Takhtajan Armen. Flowering Plants

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Superorder MAGNOLIANAE 41

parenchyma scanty paratracheal or diffuse. Rays het-

erogeneous with elongate ends. Fibers with simple or

slightly bordered pits, sometimes septate. Sieve ele-

ments long and slender, with slowly tapering ends;

sieve-element plastids of Pcs-type with several protein

crystals, one being rod-shaped, which is family spe-

ciﬁ c. Nodes multilacunar. Leaves alternate, distichous

(but spirally arranged in young plants), simple, entire,

pinnately veined, and without stipules. Prophylls in a

median position. Stomata paracytic or occasionally

actinocytic. Flowers showy, solitary, bisexual, without

perianth, enclosed when young in a deciduous calyp-

tra (a single bract, according to Baillon 1868; Eames

1961; Endress 1977) attached to the rim of an enlarged,

somewhat concave, turbinate receptacle and falling off

as a conical lid. Stamens 20–100, staminodia numer-

ous, inserted into the receptacle near its rim; the inner-

most stamens are sterile (staminodia), somewhat

ﬂ eshy, more or less petaloid, the others fertile, ribbon-

shaped, with a short, broad, laminar base. Anthers

long, basiﬁ xed, tetrasporangiate, latrorse-introrse,

opening longitudinally, with prolonged thickened con-

nective. Pollen grains globose-oblate, psilate, with

structureless, atectate-amorphous exine, with two par-

allel, equatorially encircling colpi that are differenti-

ated from the nonapertural areas of the exine by their

fossulate-rugulate surface. Gynoecium of numerous

(ca. 13–70) small carpels spirally arranged in the hol-

low receptacle, all appressed and connate, but each has

a decurrent stigma with projecting papillae and a slit-

like opening. Each carpel with two to several

(up to ca. 11) ovules arranged laminar-laterally. Ovules

anatropous with hood-shaped outer integument.

Endosperm cellular. Fruit an urceolate-turbinate berry

with a truncated, ﬂ attened, or convex apex. Seeds 1–3

in each carpel, subangular and coarsely pitted; seed

coat composed of a single-layered thin-walled exotesta

or large tanniniferous cells, but has 3–4 layers of lon-

gitudinal thin-walled ligniﬁ ed ﬁ bers in the mesotesta

and 2 layers of thin-walled parenchymatous endotesta;

embryo small, straight; endosperm copious, ruminate,

oily. Producing various types of lignans and alkaloids,

n = 10.. . . . . . . . . . . . . . . . . . . . . . .1. eupoma tia ceae

1 Vessels with simple perforations; lateral pitting alter-

nate or sometimes transitional to opposite. Evergreen

trees, shrubs, or woody lianas, bearing essential oils,

resinous or not resinous. Pith generally septate. Fibers

usually with numerous simple or narrowly bordered or

rarely distinctly bordered pits. Rays typically wide

and high, homogeneous or weakly heterogeneous,

occasionally distinctly heterogeneous. Axial paren-

chyma apotracheal or sometimes scanty paratracheal.

Sieve-element plastids of Psc- or (Xylopia) of Pcsf-

types. Nodes mostly trilacunar. Leaves entire, pin-

nately veined, often pellucid-dotted, often with a

pulvinus that is swollen and elongate or contorted for

climbing, without stipules. Stomata nearly always

paracytic. Prophylls in some genera – e.g., in Annona

and Asimina – paired and lateral (as usual in dicotyle-

dons), but usually they are solitary and median, in dor-

sal position, as in the monocotyledons. Flowers in

various types of basically cymose inﬂ orescences or

solitary in the axils, mostly entomophilous, bisexual

or rarely unisexual, with cyclic and usually 3-merous

perianth. Sepals (2)3(4), imbricate or valvate, free or

partly connate. Petals 3-6(-12), usually 6 in 2 cycles of

3, free or very rarely basally connate, imbricate or val-

vate. Stamens numerous (up to 100) and spirally

arranged or rarely 3 or 6 (up to 15) and cyclic, gener-

ally free, rarely basally connate, ﬁ laments short and

stout, with single vascular trace. Anthers linear; con-

nective prolonged and often expanded above the

anther so that the anther appears to be peltate.

