Takhtajan Armen. Flowering Plants

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A. Takhtajan, Flowering Plants, 101

Trees or shrubs, very rarely suffruticose herbs

(Pachysandra). Vessels mostly with scalariform perfo-

rations. Sieve-element plastids of S-type or rarely

(Buxales) of PCS- or Pc-type. Leaves alternate or

sometimes opposite or verticillate, simple or less often

pinnately compound. Stomata of various kinds.

Flowers mostly small, in various kinds of inﬂ ores-

cences, cyclic, bisexual or unisexual, commonly apeta-

lous and often wholly without perianth, typically

anemophilous. Stamens (1)2-several or sometimes

numerous, often with a prolonged connective, free,

mostly with elongate ﬁ laments. Pollen grains mostly

2-celled, 3-colpate to often porate. Gynoecium apocar-

pous or more often syncarpous, sometimes pseudo-

monomerous. Ovules anatropous to orthotropous,

bitegmic or less often unitegmic, crassinucellate.

Endosperm cellular or nuclear. Fruits dry (dehiscent or

indehiscent) or drupaceous. Seeds with small (some-

times very small) or more often well-developed

embryo; endosperm present or absent. Tannins, proan-

thocyanidins, ellagic acid, and especially myricetin

ﬂ avonoids are common; iridoids are very rare and

occur only in Liquidambar, Daphniphyllum, and

Didymeles (Giannasi 1986).

The subclass Hamamelididae represents a very

ancient line and one of the main branches of the phylo-

genetic tree of magnoliopsids. The lower Hamame-

lididae are linked to Magnoliidae via Trochodendrales

(Takhtajan 1966, 1987; Endress 1986, 1993). There

are also some links between the lower Hamamelididae

and the lower Dilleniidae and Rosidae (see Ehrendorfer

1989; Dickison 1989). However, higher members of

the subclass are so highly specialized that they show

no evident links with either the Dilleniidae or

Rosidae.

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Subclass III. HAMAMELIDIDAE

102 Subclass III. HAMAMELIDIDAE

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63–76.

Superorder TROCHODENDRANAE

Order 30. TROCHODENDRALES

Evergreen or deciduous trees. Elongate, often branched

idioblasts occur in the leaves and in at least some of

the parenchymatous tissues of the stem. Secondary

Superorder TROCHODENDRANAE 103

xylem mostly of tracheids and only partly of primitive

vessels with scalariform perforations with many bars

(Snigirevskaya, 2000). Tracheids very long, in early

wood with boarded scalariform pits. In late wood,

where tracheids are narrower, they have rounded

boarded pits. Rays heterogeneous. Axial parenchyma

diffuse and in short tangential lines. Nodes multilacu-

nar to unilacunar (Trochodendron) or trilacunar

(Tetracentron). Leaves alternate, simple, margin ser-

rate or serrulate, estipulate (Trocho dendraceae) or

with stipules adnate to slender petioles (Tetra-

centraceae). Stomata encyclocytic. Flowers small, in

racemelike aggregates or spikes, bisexual or polyga-

mous, actinomorphic or slightly disymmetric, apeta-

lous. Sepals present or absent. Stamens about 40–70

or only 4; anthers basiﬁ xed, tetrasporangiate, more or

less latrorse, opening by longitudinal slits extending

into two transversal lines at both ends, which results in

two valves resembling transparent wings (Endress

1986). Tapetum secretory. Micro sporogenesis simul-

taneous. Pollen grains 2-celled, small, spheroidal,

3-colpate, tectate-columellate, rugulate-reticulate

between the apertures but striate near the apertures,

the apertures coarsely granulate. Gynoecium of 4–17

laterally connate, conduplicate carpels. Stylodia free,

with decurrent stigma, which bears unicellular papil-

lae. Ovary semiinferior; dorsal part of ovary horizon-

tally extended at anthesis, with many sunken stomata

and nectariferous. Ovules 5–30 per carpel, pendulous,

anatropous, apotropous, bitegmic, with long integu-

ments and a chalazal protrusion, crassinucellate;

micropyle endostomal. Female gametophyte of

Polygonum-type. Endosperm cellular or nuclear.

