Takhtajan Armen. Flowering Plants

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282 Subclass V. DILLENIIDAE

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Superorder MALVANAE 283

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284 Subclass V. DILLENIIDAE

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Superorder EUPHORBIANAE

Order 77. EUPHORBIALES

Trees and shrubs, woody lianas, or perennial and annual

herbs. Plants often with milky or colored juice. Vessels

with scalariform or more often simple perforations; lat-

eral pitting alternate or rarely opposite, very rarely sca-

lariform. Fibers mostly with simple pits. Rays

heterogeneous of various types. Axial parenchyma

mostly abundant and apotracheal or less often paratra-

cheal, but sometimes scanty or even wanting. Sieve-

element plastids of S-type. Nodes uni-, penta-, or

multilacunar. Leaves mostly alternate, simple or less

often compound, with pinnate or palmate venation,

mostly stipulate. Stomata mostly paracytic. Flowers in

very divers types of inﬂ orescences, unisexual (monoe-

cious or dioecious) or rarely bisexual, actinomorphic or

rarely slightly zygomorphic, often apetalous, sometimes

perianthless. Perianth 5-merous, rarely 4-or 3-merous.

Sepals and petals free or less often connate at the base.

Stamens 2-many, or sometimes fewer or even solitary,

free or variously connate; anthers usually 2-locular,

opening by longitudinal slits or sometimes transversely,

rarely by apical pores. Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains 2-celled

or 3-celled, 3-colpate, 3–4-colporate, pantoporate or

sometimes inaperturate. Nectary disc often present.

Gynoecium usually of three or less often two or four and

sometimes many united carpels, rarely pseudomono-

merous; ovary usually superior, 3-locular, rarely 2- or

4-locular or plurilocular, with free or more or less con-

nate stylodia. Ovules two or one per locule, pendulous,

anatropous, epitropous (the raphe ventral) or rarely

(Panda) orthotropous, mostly bitegmic, crassinucellate

or (Pandaceae) tenuinucellate. Female gametophyte

monosporic, bisporic, or tetrasporic. Endosperm nuclear.

Fruits of diverse types, but mostly capsules. Seed coat

usually formed by both integuments; embryo straight or

rarely curved; endosperm usually copious, rarely lack-

ing or represented by one layer of cells.

The archaic members of Euphorbiales show obvi-

ous links with the Elaeocarpales and Malvales.

Evidently the gynoecium was originally typically

paracarpous (as in Elaeocarpales and Malvales) and

became multilocular as a result of fusion of intrusive

parietal placentas in the center of the ovary. On the

other hand, the Euphorbiales have much in common

with the archaic members of the Violales, particularly

with the Flacourtiaceae. Hallier (1912: 217) included

the Euphorbiaceae in his “Passionales” close to the

Flacourtiaceae and noted that they originated from the

latter “pres de Pangiees et Idesiees.” Earlier (Hallier

1911: 5) he had emphasized the existence of numerous

parallel forms between these two families, such as

Pangium and Aleurites, Hydnocarpus and Cyclostemon,

or Ryparosa and Baccaurea. There are many common

features in wood anatomy, ﬂ ower morphology, and

palynology between Flacourtiaceae and Euphorbiaceae.

According to Metcaife and Chalk (1950: 1230),

“Antidesma, Bischofﬁ a, and Phyllanthus are very simi-

lar to certain genera of the Flacourtiaceae, e.g.,

Calancoba, Erythrospermum, and Kiggelaria;

Acalypha, Aporosella, Glochidion, and Hymenocardia,

with small intervascular and vessel ray pitting, suggest

the group of genera in the Flacourtiaceae that were

formerly included in the Samydaceae. Bridelia and

Cleistanthus closely resemble some genera of the

Samydaceae, e.g., Homalium (see also Miller 1975).

Keating (1973) has observed palynological similarities

between the Flacourtiaceae and the Phyllanthaceae.