Staminodia present only in a few genera. Pollen grains

very diverse in size and structure, monocolpate to

inaperturate, inconspicuously granular to columellate,

atectate, tectate-imperforate, tectate-perforate, semi-

tectate, or intectate. Gynoecium of more or less numer-

ous, several or rarely only one carpel, spiral and

apocar pous or, less frequently, cyclic and paracarpous

(Monodoroideae). Carpels 1–400 (ca. 400 in Annona

muricata – Igersheim and Endress 1997), condupli-

cate, sometimes unsealed, whorled or irregularly

arranged, free in most genera, with 1–30 (up to 60 or

more in Monodora myristica) ovules on submarginal,

laminar-lateral, parietal or nearly basal placentas; sty-

lodia free, short, and thick. Ovary superior, 1-locular,

or 2–15-locular. Ovules 1–10, ascending, with hood-

shaped outer integument. Fruits of various types, but

mostly more or less ﬂ eshy, distinct, stipitate, indehis-

cent, or, rarely (Anaxagorea), dehiscent fruitlets. Seeds

with small embryo and copious, and ruminate, oily

endosperm and seldom with perisperm; n = 7, 8, 9,

mostly 8.. . . . . . . . . . . . . . . . . . . . . . . 2. annona ceae

1. EUPOMATIACEAE

Endlicher 1841. 1/3 Eastern and southeastern

Australia (from temperate Victoria to subtropical

42 Subclass I. MAGNOLIIDAE

New South Wales and tropical Queensland) and New

Guinea (from sea level up to 1,300 m altitude).

Eupomatia

The Eupomatiaceae have some features in common

with both the Himandandarceae and Annonaceae but

differ from them in many respects, including complete

absence of perianth, circular aperture of pollen grains,

and the basic chromosome number.

2. ANNONACEAE

A.L. de Jussieu 1789 (including Hornschuchiaceae J.

Agardh 1858; Monodoraceae J. Agardh 1858).

128/2400. Except for the temperate North American

genus Asimina, the family has a pantropical distribu-

tion, but is concentrated in the Paleotropics. Low and

more stable chromosome numbers dominate in the

Paleotropics; higher and more diversiﬁ ed ones in

the Neotropics (Morawetz 1988).

2.1 ANNONOIDEAE

Carpels spirally arranged (rarely few in I cycle), free

from one another from the beginning (rarely only 1), or

if at length united, then forming a many-locular syn-

carp. Stigmas more or less erect, rarely radiating. –

uvarieae: Uvaria, Balonga, Tetrapetalum, Ellipeia,

Ellipeiopsis, Sapranthus, Stenanona, Afroguatteria,

Rauwenhofﬁ a, Sageraea, Dendrokingstonia, Stelecho-

carpus, Dasoclema, Hexalobus, Cleistopholis, Green-

way odendron, Mkilua, Toussaintia, Enicosanthum,

Enicosanthellum; miliusieae: Fenerivia, Hetero-

petalum, Neostenanthera, Boutiquea, Marsypopetalum,

Meiogyne, Phaeanthus, Trivalvaria, Anomianthus,

Piptostigma, Cymbopetalum, Miliusa, Mezzettiopsis,

Orophea, Phoenicanthus, Alphonsea, Platymitra,

Mezzettia; xylopieae: Xylopia, Cardiopetalum,

Meiocarpidium, Polyceratocarpus, Dielsiothamnus,

Drepananthus, Anaxagorea, Artabotrys, Pseudo-

artabotrys, Cyathocalyx, Diclinanona; annoneae:

Desmos, Dasymaschalon, Cyathostemma, Polyalthia,

Polyaulax, Oncodostigma, Monocarpia, Exellia,

Unonopsis, Uvariodendron, Uvariastrum, Dennettia,

Asteranthe, Desmopsis, Guamia, Haplostichanthus,

Monocyclanthus, Chieniodendron, Enantia, Woodiel-

lantha, Cleistochlamys, Disepalum, Pseuduvaria,

Mitre phora, Goniothalamus, Richella, Schefferomitra,

Melodorum, Friesodielsia, Oreomitra, Petalolophus,

Popowia, Neo-uvaria, Papualthia, Mitrella, Pyramid-

anthe, Fissistigma, Cananga, Bocageopsis, Onychopetalum,

Monanthotaxis, Mischogyne, Atopostema, Gilbertiella,

Uvariopsis, Ambavia, Bocagea, Ophrypetalum, Trigy-

naea, Porcelia, Hornschuchia, Froesiodendron, Oxandra,

Pseudoxandra, Crematosperma, Ephedranthus, Ruizo-

dendrom, Malmea, Guatteria, Guatteriella, Guatter-

iopsis, Asimina, Deeringothamnus, Fitzalania, Tridi meris,

Lettowianthus, Annona, Anonidium, Raimondia,

Rollinia, Rolliniopsis, Pachypodanthium, Lestestudoxa,

Duckeanthus, Duguetia, Fusaea; Tetramerantheae:

Tetrameranthus.