Fruits syncarpous multifollicles (follicetum), ventri-

cidal and slightly loculicidal. Seeds small, ﬂ at, with

chalazal, apical, and lateral winglike extensions. The

raphe is in the form of a ridge along the whole length

of the seed, its vascular bundle forming a hairpin loop

in the chalazal extension and terminating below the

hypostase. Seed coat thin, formed by both integuments

(endotestal and exotegmic). Embryo straight, small,

differentiated into cotyledons, radicle, and plumule;

endosperm copious, oily, and proteinaceous. Producing

chalkons (Tetracentraceae), proanthocyanidins

(Trochodendraceae), ﬂ avonols (quercetin and kaemp-

ferol); n = 20, 24.

In many respects Trochodendrales occupy, as it

were, an intermediate position between Magnoliidae

and Hamamelididae, but in the totality of their

characters they stand nearer to the latter (Takhtajan

1966, 1969, 1980, 1987; Cronquist 1981, 1988;

Endress 1986, 1993). In Tetracentron all ﬂ oral organs

contain scattered enlarged cells resembling the “oil”

cells that are generally present in many Magnoliidae,

but absent in Hamamelididae. They are often more or

less spherical as in Magnoliidae; in Trochodendron

similar cells differentiate into scleriﬁ ed, branched idio-

blasts. However, in the gynoecium, at least at anthesis,

many of these cells are not scleriﬁ ed and presumably

also correspond to “oil” cells (Endress 1986: 315).

Key to Families

1 Evergreen trees forming long shoots only. Buds

large, with many imbricate scales. Leaves closely

clustered near the branch tips in pseudowhorls,

obovate to broadly lanceolate, crenate serrate, cori-

aceous, pinnately veined, estipulate; petiole with an

arc of three vascular bundles. Idioblasts sclerenchy-

matous, not secretory. Flowers bisexual or occa-

sionally polygamous, with long pedicels; in short,

initially terminal but soon appearing axillary,

upright racemelike structure with a terminal ﬂ ower.

Pedicels with bract and several bracteoles, expanded

above into subconical torus. Sepals essentially

absent (rudimentary tepals are recognizable in early

ontogenetic stages of the ﬂ ower, but they are oblit-

erated at anthesis and never function as protective

organs, Endress 1993). Stamens 40–70, spirally

arranged; anthers mucronate. Carpels (4-)6-11(17);

stylodia conduplicate and deeply canaliculate ven-

trally, recurved. Ovules 15–30 per carpel.

Endosperm cellular. Seeds 15–25 per locule, pen-

dulous, smooth, linear; seed coat endotestal-endote-

gmic, exotesta consists of one tanniniferous layer. n

= 20. . . . . . . . . . . . . . . . . 1. trochodendraceae.

1 Deciduous trees with slender branches bearing short

shoots. Short shoots alternately arranged, marked

with crowded concentric scars of fallen leaves and

bud scales and bearing a single subterminal leaf and

an inﬂ orescence each year. They form sympodial

systems with adaxial prophylls. Buds slender, with

two outer sheathing scales. Leaves cordate-ovate,

closely crenate, palmately curviveined from base,

with stipules adnate at the base of petiole; petiole

with a single arcuate or almost cylindrical vascular

strand. Idioblasts secretory, not sclerenchymatous.

Flowers bisexual, actinomorphic, sessile in the axil

of minute bract, in alternate groups of four on

104 Subclass III. HAMAMELIDIDAE

slender, catkinlike spike. Sepals four in two decus-

sate pairs, thin and have only a rudimentary vascu-

lar bundle. Stamens four, in two decussate pairs,

each opposite a sepal; anthers truncate-rounded at

the apex. Carpels four, in a cycle, alternate with sta-

mens; stylodia subulate, at ﬁ rst connivent, later

becoming sharply recurved and subbasal by great

overgrowth of ventral region of each carpel. Fruits

of four laterally coalescent carpels, these loculicid-

ally dehiscent along the entire exposed portion of

the ventral face to base of persistent deﬂ exed stylo-

dia. Ovules 5-8(-10) per carpel. Endosperm nuclear.

Seeds 5-8(-10) per locule, faceted, linear-oblong;

seed coat exotegmic, exotesta consists of one

colourless layer. n = 24. . . . 2. tetracentraceae.

1. TROCHODENDRACEAE

Eichler 1865. 1/1. From Korea to Japan, Ryukyu

Islands, and Taiwan.

Trochodendron.

2. TETRACENTRACEAE

A.C. Smith 1945. 1/1. Eastern Himalayas (from east-

ern Nepal to Arunchal-Pradesh), northern Burma,

southwestern and central China.

Tetracentron.

Close related to the Trochodendraceae.