There are also similarities in the leaf morphology,

ﬂ ower morphology, and seed anatomy (ﬁ brous exoteg-

men of some Flacourtiaceae and some Euphorbiaceae,

according to Corner [1976]). In his Principia botanica

(1960: 436, Fig. 50), Croizat regarded links both with

the Flacourtiaceae and the Sterculiaceae. There are

indeed some similarities between the Euphorbiaceae

and the Sterculiaceae, including petiole anatomy

(Dehay 1935), the presence of branched multicellular

hairs, the structure of the gynoecium, a uniformity in

the exotegmic palisade of Euphorbiaceae-Crotonoideae,

that, according to Corner (1976: 130), link them with

Sterculiaceae and related families.

One may presume that the Euphorbiales have arisen

either from some ancient group intermediate between

Superorder EUPHORBIANAE 285

the Flacourtiaceae and the Elaeocarpales and Malvales

or directly from some Flacourtiaceae-like lower

Violales. In both cases the Euphorbiales have only col-

lateral afﬁ nities with the Elaeocarpales and Malvales.

Key to Families

1 Ovules with obturator.

2 Ovules two per locule.

3 Vessel elements with scalariform perfora-

tion, very rarely (Glochidion) with simple

perforation.

4 Flowers in axillary or terminal spicate, rac-

emose or paniculate inﬂ orescences, or ﬂ ow-

ers solitary. Monoecious or rarely

(Antidesma) dioecious herbs to trees.

Laticifers absent. Nodes unilacunar with

one trace. Leaves usually alternate (very

rarely opposite), simple and unlobed, rarely

(Bischoﬁ a) trifoliolate, entire, stipulate.

Stomata paracytic or anisocytic. Flowers

small. Calyx often basally conntate; petals

ﬁ ve or absent, nectary disc present or

absent. Stamens 2–35, often more or less

connate; anthers extrorse. Pollen grains

2-celled, 3–4-colporate (rarely porate, pan-

toporate in Phyllanthus), semitectate, rarely

echinate. Gynoecium of 1(2–5) to 15 car-

pels; style single or branched, stigmas usu-

ally biﬁ d, fringed, wet; ovules hemitropous,

bitegmic, micropyle bistomal, outer integu-

ment 2-many and inner integument 2–3(-5)

cells across, nucellus 10 or more cells

across, protruding. Fruits septicidal cap-

sule, berries or drupes. Seeds large, without

caruncle or sometimes with rudimentary

caruncle; endosperm copious or absent.

Present cyanogenesis via the tyrosine path-

way, tropane and pyrrolizidine alkaloids,

cucurbitacins [triterpenes], nonhydrolys-

able tannins [geraniin], in Aleurites detected

inulin, n = 13, 14. . . 1. phyllanthaceae.

4 Flowers in axillary clusters, or cauliﬂ orous.

Evergreen trees or shrubs; hairs unicellular;

Leaves alternate, entire or serrulate,

2-ranked, stipulate Flowers axillary, pedi-

celled, apetalous; male ﬂ owers: sepals 3–6-

lobed, imbricate; stamens (2-)3–20

(-many); ﬁ laments free or subconnate;

anthers extrorse or introrse; pollen grains

3-colporate; disc present or absent; female

ﬂ owers: sepals 3–6-lobed, imbricate,

gynoecium of 1[-4(-9)] carpels; ovary usu-

ally 2–3-locular; style branches short or

absent, stigmas ﬂ ap-like; ovules two in each

locule, bitegmic, rarely (Drypetes mac-

rostigma) unitegmic (Corner 1976); fruit

drupaceous or dehiscing septicidally

(Lingelsheimia) into two 3-valved cocci or

six valves; seeds with sclereidal exome-

sotesta, exotegmen cells cuboidal; embryo

straight, endosperm copious, cotyledons

broad, ﬂ at and somewhat longer than the

radicle. Contain cucurbitacins glucosino-

lates, mustard oils present in Drypetes, n =

(19) 20 (21).. . . . . . . . . putranjivaceae.

