Takhtajan Armen. Flowering Plants

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Superorder MYRTANAE 333

with membranous, minutely

wing, endotesta crystallifer-

ous, endotegmen tanniniferous;

endosperm lacking. Fibers

with distinctly bordered pits.

Internal phloem characteristi-

cally present. Nodes trilacunar

with three traces. Leaves oppo-

site, glabrous, entire, with a

marked midrib and continuous

marginal vein; petioles short;

small stipules sometimes

present. Stomata paracytic or

(Dactylocladus) anomocytic.

Evergreen trees up to 50 m

tall, often with quadrangular

twigs. Plants accumulate

aluminium. . . . . . . . . . . . . . .

7. crypteroniaceae.

10 Flowers not polygamo-dioe-

cious. Pollen grains never

2-colporate. Endothelium

always absent.

11 Anthers mostly opening

by single apical pore or

seldom by two pores or by

longitudinal slits; connec-

tive usually provided by

conspicuous appendage.

12 Interxylary (included)

phloem of the foram-

inate type present.

Fibers with distinctly

bordered pits. Nodes

unilacunar with three

traces. Large shrubs or

huge trees (up to 35 m

tall); indumentum

absent or consists of

uniseriate hairs. Leaves

opposite, petiolate,

simple, entire; venation

pinnate with indistinct

lateral and intramar-

ginal veins; stipules

absent. Stomata para-

cytic. Inﬂ orescences at

nodes of young wood,

sometimes cauliﬂ o-

rous, cymose, umbel-

loid, fasciculate, or

sometimes solitary at

nodes of older wood.

Flowers generally

small, actinomorphic,

bisexual or rarely

(Lijndenia) androdioe-

cious, 4–5-merous.

Sepals 4–5, minute,

imbricate in bud. Petals

imbricate in bud, yel-

low, pink, or bluish.

Stamens 4–5 or 8–10,

with a carnose connec-

tive generally provided

with a terpenoid-pro-

ducing gland; anthers

2-locular, versatile, with

ﬁ brous endothecium,

opening by longitudi-

nal slits, sometimes

short and functioning as

pores. Pollen grains het-

erocolpate. Gynoecium

of 2–5 carpels; style

elongate, stigma punc-

tate; placentation basal

or axile; ovary inferior,

1–5-locular, with two to

numerous, ascending,

anatropous, or campy-

lotropous ovules per

locule. Fruits ﬂ eshy,

1–5-seeded berries.

Seeds usually large,

without endosperm;

seed coat with ﬁ brous

exotegmen; embryo

green, with thick or con-

volute cotyledons. n = 7.

8. memecylaceae.

12 Interxylary phloem lack-

ing. Herbs, shrubs, treelets,

or trees up to 45 m tall, less

frequently scandent shrubs

or lianas, self supporting,

or epiphytic, or climbing.

Young stems often tetrago-

334 Subclass VI. ROSIDAE

nal. Indumentum very dif-

ferentiated, the trichomes

being generally large, mul-

ticellular, and scalelike.

Vessels with simple perfo-

ration and with vestured

pits. Fibers with simple

pits, often septate. Wood

parenchyma paratracheal,

or apotracheal and paratra-

cheal. Nodes unilacunar

with three traces. Leaves

nearly always opposite,

simple, entire to dentate,

conduplicate or supervo-

lute, with 3-9(-19) main

veins separate from the

base of the lamina; stipules

lacking. Stomata anomo-

cytic, polycytic, or ency-

clocytic. Flowers in

terminal panicles or cymes,

rarely solitary, bisexual, or

rarely androdioecious,

small and nearly actino-

morphic to large and

distinctly zygomorphic,

3-5(-7)-merous. Sepals 4,

or 5(-7), imbricate, or val-

vate. Petals imbricate in

bud. Stamens 4–5, or 8, or

10(-96), with connectives

often prolonged or pro-

vided with appendages, but

without glands; ﬁ laments

often twisted, bringing all

the anthers to one side of

the ﬂ owers, free of one

another; anthers basiﬁ xed,

tetrasporangiate, without

ﬁ brous endothecium, usu-

ally opening poricidal (api-

cally, with one, two, or

rarely four pores per

anther), or by short slits, or

rarely opening longitudi-

nally. Nectaries present or

absent. Pollen grains het-

erocolpate, supplied with

pseudocolpi or with inter-

colpate depressions isomer-

ous with the usually three

apertures. Gynoecium

generally of 3-5 (-14) car-

pels; ovary superior to infe-

rior, mostly 3–5-locular,

rarely 1-locular when the

partitions are dissolved,

with axile or rarely basal

placentation. Styles one,

apical, elongate; stigma

punctate to capitate. Ovules

(2-)6–50 per locule, usu-

ally anatropous, rarely

(Rhexia) orthotropous,

bitegmic, outer integument

contributing to the micro-

pyle (in addition to the

inner one); mizropyle zig-

zag. Fruits berries or loculi-

cidal capsules. Seeds

numerous, small; seed coat

without ﬁ brous exotegmen;

embryo minute, with two

small unequal cotyledons.

Contain acylated anthocy-

anins (cyanidin and del-

phinidin), ﬂ avonols

(kaempferol, quercetin, and

myricetin), rarely ellagic

acid n = 5(-7)-8–18.