Bibliography

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Leroy J-F. 1988. L’organisation fondamentale d’une Angiosperme

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Mohana Rao PR. 1981. Seed and fruit anatomy of Trochodendron

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Oliver D. 1889. Tetracentron sinense Oliv. Hooker’s Icones

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Pan KY et al. 1993. The embryology of Tetracentron sinense

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Praglowski J. 1974. The pollen morphology of the Rocho den-

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Rao PRM. 1983. Seed and fruit anatomy of Trochodendron ara-

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Smith AC. 1945. A taxonomic review of Trochodendron and

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Tsuji SI, M Matsushita. 1991. Notes on pollen morphology of

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Wagner R. 1903. Beitrage zur Kenntnis der Gattung

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409–422.

Superorder TROCHODENDRANAE 105

Wang FH, NF Chien, and YL Zhang. 1984. A study on the pol-

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Order 31. CERCIDIPHYLLALES

Deciduous trees with branches differentiated into long

and short shoots. Some of the cells of the parenchyma-

tous tissues containing crystals of calcium oxalate.

Vessel elements very long, angular, with oblique sca-

lariform perforation plates that have 20–50 bars; lat-

eral pitting scanty, from opposite to scalariform. Fibers

with conspicuously bordered pits. Rays heterogeneous.

Axial parenchyma scanty, apotracheal. Sieve-element

plastids of S-type, large, often surrounded by endo-

plasmic reticulum membranes. Nodes on long shoots

trilacunar, those on short shoots unilacunar. Leaves

simple, dimorphic, those on long shoots pinnately

veined, elliptic or broadly ovate, entire or ﬁ nely ser-

rate, opposite, subopposite, or occasionally in irregular

whorls of three. However, those on sympodially grow-

ing short shoots, which develop from the axillary buds

of long shoot leaves and have an adaxial prophyll, are

palmately veined, broadly cordate or reniform, crenate,

alternate and solitary (each short shoot bears a single

leaf in each growing season). In buds leaves are invo-

lute and have well-developed marginal glands; stipules

present, small, adnate to the petiole and caducous.

Stomata anomocytic. Flowers dioecious, without peri-

anth, arranged in small, dense, racemose heads terminal

on the short shoot and produced before or with the

leaves. Male inﬂ orescences contain 16–35 stamens,

but the individual ﬂ owers difﬁ cult to delimit since

perianth and often subtending bracts are absent.

Stamens with long, slender, pendulous ﬁ laments and

long pink apiculate anthers; anthers tetrasporangiate,

basiﬁ xed, latrorse, opening by simple longitudinal

slits. Tapetum secretory. Microsporogenesis simulta-

neous. Pollen grains 2–2-celled, 3-colpate, tectate-

columellate, ﬁ nely reticulate, with minute tectal

perforations; the apertural exine is coarsely structured.

Female inﬂ orescences of 2–8 unicarpellate ﬂ owers;

carpels slightly stipitate, gradually narrowed to a

slender stylodium with a long, 2-ridged, decurrent red

stigma that bears unicellular papillae. The ventral

suture of each carpel is oriented abaxially with respect

to the inﬂ orescence axis, which is unusual. According

to Solereder (1899) the abaxial orientation of the suture

is due to loss of one of a pair of opposite carpels, which

is conﬁ rmed by paleobotanical data (Crane and Stockey

1986). The ovary contains 15–30 ovules with the lami-

nar lateral placentation. Ovules anatropous, apotro-

pous, bitegmic, crassinucellate. Female gametophyte

of Polygonum-type Endosperm cellular. Fruits follicles.

Seeds small, ﬂ attened, with a long chalazal extension

forming a wing that contains the ovular vascular bun-

dles forming a hairpin loop (as in Trochodendrales).

Seed coat formed mainly by the outer integument;

exotestal cells enlarged, slightly thickened; embryo

large, well-differentiated into cotyledons and long

hypocotyl; endosperm scanty, oily. Tanniniferous and

producing both phenolics and ﬂ avonols (quercetin and

kaempferol). n = 19.

Although Cercidiphyllales are related to the

Trochodendrales, particularly to Tetracentraceae, they

differ from them markedly in several characters, such

as dimorphic leaves, extremely congested inﬂ ores-

cences, much more reduced dioecious and unicarpel-

late ﬂ owers, and the presence of vessels.

1. CERCIDIPHYLLACEAE

Engler 1907. 1/2. Japan and China (Hubei, Sichuan,

and Shaanxi).

Cercidiphyllum.

Bibliography

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Cercidiphyllum. J. Palaeontol. 13: 485–499.