3 Vessels with simple perforations. Monoecious

or dioecious trees. Mucilage cells present in

epidermis. Axial parenchyma apotracheal and

paratracheal; prismatic crystals in wood

parenchyma present. Leaves alternate, long

petiolate, trifoliolate, leaﬂ ets oblong- elliptic,

entire; stipules very minute, with colleters,

caduceus or persistent. Stomata paracytic.

Flowers small, axillary, solitary (the female

ﬂ owers, each on an apically expanded, long

peduncles), or aggregated in axillary panicles

or in catkins; perianth in male ﬂ owers absent,

stamens 3–54, loosely clustered and forming

a globose head; ﬁ laments free, short; anthers

minutely pubescent, 2-locular, slightly

extrorse, opening longitudinally. Pollen grains

spinulose, 6-colporate. Female ﬂ owers on

slender pedicels; sepals 4–5, narrowly lance-

olate, unequal, valvate or imbricate; nectarif-

erous weak disk around the base of the ovary.

Gynoecium of two carpels; ovary superior,

2-locular; stigmas stout, 2-lobed; ovules two

per locule, pendulous, anatropous, with obtu-

rators. Walls of the ovary, the style, the peri-

anth and pedicel have large cavities with a

viscous substance. Fruits globous drupes,

with a thin, ﬂ eshy, orange pericarp containing

numerous vesicles of bitter juice; seeds two

or one by abortion. Seeds massive, embryo

bent, with rather large and leaf-like cotyle-

dons without a plumule, much corrugated,

with very scanty endosperm. Present picro-

toxanes. . . . . . . . . . . . 3. picrodendraceae.

286 Subclass V. DILLENIIDAE

2 Ovules one. Trees, shrubs, subshrubs, or herbs

(some climbing or twining), very rarely ﬂ oating

aquatics. Indumentum simple, sometimes urticat-

ing, amalpighiaceous, dendritic, stellate, lepidote,

or absent. Stems sometimes succulent and/ or with

latex. Vessels usually with simple perforations,

but sometimes with scalariform ones. Fibers with

bordered or simple pits. Rays uniseriate and mul-

tiseriate. Axial parenchyma apotracheal, some-

times wanting. Nodes mostly pentalacunar with

ﬁ ve traces. Leaves usually alternate, sometimes

opposite, rarely verticillate, entire to dentate or

palmately lobed or compound, with pinnate or

palmate venation; stipules usually present, some-

times transformed into glands or spines, or absent.

Stomata of various types, but mostly paracytic.

Flowers in terminal or axillary, basically cymose

inﬂ orescences, always unisexual (monoecious or

dioecious), sometimes individually very much

reduced and grouped into pseudanthia, often apet-

alous, actinomorphic (but pseudanthia often zygo-

morphic). Perianth segments free or connate,

valvate, imbricate, or open, sometimes reduced or

absent; sepals and petals (1-)3–6(-8), sometimes

distinctly colored. Nectary disc mostly present,

intrastaminal or extrastaminal, annular, cupular or

of separate glands. Stamens one (Euphorbieae),

3–100 (up to 1,000 in Ricinus); ﬁ laments free or

connate; anthers introrse, latrorse or extrorse.

Pollen grains 2- or 3-celled, tectate or semitectate,

often reticulate, mostly 3-colporate, very diverse

in exine ornamentation. Gynoecium of (2)3(4-

many) carpels, rarely pseudomonomerous; stylo-

dia free or more or less united, entire, biﬁ d,

multiﬁ d or laciniate; ovary superior, sessile or

rarely stipitate, (1-)2–5(-20)-locular, most com-

monly 3-locular, with ovule per locule; styles

(1)3–4(20), free or connate, erect or spreading,

entire, 2-lobed, -ﬁ d or –partite to multiﬁ d or lac-

iniate; the adaxial surface usually stigmatic

throughout, smooth, granulate, papillose, plumose

or ﬁ mbriate. Ovules anatropous or less often

hemitropous or amphitropous, epitropous, biteg-

mic; the nucellus usually forms a beak that extends

beyond the inner integument and comes in contact

with the placental obturator, which forms a roof

over the exo(bi)stomal, micropyle. Female game-

tophyte of diverse types, but generally of

Polygonum-type. Fruits typically capsular schizo-

carps, primarily septicidal, with carpels (cocci)