2.2 MONODOROIDEAE

Carpels cyclically arranged, united from the beginning

into I-locular ovary with parietal placentas. Tropical

Africa and Madagascar. – Monodora, Isolona.

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nal) basic chromosome number of angiosperms. Taxon 21:

57–65.

Walker JW. 1976. Evolutionary signiﬁ cance of the exine in the

pollen of primitive angiosperms. In: K Ferguson and J Müller,

eds. The evolutionary signiﬁ cance of the exine, pp. 251–308.

Linn. Soc. Symposium, No. 1. Academic Press, London/

New York.

Woodland PS and PR Garlick. 1982. The ﬁ ne structure of the

pollen of Eupomatiaceae. Austral. J. Bot. 30: 297–301.

Woodland PS and N Prakash. 1993. Floral morphology, embry-

ology and relationships of the Eupomatiaceae. Abstr. XV

Inter. Bot. Congr., p. 95. Yokohama.

Young DA. 1983. Leaf ﬂ avonoids of Eupomatiaceae. Biochem.

Syst. Ecol. 11(3): 209–210.

Order 12. MYRISTICALES

Trees or seldom shrubs, usually aromatic and resinous.

Trichomes of various kinds, eglandular ones elongate,

unicellular and colourless, glandular ones with a multi-

cellular, basally embedded stalk and a multicellular

peltate, later balloon-shaped head with golden-yellow

content. Nodes trilacunar with three traces. Pith sep-

tate. Vessels typically with both simple and scalariform

perforations, ranging from mostly simple to all scalari-

form, with occasional reticulate perforations in most

genera; scalariform plates moderately oblique and usu-

ally with 1–10 (occasionally up to 20) broad bars; lat-

eral pitting from scalariform to alternate. Fibers with

simple or sometimes narrowly bordered pits. Rays uni-

seriate or biseriate, heterogeneous. Axial parenchyma

apotracheal banded and diffuse to paratracheal. In the

axial parenchyma and the rays as well as in the phloem

there are tanniniferous tubes containing pale or red

resin. Sieve-element plastids of S-type and Psc-type,

the latter with tiny protein crystals. Leaves alternate,

often distichous, sometimes falsely verticillate, simple,

entire, pinnately veined, occasionally pellucid-dotted,

estipulate. Stomata paracytic. Flowers rather small, in

axillary or rarely terminal, cymose or racemose inﬂ o-

rescences, unisexual, dioecious or rarely (Endocomia)

monoecious, actinomorphic, apetalous, basically

3-merous. Calyx cupulate or campanulate, (2)3(-5)-

lobed, the lobes valvate. Male ﬂ owers lacking even a

vestigial gynoecium; stamens 2–45, with ﬁ laments

partially or completely fused; anthers free or more

often laterally connate, tetrasporangiate, extrorse or

seldom latrorse, opening longitudinally, mutinous or

shortly apiculate. Tapetum secretory. Microsporogenesis

successive. Pollen grains 2-celled, boat-shaped to sub-

globose, 1-colpate to inaperturate, tectate to intectate,

granular to columellate. Female ﬂ owers without sta-

minodia; gynoecium of a single conduplicate, unsealed,

sometimes short stipitate carpel with solitary, subbasal

to basal ovule; stigma sessile or subsessile or rarely

(Brocboneura) on a long stylodium, small, simple, or

more or less 2-lobed. Ovule solitary, anatropous or

rarely hemitropous or suborthotropous, bitegmic,

crassinucellate. Female gametophyte of Polygonum-

type. Endosperm nuclear. Fruits ﬂ eshy to coriaceous or

ligneous capsule, generally dehiscent both ventrally

and dorsally. Seeds usually ellipsoid to subglobose,

1–5 cm long, covered mostly with crustaceous to ﬂ eshy,

laciniate or entire funicular aril; seed coat formed by

both integuments, generally multiplicative, with well-

developed vascular system and has the most elaborate

construction among the ﬂ owering plants; inner epider-

mis of the outer integument strongly developed, trans-

formed into the palisade mechanical layer of the testa.