Crane PR. 1984. A reevaluation of Cercidiphyllum-like plant

fossils from the British early Tertiary. Bot. J. Linn. Soc. 89:

199–230.

Crane PR and RA Stockey. 1985. Growth and reproductive biol-

ogy of Joffrea speirsii gen. et sp. nov., a Cercidiphyllum-like

plant from the Late Paleocene of Alberta, Canada. Canad.

J. Bot. 63: 340–364.

Crane PR and RA Stockey 1986. Morphology and development of

pistillate inﬂ orescences in extant and fossil Cercidiphyllaceae.

Ann. Missouri Bot. Gard. 73: 382–393.

Endress PK. 1986. Floral structure, systematics, and phylogeny

in Trochodendrales. Ann. Missouri Bot. Gard. 73: 297–324.

106 Subclass III. HAMAMELIDIDAE

Endress PK. 1993. Cercidiphyllaceae. In: K Kubitzki, ed., The

families and genera of vascular plants, vol. 2, pp. 250–252.

Springer, Berlin/Heidelberg/New York.

Feng Y-X, X-Q Wang, K-Y Pan, D-Y Hong. 1998. A reevalua-

tion of the systematic positions of the Cercidiphyllaceae and

Daphniphyllaceae based on rbcL gene sequence analysis,

with reference to the relationship in the ‘Lower’ Hamamelidae.

Acta Phytotax. Sinica 36: 411–422.

Harms H. 1916. Über die Blütenverhältnisse und die system-

atische Stellung der Gattung Cercidiphyllum Sieb. and Zucc.

Ber. Deutsch. Bot. Ges. 34: 272–283.

Heel WAV. 1987. Note on the morphology of the male inﬂ ores-

cences in Cercidiphyllum (Cercidiphyllaceae). Blumea 32:

303–309.

Jähnichen H, DH Mai, and H Walther. 1980. Blätter und Früchte

von Cercidiphyllum Siebold et Zuccarini im mitteleuropais-

chen Tertiär. Schriftenreihe Geol. Wiss. 16: 357–399.

Leroy JE. 1980. Développement et organogenese chez le

Cercidiphyllum japonicum: Un cas semblant unique chez les

Angiospermes. C. R. Acad. Sci. Paris., ser. D, 290: 679–682.

Mohana Rao PR. 1986. Seed and fruit anatomy in Cercidiphyllum

japonicum with a discussion on the afﬁ nities of

Cercidiphyllaceae. Flora 178: 243–249.

Rowley JR. 1992. Pollen of Cercidiphyllum (Cercidiphyllaceae).

Bot. Zhurn. 77(11): 13.

Solereder H. 1899. Zur Morphologic und Systematik der Gattung

Cercidiphyllum Sieb. and Zucc. mit Berücksichtigung der

Gattung Eucommia Oliv. Ber. Deutsch. Bot. Ges. 17: 387–406.

Spongberg SA. 1979. Cercidiphyllaceae hardy in temperate

North America. J. Arnold Arbor. 60: 367–376.

Swamy BGL and IW Bailey. 1949. The morphology and rela-

tionships of Cercidiphyllum. J. Arnold Arbor. 30: 187–210.

Van Heel WA. 1986. A deviating female ﬂ ower of Cercidiphyllum

japonicum (Cercidiphyllaceae). Blumea 31: 273–276.

Van Heel WA. 1987. Note on the morphology of the male inﬂ o-

rescences in Cercidiphyllum (Cercidiphyllaceae). Blumea

32: 303–309.

Wang D and SZ Gao. 1990. A study on the relationships of

Cercidiphyllaceae. I. Leaf architecture and vasculature

changes of petiole. Acta Bot. Bor.-Occid. Sinica 10: 37–41.

Superorder MYROTHAMNANAE

Order 32. MYROTHAMNALES

Small, sympodially branching, ascending, rigid, xero-

phytic shrubs with an extreme poikilohydrous capacity

(“resurrection plants”) and characterized by the pro-

duction of balsam (resin) in large, inﬂ ated, thin-walled

epidermal cells. Branches clearly differentiated into

long and short shoots, and lateral branches largely

developed as short shoots. Vessel elements long, nar-

row, angular, with oblique scalariform or reticulate sca-

lariform perforation plates, with numerous (ca. 45)

bars; lateral pitting alternate. Fibers with bordered pits.

Rays exclusively uniseriate and composed mostly of

upright cells (and a few square cells), without crystals.

Axial parenchyma absent. Sieve-element plastids of

S-type. Ultrastructurally the chloroplasts possess a

unique type of granum (Welburn and Welburn 1976).