elastically dehiscent from a persistent columella,

but sometimes drupaceous or baccate, very rarely

samaroid. Seeds one or ttwo in each locule, exote-

gmic, mostly with a micropylar caruncle; embryo

straight to curved or folded, usually with broad

and ﬂ at cotyledons; endosperm mostly well devel-

oped, less often wanting, Plants Al-accumulators,

producing cucurbitacins (triterpenes), ellagitan-

nins (geraniin and mallotussic acid), lectins

(hemagglutinins), cocarcinogens (phorbol ester

diterpenes), antidesmone and related compounds;

n = (5)6–9(-11), 12–14, 19. . .4. euphorbiaceae.

1 Ovules without obturator. Seeds not carunculate.

Dioecious trees or shrubs, indumentum simple.

Vessels with scalariform or both scalariform and

simple perforations. Leaves alternate, distichous,

simple, often serrate, entire or toothed; stipules

small, generally persistent. Inﬂ orescences terminal

or cauliﬂ orous and thyrsiform, or axillary and fas-

ciculate; bracts minute. Sepals free or connate,

imbricate or open. Petals slightly imbricate to val-

vate, often hooded. Stamens 5–15, free or nearly so;

anthers usually introrse. Nectary disc very small or

absent, rarely developed (Centroplacus). Gynoecium

of 2–5 carpels with 2–5(10) lobed style. Ovules

pendulous, anatropous and epitropous (the raphe

ventral) or (Panda) orthotropous, two (Centroplacus)

or one in each locule. Fruits capsular (Centroplacus)

or drupaceous; exocarp ﬂ eshy, tartareous or woody;

seeds ecarunculate; exotesta and endotegmen tan-

niniferous, endosperm copious; cotyledons thin and

ﬂ at, oily. n = 15. . . . . . . . . . . . . . . . 5. pandaceae.

1. PHYLLANTHACEAE

Martynov 1820 (including Antidesmataceae Loudon

1830, Aporosaceae Lindley ex Planchon 1854,

Bischoﬁ aceae Airy Shaw 1965, Hymenocardiaceae

Airy Shaw 1965, Porantheraceae Hurusawa 1954,

Scepaceae Lindley 1836, Stilaginaceae C.A. Agardh

1824, Uapacaceae Airy Shaw 1965). 59/1745.

Pantropical, but mainly in Malesia.

wielandieae: Heywoodia, Savia, Gonatogyne,

Petalo discus, Blotia, Actephila, Discocarpus, Lachnostylis,

Chonocentrum, Wielandia; amanoeae: Pentabrachion,

Amanoa; bridelieae: Cleistanthus, Bridelia; phyllant-

heae: Astrocasia, Leptopus, Chascotheca, Zimmermannia,

Superorder EUPHORBIANAE 287

Zimmermanniopsis, Meineckia, Pseudolachnostylis, Keayo-

dendron, Securinega, Andrachne, Flueggea, Richeriella,

Margaritaria, Phyllanthus, Reverchonia, Sauropus,

Breynia, Glochidion; antidesmeae: Spondianthus,

Uapaca, Protomegabaria, Maesobotrya, Richeria,

Jablonskia, Baccaurea, Ashtonia, Aporusa, Thecacoris,

Phyllanoa, Celianella, Leptonema, Antidesma, Aerisil-

vaea, Meborea, Hyeronima, Poranthera, Oreoporanthera;

tacarcuneae : Tacarcuna; hymenocardieae: Didy mo-

cistus, Hymenocardia; bischofieae: Bischoﬁ a; martre-

tieae: Martretia; dicoelieae: Dicoelia; croiza tieae:

Croizatia.

The family Phyllanthaceae is the most archaic

member of the order.

2. PUTRANJIVACEAE

Endlicher 1841. 4/215. Tropical America, tropical and

South Africa, South, Southeast and East Asia, Malesia,

Australia.