9. melastomataceae.

11 Anthers always opening by

longitudinal slits. Mostly

herbs, less often subshrubs,

shrubs, or sometimes trees,

rarely very large and tall; bark

ﬂ aky, mucilage cells com-

mon; stems often 4-angled or

narrowly 4-winged when

young. Hairs uni- or bi-,

sometimes multicellular, or

(Cuphea) with cystolitic,

malpighiaceous, or glandular

hairs. Branched foliar sclere-

ids mostly absent. Internal

phloem usually present.

Vestured pits present in the

Superorder MYRTANAE 335

secondary xylem and internal

phloem. Axial parenchyma

scanty paratrachel. Leaves

usually opposite, less often

verticillate, rarely subalter-

nate, generally decussate,

simple, commonly entire, ﬂ at

to conduplicate, stipules ves-

tigial or none. Stomata mostly

anomocytic. Inﬂ orescences

axillary or terminal, thyrses,

cymes, umbelliform clusters,

of ﬂ owers solitary; epicalyx

of small teeth often present,

the teeth alternating with

calyx lobes. Flowers bisexual,

actinomorphic or sometimes

more or less zygomorphic,

with conspicuous, sometimes

spurred ﬂ oral tube, usually

(3)4-6(-8-16)-merous. Sepals

rather thick, valvate, appear-

ing as valvate lobes of the ﬂ o-

ral tube, often alternating

with external appendages at

the sinuses. Petals free,

attached at the summit or

within the ﬂ oral tube, pin-

nately veined, crumpled in

bud, sometimes wanting.

Stamens usually twice as

many as the sepals or petals,

in two cycles, rarely (Rotala)

solitary or numerous and cen-

trifugal as in Lagerstroemia;

ﬁ laments more or less elon-

gate, straight or incurved (the

long ﬁ laments of Lafoensia

inrolled) in bud, usually

inserted on the inside of the

ﬂ oral tube. Anthers versatile,

introrse, dorsiﬁ xed or seldom

basiﬁ xed. Pollen grains 3-col-

porate (some heterocolpate

with either three or six sub-

sidiary colpi). Nectary often

present, annular or unilateral

(Cuphea). Gynoecium of

2–4(-8), rarely (Punicoideae)

7–9(-15) carpels; style ﬁ li-

form, commonly elongate,

sometimes bent in bud, mostly

with capitate stigma; ovary

superior to inferior, often sur-

rounded at the base by an

annular nectary disc (or the

nectary unilateral), multiloc-

ular with as many locules as

carpels, but sometimes the

septa not reaching the sum-

mit; rarely the ovary

pseudomonomerous; placen-

tation axile or rarely free-cen-

tral. Ovules usually more or

less numerous in each locule,

anatropous or very rarely

amphitropous. Fruits usually

capsular, dehiscing variously

or seldom indehiscent, rarely

berrylike. Seeds generally

more or less numerous, ﬂ at-

tened, sometimes winged,

nearly or quite without

endosperm, with straight oily

embryo; cotyledons folded.

Often producing quinoliz-

idine alkaloids. n = 5, 7, 8,

15, 24, 28. 10. lythraceae.

7 Female gametophyte of Oenothera-

type. Annual or perennial herbs or

sometimes shrubs, lianas (Fuchsia),

rarely trees to 30 m tall (Hauya, some

Fuchsia), often with epidermal oil

cells. Included phloem of foraminate

type often present. Fibers with sim-

ple pits, often septate. The vegetative

parts are rich in oxalate raphides.

Leaves alternate, opposite, or more

rarely verticillate, usually entire or

dentate, but sometimes lobed or

lyrate-pinnatiﬁ d; small stipules

sometimes present, but in most gen-

era they are vestigial or wanting.

Stomata with three or more subsid-

iary cells, often anisocytic. Flowers

solitary in the axils, or in spikes, or

panicles, bisexual or seldom (spp. of

Fuchsia) unisexual, mostly with long

336 Subclass VI. ROSIDAE

ﬂ oral tube, which is nectariferous

within, actinomorphic or less often

more or less zygomorphic (strong

zygomorphy in Lopezia), mostly

4-merous or sometimes 2-merous

(Circaea), 5-merous, or 7-merous

(Ludwigia). Sepals 2–7, valvate,

commonly appearing as lobes on the

ﬂ oral tube. Petals 2–7, imbricate,

valvate, or contorted, often clawed or

stipitate, rarely wanting. Stamens

attached within the ﬂ oral tube, or

surrounding the epigynous nectary

disc, generally 2-cyclic or 1-cyclic,

or rarely reduced to 1–2 (and one of

these sterile in Lopezia). Staminodes

sometimes present in Lopezia and

Fuchsia. Pollen grains borne in

monads or sometimes in tetrads, or

polyads (Ludwigia), (2)3(-6)-apertu-

rate (colpate, colporate, or often

porate), with viscin threads on the

proximal surface in all but one spe-

cies (Circaea alpina), with dominant

circular or triangular central body

and usually markedly to slightly

protruding apertures; pseudocolpi

lacking. Gynoecium generally of as

many carpels as the sepals or petals;

style with a capitate, cylindrical, clav-

ate, or 4-lobed stigma; ovary inferior

or rarely semi-inferior, with as many

locules as carpels or the partitions

sometimes imperfect, so placentation

accordingly axile or parietal. Ovules

numerous or several (seldom 1 or 2)

in each locule or on each placenta,

anatropous. Fruits loculicidal (or both

loculicidal and septicidal) capsules or

sometimes berries (Fuchsia) or small

nuts (Gaura and Circaea). Seeds

numerous, several, or only one;

exotesta often hairy or papillate, inner

walls thickened and ligniﬁ ed; embryo

oily, straight or nearly so; endosperm

absent. Contain ﬂ avonoid sulphates,

n = 7, 9, 11. . . . . . 12. onagraceae.

