂ avonols (kaempferol and quercetin),

producing alkaloids. . . . . . . . . . . . 1. akaniaceae.

1 Flowers slightly zygomorphic, nectary disc pres-

ent. Myrosin cells and mustard oils present in the

bark and inﬂ orescences. Small deciduous plants.

Leaves petiolate, leaﬂ ets entire, estipulate. Flowers

rather large, pinkish, in erect terminal racemes.

Calyx of connate sepals, campanulate, 5-toothed.

Petals ﬁ ve, free, unequal, basally clawed, imbri-

cate. Stamens eight, in two cycles, with slender,

hairy ﬁ laments attached to the rather thin, annular

nectary disc; anthers dorsiﬁ xed, introrse, connec-

tive is slightly prolonged, dehiscence is lateral, but

starts at the apex. Pollen grains 3-colpate or 2-col-

pate. Gynoecium of three (sometimes ﬁ ve) carpels,

with elongate, curved style and six-lobed stigma;

ovary 3-locular or sometimes 5-locular, with 2–3

campylotropous, apical-axile and pendulous ovules

per locule. Fruits thick-walled, 3–5-locular pubes-

cent capsules. Seeds orange to red; embryo large,

curved, endosperm lacking. Contain glucosino-

lates.. . . . . . . . . . . . . . . .2. bretschneideraceae.

1. AKANIACEAE

Stapf 1912. 1/1. Eastern Australia from southern coast

of Queensland to northeastern New South Wales.

Akania.

2. BRETSCHNEIDERACEAE

Engler et Gilg 1924. 1/1. Western and southwestern

China, northern Vietnam, Thailand, Taiwan.

Bretschneidera.

Closely related to Akaniaceae.

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240 Subclass V. DILLENIIDAE

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Yang D-Q and C-M Hu. 1985. The chromosomes of

Bretschneidera. Hems. Notes Roy. Bot. Gard. Edinb. 42:

347–349.

Order 70. TROPAEOLALES

More or less succulent perennial or annual herbs,

mostly climbing (petioles twining); perennial species

with a rhizome, a tuber or corms. Leaves and stems are

often covered by waxes. Stem with a ring of vascular

bundles surrounding the large pith and eventually

becoming connected through the activity of intravas-

cular cambium. Vessels with simple perforations, but

sometimes also with reticulate or intermediate perfora-

tions. Sieve-element plasdtids of S-type. Nodes trila-

cunar. Leaves alternate (or the lower opposite), more

or less peltate, entire, more or less lobed to palmately

divided, with palmate venation, stipules present or

absent, sometimes present only in the seedling stage,

stipules mostly small and caducous, but large and leaf-

like in a few species; petioles usually longer than the

lamina, more or less cirrhose in climbing species.

Flowers large, solitary, and axillary or rarely in umbel-

late or umbelliform inﬂ orescences, usually with long

and commonly pendant or subpendant, or rarely erect

(Magallana) pedicels, usually without bracteoles,

bisexual, usually strongly zygomorphic, 5-merous.

Calyx 5-lobed, imbricate or valvate, the adaxial one or

the adaxial and two lateral ones extended into usually

long nectariferous spur (much reduced or absent in

Trophaeastrum). Petals clawed, the three abaxial

(lower) ones usually different from the other two, often

with a hairy claw, rarely wanting. Stamens eight in a

single whorl; ﬁ laments free; anthers small, basiﬁ xed,

introrse, opening longitudinally. Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains

3-celled, 3-colporate or less often 2-colporate, with

reticulate ornamentation. Gynoecium of three united

carpels, with slender, terminal, apically triﬁ d style;

ovary superior, 3-locular, with a solitary, apical-axile,

pendulous ovule per locule. Ovules anatropous or

epitropous, bitegmic, tenuinucellate. Female gameto-

phyte or Polygonum-type. Endosperm nuclear. The

derivatives of the two uppermost cells of the proem-

bryo divide repeatedly to produce the basal mass of

cell that gives rise to very aggressive haustorial sys-

tem, at least in Tropaeolum majus. The peripheral cells

of the basal cell mass, close to the suspensor and on the

opposite side of the funicle, give rise to a rootlike haus-

torium that penetrates the integuments below the

micropyle and grows around the ovule in the carpel

cavity and into the carpel wall; shortly another hausto-

rium arises from the peripheral cells of the basal cell

masses, close to the funicle, grows through the integu-

ment and funicle, reaches the vascular bundle of the

placenta and grows along its curvature (Walker 1947;

Bhalla et al. 1979). Fruits separating into 1-seeded

ﬂ eshy or dry indehiscent mericarps or (Magellana) not

separating, broadly 3-winged, with only one mericarp

maturing; a short carpophore is found in Magellana

and a rather conspicuous one in Tropaeolum sect.

Schizotrophaeum. Seeds endotestal; embryo large,

straight, rich in oil, amyloid and protein, with thick

cotyledons; endosperm lacking. Contain myrosin cells

and tard oil; present glucosinolates, erucic and eicose-

nic acids, n = 12–15, 21.

Related to the Capparales.

1. TROPAEOLACEAE

Berchtold et J.Presl 1820. 3/95. Mountains of America

from Mexico to cool-temper-Chile (Tierra del Fuego).

Tropaeolum, Tropaeastrum, Magellana.

On the basis of the analysis of the elementary

cycles of the ﬂ ower of Tropaeolum majus, Chadefaud

(1974) concluded that Tropaeolaceae belong to the

Capparales. Dahlgren (1989) placed his Tropaeolales

(Tropaeolaceae and Limnanthaceae). Tropaeolaceae

shares with Capparales such common characters as

the absence of bracteoles, racemose inﬂ orescences,

the presence of a gynophore, strongly developed mar-

ginal bundles extending into the style, and a few other,

Superorder CAPPARANAE 241

more generalized characteristics (Ronse Decraene and

Smets 2001). However, strongly differ from them in

the axile placentation of the tenuinucellate ovules,

fruit morphology and dehiscence, the absence of pari-

etal cells, and very aggressive haustoria.

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Order 71. LIMNANTHALES

Small, delicate annuals of damp places. Vascular system

of the stem consists of a ring of 8–10 separate bundles.

Vessels with simple perforation. Phloem weakly devel-

oped. Sieve-element plastids of S-type. Nodes unilacu-

nar. Leaves alternate, simple or compound (pinnatisect

to pinnate), estipulate. Stomata anomocytic. Flowers

solitary on long, axillary pedicels, bisexual, actinomor-

phic, (4)5-merous (Limnanthes) or 3-merous (Floerkea),

Sepals more or less free, valvate (Limnanthes), or