Takhtajan Armen. Flowering Plants

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212 Subclass V. DILLENIIDAE

Niemeyera, Pradosia, Leptostylis, Pycnandra,

Synsepalum, Englerophytum, Xantolis, Capurodendron;

omphalocarpeae: Tsebona, Magodendron, Ompha-

locarpum, Tridesmostemon.

The delimitation of genera in the Sapotaceae has

always been very controversial. Here I follow

Pennington’s (2004) system of classiﬁ cation.

Bibliography

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ships in the order Primulales, with special emphasis on the

family circumscriptions. Canad. J. Bot. 73: 1699–1730.

Anderberg AA and U Swenson. 2003. Evolutionary lineages in

Sapotaceae (Ericales): a cladistic analysis based on ndhF

sequence data. Int. J. Plant Sci. 164: 763–773.

Aurbeville A. 1964. Système de classiﬁ cation des Sapotacèes.

Adansonia, ser. 2, 4: 38–42.

Baehni C. 1938. Memoires sur les Sapotacees: I. Systeme de

classiﬁ cation. Candollea 7: 394–508.

Baehni C. 1965. Memoires sur les Sapotacees: III. In-ventaire

des genres. Boissiera 11: 1–262.

Bartish IV, U Swenson, J Munzinger, and AA Anderberg. 2005.

Phylogenetic relationships among New Caledonian

Sapotaceae (Ericales): molecular evidence for generic poly-

phyly and repeated dispersal. Am. J. Bot. 92: 667–673.

Bokdam J. 1977. Seedling morphology of some African

Sapotaceae and its taxonomical signiﬁ cance. Meded. Land.

Wagen. 77/20: 1–84.

Cronquist A. 1946a. Studies in the Sapotaceae: II. Survey of the

North American genera. Lloydia 9: 241–292.

Cronquist A. 1946b. Studies in the Sapotaceae: VI. Miscellaneous

notes. Bull. Torrey Bot. Club 73: 465–471.

Govaerts R, DG Frodin, and TD Pennington. 2001. World

checklist and bibliography of Sapotaceae. Royal Botanic

Gardens, Kew.

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S Balasubramaniam. 1977. Triterpenoids and steroids of

some Sapotaceae and their chemotaxonomic signiﬁ cance.

Phytochemistry 16: 923–926.

Harley MM. 1986. Distinguishing pollen characters for the

Sapotaceae. Canad. J. Bot. 64: 3091–3100.

Harley MM. 1991a. The pollen morphology of the Sapotaceae.

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TD Pennington, The genera of Sapotaceae, pp. 23–50. Royal

Botanic Gardens, Kew, New York.

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of Sapotaceae, pp. 15–22. Royal Botanic Gardens, Kew,

New York.

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seed anatomy, 4: 49–53. Nauka, St. Petersburg (in Russian).

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Neotropical Sapotaceae: I–XIX. US Forest Service Research

Paper. FPL 325–331, 1978; 349–354, 1979; 358–363,

1980.

Lam HJ. 1925. The Sapotaceae, Sarcospermaceae, and

Boerlagellaceae of the Dutch East Indies and surrounding

countries (Malay Peninsula and Philippine Islands). Bull.

Jard. Bot. Buitenzorg 7: 1–289.

Lam HJ. 1939. On the system of the Sapotaceae, with some

remarks on taxonomical methods. Rec. Trav. Bot. Neerl. 36:

509–525.

Lam HJ and WW Varossieau. 1938. Revision of the

Sarcospermataceae. Blumea 3: 183–200.

Ng FSP. 1991. The relationships of the Sapotaceae within the

Ebenales. In TD Pennington. The genera of Sapotaceae,

pp. 1–13. Royal Botanic Gardens, Kew.

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1–770.

Pennington TD. 1991. The genera of Sapotaceae. Royal Botanic

Gardens, Kew.

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lies and genera of vascular plants, 6: 390–421. Springer,

Berlin/Heidelberg/New York.

Record SJ. 1939. American woods of the family Sapotaceae.

Trop. Woods 59: 21–51.

Smedmark JEE, U Swenson, and AA Anderberg. 2006.

Accounting for variation in substitution rates through time in

Bayesian phylogeny reconstruction of Sapotoideae

(Sapotaceae). Molec. Phyl. Evol. 39: 706–721.

Swenson U and AA Anderberg. 2005. Phylogeny, character evo-

lution, and classiﬁ cation of Sapotaceae (Ericales). Cladistics

21: 101–130.

Waterman PG and EN Mahmoud. 1991. Chemical taxonomy of

the Sapotaceae: patterns in the distribution of some simple

phenolic compounds. In: TD Pennington, ed. The genera of

Sapotaceae, pp. 51–74. Royal Botanic Gardens, Kew,

New York.

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1. Sapotaceae. J. Jpn. Bot. 46: 161–165.