3 Characteristic, schizogenous secretory

cavities lined with epithelium when

young and ﬁ lled with essential oils pres-

ent in leaves and other aerial organs.

Calyx and corolla imbricate.

13 Stamens usually numerous. Trees

(often rather large), shrubs or sub-

shrubs, with unicellular or sometimes

bicellular or multicellular hairs, usu-

ally with ectotrophic mycorrhizae.

Bark fairly smooth or rough and ﬁ s-

sured, sometimes peeling in large

plates (e.g., Psidium guajava). Internal

phloem characteristically present,

next to the pith. Secondary phloem of

young stems usually stratiﬁ ed. Vessels

usually with simple perforations,

rarely (Neomyrtus, Myrceugenia,

Luma, Ugni, Myrteola, and several

other Central and South American

genera) with scalariform perforations.

Fibers mostly with distinctly bordered

pits (ﬁ ber-tracheids), very rarely sep-

tate. Axial parenchyma typically

apotracheally diffuse or diffuse-in-

aggregates. Leaves opposite or less

often alternate, rarely verticillate,

simple, entire, often coriaceous, glan-

dular-punctate, often with a continu-

ous intramarginal vein; stipules

vestigial or wanting. Stomata anomo-

cytic or seldom paracytic. Flowers in

various types of cymose or racemose,

often complex inﬂ orescences, rarely

solitary and axillary, bisexual, nearly

always actinomorphic, mostly 4- or

5-merous, with a ﬂ oral tube of vari-

able length often prolonged above the

ovary, on the margin of which the

sepals, petals, and normally numerous

stamens are all inserted. Sepals (3)4–

5(-8), usually imbricate, or the calyx

sometimes very reduced or splitting at

anthesis or forming a deciduous calyp-

tra. Petals similar, sometimes (as in

spp. of Eucalyptus) connivent to form

a calyptra, or sometimes wanting.

Stamens usually numerous, develop-

ing centripetally, free or basally united

into four or ﬁ ve fascicles (nearly

always antepetalous) each supplied by

Superorder MYRTANAE 337

one vascular trunk bundle, usually

inﬂ exed in bud; anthers small, versa-

tile, opening by longitudinal slits or

(as in certain sections of Eucalyptus

and in Chamelaucium and related

genera) by pores, the connectives usu-

ally have one or more apical secretory

cavities (glands). Pollen grains

2-celled, generally triangular, usually

syncolporate, and lack pseudocolpi.

Staminodia are rarely present. Nectary

disc borne on the summit of the ovary

or lining the prolonged ﬂ oral tube.

Gynoecium of 2–5(18) carpels; style

generally elongate, with capitate or

punctiform stigma; ovary with as

many locules as carpels or seldom

(some Leptospermoideae) the ovary

pseudomonomerous, with a single

locule, semi- inferior or inferior, or

rarely superior; ovules usually numer-

ous, occasionally few, very rarely soli-

tary, anatropous or campylotropous,

bitegmic or (some ssp. of Eugenia and

Syzygium) unitegmic. Female game-

tophyte of Polygonum-type or rarely

(Heteropyxideae) Allium-type. Fruits

baccate or loculicidal (very rarely cir-

cumscissile) capsules, sometimes

drupes, schizocarps, or nutlike,

crowned by persistent sepal-lobes or

scars. Seeds one to few, very occa-

sionally many; seed coat usually

membranous or chartaceous, some-

times hard and bony; embryo of

diverse shape and size, rarely

(Eugenia) undifferentiated, green or

white; endosperm usually absent,

occasionally in a scanty amount; coty-

ledons often connate, intricately

folded. Contain ethereal oils (usually

terpenes) and polyhydroxyalkaloids

(Porter et al. 2000), n = 5–11(12), less

often 6–9. . . . . . . . . 13. myrtaceae.

13 Stamens 1–5(-7), usually only one

fertile. Trees, sometimes very large,

often with well- developed buttresses,

shrubs, or somewhat woody vines,

rarely herbs; young parts often

densely pubescent with simple or

stellate (Erisma) hairs. Intraxylary

phloem usually present in the pre-

medullary region. Indumentum often

brown, hairs T-shaped, unicellular,

or stellate. Vessels exclusively with

simple perforations; lateral pitting

alternate and vestured. Fibers with

few, small, simple or sometimes with

very small borders in Vochysia, a few

cells sometimes septate in Erisma

and Vochysia. Rays heterogeneous to

homogeneous. Axial parenchyma

predominantly paratracheal. Nodes

trilacunar or unilacunar. Leaves ever-

green, opposite, or sometimes ver-

ticillate, leathery; stipules small,

sometimes represented by large

glands, or colleter-like, or wanting.