Nodes trilacunar with three traces. Leaves small,

opposite-decussate, with broad sheathing base, cuneate,

with distal part rounded and dentate, plicate and folding

up like a fan during the dry season, with palmately-

ﬂ abellate venation. The leaf bases of opposite leaves are

fused and form a sheath that remains on the shoot after

the leaf has abscised. Stipules intrapetiolar, small, tooth-

like. Stomata encyclocytic. Inﬂ orescences terminal on

short shoots, erect, spikelike, with or without terminal

ﬂ ower, the ﬂ owers (or ﬂ oral triads at the base of the

inﬂ orescence) opposite and subtended by foliose (the

lowest) or bracteose, cucullate leaves, unisexual, with

either male or female ﬂ owers (Dahlgren and van Wyk

1988). Flowers sessile, bracteolate or without bracte-

oles, unisexual, actinomorphic or somewhat zygomor-

phic, apetalous, with 0–4 (or more) lanceolate scales

interpreted as “tepals” (Dahlgren and van Wyk 1988)

(vestigial sepals?). Stamens 3–8, when 4 in terminal

ﬂ owers with 4 “tepals” alternating with them. Filaments

cylindrical, distinct or connate at the base. Anthers

large, tetrasporangiate, basiﬁ xed, with a connective

prolonged into a beak, latrorse, opening longitudinally

almost slitlike but with a stomial bifurcation at the

proximal end (Hufford and Endress 1989). Tapetum

secretory. Microsporogenesis simultaneous. Pollen

grains 2-celled, in tetrahedral tetrads, each with three

ill-deﬁ ned pores and intectate. The exine sculpturing

consists of clavate columellae that are ornamented with

minute papillae (Zavada and Dilcher 1986). Gynoecium

of 3–4 united carpels with free, recurved, short, broad

stylodia and enlarged, winglike, ventrally decurrent,

marginally more or less ungulate stigma bearing unicel-

lular papillae. Ovary 3–4 - locular, with 28–32 submar-

ginal ovules in each locule. Ovules anatropous,

bitegmic, crassinucellate, micropyle bistomal. Female

gametophyte of Allium-type. Endosperm nuclear. Fruits

small, leathery capsules; the carpels separating above

and opening ventrally. Seeds numerous, small, ovoid,

with thin, reticulate seed coat formed mainly by the

inner integument, exotestal cells with somewhat thick-

ened outer walls. Embryo small, weakly differentiated;

endosperm copious, oily. Plants strongly tanniniferous

and containing ellagic acid, chalcones, ﬂ avonols (quer-

cetin and kaempferol), p-coumaric acid, sinapinic acid,

Superorder HAMAMELIDANAE 107

and p-hydrobenzoic acid as well as vanillinic and syrin-

gic acids and cyanidin. Crystal druses of calcium

oxalate are abundant in the leaves, n = 10.

Although Myrothamnus somewhat resembles the

genus Distylium of the Hamamelidaceae (Endress 1977,

1989), it differs from Hamamelidaceae and related fam-

ilies in many respects, including its vegetative morphol-

ogy (particularly leaf morphology and anatomy),

uniseriate rays, absence of the axial parenchyma, pollen

morphology, morphology of the gynoecium, female

gametophyte of Allium-type, and copious endosperm.

It is therefore more correct to separate the Myro-

thamnaceae into an order of their own (Nakai 1943;

Takhtajan 1987, 1997). Endress (1989: 199) has come

to the conclusion that features Myrothamnus shares

with Cercidiphyllum and Trochodendrales s. 1.

“strongly point to a closer phylogenetic relationship of

the Myrothamnaceae with the Trochodendrales than

with the Hamamelidales, and especially with the

Cercidiphyllaceae.” I agree that Myrothamnaceae are

relatively closer to the Trochodendranae (Trocho-

dendrales s.1.) than to the hamamelides. However, tak-

ing into consideration so many important differences

between Trochodendranae (including Cercidiphyllales)

and the Myrothamnaceae, it seems more appropriate to

consider Myrothamnales a separate superorder. The

Myrothamnales differ from the Trochodendranae by

their poikilohydrous capacity, leaf morphology (includ-

ing very peculiar and unique ﬂ abellate venation), uni-

seriate rays, absence of axial parenchyma, tetradinous

and intectate 3-porate pollen grains, Allium-type

female gametophyte, nuclear endosperm, and septi-

cidal capsules. I consider Myrothamnanae a separate

evolutionary lineage and a very advanced side branch

of the phylogenetic tree of the Hamamelidae (1997).