Order 63. PRIMULALES

Trees or shrubs, sometimes woody lianas, rarely

sub-shrubs, mostly perennial or less often annual

herbs, sometimes aquatic (Hottonia). In Myrsinaceae,

Theophrastaceae and in some genera of Primulaceae,

particularly in Lysimachia, occur secretory cells or

cavities containing red or reddish substance. Vessels

with simple perforations or occasionally some of

them with scalariform perforations; lateral pitting

alternate. Fibers with simple or sometimes more or

less indistinctly bordered pits. Rays heterogeneous to

homogeneous, typically multiseriate. Axial paren-

chyma paratracheal and usually scanty, often lacking.

Sieve-element plastids of S-type. Nodes unilacunar

or sometimes trilacunar. Leaves alternate, opposite,

or less often verticillate, often gland-dotted or

Superorder PRIMULANAE 213

farinose, simple, entire or variously lobed, rarely

(Hottonia) pectinately pinnatisect, estipulate. Stomata

mostly anomocytic, or anisocytic. Flowers large to

small, in various kinds of inﬂ orescences or solitary,

bracteate but without bracteoles, bisexual or unisex-

ual, actinomorphic or rarely (Coris) somewhat zygo-

morphic, mostly 5-merous, often heterostylic. Sepals

3–6, mostly ﬁ ve, free or basally connate, imbricate,

valvate or contorted, often glandular-punctate, usu-

ally persistent. Corolla sympetalous, with imbricate

lobes, rarely petals secondarily free (in 3-merous

genus Pelletiera) or lacking (Glaux). Stamens as

many as and opposite the corolla lobes, attached to

the corolla tube or rarely almost free; sometimes there

is also an outer cycle of scalelike staminodia (as in

Theophrastaceae and Samolus and Soldanella) alter-

nating with the functional stamens; anthers tetraspo-

rangiate, sometimes with a prolonged connective,

introrse, opening longitudinally, or by apical pores.

Tapetum secretory. Microsporogenesis simultaneous.

Pollen grains 2-celled, (3-)-5-8-colpate, or 3(-5)-col-

porate, tectate- columellate, with tectum ﬁ nely granu-

lar to perforate. Gynoecium mostly of ﬁ ve united

carpels, with a style terminated by a capitate or punc-

tate, sometimes shallowly lobed stigma; ovary supe-

rior, sometimes semi-inferior, unilocular but often with

vestigial partitions at the base, mostly with numerous

ovules on a free-central placenta. Ovules anatropous or

hemitropous, sometimes campylotropous, bitegmic or

rarely unitegmic (as in Aegiceras, Cyclamen and

Douglassia), tenuinucellate, with endothelium.

Integumentary tapetum present or less often (some

Myrsinaceae and Theophrastaceae, Coris and

Cyclamen) absent. Female gametophyte of Polygonum-

type. Endosperm nuclear. Fruits berries, drupes or cap-

sules. Seeds large to small, with straight embryo and

usually copious endosperm; seed coat formed by the

outer integument. Commonly containing various sapo-

nine compounds, and many genera also produce qui-

noid compounds, notably primulagenin. n = 5, 8–15,

17, 19, 22.

Primulales have many common features with the

Ericales and Sapotales (conﬁ rmed by serological data,

Frohne and John [1978]), which supports their com-

mon ancestry. All this groups originated from the some

thealean ancestor. Hallier (1912) derived Primulales

from the Ochnaceae. But in Hutchinson’s opinion

(1959, 1969) the Myrsinaceae (but not the order

Primulales) linked with the Sapotaceae.

Key to Families

1 Fruits not viviparous.

2 Flowers actinomorphic. Calyx not spiny.

3 Mostly evergreen trees and shrubs, sometimes

subshrubs, very rarely subherbaceous.

4 Plants with secretory cells, cavities, or canals

with yellow or reddish-brown contents in the

stem and leaves. Staminodia absent.

5 Ovary superior. Pedicel without bracte-

oles. Vessels with simple or occasionally

scalariform perforations with 2–5 bars.

Fibers typically septate. Nodes trilacunar

with three traces. Leaves gland-dotted.

Sclerenchymatous subepidermal ﬁ bers

and submarginal strands in leaves want-

ing. Flowers small, without staminodia.

Sepals 5-4(3-6, very rarely 9). Petals

5-4(3-6, very rarely 7). Anthers introrse,

sometimes transversely locellate (Ardisia

elliptica). Gynoecium of 3–5(6) carpels.

Fruits usually berries or drupes, rarely

capsule, opening with valves. Seeds rela-

tively small to relatively large; seed coat

undistinguished, tegmen thickened before

becoming crushed, endotegmen crystal-

liferous; embryo straight or slightly

curved; endosperm copious, bony, some-

times ruminate, rarely well preserved only

on the funicular and placental side. Mostly

evergreen small trees or shrubs, some-

times lianas or epiphytes, rarely subshrubs

and very rarely (Ardisia primulifolia)

subherbaceous. Present benzoquinones,

n = 10–13, 15, 17, 23. . .1. myrsinaceae.