Stomata paracytic or anomocytic.

Inﬂ orescence terminal, axillary

racemes (panicles) or cincinni.

Flowers obliquely zygomorphic.

Sepals ﬁ ve, basally connate, persis-

tent, imbricate, one of them usually

the largest and with gibbous or

spurred base, rarely (Korupodendron)

calyx petaloid. Petals 1–3, very

rarely 5, or sometimes wanting, more

or less unequal, contorted, or imbri-

cate. Stamens 1-5(-7), usually only

one fertile; ﬁ laments tree or rarely

connate in two groups; anthers

introrse, sometimes with an expanded

connective, opening longitudinally.

Pollen grains 3-colporate, with retic-

ulate or striate exine. Nectaries want-

ing. Gynoecium usually of three

carpels, sometimes pseudomono-

merous; style with a capitate or lat-

eral stigma, sometimes very short;

ovary superior and 3-locular or infe-

rior and 1-locular, ovules (1)2-many

in each locule, pendulous, epitrop-

ous or anatropous, bitegmic, crassi-

nucellate. Fruits loculicidal capsules

or sometimes samaras winged with

accrescent calyx. Seeds often winged

or hairy, testa thin, endotestal cells

338 Subclass VI. ROSIDAE

more or less thickened, pectic,

mesotegmic cells ﬁ brous, thick-

walled or nor, or testa multiplicative,

exotesta with thickened hairs;

embryo straight (longitudinal to the

seed), endosperm wanting or rarely

developed; cotyledons folded. Plants

5-deoxyﬂ avonoids, usually accumu-

late aluminum,

n = 11 or 12. . . . 14. vochysiaceae.

2 Plants annual aquatics, sometimes free-ﬂ oat-

ing. Raphides wanting. Submerged part of

the stem with elongate internodes and more

or less opposite or ternate, ﬁ liform-dissected,

leaﬂ ike green organs ( transformed stipules

according to Vasiliev [I960]) and also with

ﬁ liform adventitious roots from some of the

nodes; aerial part of the stem short, densely

crowded leaves. Leaves with more or less

rhombic laminas, the elongate petiole usu-

ally with an inﬂ ated, aerenchymatous ﬂ oat

near midlength and small, cleft, deciduous

stipules; the ﬂ oating leaves have marginal

teeth with a unique double apex. Stomata

anomocytic. Flowers solitary on short pubes-

cent pedicels, in the axils, slightly elevated

above the water, bisexual, actinomorphic,

4-merous. Sepals basally connate into a short

hypanthial tube, lobes valvate, two or four of

them persistent and accrescent in fruit as

hornlike or spinelike projections. Petals

alternate with the sepals, imbricate. Stamens

alternate with the petals, with short ﬁ la-

ments; anthers dorsiﬁ xed, opening longitu-

dinally. Pollen grains 2-celled, triangular,

3-colporate, with three prominent meridi-

onal ridges formed by the uplifting and fold-

ing of the ectexine and enclosing a cavity;

the ridges pass over the colpi as a wide ver-

rucate-granular band; the apertures are prod-

ding and swollen as elongated domes.

Gynoecium of two carpels, with subulate

style and capitate stigma; ovary partly infe-

rior, surrounded at the base by 8-lobed cupu-

lar disc, 2-locular, with usually one,

pendulous, apical-axile, anatropous ovule

per locule, but one locule and its ovule abort-

ing after anthesis. Endosperm formation

hardly takes place at all. Fruits mostly

1-seeded, horned, drupelike, with thin, soon

disappearing, ﬂ eshy exocarps and persistent,

stony endocarp. Seed large, without

endosperm; embryo straight, the cotyledons

very unequal in size, shape, and behavior,

one large, starchy, kept within the fruit, the

other small, scalelike, growing out with plu-

mule through the pore left by dehiscence of

the style, n = 20, 24. . . . . . . 11. trapaceae.

1. ALZATEACEAE

S. Graham 1985. 1/1. Distributed along the lower

slopes of the Andean Cordillera in Colombia, Ecuador,

Peru and Bolivia, in humid forests of the upper

Amazonian basin, and disjunctly in the cloud forests of

Costa Rica and Panama.

Alzatea.

“Comparison of specialized morphological attri-

butes and sequence data from the rbcL gene indicate

that Alzatea belongs near the small myrtalean families

Rhynchocalycaceae, Oliniaceae, and Penaeaceae”

(Graham 2004). From all other myrtalean families the

Alzateaceae differ in their trilacunar node.

2. RHYNCHOCALYCACEAE

L. Johnson et B. Briggs 1985. 1/1. Eastern parts of

South Africa (endemic to the sandstone region of

southern Natal and Pondoland).

Rhynchocalyx.

In the cladistic analysis of Johnson and Briggs

(1984:732), Rhynchocalyx comes out on a common

stem with Alzatea. According to Dahlgren and van

Wyk (1988: 47), Rhynchocalyx is clearly related to a

group of families consisting of Crypteroniaceae,

Penaeaceae, Oliniaceae, Alzateaceae, and Lythraceae

without, however, particularly close living relatives.