Derivatives of the inner integument transformed into

the complex tegmen that is also vascularized and par-

ticipates in the rumination of the endosperm; chalaza

large; endosperm copious, ruminate, oily, and also con-

tains large protein crystals and starch grains; embryo

small, straight, nearly basal, differentiated into very

short radicle, weakly developed hypocotyl and partly

or completely connate cotyledons. Produce essential

oils, ﬂ avonoids, legnans, polyketides (acetogenins),

tryptamine alkaloids. n = 19, 21, 22, 25, 26.

The Myristicaceae deﬁ nitely belong to the

Magnolianae, but differ markedly from all of the other

families in their very elaborate seed structure and tan-

niniferous tubes in rays, axial parenchyma, and phloem.

The seed structure somewhat resembles the Aristo-

lochiaceae, but the latter has starch-free and not rumi-

nate endosperm and lacks an aril. The Myristicaceae

resemble Annonaceae in their ruminate endosperm.

According to Hegnauer (2000) Myristicaceae resemble

Lauraceae in their seed fats and in their secondary

metabolism, in their producing of noelignans and lign-

ans, and of polyketides.

1. MYRISTICACEAE

R. Brown 1810. 21/500. Widely distributed in tropical

countries, especially in Asia.

46 Subclass I. MAGNOLIIDAE

Mauloutchia, Brochoneura, Cephalosphaera,

Haematodendron, Scyphocephalium, Pycnanthus,

Staudtia, Coelocaryon, Compsoneura, Osteophloeum,

Otoba, Virola, Bicuiba, Iryanthera, Knema, Horsﬁ eldia,

Endocomia, Gymnacranthera, Myristica, Paramy-

ristica, Doyleanthus.

Bibliography

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Myristica fragrans Houtt.: The nutmeg of commerce.

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Armstrong JE and AK Irvine. 1989. Floral biology of Myristica

insipida (Myristicaceae): A distinctive beetle pollination

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Armstrong JE and SC Tucker. 1986. Floral development in

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Armstrong JE and TK Wilson. 1978. Floral morphology of

Horsﬁ eldia. Am. J. Bot. 65: 441–449.

Baas P. 2000. Wood anatomy of Myristicaceae. In: PF Stevens,

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Suborder LAURANAE 47

Suborder LAURANAE

Order 13. LAURALES

Trees or shrubs, sometimes woody lianas, rarely twin-

ing parasitic herbs (Cassytha), usually with oil or muci-

lage cells in the parenchymatous tissues. Nodes

unilacunar, with 1–7 traces. Xylem usually with well-

developed vessels. Vessels with scalariform or simple

perforations; lateral pitting scalariform to alternate or

mostly alternate. Fibers ranging from tracheids through

ﬁ ber-tracheids to libriform-ﬁ bers. Rays mostly hetero-

geneous. Axial parenchyma apotracheal or paratra-

cheal, rarely absent. Sieve-element plastids of S-, Psc-,

or Pest-type. Leaves alternate or opposite, sometimes

verticillate, simple, entire or sometimes lobed or den-

tate, pinnately or rarely palmately compound, without

stipules. Stomata paracytic or less often anomocytic.

Flowers usually in cymose or racemose inﬂ orescences,

rarely solitary, actinomorphic or very rarely oblique,

spiral, spirocyclic or more often cyclic, bisexual or uni-

sexual, mostly with more or less developed and usually

cup-shaped hypanthium (ﬂ oral cup) of receptacular

origin. Perianth spiral or cyclic (often 3-merous), usu-

ally not differentiated into sepals and petals. Androecium

of (3-) 5 to many stamens, cyclic or spiral; stamens

more or less ribbon-shaped or much more often differ-

entiated into ﬁ lament and anther; ﬁ laments frequently

with two lateral glands; anthers tetrasporangiate or dis-

porangiate, opening longitudinally or more often by

valves, rarely by pores or very rarely circumscissile

(Hennecartia). Tapetum secretory or amoeboid.

Microsporogenesis successive or simultaneous

(Calycanthaceae). Pollen grains 2-celled or rarely

3-celled, mostly inaperturate, frequently 2-colpate,

rarely with 2–12 slightly irregular porate apertures,

mostly tectate. Gynoecium apocarpous, spiral

(Gomortegaceae, Calycanthaceae and some

Monimiaceae) or whorled (polymerous or more often

monomerous) or less frequently syncarpous; carpels

1–2,000, glabrous or rarely hairy, fully sealed and with

a localized stigma on a distinct stylodium or rarely

unsealed above and with shortly decurrent stigma.