1. MYROTHAMNACEAE

Niedenzu 1891. 1/2. Tropical Africa and South Africa

and Madagascar.

Myrothamnus.

Bibliography

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(Myrothamnaceae) and the problem of multiperforiate perfo-

ration plates. J. Arnold Arbor. 57: 119–126.

Carlquist S. 1990. Leaf anatomy of Geissolomataceae and

Myrothamnaceae as a possible indicator of relationship to

Bruniaceae. Bull. Torrey Bot. Club 117: 420–428.

Dahlgren R and AE van Wyk. 1988. Structures and relationships

of families endemic to or centered in southern Africa.

Monogr. Syst. Bot. Missouri Bot. Gard. 25: 1–94.

Endress PK. 1989. The systematic position of the Myrothamnaceae.

In: PR Crane and S Blackmore, eds. Evolution, systematics,

and fossil history of the Hamamelidaceae, vol. 1, pp. 193–

200. Claredon Press, Oxford.

Grundell R. 1933. Zur Anatomic von Myrothamnus ﬂ a-bellifolia

Welw. Symbolae Botanicae Upsaliensis 2: 1–17.

Jäger-Zurn I. 1966. Inﬂ oreszenz- und blütenmorpholog-ische,

sowie embryologische Untersuchungen an Myrothamnus

Welw. Beitr. Biol. Pﬂ . 42: 241–271.

Kubitzki K. 1993. Myrothamnaceae. In: K Kubitzki, ed. The

families and genera of vascular plants, vol. 2, pp. 468–469.

Springer, Berlin/Heidelberg/New York.

Puff C. 1978a. The nodal anatomy of Myrothamnus ﬂ a- bellifolius

(Myrothamnaceae): Another example of a ‘split-lateral’

condition. J. Arnold Arbor. 59: 192–196.

Puff C. 1978b. Zur Biologic von Myrothamnus ﬁ abelli-folius

Welw. (Myrothamnaceae). Dinteria 14: 1–20.

Vieweg GH and H Ziegler. 1969. Zur Physiologic von

Myrothamnus ﬂ abellifolia. Ber. Deutsch. Bot. Ges. 82:

29–36.

Weimarck H. 1936. Myrothamnus ﬂ abellifolius Welw.: Eine

polymorphe Pﬂ anzenart. Bot. Not. 1936: 451–462.

Wellburn FAM. and AR Wellburn. 1976. Novel chloroplasts and

unusual cellular ultrastructure in the “resurrection” plant

Myrothamnus ﬂ abellifolius Welw. (Myrothamnaceae). Bot.

J. Linn. Soc. 72: 51–54.

Superorder HAMAMELIDANAE

Order 33. HAMAMELIDALES

Evergreen or deciduous trees and shrubs. Crystals of

calcium oxalate usually present in some of the cells of

the parenchymatous tissues. Vessel elements narrow

and sometimes very long, with very oblique end walls,

mostly with scalariform perforations that have few to

numerous bars (more than 50 in Corylopsis and

Exbucklandia and 50–100, sometimes 130 or even 160

in Ostrearia lateral pitting from opposite to scalari-

form, rarely alternate. Fibers with bordered pits. Rays

heterogeneous. Axial parenchyma apotracheal and

typically diffuse. Sieve-element plastids of S-type with

5–10 starch grains. Nodes trilacunar to pentalacunar.

Leaves alternate, simple, often palmately lobed, pin-

nately or palmately veined, stipulate. Stomata para-

cytic. Flowers in inﬂ orescences of various types, of

medium size or small, bisexual, polygamous or

108 Subclass III. HAMAMELIDIDAE

unisexual, actinomorphic or rarely zygomorphic, with

double perianth or seldom apetalous, sometimes with-

out perianth or with perianth extremely reduced. Sepals

(3)4-5(-10), small, distinct or more often more or less

connate, sometimes very reduced or lacking. Petals

usually 4–5, commonly small or narrow, sometimes

wanting. Stamens (3)4-5 (up to 10), in one cycle, alter-

nating with petals or staminodia, or sometimes more

numerous (up to 32) and initiated in centripetal or cen-

trifugal sequence. Anthers frequently with a shortly

prolonged connective, mostly tetrasporangiate,

latrorse, opening by two valves like two window shut-

ters (Endress 1986, 1989a, 1993; Hufford and Endress

1989). Only in ﬁ ve genera of tribe Hamamelideae do

the thecae open by a single valve that serves both

pollen sacs (Endress 1989b). Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains

2-celled, from 3-colpate to pantoporate and pantocol-

pate, tectatecolumellate, reticulate. Gynoecium of 2 or

less often 3-8(9) carpels, or more frequently united at

the base; stylodia always free, with decurrent stigma.