They conclude that Oliniaceae, Rhynchocalycaceae,

and Penaeaceae seem to have evolved from the same

ancestral stock, which is to some extent also reﬂ ected

in their present geographical distribution.

3. GEISSOLOMATACEAE

Endlicher 1841. 1/1. Restricted to the Cape Province,

occurring in the Langeberg Mountains of the Swellendam

and Riverside districts, where they grow in fynbos.

Superorder MYRTANAE 339

Geissoloma.

Related to the Pennaeaceae. Differing mainly in

female gametophyte. Lindley (1853) and Gundersen

(1950) even included Geissoloma in Penaeaceae.

4. PENAEACEAE

Sweet ex Guillemin 1828. 7/25. Conﬁ ned to the Cape

Province and concentrated in the southwest. Penaea

has the widest distribution toward the east and extends

to Port Elizabeth.

Endonema, Glischrocolla, Sonderothamnus,

Saltera, Brachysiphon, Stylapterus, Penaea.

Penaeaceae are closely related to the Memecylaceae,

Cryptoreniaceae, and Alzateaceae, and especially to

Oliniaceae and Rhynchocalycaceae (Dahlgren and van

Wyk 1988: 45). According to Dahlgren and Thorne

(1984: 670), the ancestors of Penaeaceae could have

had a common origin with Rhynchocalycaceae.

5. OLINIACEAE

Arnott 1839. 1/8. Southern and eastern Africa (Olinia

ventosa on St. Helena is most probably an introduction

from South Africa – Dahlgren and van Wyk [1988]).

Olinia.

Probably related to the Penaeaceae (Johnson and

Briggs 1984). “Penaeaceae, despite their rather different

type of embryo sac, appear to be especially close to

Oliniaceae,” state Dahlgren and van Wyk (1988: 43).

Molecular studies clearly support a placement

Oliniaceae in the order Myrtales (Conti et al. 1996).

6. COMBRETACEAE

R. Brown 1810 (including Myrobalanaceae Martynov

1820, Terminaliaceae Jaume Saint-Hilaire 1805).

14/500. Mainly tropical regions, with a few species

extending into subtropical areas; particularly common

in Africa. Strephonema (3) is endemic to tropical West

Africa.

6.1 STREPHONEMATOIDEAE

Ovary semi-inferior. Pollen grains without pseudo-

colpi. Cotyledons massive, hemispheric. Stomata

paracytic. Fibers with clearly bordered pits. –

Strephonema.

6.2 COMBRETOIDEAE

Ovary inferior. Pollen grains with pseudocolpi.

Cotyledons folded, spirally twisted. Stomata

anomocytic or encyclocytic. Fibers with simple pits. –

laguncularieae: Laguncularia, Lumnitzera,

Macropteranthes, Dansiea; combreteae: Combretum

(including Quisqualis, Calopyxis, Thiloa, Melostemon),

Guiera, Calycopteris, Pteleopsis, Terminalia ( including

Terminaliopsis, Bucida), Buchenavia, Anogeissus,

Finetia, Conocarpus.

Combretaceae occupy a somewhat isolated position

within the Myrtales. Johnson and Briggs (1984) place

them between Penaeaceae-Oliniaceae and Cryptore-

niaceae-Melastomataceae.

7. CRYPTERONIACEAE

A. P. de Candolle 1868 (including Henslowiaceae

Lindley 1835). 3/9–10. Sri Lanka, India (Assam and

Bengal), southwestern China, Andaman Islands, Malay

Peninsula, Philippine Islands, Malay Archipelago.

Crypteronia, Dactylocladus, Axinandra.

Crypteroniaceae exhibit relationships to

Alzateaceae-Rhynchocalycaceae and especially to

Memecylaceae. Some authors, including van Vliet,

Koek-Noorman, and ter Welle (1981) and van Vliet

and Baas (1984) even include Crypteronioideae and

Memecyloideae in the Melastomataceae sensu lato.

But from all other Myrtales the Crypteroniaceae (at

least the genus Axinandra) differ in the presence of

endothelium (Tobe and Raven 1983b).

8. MEMECYLACEAE

A. P. de Candolle 1828 (including Mouririaceae

Gardner 1840). 6/450. Pantropical.

Memecylon, Warneckea, Lijndenia, Spathandra,

Mouriri, Votomita.

Related to Melastomataceae, but differ in the leaf

venation, the number and size of the seeds and the

presence of an elliptic terpenoid-producing gland on

the staminal connective (Renner 2004).

9. MELASTOMATACEAE

A. L. de Jussieu 1789 (including Blakeaceae Barnhart

1895, Miconiaceae C. Martius 1835, Rhexiaceae

340 Subclass VI. ROSIDAE

Dumortier 1822). 163/4500–4570. Mainly in tropical

and subtropical regions, especially in South

America.