Ovule solitary in each carpel or in each locule, anatro-

pous, rarely orthotropous or hemitropous, bitegmic or

rarely unitegmic, crassinucellate. Female gametophyte

usually of Polygonum-type. Endosperm cellular or

nuclear (Lauraceae-Lauroideae). Fruits of various

types. Seeds with endotestal seed coat, embryo small to

large; endosperm well developed, scanty or absent.

Produces proanthocyanins but not ellagic acid, and very

often with benzyl isoquinoline or aporphine alkaloids.

Close to the Magnoliales but more advanced.

Evidently derived from some vesselless ancestors.

Key to Families

1 Seeds with well-developed endosperm and small or

medium sized embryo.

2 Gynoecium apocarpous.

3 Anthers opening by slits. Trees or shrubs,

rarely woody lianas. Spherical ethereal cells

present; mucilage cells absent. Nodes unilacu-

nar with 1–7 traces. Nodes unilacunar with

1–7 traces. Vessels with simple perforation.

Sieve-element plastids of S-, Psc-, or Pcsf-

type. Leaves alternate or opposite. Stomata

paracytic or anomocytic. Flowers in cymose

inﬂ orescences or rarely solitary, unisexual in

most genera, bisexual in Hortonia, actinomor-

phic or very rarely oblique. Receptacle trans-

formed into more or less well- developed ﬂ oral

cup, more or less globose or urceolate to

widely campanulate. Perianth segments spiral

or in cycles, often 4-merous or of decussate

pairs, sometimes absent. Stamens usually

numerous and dispersed over the inner surface

of the receptacle or fewer and sometimes in

one or more 4- merous cycles; ﬁ laments in

Hortonia, Peumus, and Monimia with two

nectariferous glands; anthers tetrasporangiate

or disporangiate, opening longitudinally or by

valves. Pollen grains inaperturate or dicolpate.

Carpels 1–2,000 (1 in Xymalos, 1–2 in

Hennecartia, 1,000 in Decaryodendron and

up to 2,000 in Tambourissa ﬁ cus, the largest in

Magnoliidae), small, whorled or spiral or

irregular, free (in Tambourissa congenitally

united and congenitally immersed in the ﬂ oral

cup (Endress et al. 1997); the carpel is covered

with unicellular, often ligniﬁ ed or stellate

hairs, or glabrous; ovary superior (inferior in

Tambourissa – Monimiaceae); stylodia short

or elongate, with terminal stigma. Ovules soli-

tary, anatropous or (Kibara, Kibaropsis,

Xymalos) hemitropous, erect or pendulous,

bitegmic. Fruits drupes. Seeds endotestal,

the tegmen eventually crushed; endosperm

48 Subclass I. MAGNOLIIDAE

copious, oily; embryo very small to moder-

ately large, straight, differentiated, with two or

very rarely four (Kibaropsis) cotyledons.

Hippocrepiform sclereids present. Benzyl

isoquinolines present. n = 19–22, 39, 43, 57,

etc.. . . . . . . . . . . . . . . . . . . . 1. monimiaceae.

3 Anthers opening by valves.

4 Fruits of long-stylodiated plumose achenes.

Trees or shrubs. Nodes with 1 trace. Sieve-

tube plastids P-type. Leaves opposite, petio-

late, gland-dotted or not, simple, entire or

dentate, estipulate. Stomata anomocytic.

Flowers solitary or in racemes or cymes,

bisexual or monoecious or polygamo-dioe-

cious, actimo norphic or somewhat zygomor-

phic. Perianth segments with distinct calyx

and corolla, or sepaloid, or vestigial, or absent.

Corolla when present 7–20, or more. Stamens

(4-)6-12(-100). Filaments appendaged, each

with a pair of glandular scales at the base.

Anthers adnate, extrorse, disporangiate, open-

ing by two valves. Pollen grains reticulate.

Gynoecium of 3–100 carpels with a lateral

style, or with a gynobasic style. Ovary supe-

rior to inferior; ovule 1, anatropous, or rarely

orthotropous, micropyle endostomal, directed

downward. Fruits plumose, with woody

hypanthium. Seeds with small, well differenti-

ated, straight embryo; endosperm oily.

Flavonols present; kaempferol and quercetin,

n = 22, 57.. . . . . . . 4. atherospermataceae.