Each locule and each free carpel mostly with one

ovule, less often two ovules, and in each locule of syn-

carpous gynoecia sometimes more. Ovules anatropous,

more or less pendulous, bitegmic, crassinucellate.

Endosperm cellular (Parrotiopsis) or nuclear. Fruits of

small achenes, or woody capsules with a leathery exo-

carp and a bony endocarp. Seed coat formed mainly by

the outer integument; embryo long, straight; endosperm

copious to scanty, oily, and proteinaceous. More or

less tanniniferous, producing phenolics ﬂ avonols

(quercetin, kaempferol, and frequently also myricetin)

and also iridoid compounds.

This order in many ways serves as a connecting link

between Trochodendrales on the one hand and the

“amentiferous” orders Casuarinales, Fagales, etc., on

the other. The order Hamamelidales is evidently derived

from an immediate ancestor of Trochodendrales.

Key to Families

1 Pollen grains commonly 3-colpate, seldom pantocol-

pate (Matudaea), 4-colpate to pantoporate (Distyilum)

or pantoporate (Sycopsis). Petioles with a trough-

shaped or cylindrical vascular strand. Resin ducts

absent (except Mytilaria, which has resin canals in

the pith of stem). Flowers small, bisexual, in some

genera bisexual and male, rarely unisexual

(Fortunearia, Sinowilsonia). Sepals usually imbri-

cate. Petals present or absent, when present (2-)4, or

5, gamopetalous, valvate. Stamens 4–32; anthers bas-

iﬁ xed, introrse, tetrasporangiate or rarely (Hamamelis)

disporangiate, opening by pores, or by short or longi-

tudinal slits, of by longitudinal valves, usually append-

ages. Carpels two or rarely three, small or medium;

ovary more or less inferior, ovules mostly solitary,

sometimes 6–15; outer integument 6–7 cells across,

micropyle zig-zag. Fruits loculicidal and septicidal

capsules with woody exocapr and horny endocarp.

Seeds winged or wingless, with thick, hard, and con-

spicuous testa, consisting of ﬁ berlike cells and form-

ing the main protective part of the seed coat; hilum

large. The tegmen does not contribute to the construc-

tion of the mature seed coat; only the inner epidermis

persists; the nucellus persists in ripe seed; endosperm

oily; embryo straight. Trees or shrubs, often with

stellate indumentum. Vessels with scalariform or sca-

lariform and reticulate perforation. Vestured pits pres-

ent. Nodes pentalacunar with ﬁ ve traces. Leaves

dissected or entire, often palmatiﬁ d. Contain ﬂ avonols

kaempferol, quercetin, and myricetin, producing

C-glycosylﬂ avones, sometimes (Corylopsis) ellagic

acid.. . . . . . . . . . . . . . . . . . . . . . 1. hamamelida ceae

1 Pollen grains always pantoporate, the pores circu-

lar, often with islands of ectexine on the pore mem-

brane. Petioles with 3–5 vascular bundles. Resin

ducts present in roots, stem, leaves, and ﬂ oral

organs. Flowers in terminal racemes (male) or in

globular heads. Perianth sepaline or vestigial, absent

in male ﬂ owers, or 12–50 minute lobes or scales in

female ﬂ owers. Stamens 12–50, anthers opening by

longitudinal slits or valves. Staminodes 3–10.

Carpels 2; ovary inferior, 2-locular. Ovules 20–50

per locule, horizontal. Fruit a septicidal capsule.

Seeds small, winged; the wing formed by the longi-

tudinal elongation and ﬂ attening of the integument

around the micropyle; the ripe seed with the 2-celled

thick testa, but the protective part of the seed coat

contributed by the outer epidermis of the inner

integument with ligniﬁ ed and ﬁ brous cells; the

nucellus does not persist in ripe seed; endosperm

slight, embryo long. Large, monoecious, deciduous

trees. Vessels without vestured pits. Leaves ovate,

palmatiﬁ d or tricuspidate, entire, serrate, or dentate;

domatia (Liquidambar) presented by pockets, or

hair tufts. Iridoids detected (Liquidambar), present

proanthocyanidins (cyanidin and delphinidin), ﬂ a-

vonols (quercetin and myricetin), and ellagic acid,

n = 15, 16 . . . . . . . . . . . . . . . . . . 2. altingiaceae

Superorder HAMAMELIDANAE 109

1. HAMAMELIDACEAE

R. Brown 1818 (including Disanthaceae Nakai 1943,

Exbucklandiaceae Reveal et Doweld 1999,

Fothergillaceae Nuttall 1818, Parrotiaceae Horaninow

1834, Rhodoleiaceae Nakai 1943). 28/100-110.