5 Ovary semi-inferior. Pedicel with a single

bract and two bracteoles. Evergreen small

trees, shrubs or sometimes lianas; secre-

tory canals well developed. Vessels with

simple perforation, but sometimes perfo-

ration scalariform with 5–7 bars; axial

parenchyma scanty paratracheal (Lens et

al. 2005). Leaves without immersed glan-

dular hairs, prismatic crystal druses pres-

ent. Inﬂ orescence terminal or axillary,

paniculate or racemose. Flowers small.

Corolla campanulate or sometimes urceo-

late, with a well-demarcated tube. Stamens

not exerted, adnate to the corolla at about

the middle or higher. Anthers not trans-

214 Subclass V. DILLENIIDAE

versely locellate, dorsiﬁ xed. Gynoecium

with somatic nectaries, stigma truncate or

capitate, entire or 2–5-lobed. Ovules many

to numerous, densely set on the placenta

and have an inner integument with a tape-

tum. Fruits many-seeded baccate, formed

by the ovary as well as the surrounding

tissue, with a somewhat woody endocarp,

externally punctate or lineate. Seeds small,

testa reticulate, two-layered with rhom-

boid crystals; embryo with short hypocotyl

and short, narrow cotyledons; endosperm

copious. n = 10. . . . . . . . . . 2. maesa ceae.

4 Plants without secretory system in the stem

and leaves. Vessels exclusively with simple

perforations; rays broad. Fibers not septate;

rays exclusively multiseriate. Nodes unila-

cunar with one trace (Jacquinia) or penta-

lacunar with ﬁ ve traces (Clavija). Leaves

commonly crowded at the ends of the

branches, with long strands of sclerenchyma-

tous tissue beneath the epidermis of the

leaves, estipulate. Flowers rather large, bisex-

ual, or unisexual (dioecious), with 4–5 peta-

loid or glandular staminodia alternating with

the corolla lobes and attached to the corolla

tube. Sepals 5(4), free or basally connate

(Clavija), persistent, imbricate. Petals 5(4),

corolla rotate, urceolate or funnel-shaped,

somewhat ﬂ eshy, gland dotted or streked.

Stamens 5(-4), inserted near the base of

the corolla tube; anther extrorse or introrse,

mostly with a prolonged connective.

Staminodia present. Pollen grains 3-colporate

or (Clavija) 4-colporate. Gynoecium of

5(-4) carpels. Ovary superior or semi-inferior,

unilocunar, the cavity ﬁ lled with mucilage.

Ovules numerous, imbedded in mucilage, on

central or basal placenta, ascending, anatro-

pous or campylotropous. Fruits berries or

rarely drupes, or rarely (Samolus) capsules,

containing many to few seeds (very rarely

only one). Seeds often fairly large; exotestal

cells ﬂ attened, thick-walled, hypo dermal

cells often crystalliferous; embryo straight

or slightly curved; endosperm copious,

bony; testa yellow, orange to red,. n = 9,

12–14, 19. . . . . . . . . . 4. theophrastaceae.

3 Perennial or annual herbs, rarely subshrubs.

Nodes unilacunar. Leaves alternate or more

often opposite or whorled, often all basal, often

glandular-punctate, mostly simple and entire or

merely toothed or lobed, in the aquatic genus

Hottonia pectinately pinnatisect. Stomata ano-

mocytic. Flowers variously in panicles, umbels,

racemes, of heads or solitary, individually brac-

teate but without bracteoles. Sepals 5(4-9), con-

nate, persistent, imbricate, or contorted; petals

5(4-9), (0 in Glaux), connate, imbricate or con-

torted; petals deeply biﬁ d to bilobed, or entire.

Stamens 5(4-9), inserted near the base of the

corolla tube, or midway down the corolla tube,

or in the throat of the corolla tube; ﬁ laments

long or short, rarely alternating with stamin-

odes (scales); anthers dorsiﬁ xed, opening by

longitudinal slits or sometimes by terminal

pores. Pollen grains 3(2-4)-colporate or colpor-

oidate, or 3–10-colpate. Gynoecium of 5(4-9)

carpels. Ovary superior, 1-locular. Ovules

numerous, ascending, anatropous to hemianat-

ropous. Fruits capsular, dehiscent by valves or

sometimes circumscissile, or rarely indehiscent.

Seeds small, usually more or less numerous;

seed coat rather thin, formed by both the integu-

ments, endotegmen often crystalliferous;

embryo straight, surrounded by usually copious,

oily endosperm, rarely (Soldanella, Hottonia)

endosperm lacking. Cotyledons one (Cyclamen)

or two, semi-cylindric. Present ﬂ avonols (kae-

mpferol, quercetin, myricetin), cucurbitacins,

n = 8–12(13). . . . . . . . . . . . . .5. primulaceae.

2 Flowers zygomorphic. Calyx spiny. Small,

evergreen, thyme-like subshrubs, stem often red-

dish-tinged. Leaves alternate, linear, coriaceous,

persistent, entire or obscurely dentate, gland-dot-

ted. Flowers in terminal spike-like racemes. Sepals

ﬁ ve, membranous, 10-nerved, teeth short, persis-

tent, valvate, each bearing a large black dorsal

gland, and an outer ring of 10–15 spreading aculei

below the teeth. Corolla campanulate, sub-bilabi-

ate (three upper lobes longer), lobes unequally

biﬁ d, bright magenta to rose or white. Stamens

ﬁ ve, antepetalous, ﬁ laments ﬁ liform, exerted,

glandular at the base; anthers small, globose,

introrse. Pollen grains relatively large and have a

unique kind of reticulate pattern in addition to the

tectum perforations (Carrion et al. 1993).