4 Fruitlets drupes. Shrubs or small trees bearing

essential oils. Nodes with one trace. Vessels

with simple or scalariform and simple perfora-

tions. Xylem without ﬁ bre tracheids. Axial

parenchyma apotracheal. Leaves evergreen,

opposite or verticillate, simple, entire, gland-

dotted, or not gland-dotted, estipulate. Stomata

paracytic. Flowers in cymes, spikes, or in pan-

icles, monoecious or dioecious, actinomorphic

to zygomorphic. Free hypanthium present.

Perianth with distinct calyx and corolla, peri-

anth segments 4-6(-8), or obscure, calyptrate.

Stamens (1)2-100, ﬁ laments free, without nec-

tariferous appendages. Anthers often with one

ﬂ ap, introrse, bisporangiate (Siparuna), or tet-

rasporangiate (Glossocalyx). Pollen grains

inaperturate. Gynoecium of (3-)4-100 carpels;

style free or connate at the apex; ovule solitary

in each carpel, ascending, anatropous, uniteg-

mic or bitegmic (Glossocalyx – Philipson

1993). Fruits druplets, with ﬂ eshy appendage,

enclosed in the ﬂ eshy hypanthium. Seeds

erect, endosperm not oily. Plants accumulate

Aluminum; n = 22. . . . . . . . 5. siparunaceae

2 Gynoecium syncarpous. Aromatic trees with

secretory cells containing a yellowish, resinous

substance. Nodes with two traces. Vessels with

scalariform perforations with 9–19 bars. Sieve-

element plastids of Pcsf-type. Leaves opposite,

decussate, simple, entire, pinnately veined, estip-

ulate. Stomata paracytic. Flowers rather small, in

axillary or terminal racemes, subtended by two

inconspicuous, caducous bracteoles, bisexual.

Perianth segments 7–10, more or less spirally

arranged, intergrading with the stamens. Stamens

7–13, the outer 1–3 petaloid, the next 5–10 dif-

ferentiated into ﬁ lament and anther, basally pro-

vided with two short-stalked nectariferous

glands; anthers disporangiate, the outer stamens

introrse, the middle ones latrorse, opening from

the base upward by two valves. Pollen grains

globose, inaperturate, tectate, with a delicate

exine and a thicker intine. Gynoecium of (2)3(-5)

glabrous carpels; style very short, with 2(3) stig-

matic branches; ovary deeply inferior,

(2)3(-5)-locular, each locule with a single ortho-

tropous to hemianatropous (Endress et al. 1997)

ovule pendulous from the apex. Fruits drupa-

ceous, mostly one-seeded. Seeds thin-walled;

endosperm copious, oily; embryo rather large, well

differentiated, n = 21. . . . . 6. gomortegaceae.

1 Seeds without endosperm and with large or medium-

sized embryo.

5 Leaves alternate.

6 Ovary superior, sometimes semi-inferior, very

rarely inferior (Hypodaphnis). Aromatic trees or

shrubs or rarely (Cassytha) twining parasitic

herb. Vessels mostly with simple perforations,

but simple and scalariform in some genera and

sometimes scalariform (commonly with few

bars, rarely up to 25 in some species of

Actinodaphne, Aiouea, and Litsea), lateral pit-

ting alternate. Fibers predominantly libriform.

Rays heterogeneous to homogeneous, in

Hypodaphnis exclusively homogeneous (com-

posed solely of procumbent cells). Axial paren-

chyma basically paratracheal. Oil cells and

Suborder LAURANAE 49

mucilage cells present. Nodes unilacunar, with

1–3 traces. Leaves alternate to opposite or appar-

ently verticillate, simple, entire or sometimes

(Sassafras) lobed, often coriaceous, gland-dot-

ted, aromatic; venation pinnate or rarely pal-

mate. Stomata paracytic or sometimes

anomocytic. Flowers usually in axillary and

commonly cymose inﬂ orescences, rarely soli-

tary, bisexual or less often unisexual, generally

actinomorphic, mostly 3-merous. Perianth seg-

ments commonly six and mostly in two usually

equal cycles. Stamens usually in four cycles of

three, of which the innermost cycle is sterile or

often lacking; sometimes also one or two of the

outer three cycles sterile or absent; usually the

third cycle with a pair of nectariferous glands at

the base; increased number of stamens are

found in some genera (up to 32 in Cinnadenia);