Lingelsheimia, Drypetes, Sibangea, Putranjiva.

Related to the Phyllanthaceae. Radcliffe-Smith

(2001) included all four genera in tribe Drypeteae

(Phyllanthoideae).

3. PICRODENDRACEAE

J.K. Small 1917. 1/3. West Indies (Greater Antilles,

Bahamas, Cayman and Swan Islands).

Picrodendron.

“The anomalous structure of the fruit and seed, as

well as the male inﬂ orescence, seems to indicate that,

despite pollen and other similarities, Picrodendron

does not really sit comfortably in the Oldﬁ eldioideae,

nor indeed in the Euphorbiaceae.” (Radcliffe-Smith

2001).

4. EUPHORBIACEAE

A.L. de Jussieu 1789 (including Acalyphaceae A.L. de

Jussieu ex Menge 1839, Androstachyaceae Airy Shaw

1965, Bertyaceae J.G. Agardh 1858, Cheilosaceae

Doweld 2001, Crotonaceae J.G. Agardh 1858,

Hippomanaceae J.G. Agardh 1858, Mercurialaceae

Martynov 1820, Micrantheaceae J.G. Agardh 1858,

Paivaeusaceae A. Meeuse 1990, Peraceae Klotzsch

1859, Pseudanthaceae Endlicher ex Pfeiffer 1873,

Ricinaceae Martynov 1820, Ricinocarpaceae Hurusawa

1954, Tithymalaceae Ventenat 1799, Tragiaceae

Raﬁ nesque 1838, Trewiaceae Lindley 1836). 276/

c.6000. Subcosmopolitan, although predominantly

tropical; there are strong local concentrations (particu-

larly of the genus Euphorbia) in South Africa, the

Mediterranean and Irano-Turanian regions, and south-

ern North America.

4.1 STACHYSTEMONOIDEAE

(OLDFIELDIOIDEAE)

Laticifers absent. Leaves stipulate or estipulate, alter-

nate or more often opposite or verticillate, simple or

palmately compound, entire or dentate. Petals absent

(except in Croizatia). Nectary disc present or absent.

Pollen grains 2-celled, tectate, brevicolporate or porate,

usually distinctly spinulose. Ovules two per locule

(except in Scagea). Seeds often carunculate. Endosperm

usually copious (except in Hyaenanche). – podoca-

lyceae: Podocalyx, Tetracoccus, Paradrypetes;

oldfieldeae: Piranhea, Parodiodendron, Oldﬁ eldia,

Aristogeitonia, Mischodon, Voatamalo, Androstachys,

Stachyandra; caletieae: Hyaenanche, Austrobuxus,

Myladenia, Canaca, Dissiliaria, Sankowskia, Whyanbeelia,

Choriceras, Longetia, Petalostigma, Kairothamnus,

Celaenodendron, Scagea, Neoroepera, Micrantheum,

Pseudanthus, Stachystemon.

4.2 PEROIDEAE

Laticifers absent. Trees, shrubs or herbs, with stel-

late or lepidote or occasionally simple indumentum,

more or less glabrous. Leaves alternate or very rately

opposite (in Pera), sometimes pellucid-punctate,

stipulate or estipulate. Inﬂ orescences axillary, with-

out or rarely with elongated axed, surrounded by

involucral bracts in Pera

only. Petals and nectary

disc absent or present. Stamens 2–20, free to con-

nate. Pollen grains 3–4-colporate, mostly tectate-

perforate, sometimes almost tectate or ﬁ nely

reticulate. Ovary 3(-4)-locular, styles biﬁ d to bipar-

tite. Ovule one per locule. Fruits dehiscent, septa

membranous, fragile, valves often remaining attached

to the base of the columella after dehiscence. Seeds

carunculate or arillate; endosperm copious, rarely

(Trigonopleura) scanty; cotyledons longer and wider

than radicle. – clutieae: Clutia; pogonophoreae:

Pogonophora; chaetocarpeae: Trigonopleura,

Chaetocarpus; pereae: Pera.