kibessieae: Pternandra (including Kibessia);

astronieae: Astrocalyx, Astronia, Astronidium,

Becca-rianthus; sonerileae: Anerincleistus,

Barthea, Blastus, Bredia, Oxyspora, Poikilogyne,

Driessenia, Neodriessenia, Ochthocharis,

Poilannommia, Sporoxeia, Stussenia, Amphiblemma,

Aschistanthera, Bertolonia, Boyania, Brittenia,

Calvoa, Centradenia, Cincinobotrys, Cyanandrium,

Dicellandra, Diplarpea, Enaulophyton, Fordiophyton,

Gravesia, Kerriothyrsus, Macrocentrum, Monolea,

Opisthocentra, Phyllagathis, Preussiella, Salpinga,

Sarcopyramis, Scorpiothyrsus, Sonerila, Tateanthus,

Triolena, Tryssophyton, Maguireanthus; merian-

ieae: Adelobotrys, Axinaea, Behuria, Benevidesia,

Bisglaziovia, Centronia, Dolichoura, Graffenrieda,

Huberia, Meriania, Merianthera, Neblinanthera,

Ochthephilus, Pachy loma, Phainantha,

Tessmannianthus; rhexieae: Rhexia; melastomeae:

Dissotis, Acanthella, Acotis, Acisanthera,

Amphorocalyx, Antherotoma, Appen dicularia,

Brachyotum, Cailliella, Chaetolepis, Comolia,

Comoliopsis, Desmoscelis, Dichaetanthera,

Dinophora, Dionycha, Ernestia, Fritzschia, Guyonia,

Heterocentron, Heterotis, Loricalepis, Macairea,

Mallophyton, Marcetia, Melastoma, Melastomastrum,

Microlepis, Monochaetum, Nepsera, Nerophila,

Otanthera, Osbeckia, Pilocosta, Poteranthera,

Pseudoosbeckia, Pterogastra, Pterolepis,

Sandemania, Schwackaea, Siphanthera, Svitramia,

Tibouchina, Tibouchinopsis, Tristemma; microl-

icieae: Lavoisiera, Bucquetia, Cambessedesia,

Castratella, Chaetostoma, Eriocnema, Lavoisiera,

Lithobium, Microlicia, Rhynchanthera, Stenodon,

Trembleya; miconieae: Medinella, Alloneuron,

Allomaieta, Anaectocalyx, Bellucia, Boerlagea,

Calycogonium, Catanthera, Catocoryne, Chalybea,

Charianthus, Clidemia, Conostegia, Creochiton,

Cyphostyla, Diplectria, Dissochaeta, Henriettea,

Henriettella, Huilaea, Killipia, Kendrickia,

Kirkbridea, Leandra, Llewelynia, Loreya, Macrolenes,

Maieta, Mecranium, Medinilla, Miconia, Myriaspora,

Myrmidone, Necramium, Ossaea, Pachyanthus,

Pachycentria, Pleiochiton, Pleithiandra,

Pogonanthera, Tetrazygia, Tococa; blakeeae:

Blakea, Topobea.

Closely related to the Memecylaceae.

10. LYTHRACEAE

Jaume Saint-Hilaire 1805 (including Ammaniaceae

Horaninow 1834, Blattiaceae Engler 1892,

Duabangaceae Takhtajan 1986, Lagerstroemiaceae J.

Agardh 1858, Lawsoniaceae J. Agardh 1858,

Punicaceae Horaninow 1834, Sonneratiaceae Engler

1897). 31/600. Subcosmopolitan, but mainly tropical

and subtropical regions, especially in America;

relatively few species (mostly annual or perennial

herbs) in temperate and cold areas.

10.1 LYTHROIDEAE

Fruits usually capsular, dehiscing variously or seldom

indehiscent. Seeds generally more or less numerous,

ﬂ attened, sometimes winged, nearly or quite without

endosperm, with straight oily embryo; cotyledons

folded. Mostly herbs, less often subshrubs, shrubs, or

sometimes trees; bark ﬂ aky, mucilage cells common.

Hairs uni- or bi-, sometimes multicellular. Branched

foliar sclereids mostly absent. Internal phloem usually

present. Leaves opposite, less often verticillate, rarely

alternate, commonly entire, ﬂ at to conduplicate,

stipules vestigial or none. Stomata mostly anomocytic.

Flowers bisexual, actinomorphic or sometimes more

or less zygomorphic, with conspicuous, sometimes

spurred ﬂ oral tube, usually (3)4-6(-8-16)-merous.

Sepals rather thick, valvate, appearing as valvate lobes

of the ﬂ oral tube, often alternating with external

appendages at the sinuses. Petals free, attached at the

summit or within the ﬂ oral tube, pinnately veined,

crumpled in bud, sometimes wanting. Stamens usually

twice as many as the sepals or petals, in two cycles,

rarely (Rotala) solitary or numerous and centrifugal as

in Lagerstroemia; ﬁ laments more or less elongate,

straight or incurved (the long ﬁ laments of Lafoensia

inrolled) in bud, usually inserted on the inside of the

ﬂ oral tube. Anthers versatile or seldom basiﬁ xed.

Pollen grains 3-colporate (some heterocolpate with

either three or six subsidiary colpi). Gynoecium of

2-4(-6) carpels; style ﬁ liform, commonly elongate,

sometimes bent in bud, mostly with capitate stigma;

ovary superior to inferior, often surrounded at the base

by an annular nectary disc (or the nectary unilateral),

multilocular with as many locules as carpels, but some-

times the septa not reaching the summit; rarely the

ovary pseudomonomerous; placentation axile or rarely

free-central. Ovules usually more or less numerous in

each locule, anatropous or very rarely amphitropous.