Gynoecium of ﬁ ve carpels. Ovary superiorn 1-loc-

ular, ovules 5–6, hemianatropous. Style ﬁ liform.

Capsule globose, enclosed by the persistent calyx,

Superorder PRIMULANAE 215

dehiscent by ﬁ ve valves. Seeds 4–6, papillose;

embryo well differentiated, straight; endosperm

present. Contain C-glycosyl ﬂ avonoid in the

leaves (Frezet et al. 1975). . . . . . . 6. coridaceae.

1 Fruits viviparous. Mangrove shrubs or small trees.

Vessel elements storied (Lens et al. 2005). Leaves

alternate, coriaceous, entire, minutely punctuate,

estipulate. Flowers in sessile terminal or axillary

umbels of very short branches racemes, without

bracts and bracteoles, bisexual. Sepals ﬁ ve, strongly

and dextrorsely contorted, imbricate, very asym-

metric; corolla shortly tubular, lobes ﬁ ve, contorted.

Stamens ﬁ ve, opposite the corolla-lobes, inserted in

the densely hairy tube of the corolla; ﬁ laments

shortly connate, villous in the lower part; anthers

2-locular, introrse, opening longitudinally. Ovary

superior, 1-locular, fusiform, narrowed into a subu-

late style, stigma acute. Ovules numerous (30–100),

but only one fertile, anatropous, unitegmic;

endosperm haustoria present Fruit 1-seeded, cap-

sule, curved and elongate. Seed large, pachychala-

zal; seed-coat thin; endosperm present

(Nemirovich-Danchenko 1992: 71); embryo well

differentiated, n = 23. . . . . . . 3. aegicerataceae.

1. MYRSINACEAE

R. Brown 1810 (including Ardisiaceae Jussieu 1810,

Embeliaceae J. Agardh 1858). 38/1000. Mainly tropi-

cal and subtropical regions, extending to South Africa

and New Zealand in the south, and North China, Korea,

Japan, and Mexico in the north.

Embelia, Grenacheria, Heberdenia, Pleiomeris, Myrsine,

Rapanea, Ardisia, Afrardisia, Synardisia, Hymenandra,

Tetrardisia, Solonia, Geissanthus, Emble mantha, Sadiria,

Gentlea, Stylogyne, Ctenardisia, Yunckeria, Antistrophe,

Parathesis, Amblyanthus, Ambly anthopsis, Elingamita,

Wallenia, Loheria, Cybianthus, Grammadenia, Vegaea,

Oncostemum, Badula, Tapeino sperma, Discocalyx, Labisia,

Systellantha, Monoporus, Fittingia, Conandrium.

2. MAESACEAE

A. Anderberg, B Stähl and M Källersjö 2000. 1/c.150.

trop. Africa, Madagascar, India, Sri Lanka, Burma,

China, Japan, Indochina, Malesia, Australia, Melanesia

eastwards to Fiji, Samoa, and Tonga.

Maesa.

Anderberg and Stahl (1995) came to the deﬁ nite

conclusion that Maesa should be raised to the rank of

family, which is supported by both morphological and

molecular date (see Anderberg et al. 2000).

3. AEGICERATACEAE

Blume 1833. 1/2. Mangroves South and South-East

Asia, New Guinea, Bismark Archipelago, Solomon

Is., northeastern Australia, Lord Howe I.

Aegiceras.

Aegicerataceae are related to Myrsinaceae but mark-

edly differ from them in the presence of versa tile anthers,

viviparous fruits with exalbumbinous seeds, and uniteg-

mic ovules (Stähl and Anderberg 2004), and very impor-

tant are also differences in the wood anatomy, such as

“Aegiceras can easily be distinguished from other

Myrsinaceae by the presence of relatively narrow ves-

sels, a relatively high vessel density, short vessel ele-

ments and ﬁ bers (non-septate ﬁ bers), and multiseriate

rays with exclusively procumbent body ray cells and

without sheath cells” (Lens et al. 2005).

4. THEOPHRASTACEAE

Link 1829. 7/125. America from Mexico and southern

Florida to West Indies and northern Paraguay.

Theophrasta, Neomezia, Deherainia, Votschia,

Clavija, Bonellia, Jacquinia.

Related to the Myrsinaceae.

5. PRIMULACEAE

Batsch ex Borkhausen 1797 (including Anagallidaceae

Batsch ex Borkhausen 1797, Lysimachiaceae A.L.

Jussieu 1789, Samolaceae Raﬁ nesque 1820). 24/1000.

Nearly cosmopolitan, but mainly in temperate and cold

regions of the Northern Hemisphere.