anthers 2- or 4-locular, basiﬁ xed, tetrasporan-

giate or disporangiate, opening by two or four

valves, or rarely (Hexapora) by pores, mostly

from the base upward, in the two outer cycles

mostly introrse and in the third cycle introrse to

extrorse. Pollen grains 2-celled, more or less

spheroidal, inaperturate, with rather thick intine

and extremely thin and entirely ectexinous

exine, mostly echinate. Gynoecium monocarpe-

late. The stylodium often shows a deep furrow

at the ventral side; stigma papillate, capitate to

disciform, lobed, or decurrent. Ovule solitary,

large to very large, anatropous, pendulous, api-

cal or subapical, with dorsal raphe; outer integ-

ument not contributing to the micropyle;

endosperm haustoria present or absent. Fruits

baccate (drupes with weakly developed endo-

carp), 1-seeded; fruit enclosed in the ﬂ eshy

receptacle, or enclosed in the ﬂ eshy hypan-

thium, or without ﬂ eshy investment. Seed with

thin testa; embryo well differentiated, straight,

with very large, occasionally ruminate cotyle-

dons; endosperm reduced. Plants producing ﬂ a-

vones, 5-O-methyl ﬂ avonols, polyketides

(acetogenins), tryptamine alkaloids, kaempferol

and quercetin. n = 12(15). . . . . . 8. lauraceae

6 Ovary inferior. Fruits dry and indehiscent.

Trees, shrubs, or woody lianas. Vessels with

simple perforations; lateral pitting mostly alter-

nate. Fibers with simple or bordered pits. Rays

heterogeneous to homogeneous. Axial paren-

chyma usually para-tracheal. Nodes unilacu-

nar, with several traces. Leaves simple

(sometimes 3-lobed) or palmately compound,

pinnately, palmately, or pedately veined; glan-

dular hairs in leaf epidermis. Stomata paracytic

or anomocytic. Flowers in cymose inﬂ ores-

cences, bisexual or unisexual, more or less acti-

nomorphic. Perianth segments in two cycles of

3-4(-6) or in one cycle of 4–8, imbricate or val-

vate. Stamens 3-5(-7) in a single cycle; ﬁ la-

ments commonly with a pair of dorso-basal or

basilateral nectariferous glands; anthers dispo-

rangiate, opening by valves. Pollen grains

spheroidal, inaperturate, with massive intine

composed of unstratiﬁ ed inner and thick, radi-

ally channeled outer layer, and very thin exine

ornamented by spines and globules. Gynoecium

monocarpellate; stylodium with a ventral fur-

row decurrent from the peltate stigma. Ovule

solitary, large to very large, anatropous, pendu-

lous. Seeds ruminate (Hernandia), testa vascu-

larised, spongy, tanniniferous; n = 10, 12, 15,

20.. . . . . . . . . . . . . . . . . . .7. hernandiaceae.

5 Leaves opposite. Fibers with minute bordered

pits. Rays predominantly uniseriate or predomi-

nantly multiseriate. Sieve-element plastids

P-type. Leaves opposite, simple, entire, pinnately

veined, without stipules. Stomata paracytic.

Stamens spirally arranged at the rim of the recep-

tacle, usually more or less ribbon-shaped, with

short or no ﬁ lament, with prolonged connective,

the inner ones staminodial; anthers extrorse. Pollen

grains more or less globose, with two distal colpi,

tectate-columellate, psilate. Carpels with usually

elongate stylodium; ovules solitary, basal or two

superposed ovules, the upper one abortive. Fruits

usually of numerous achenes enclosed in the

enlarged, ﬂ eshy receptacle. Very rich in common

mono- and sesquiterpenes. Characterized by the

presence of calycanthidine and calycanthine types

of alkaloids.

7 Carpels 5–35; cotyledons 2, spirally twisted.

Deciduous or sometimes evergreen small trees

or shrubs. Nodes with two traces. Vessels with

simple perforation. Parenchyma scanty vasi-

centric with some diffuse. Flowers, bisexual,

solitary at the ends of specialized short leafy

branches. Perianth sequentially intergrading

from sepals to petals; perianth segments 15–30,

50 Subclass I. MAGNOLIIDAE

free. Stamens 5–30. Anthers adnate, extrorse,

tetrasporangiate, appendaged. Pollen grains

aperturate; 2(–3) colpate, sulculate, 2-celled.