Superorder MYRTANAE 341

Often producing quinolizidine alkaloids. n = 5, 7, 8,

15, 24, 28. – lythreae: Rotala, Ammannia,

Hionanthera, Peplis, Didiplis, Lythrum, Woodfordia,

Cuphea, Pleurophora, Galpinia, Pemphis, Capuronia,

Diplusodon, Physocalymma, Lafoensia; nesaeeae:

Crenea, Nesaea, Heimia, Decodon, Pehria, Adenaria,

Koehneria, Lourtella, Tetrataxis, Ginoria (including,

Haitia); lagerstroemieae: Lagerstroemia (includ-

ing Orias), Lawsonia.

10.2 PUNICOIDEAE

Fruits with 2–3 layers of locules and a leathery pericarp

and persistent calyx. Seeds embedded in a pulpy mass;

endosperm wanting, embryo with spirally rolled, large

cotyledons. Small trees or shrubs with quadrangular

twigs, often thorny; scattered secretory cells in the cor-

tex and pith. Vessel ray and vessel-parenchyma pits

half-bordered. Fibers septate. Axial parenchyma scanty

paratracheal to virtually absent. Nodes unilacunar.

Leaves alternate to opposite, sometimes crowded at

the tips of the twigs, estipulate. Flowers solitary or in

fascicles terminating axillary branches, bisexual, acti-

nomorphic, with colored ﬂ oral tube prolonged well

beyond the ovary. Sepals 5–8, appearing as valvate

lobes on the ﬂ oral tube, persistent on the leathery

hypanthium. Petals imbricate and crumpled in bud.

Stamens numerous at the rim of the ﬂ oral tube, devel-

oping centrifugally; ﬁ laments slender; anthers dorsift-

xed, versatile. Pollen grains 3-colporate, with slightly

angular mesocolpia. Gynoecium of 7-9(-15) carpels.

Ovary inferior. Ovules more or less numerous on each

placenta. Placentation axile (Punica protopunica) or

the axile placentation at the base of the ovary differen-

tiates into two or three superposed upper layers that

appear to have parietal placentation as a result of asym-

metric growth. Ovules 20–50 per locule, anatropous.

Produing pyridine alkaloids an ellagic acid, n = 7 in

P. protopunica and 8 in P. granatum. 1/2. From the

Balkan Peninsula to the western Himalayas (Punica

granatum) and on Socotra (P. protopunica). – Punica

(including Socotria ?).

10.3 DUABANGOIDEAE

Fruits loculicidal capsules, coriaceous, more or less

perfectly 4–8-locular, seated on the spreading leathery

calyx. Seeds numerous, ellipsoid, testa prolonged at

both ends into longish tails; endosperm lacking.

Flowers in terminal umbelliform cymes, rather large,

4-merous (Duabanga moluccana) or 5–8-merous

(D. grandiﬂ ora). Petals shortly clawed, broadly

obovate, crisped, and undulate. Stamens 12 (D.

moluccana) or more than 50, biseriate (D. grandiﬂ ora),

inserted on the ﬂ oral tube in one or two cycles, inﬂ exed

in bud; anthers recurved or replicate over one end of

the connective. Pollen grains 3-porate. Gynoecium

of 4–8 carpels; style curved, with 4–8-lobed stigma;

ovary 4–8-locular. Ovules numerous, on diffuse

placentas, covering nearly the whole inner walls of

locules. Tall, buttressed trees with drooping, 4-an-

gled ultimate branches. Leaves opposite, glaucous

beneath, estipulate. Vessel ray and vessel-parenchyma

pits large and simple. Fibers not septate. Axial paren-

chyma paratracheal. n = 12. 1/2. Southeast Asia and

New Guinea. – Duabanga.

10.4 SONNERATIOIDEAE

Fruits indehiscent berries crowned by the style base.

Seeds embedded in evil-smelling pulp, not tailed at the

ends; embryo green, endosperm lacking. Flowers 1–3

together at the tips of the ultimate, mostly pendulous

branchlets. Petals linear or linear-lanceolate or quite

absent. Stamens numerous, inserted on the ﬂ oral tube

in several cycles; anthers reniform. Disc saucer-shaped.

Pollen grains 3-colporate. Gynoecium of 10–20 car-

pels; style with capitate stigma; ovary 10–20-locular.

Ovules many, ascending, on axile placentas. Trees

with trunk surrounded by aerial roots arising vertically

as lateral, negatively geotropic branches upon often

very long horizontal roots buried in the substratum.

Vessel ray and vessel-parenchyma pits mostly half-

bordered. Fibers septate. Axial parenchyma absent,

n = 12. 1/5. From coasts of East Africa and Madagascar

to Micronesia, New Hebrides, and the Solomons,

northern Australia and New Caledonia. – Sonneratia.

Lythraceae are a rather heterobathmic family

( primitive characters, such as superior ovary and

monosporic female gametophyte of Polygonum-type,

are combined with advanced characters, such as pin-

nate venation of the petals and the dominance of herbs).

Probably are nearest to the Crypteroniaceae and

Memecylaceae-Melastomataceae.

11. TRAPACEAE

Dumortier 1829. 1/3 or about 15. Tropical, subtropi-

cal, and temperate regions of the Old World, except

Australia.