5.1 PRIMULOIDEAE

Ovary superior, capsule valvular, corolla lobes imbricate

or quineuncial in the bud. – primuleae: Omphalogramma,

Bryocarpum, Primula, Dionysia, Cortusa, Kaufmannia,

Hottonia, Sredinskya, Dodecatheon, Soldanella; and-

rosaceae: Androsace (including Douglasia), Vitaliana,

Stimpsonia, Pomatosace; ardisiandreae: Ardisiandra;

samoleae: Samolus.

216 Subclass V. DILLENIIDAE

5.2 LYSIMACHIOIDEAE

Ovary superior, capsule valvular or a pyxidium,

corolla-lobes contorted in bud. – lysimachieae:

Lysimachia (including Naumburgia), Trientalis,

Asterolinon, Pelletiera; GLAUCEAE: Glaux; anagal-

lideae: Anagallis (including Centunculus).

5.3 CYCLAMINOIDEAE

Ovary superior, capsule valvular. Tuberous plants; ﬂ owers

with reﬂ exed petal-limbs, twisted in bud. – Cyclamen.

Related to the Myrsinaceae, differing mainly in pre-

dominantly herbaceous habit, capsular fruits, seed coat

formed by both the integuments, and Caryophyllad-type

embryogenesis (Onagrad-type in the Myrsinaceae).

Although the Primulaceae are in general more advanced

than Myrsinaceae and Theophrastaceae, in some respects

they are more archaic (Wettstein [1935] considered

Myrsinaceae as an advanced family in the Primulales).

We may therefore conclude that the Primulaceae are

derived not directly from the Myrsinaceae or Theo-

phrastaceae but rather from the common ancestor of

these three families (Takhtajan 1959). The genus

Cyclamen is the most distinctive in the Primulaceae and

the systematic position of Cyclamen was discussed by

Rosvik (1966), who found it isolated in the family.

Cyclamen differs from all of them by the absence of the

integumentary tapetum.

6. CORIDACEAE

J.G. Agardh 1858. 1/2. Mediterranean, Northeastern

Africa.

Coris.

Usually included in the Primulaceae, but differs in

the zygomorphic ﬂ owers, spinose sepals and the major-

ity of them (except for Cyclamen) by the absence in

the integumentary tapetum.

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Superorder VIOLANAE

Order 64. BERBERIDOPSIDALES

Scandent, evergreen shrubs with sympodial branching

and collateral axillary buds. Vessel elements very long to

extremely long; perforations exclusively scalariform with

6–15 or more bars (mostly over 15 in Berberidopsis),

lateral pitting almost absent; vessel-to-ray pitting

scalariform, vessel to tracheid pitting composed of axial

rows of circular pits. Fibrous elements are nonseptate,

with distinctly bordered pits. Rays heterogeneous, over

1 cm in height. Axial parenchyma inconspicuous but pres-

ent. Sieve-element plastids of S-type. Nodes trilacunar

with three traces. Leaves alternate, entire to coarsely den-

tate, petiolate, coriaceous, estipulate. Stomata cyclocytic

and bicyclis. Flowers in terminal racemes (Berberidopsis)

or axillary, solitary, bisexual, actinomorphic. Sepals and

petals (9-)12(-15), and spirally arranged, gradually merg-

ing from one to the other (Berberidopsis) or the perianth

with ﬁ ve petals distinct from the ﬁ ve sepals

(Streptothamnus). Stamens 6–13 (Berberidopsis) or

numerous (Streptothamnus), free. Filaments very short,

connective broad (Berberidopsis) or ﬁ laments longer

than the anthers, ﬁ liform, connective inconspicuous

(Streptothamnus). Anthers basiﬁ xed, shortly apiculate,

opening longitudinally. Tapetum secretory. Microsporo-

genesis simultaneous. Pollen grains 3-colpate, tectum

imperforate (Berberidopsis), or perforate (Streptothamnus).

Disc dentate (Berberidopsis) or absent (Streptothamnus).

Gynoecium of three, ﬁ ve carpels. Ovary superior,

1-locular, ovules two to many on each of 3–5 parietal pla-

centae. Style stout, stigma punctuate to slightly lobed.

Ovules anatropous, bitegmic, crassinucellate. Female

gametophyte of Polygonum-type. Endosperm nuclear.

Fruits berrylike crowned by the persistent style base.

Seeds endotestal, with ﬂ eshy (Berberidopsis) or leathery

(Streptothamnus) exotesta; endotestal cells crystalliferous,

palisade, ligniﬁ ed; embryo minute and endosperm copi-

ous. Producing cyclopentenoid cyanogenic glycosides.

Evidently a very ancient family as conﬁ rmed by its

transpaciﬁ c disjunction (Berberidopsis in Chile and

Streptothamnus in Australia). Berberidopsidaceae are

probably directly related to the Dilleniales and are pos-

sible a link between the Dilleniales and Violales.

1. BERBERIDOPSIDACEAE

Takhtajan 1985. 2/3. Chile (from Talca to Osorno) and

eastern Australia (Queensland and New South Wales).

Berberidopsis, Streptothamnus.