288 Subclass V. DILLENIIDAE

4.3 CHEILOSOIDEAE

Laticifers absent. Indumentum simple and/or stellate.

Leaves alternate, simple, unlobed, usually glandular at

the base of the lamina, stipulate. Inﬂ orescences pseudo-

terminal or axillary. Petals absent; nectary disc present

in male ﬂ owers, present or absent in female ﬂ owers.

Stamens 5–12, free; anthers introrse. Pollen grains

3-colporate, echinate. Ovary 2–4-locular, styles biﬁ d.

Ovule one per locule. Fruits tardily loculicidally dehis-

cent. Seeds not carunculate but with sarcotesta,

endosperm present. – Cheilosa, Neoscortechinia.

4.4 ACALYPHOIDEAE

Laticifers generally absent (if present they are inarticu-

late). Leaves usually stipulate, simple and entire, den-

tate or palmately lobed, rarely compound, alternate or

rarely opposite, often with foliar glands. Petals and

nectary disc present or absent. Pollen grains 2-celled,

mostly 3–4-colporate, semitectate, very rarely echi-

nate. Ovules one per locule. Seeds carunculate or not.

Endosperm usually copious. – erismantheae:

Erismanthus, Moultonianthus, Syndyophyllum;

ampereae: Monotaxis, Amperea; agrostis-

tachydeae: Agrostistachys, Pseudagrostistachys,

Cyttaranthus, Chondrostylis; sphyranthereae:

Sphyranthera; chrozophoreae: Speranskia,

Caperonia, Philyra, Ditaxis, Argythamnia, Chiro-

petalum, Doryxylon, Sumbaviopsis, Thyrsanthera,

Melanolepis, Chrozophora; caryodendreae: Caryo-

dendron, Discoglypremna, Alchorneopsis; bernar-

dieae: Bernardia, Necepsia, Amyrea, Paranecepsia,

Afrotrewia, Discocleidion, Adeno phaedra; pycno-

comeae: Pycnocoma, Droceloncia, Argomuellera,

Blumeodendron, Podadenia, Ptychopyxis, Botryophora;

epiprineae: Epiprinus, Symphyllia, Cleidiocarpon,

Koilodepas, Cladogynos, Cephalocrotonopsis, Cepha-

locroton, Adenochlaena, Cephalomappa; adelieae:

Adelia, Crotonogynopsis, Enriquebeltrania, Lasio-

croton, Leucocroton; alchorneeae: Orﬁ lea, Bossera,

Alchornea, Caelebogyne, Aparisthmium, Bocquillonia,

Conceveiba, Gavarretia, Polyandra; acalypheae:

Ricinus, Adriana, Mercurialis, Seidelia, Leidesia,

Dysopsis, Wetria, Cleidion, Sampantaea, Macaranga,

Erythrococca, Claoxylon, Claoxylopsis, Discocla-

oxylon, Micrococca, Lobanilia, Mareya, Mareyopsis,

Mallotus, Deuteromallotus, Cordemoya, Coccoceras,

Avellanita, Trewia, Neotrewia, Rocking hamia,

Octospermum, Acalypha, Lasiococca, Spathiostemon,

Homonoia; plukenetieae: Haema tostemon, Astro-

coccus, Angostylis, Romanoa, Eleuthe rostigma, Pluke-

netia, Vigia, Cnesmone, Megistostigma, Sphaerostylis,

Tragia, Tragiella, Platygyne, Acidoton, Gitara,

Pachystylidium, Dalechampia; omphaleae: Omphalea.

4.5 CROTONOIDEAE

Articulated and/or inarticulated laticifers usually pres-

ent. Leaves stipulate or estipulate, simple and entire,

dentate or palmately lobed or compound, alternate,

opposite, or rarely verticillate. Petals and nectary disc

present or absent. Pollen grains 2-celled or 3-celled,

3-colporate to more commonly porate or inaperturate,

mostly with ornamentation of knobs in hexagonal

pattern. Ovule one per locule. Seeds often carunculate.