342 Subclass VI. ROSIDAE

Trapa.

Related to the Onagraceae and especially to the

Lythraceae. From the Onagraceae they differ mark-

edly in embryological characters and the absence of

the viscin threads on the pollen grains. From the

Lyrthraceae they differ in semi-inferior ovary, almost

4-merous ﬂ owers, and the endosperm-nucleus degen-

erating soon after its formation. According to Miki

(1959), Trapa evolved from Lythrum through the

sintermediate fossil genus Hemitrapa.

12. ONAGRACEAE

A. L. de Jussieu 1789 (including Circaeaceae Ruthe

1827, Epilobiaceae Ventenat 1799, Fuchsiaceae Lilja

1870, Isnardiaceae Martynov 1820, Jussiaeaceae

Martynov 1820, Lopeziaceae Lilja 1870, Oenotheraceae

C. C. Robin 1807). 17/680. Widespread in tropical,

subtropical, and temperate regions, especially in the

New World, where they are centered in southwestern

North America.

12.1 JUSSIAEAOIDEAE

Flowers 4–5-merous, hypanthium absent. Nectary on

gynoecium; style short, stigma capitate. – jussiaeeae:

Ludwigia (including Jussiaea).

12.2 ONAGROIDEAE

Flowers 4-merous (2-merous in Circaea), hypanthium

long. Style short, stigma divided, dry, n = 9–11. – hau-

yeae: Hauya; fuchsieae: Fuchsia; circaeeae:

Circaea; lopezieae: Lopezia; gongylocarpeae:

Gongylocarpus; epilobieae: Chamerion, Epilobium

(including Boisduvalia, Chamaenerion and

Zauschneria); onagreae: Xylonagra, Clarkia,

Gayophytum, Camissonia, Oenothera, Calylophus,

Stenosiphon, Gaura, Megacorax.

A very distinctive family (Dahlgren and Thorne

1984) that reveals some similarities with the

Lythraceae s. str.

13. MYRTACEAE

A. L. de Jussieu 1789 (including Chamelauciaceae

F. Rudolphi 1830; Heteropyxidaceae Engler et Gilg

1920; Kaniaceae Nakai 1943; Leptospermaceae

F. Rudolphi 1830; Melaleucaceae Vest 1818;

Myrrhiniaceae Arnott 1839; Psiloxylaceae Croizat

1961). 131/4000–4600. Tropical and partly subtropical

and warm-temperate regions, mainly in Australia, but

also in South America.

Classiﬁ cation after P. G. Wilson et al. (2005).

13.1 PSILOXYLOIDEAE

Plants dioecious. Leaves alternate or pseudo- alternate

(disjunct-opposite), secretory cavities present (but

not containing essential oils in Psiloxylon). Stamens

not inﬂ exed in bud, anthers 4-locular. Stigma in

female ﬂ owers (but not in pistillodia) with ﬂ eshy,

ﬂ attened lobes or capitate. Female gametophyte of

Allium-type. Fruits punctate, many-seeded berries,

seed unwinged (Psiloxylea) or 2(3)-valved loculicidal

capsules, seed winged (Heteropyxideae), n = 12. –

psiloxylea: Psiloxylon; heteropyxideae:

Heteropyxis.

13.2 MYRTOIDEAE

Plants with bisexual ﬂ owers, rarely andromonoecious.

Leaves opposite or spirally arranged; secretory cavi-

ties present, containing essential oils. Stamens inﬂ exed

in bud, anthers 2-locular, terpenoid-containing glands

in the apex of the connective, stigma dry. Female

gametophyte of Polygonum

-type. Fruits indehiscent,

pericarp usually ﬂ eshy; seeds usually with sclerotic

testa. Polyhydroxyalkaloids present, n = 11. –xan-

thostemoneae: Xanthostemon, Pleurocalyptus,

Purpureostemon; lophostemoneae: Lophos temon,

Welchiodendron, Kjellbergiodendron, Whiteo dendron;

osbornieae: Osbornia; melaleuceae: Beaufortia,

Callistemon, Calotham nus, Eremaea, Lamarchea,

Melaleuca, Petraeomyrtus, Phymatocarpus, Regelia;

kanieae: Barongia, Basisperma, Kania, Lysicarpus,

Mitrantia, Ristantia, Sphaerantia, Tristaniopsis; back-

housieae: Backhousia, Choricarpia; metro sidereae:

Carpolepis, Mearnsia, Metrosideros, Tepualia; trist-

anieae: Tristania, Thaleropia, Xanthomyrtus; syzy-

gieae: Acmena, Acmenosperma, Anetholea, Syzygium

(including Aphanomyrtus), Waterhousea; myrteae:

Acca, Feijoa, Accara, Amomyrtella, Amomyrtus,

Archirhodomyrtus, Austromyrtus, Blephar oxalyx,

Calycolpus, Calycorectes, Calyptranthes, Campo-

manesia, Chamguava, Decaspermum, Eugenia

(including Monimiastrum), Gomidesia, Gossia,

Hexachlamys, Legrandia, Lithomyrtus, Lophomyrtus,

Luma, Marlierea, Meteo romyrtus, Mosiera,

Myrceugenia, Myrcia, Myrcianthes, Myrciaria,