Bibliography

Baas P. 1984. Vegetative anatomy and taxonomy of Berberidopsis

and Streptothamnus (Flacourtiaceae). Blumea 30: 39–44.

Carlquist S. 2003. Wood anatomy of Aextoxicaceae and

Berberidopsidaceae is compatible with their inclusion in

Berberidopsidales. Syst. Bot. 28: 317–325.

Heel WA. van. 1984. Flowers and fruits in Flacourtiaceae. V.

The seed anatomy and pollen morphology of Berberidopsis

and Streptothamnus. Blumea 30: 31–37.

Jaroszewski JW, PS Jensen, C Cornett, and JR Byberg. 1988.

Occurrence of lotaustralin in Berberidopsis Beckeri and its

relation to the chemical evolution of Flacourtiaceae.

Biochem. Syst. Ecol. 16: 23–28.

Kubitzki K. 2007. Berberidopsidaceae. In: K Kubitzki, ed. The

families and genera of vascular plants, vol. 9, pp. 33–35.

Springer, Berlin/Heidelberg/New York.

Ronse Decraene L-P. 2004. Floral development in Berberidopsis

corallina: a crucial link in the evolution of ﬂ owers in the core

eudicots. Ann. Bot. N. S. 94: 741–751.

Trifonova VI. 1992. Berberidopsidaceae. In: AL Takhtajan, ed.

Comparative seed anatomy, vol. 4, p. 78. Nauka, St.

Petersburg (in Russian).

Superorder VIOLANAE 219

Van Heel WA. 1979. Flowers and fruits in Flacourtiaceae IV.

Hydnocarpus, Kiggelaria africana L., Casearia,

Berberidopsis corallina Hook. f. Blumea 25: 513–529.

Van Heel WA. 1984. Flowers and fruits in Flacourtiaceae. V. The

seed anatomy and pollen morphology of Berberidopsis and

Streptothamnus. Blumea 30: 31–37.

Veldkamp JF. 1984. Berberidopsis (Flacourtiaceae) in Australia.

Blumea 30: 21–29.

Order 65. AEXTOXICALES

Large, evergreen trees. Twigs, leaves (below), and

inﬂ orescences lepidote, with peltate scales all over.

Vessels with scalariform perforations that have rather

numerous bordered bars. Axial parenchyma diffuse,

moderately common. Rays heterogeneous, droplets

or amorphous deposits of a dark-staining substance

common in ray cells (especially procumbent cells).

Leaves opposite, simple, conduplicate, entire, with-

out stipules. Stomata encyclocytic. Flowers in axil-

lary, pendulous racemes, dioecious, (4)5(6)-merous,

completely enveloped in bud by a calyptrate bracte-

ole. Sepals free, strongly imbricate. Petals broadly

clawed, incurved in buds, with a thick midrib, imbri-

cate. Stamens ﬁ ve, antesepalous, alternating with

well-developed, ﬂ eshy, reniform nectary glands; ﬁ la-

ments thick and ﬂ eshy; anthers introrse, opening by

porelike slits toward the summit. Pollen grains 3-col-

porate. Tapetum secretory. Microsporogenesis simul-

taneous. Female ﬂ owers with ﬁ ve ﬂ eshy staminodia

alternating with the nectary glands. Gynoecium of

two carpels, with a short style strongly deﬂ exed to

one side and appressed to the ovary, shortly biﬁ d;

ovary superior, 2-locular, but only one loculus fertile.

Ovules two, pendulous, anatropous and apotropous,

with a massively beaked nucellus protruding beyond

the integuments, bitegmic, crassinucellate. Fruits dry,

small drupes with a single 1-seeded pyrene. Seeds

with ruminate, copious, ﬂ eshy endosperm and a well-

developed embryo that has ﬂ attened, cordate-orbicular

cotyledons. n = 16.

Both molecular data and wood anatomy show the

close afﬁ nity with the Berberidopsidales. However

Aextoxicales markedly differ from the Berberidop-

sidales in opposite leaves, their dioecious ﬂ owers,

3-colporate pollen grains, the 2-carpellate gynoecium,

drupaceous fruits, ruminate endosperm, and well-

developed embryo.

1. AEXTOXICACEAE

Engler et Gilg 1920. 1/1. Chile.

Aextoxicon.

Bibliography

Carlquist S. 2003. Wood anatomy of Aextoxicaceae and

Berberidopsidaceae is compatible with their inclusion in

Berberidopsidales. Syst. Bot. 28: 317–325.

Hooker WJ. 1836. Aextoxicum functatum. Hooker’s Icones

Plantarum 1: t. 12.

Kubitzki K. 2006. Aextoxicaceae. In: K Kubitzki, ed. The fami-

lies and genera of vascular plants, vol. 9, pp. 23–25. Springer,

Berlin/Heidelberg/New York.

Komar GA. 1992. Aextroxicaceae. In: AL Takhtajan, ed. Com-

parative seed anatomy, vol. 4, p. 404. Nauka, St. Petersburg

(in Russian).