Endosperm usually copious, often oily. – micran-

dreae: Micrandra, Micrandropsis, Cunuria, Hevea;

manihoteae: Manihot, Cnidoscolus; adenoclineae:

Glycydendron, Klaineanthus, Tetrorchidium, Adeno-

cline, Ditta, Endospermum; gelonieae: Suregada,

Cladogelonium;

elateriospermeae: Elaterio spermum;

jatropheae: Jatropha, Vaupesia, Oligoceras,

Deutzia nthus, Loerzingia, Joannesia, Leeuwenbergia,

Annesijoa; codiaeae: Baloghia, Hylandia, Ostodes,

Pausandra, Dodecastigma, Pantadenia, Parapan-

tadenia, Dimorphocalyx, Fontainea, Codiaeum, Acido-

croton, Ophellantha, Blachia, Strophioblachia, Sagotia,

Baliospermum; trigonostemoneae: Trigonostemon;

ricinocarpeae: Ricinocarpos, Alphandia, Beyeria,

Bertya, Myricanthe, Cocconerion, Borneodendron;

crotoneae: Mildbraedia, Chylamydojatropha, Para-

croton, Moacroton, Apodiscus, Croton, Crotonopsis,

Eremocarpus, Julocroton; ricinodendreae: Givotia,

Ricinodendron, Schinziophyton; aleuritideae:

Aleurites, Reutealis, Vernicia, Garcia, Cavacoa,

Grossera, Tapoides, Anomalocalyx, Sandwithia,

Tannodia, Neoholstia, Domohinea, Cyrtogonone,

Crotonogyne, Manniophyton, Neoboutonia, Benoistia.

4.6 EUPHORBIOIDEAE

Laticifers inarticulate. Leaves mostly stipulate (except

in Euphorbia), simple entire, dentate or lobed, alternate

(rarely opposite). Sepals mostly valvate (rarely imbri-

cate). Petals and usually nectary disc absent. Pollen

grains 2-celled or 3-celled, 3-colporate. Ovules one

per locule. Seeds carunculate or not. Endosperm copi-

ous. – stomatocalyceae: Plagiostyles, Pimelo-

dendron, Hamilcoa, Nealchornea; hippomaneae:

Homalanthus, Dendrocousinsia, Excoecaria, Spiro-

stachys, Sebastiania, Colliguaja, Microstachys,

Conosapium, Grimmeodendron, Bonania, Adenopeltis,

Stillingia, Spegazziniophytum, Falconeria, Sapium,

Superorder EUPHORBIANAE 289

Hippomane, Dendrothrix, Senefelderopsis, Plera-

denophora, Sclerocroton, Balakata, Triadica,

Shira kiopsis, Mabea, Gymnanthes, Ditrysinia,

Anomostachys, Neoshirakia, Pseudosenefeldera,

Actinostemon, Senefeldera, Rhodothyrsus, Dalembertia,

Maprounea; pa chystroma teae: Pachystroma; hureae:

Alger nonia, Tetraplandra, Ophthalmoblapton, Hura;

euphorbieae: Anthostema, Dichostemma, Neoguil-

lauminia, Calycopeplus, Euphorbia, Elaeophorbia,

Chamaesyce, Cubanthus, Monadenium, Synadenium,

Endadenium, Pedilanthus.

The Euphorbiaceae are one of the most diversiﬁ ed

angiosperm families.

5. PANDACEAE

Engler et Gilg 1912–1913. 4/18. Tropical Africa,

Southeast Asia to Solomon Islands, New Guinea.

cenrtoplaceae: Centroplacus; galearieae:

Galearia, Microdesmis, Panda

It is widely accepted that the Pandaceae belong to

the order Euphorbiales and Webster (1994), Thorne

(2006) and Radcliffe-Smith (2001) even include them

in the Euphorbiaceae in the subfamily Acalyphoideae.

However Stuppy (1996) considers that from the ovule-

and seed-morphology, the Pandaceae should be

re-instated.

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