Ovules two, ascending, anatropous or apotro-

pous, crassinucellate or subcrassinucellate. Fruits

achenes, enclosed in the ﬂ eshy hypanthium;

seeds with well differentiated, large embryo;

endosperm wanting. Producing tryptamine

(calycanthidine) and benzylisoquinoline alka-

loids; n = 11. . . . . . . . . . . 3. calycanthaceae

7 Carpels 1 or 2(3); cotyledons 3 or 4, ﬂ eshy,

peltate. Evergreen trees. Vessels with scalari-

form perforation. Axial parenchyma abundant,

predominantly apotracheal. Flowers, bisexual

and male, solitary (3) on bracteate, axillary

peduncles, or in few-ﬂ owered terminal inﬂ o-

rescences. Perianth sequentially intergrading

from sepals to petals, or petaloid; perianth

segments 30–40, free. Stamens 13–15. Anthers

adnate, extrorse, appendages. Pollen grains 2

colpate. Gynoecium of 1 or 2(3) carpels.

Ovules 1–2, anatropous. Stigma multiseriate-

multicellular-papillate, ﬂ eshy, subsessile,

broad, obliquely terminal. Fruits strictly non-

ﬂ eshy, but enclosed in the ﬂ eshy receptacle.

Seeds without endosperm. Embryo with three

or more often four massive, ﬂ eshy-ﬁ rm, coty-

ledons attached in a single whorl. Producing

luteolin; n = 11.. . . . . . . . 2. idiospermaceae

1. MONIMIACEAE

A. L. de Jussieu 1809 (Hortoniaceae A. C. Smith

1971). 23/200. Tropical and subtropical regions,

especially of Southern Hemisphere, where they reach

temperate zones; concentrated in Malesia, in the

islands of the southwestern Indian Ocean, including

Madagascar, and in South America; poorly represented

in Africa and absent in India.

1.1 HORTONIOIDEAE

Stomata paracytic. Flowers bisexual. Receptacle con-

cave. Perianth spiral, with no clear distinction between

outer sepaloid and inner petaloid members. Stamens

with two appendages on the ﬁ lament. Anthers tetraspo-

rangiate, opening longitudinally. Staminodia present.

Pollen grains inaperturate, with semihelical bands.

Carpels free. Ovules bitegmic, the micropyle directed

upward. Hippocrepiform sclereids present. – Hortonia

(3, Sri Lanka).

1.2 MOLLINEDIOIDEAE

Stomata paracytic. Flowers unisexual (monoecious or

dioecious). Receptacle ﬂ at, concave, urceolate or glo-

bose, sometimes pollinated at the upper margin of the

globose ﬂ oral cup, at a “hyperstigma” (Endress 1979,

1980b, 1983). Perianth segments sepaloid or absent.

Stamens without appendages on the ﬁ laments. Anthers

tetrasporangiate, opening longitudinally or rarely cir-

cumscissile (Hennecartia). Staminodia absent in female

ﬂ owers, but sometimes the inner stamens sterile in male

ﬂ owers. Pollen grains inaperturate, rarely in tetrads or

dyads. Carpels free, or inferior in Tambourissa, solitary

in Xymalos. Stylodia subulate or stigmas sessile. Ovules

bitegmic, with the micropyle directed upward (sideways

in Kibaropsis). – hedycaryeae: Decarydendron,

Ephippiandra, Hedycarya, Kibaropsis, Levieria,

Tambourissa, Xymalos; mollinedieae: Austromatthaea,

Faika, Kairoa, Kibara, Macropeplus, Matthaea,

Mollinedia, Parakibara, Steganthera, Tetrasynandra,

Wilkiea; hennecartieae: Hennecartia.

1.3 MONIMIOIDEAE

Stomata anomocytic. Flowers unisexual (dioecious).

Receptacle a shallow cup or urceolate. Perianth seg-

ments sepaloid. Filaments appendaged (except in

Palmeria). Anthers tetrasporangiate or disporangiate,

opening longitudinally. Staminodia absent. Pollen

grains inaperturate. Carpels free, stylodia elongate,

projecting through the ostiole. Ovules bitegmic, the

micropyle directed upward. – palmeryeae: Palmeria;

monimieae: Monimia; peumeae: Peumus.

The most archaic member of the family is the genus

Hortonia. The Monimiaceae are divided into a number

of well-deﬁ ned subfamilies (Schodde 1970; Thorne

1974, 1992, 2000).

2. IDIOSPERMACEAE

S. T. Blake 1972. 1/1. Northeastern Queensland.

Idiospermum.

Very closely related to the Calycanthaceae, but dis-

tinctive nature of the ﬂ avonoids of Idiospermum,

together with the morphological and anatomical differ-

ences between Calycanthaceae and Idiospermum

sup-

port removing Idiospermum from the Calycanthaceae