Order 66. VIOLALES (PASSIFLORALES)

Trees, shrubs, and herbs, often lianas. Vessels mostly

with simple perforations, but in some archaic members

of the order some or all of them with scalariform perfo-

rations, sometimes with many bars; lateral pitting from

scalariform to alternate or rarely (Sayauxia) scalariform

and opposite. Fibers with bordered or simple pits, often

septate. Rays from heterogeneous to homogeneous.

Axial parenchyma of various types or absent. Sieve-

element plastids of S-type. Nodes mostly trilacunar.

Leaves alternate or opposite, rarely verticillate, simple,

entire to dentate or serrate, sometimes (some Violaceae)

lobed or even dissected, commonly pinnately veined;

stipules present, often caducous, or absent. Stomata

paracytic, anisocytic or anomocytic. Flowers small to

large, mostly in various kinds of inﬂ orescences or rarely

solitary, bisexual or less often unisexual, actinomorphic

or sometimes more or less zygomorphic, cyclic or rarely

with a spiral perianth, mostly 5-merous, sometimes

apetalous. Sepals 6–3, rarely up to 15, mostly 5, free or

more or less connate, usually imbricate. Extrastaminal

corona sometimes present. Petals 6–3, rarely up to 15,

mostly 5, free or shortly connate at the base (in the

Achariaceae connate into a campanulate 5–3-lobed

tube), mostly imbricate. Stamens mostly the same num-

ber as the number of petals, free or more or less connate;

anthers tetrasporangiate, opening longitudinally or rarely

220 Subclass V. DILLENIIDAE

by terminal pores (as in Kiggelaria). Tapetum secretory.

Micro sporogenesis simultaneous. Pollen grains 2-celled,

3-colporate to 10–12-colporate, generally reticulate,

rarely inaperturate. Gynoecium syncarpous, of 2–12

carpels, rarely of one carpel, with free or more or less

united stylodia; ovary superior, rarely inferior (Bembicia)

with numerous or less often several or only 1–2 ovules

mostly on parietal placentas. Ovules mostly anatropous,

bitegmic, crassinucellate. Female gametophyte of

Polygonum-type. Endosperm nuclear. Fruits of various

types, mostly capsules or berries. Seed of various sizes

and shapes, arilate or not; seed coat formed by both

integuments, mostly exotegmic; embryo straight to

curved, endosperm copious to scanty. Certain members

of the order are cyanogenic (Malesherbiaceae,

Passiﬂ oraceae, Flacourtiaceae, and Turneraceae) and

produce elaborate cyanogens with cyclopentenoid ring

structures.

Violates are close to the Theales and share a com-

mon origin from a dillenialean ancestor.

Key to Families

1 Gynoecium of one carpel. Evergreen trees and shrubs

with pericyclic cork. Vessels usually with exclusively

scalariform perforations. Fibers with bordered pits,

septate. Axial parenchyma rare to frequent, vasicen-

tric and apotracheal diffuse. Nodes trilacunar with

three traces. Leaves alternate, simple, entire, serrate

or on the same specimen incised, estipulate. Stomata

anisocytic. Flowers axillary, small, sessile or pedicel-

late, bisexual, actinomorphic. Sepals (4-)5(-6),

broadly imbricate. Petals absent. Stamens 65–110,

persistent, inserted outside ﬂ oral cup; ﬁ laments are

longer than anthers, narrow; anthers very small,

basiﬁ xed, introrse; the connective broad and without

protrusion, opening longitudinally. Nectary is located

on the ﬂ oral cup, between androecium and gynoe-

cium. The carpel is elevated above the ﬂ oral base on a

short stalk; ovary superior, thick-walled, with one

short lateral placenta. Stigma large, almost sessile

forms two recurved crests and appears capitate-bilobed

and somewhat decurrent on the ventral side. Ovules

6–8, biseriate, campylotropous and syntropous; pla-

centation lateral; micropyle formed by the inner integ-

ument, which protrudes from the outer integument.

Fruits baccate. Seeds incurved, with an hippocrepi-

form embryo and scanty endosperm. Contain

mangiferin (Bate-Smith 1965).. . . 12. aphloiaceae.

1 Gynoecium mostly of 2–5 carpels.

2 Perianth present.

3 Petals present.

4 Petals more or less free.

5 Flowers with an extrastaminal corona.

6 Seeds exarillate. Xerophitic shrubs up

to 3.5 m tall, or small shrublets, peren-

nial or annual herbs with foetid smell,

erect or procumbent, bearing branches

sparsely to densely pubescent with

both simple and glandular hairs.

Vessels usually with simple perfora-

tions, but sometimes scalariform with

a few bars. Fibers rather short, with

very small pits. Nodes unilacunar

with one trace. Leaves alternate, sim-

ple, entire, sessile or petiolated,

toothed or sometimes pinnatiﬁ d;

stipule-like leaves usually present,

often 1–5-lobed. Stomata anomo-

cytic. Inﬂ orescences axillary or ter-

minal racemes, panicles or cymes, or

ﬂ owers solitary. Flowers bisexual,

actinomorphic, with a long, slender,

straight or curved hypanthium, blue,

purple, white, yellow, red to orange.