Tropical and South Africa, Madagascar, southeastern

Transcaucasia, northern Iran, western and eastern

Himalayas, Assam, Manipur, eastern Asia and

Southeast Asia, New Guinea, northeastern Australia,

eastern North America, and Central America.

1.1 DISANTHOIDEAE

Leaves suborbiculate, cordate or truncate at the base,

entire, palmately 5-nerved. Stipules scarious, linear,

caducous. Flowers two in a head, with short bracts at

the base, bisexual, entomophilous. Perianth present,

petals ﬁ ve, circinate in bud, ﬂ abellate, stellately spread-

ing. Anthers opening by simple longitudinal slits. Each

locule with about ﬁ ve ovules; ripe locule with 4–5

seeds. Seeds not ejected out of capsule, n = 8, chromo-

somes the largest in the family and nuclei similar to the

Tetrameranthus type (Morawetz and Samuel 1989). –

Disanthus (1 sp. in eastern China and Japan).

1.2 HAMAMELIDOIDEAE

Leaves mostly ovate, sometimes cordate, pinnately

veined, entire or toothed. Stipules mostly small, com-

monly caducous. Stellate hairs present. Flowers usually

in spikes or heads, commonly bisexual entomophilous or

anemophilous. Perianth mostly present, but petals often

lacking. Each locule mostly with 1 ovule, rarely with one

fertile and two sterile; ripe locule with one seed. Fruit

with ballistic dispersal. The seeds ejected at maturity,

n = 12. – hamamelideae: Hamamelis, Loropetalum,

Tetrathyrium, Maingaya, Embolanthera, Dicoryphe,

Trichocladus, Ostrearia, Neostrearia, Noahdendron;

corylopsideae: Corylo psis; eustigma teae: Eustigma,

Fortunearia, Sinowil sonia; fothergilleae:

Molinodendron, Fothergilla, Parrotiopsis, Shaniodendron,

Parrotia, Sycopsis, Distyliopsis, Distylium, Matudaea.

I.3 EXBUCKLANDIOIDEAE

Leaves ovate, often tricuspidate, mostly cordate, pal-

mately veined. Stipules mostly large and covering sub-

sequent bud. Flowers aggregated in dense spikes,

bisexual. Perianth present or absent. Each locule with

5–8 ovules. The seed not ejected, n = 8–12. —

Exbucklandia, Mytilaria, Chunia.

1.4 RHODOLEIOIDEAE

Leaves ovate, rarely cordate, entire, pinnately veined,

vestigially stipulate. Inﬂ orescence a pseudanthium of

strongly zygomorphic ﬂ owers. Flowers bisexual,

ornitophilous. Perianth present. Each locule with 1–2

fertile ovules plus 10–20 sterile ones; ripe locule with

2 or more seeds. Seeds not ejected, n = 12. — Rhodoleia

(1–10 spp. in eastern Asia and Malesia).

Disanthoideae are in general the least specialized

subfamily (Takhtajan 1980, 1987, 1997; Cronquist

1981; Wolfe 1989; Pan et al. 1991) while Rhodoleioideae

are the most specialized.

2. ALTINGIACEAE

Horaninow 1841. 3/17. Eastern Mediterranean (islands

of the Aegean Sea-Rhodes and Cyprus), eastern

Himalayas (Bhutan and Assam), continental China,

Taiwan, Southeast Asia, eastern and northeastern

North America, northeastern Mexico, and Central

America to Costa Rica.

Liquidambar, Altingia, Semiliquidambar.

Usually included in the Hamamelidaceae, but the

combination of characteristics “sets the subfamily quite

apart from the rest of the family” (Bogle 1986: 328).

Although speciﬁ c characteristics, including resin ducts

and periporate pollen grains, do occur in certain genera

of the other subfamilies (Bogle 1986), in general the

Altingiaceae are so distinct that they deserve the status

of a separate family. Chemistry also underscores

Altingiaceae’s distinctive nature (Ferguson 1989).

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