Sepals ﬁ ve, persistent, imbricate or

valvate; the margins sometimes lined

with glandular hairs. Petals ﬁ ve, val-

vate or with open aestivation, corolla

appendiculate, with a membranous,

denticulate corona. Both the androe-

cium and gynoecium borne on a vil-

lous, lobed androgynophore. Stamens

ﬁ ve, free and arising from the apex of

the androgynophore, extending beyond

the throat of the ﬂ oral tube and corona;

ﬁ laments ﬁ liform; anthers 2-locular,

dorsiﬁ xed. Pollen grains 3-colporate.

Gynoecium of 3(-4) carpels; stylodia

three, free, ﬁ liform, emerging near

apex of ovary, generally extending

stigmas beyond anthers. Ovary supe-

rior, 1-locular, stipitate, with numer-

ous, anatropous ovules on parietal

placentas. Fruits loculicidal capsules

enclosed by the persistent hypanthium,

stipitate. Seeds one to many, crusta-

ceous, pitted seed coat and with

straight, medium-sized embryo and

Superorder VIOLANAE 221

copious, ﬂ eshy, dotted endosperm.

Producing cyclopentenoid cyanogens;

n = 14.. . . . . . 10. malesherbiaceae.

6 Seeds arillate. Woody or herbaceous

lianas, climbing by axillary tendrils that

represent modiﬁ ed axes of the inﬂ ores-

cences or part of inﬂ orescences, less

often erect shrubs or trees. Woody

stems often with anomalous secondary

growth. Vessels mostly with simple

perforations, but in Soyauxia exclu-

sively with scalariform perforations,

and in Androsiphonia and Mitostemma

glasiovii with some amount of scalari-

form plates; lateral pitting usually alter-

nate, but in Soyauxia scalariform and

opposite. Fibers with bordered pits.

Rays of various types. Axial paren-

chyma usually apotracheal. Nodes tri-

lacunar. Leaves alternate, entire or often

palmately lobed, rarely compound,

often with nectary glands on the peti-

ole; stipules often small, sometimes

foliose, then to 7 cm long, often cadu-

cous. Stomata mostly anomocytic.

Inﬂ orescences axillary cymes or

racemes, or cauliﬂ orous, or the ﬂ owers

solitary, rarely the middle ﬂ ower

replaced by a hook. Flowers bisexual or

rarely unisexual, actinomorphic, mostly

with ﬂ at or tubular hypanthium, often

with elongate androgynophore. Sepals

(3-)5(-8), free or sometimes basally

connate, imbricate, persistent. Petals as

many as sepals, free or basally connate

(connate in tube around the gynoecium

in Androsiphonia), imbricate, rarely

wanting or very rudimentary. Extra-

staminal corona (poorly developed in

Adenia) of one or more series of ﬁ la-

ments or scales present. Stamens mostly

ﬁ ve, less often more (numerous in

Smeathmannia, Soyauxia, and Barteria)

or four, usually alternate with the pet-

als; anthers dorsiﬁ xed, introrse; anther

epidermis persistent. Pollen grains

3–12-colporate. Nectary disc of sta-

minodial origin often present around

the ovary. Gynoecium of (2)3(-5) car-

pels; stylodia mostly free or connate

only at the base, each with a capitate to

clavate or discoid stigma, rarely

(Barteria and Crossostemma) stylodia

united into a simple style; ovary supe-

rior, unilocular with more or less

numerous, orthotropous to anatropous

ovules borne on parietal placentas;

micropyle bitegmic or zig-zag. Fruits

loculicidal or irregularly decaying cap-

sules or berries, rarely ﬂ eshy with

irregular, apical dehiscence. Seeds

usually much-compressed, with bony,

pitted testa and mostly with a ﬂ eshy,

apical aril; endotesta short-palisade,

crystalliferous, ligniﬁ ed or not; embryo

large, straight, spatulate, embedded in

the copious, oily, rarely ruminate

(Passiﬂ ora foetida) endosperm.

Usually producing cyclopentenoid

cyanohydrin glycosides dominated by

transdioxygenes; alkaloids, ellagic

and gallic acids present or absent,

n = 6–7, 9–12.. . . 8. passifloraceae.

5 Flowers without extrastaminal corona.

7 Stamens usually 5.

8 Ovules at the top of the columnar.

Deciduous shrubs or small trees.

Vessel elements narrow and long,

with scalariform plates that have

numerous (up to 20 or more) bars.

Fibers with bordered pits. Lateral

pitting scarce. Rays uniseriate,

homogeneous. Axial parenchyma

absent. Leaves alternate, simple,

petiolate, entire, ovate, with small

caduceus (lanceolate) stipules.

Stomata anomocytic. Flowers in

axillary, umbellate, long peduncu-

late, globose inﬂ orescence, small,

actinomorphic. Extrastaminal disc

present; 5–7 glands opposite the

corolla members. Sepals 5–7, lin-

ear, pubescent sepals shortly con-

nate at the base, valvate, persistent.

Petals 5–7, very similar to the

sepals, valvate, persistent. Stamens

as many as and alternate with the

sepals, persistent, erect in bud, free;