Takhtajan Armen. Flowering Plants
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272 Subclass V. DILLENIIDAE
Dendristellera, Restella, Stelleropsis, Stellera, Dais,
Struthiola, Passerina, Drapetes, Kelleria, Pimelea,
Lasiadenia.
Evidently related to the Malvalean families (Hallier
1922; Domke 1934). They have many common fea-
tures with the Tiliaceae. According to Metcalfe and
Chalk (1950: 1181), general anatomical evidence
“appears to indicate that the Gonystylaceae,
Thymelaeaceae, and Tiliaceae have defi nite affi nities
with each other. The occurrence of secretory elements
containing mucilage is common to Gonystylus, the
Tiliaceae, and Thymelaeaceae”. The occurrence of
cyclopropenyl fatty acids (Vickery 1980), and exoteg-
mic seeds (Corner 1976) also support the affi nities.
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Order 76. URTICALES
Trees, shrubs, or herbs, often with cystoliths or
laticifers or both. Vessels with simple perforations;
lateral pitting alternate. Fibers mostly with simple pits.
Rays heterogeneous or sometimes homogeneous.
Axial parenchyma usually paratracheal. Sieve-element
plastids of S-type or (some Ulmaceae) of Pc-type.
Nodes trilacunar or (Ficus) pentalacunar. Leaves alter-
nate or less often opposite, almost always stipulate
(absent in some Parietarieae), mostly simple, very often
oblique at the base. Stomata anomocytic or sometimes
of other types. Flowers small and inconspicuous, in
cymose or racemose infl orescences of sometimes very
complex structure, rarely solitary, unisexual or less
often bisexual, cyclic, apetalous, anemophilous or
secondarily entomophilous. Sepals 2–8 (mostly 4–5), in
one or sometimes two cycles, usually valvate, sometimes
more or less connate, sometimes wanting. Stamens
mostly as many as sepals and opposite them, rarely
fewer or more numerous; fi laments free or sometimes
connate; anthers usually 2-locular (1-locular in Ficus
subsect. Malvanthera). Tapetum secretory. Micro-
sporogenesis simultaneous. Pollen grains 2-celled or
less often 3-celled, 2-pantoporate. Gynoecium usually
pseudomonomerous, of two carpels (rarely of three
carpels in some Moraceae), with one or two separate
stylodia, stigmas de-current. Ovary superior or inferior,
usually 1-locular, but occasionally 2-locular (e.g., in
Ulmus). Ovules solitary (or solitary in each of two
locules), anatropous, hemitropous, or (Cecropiaceae
and Urticaceae) orthotropous, bitegmic, crassinucellate.
Female gametophyte of Polygonum- or less often
Allium-, Adoxa-, or Drusa-types. Endosperm nuclear.
Fruits of various types. Seeds with straight or curved
embryo, with or without endosperm.
Weddell (1856) was the fi rst who discussed several
characters indicating affi nity of the Urticaceae with the
Malvales, especially with the Tiliaceae. Hallier (1903,
1905) included Urticaceae s.1. in the Malvales but later
(Hallier 1908, 1912) changed his opinion and moved
them in his “Terebinthines.” Bessey (1915) also
included Ulmaceae, Moraceae, and Urticaceae in his
Malvales s.l. More recently Thorne (1968, 1974, 1976,
1983, 1992a, b, 2000), Stebbins (1974), Dahlgren
(1975, 1980, 1983), and Friis (1993: 617) came to the
conclusion that the Urticales are closely related to the
Malvales but are distinct enough to merit their own
order. The affi nity between them is confi rmed by
anatomical data, including wood anatomy and the
occurrence of mucilage cells and canals (Guerin 1923;
Holm 1927; Metcalfe and Chalk 1950; Hegnauer
1973), as well as by serological studies (Petersen and
Fairbrothers 1985) and phytochemical data (Hegnauer
Superorder MALVANAE 279
1964, 1969, 1973; Soepadmo 1977), as well seed anatomy
(Kravtsova 2003). According to Berg (1977b:371),
“The affi nities of the Urticales with the Malvales are
closest between the Ulmaceae and Tiliaceae, the least
derived families of the orders.”
The Urticales have most probably originated from an
ancient tiliaceous ancestor.
Key to Families
1 Ovules subapical, pendulous, anatropous to campy-
lotropous. Ovary with two stylodia, unilocular or
bilocular, but one of them sometimes more or less
reduced. Milky latex present or absent. Fruits dry or
drupaceous.
2 Plants without milky latex. Stamens erect in bud.
Flowers unisexual, polygamous, or rarely
bisexual.
3 Trees and shrubs, usually with scattered muci-
lage cells or even mucilage canals, and with
strong tendency toward mineralization of the
cell walls with calcium carbonate or silica.
Leaves simple, often oblique at the base, usually
distichous, rarely opposite, with typically cadu-
cous stipules. Flowers in cymose or fasciculate
infl orescences, monoecious or less often bisex-
ual. Pollen grains 4–7-porate, exine rugulose.
Ovary unilocular or rarely bilocular. Fruits dry
or drupaceous. Seed fl attened, coat undistin-
guished, exotestal cells elongated, unthickened;
embryo straight or curved (Zelkova); endosperm
scanty or absent. Contain sesquiterpene lac-
tones, n = 10, 14. . . . . . . . . . . . . . 1. ulmaceae.
3 Perennial or annual herbs with secretory canal
in the phloem, but without mucilage cells.
Stems erect or twining, either scabrous or
armed with rigid climbing hairs and often with
glandular trichomes. Leaves palmately lobed
(Humulus) or palmately compound (Cannabis),
sometimes simple, petiolate, serrate, generally
decussate, near the stem apex often alternate,
opposite at least below, with triangular, persis-
tent stipules. Flowers in complex, basically
cymose infl orescences, unisexual (monoe-
cious or more often dioecious). Ovary uniloc-
ular with solitary anatropous ovule, pendent
from near the apex of the locule. Fruits nutlike
or achenes covered by the perianth. Seeds with
curved (Cannabis) or coiled (Humulus)
embryo and a small amount of fl eshy, oily
endosperm. Accumulating quebrachitol and
producing pyridine alkaloids, and sometimes
tanniferous, with proanthocyanins, n = 7,
9, 10. . . . . . . . . . . . . . . . . . .3. cannabaceae.
2 Plants nearly always with milky latex (wanting in
Fatoua) borne in laticifers widely distributed in the
parenchymatous parts of the stem and often in the
leaves. Trees, shrubs, woody lianas (including
stranglers), subshrubs, succulent or tuberous geo-
phytes, and rarely annuals (Fatoua). Leaves simple
or rarely compound, alternate or opposite, often
with cell walls more or less mineralized with cal-
cium carbonate or silica; stipules present, some-
times much reduced. Stomata anisocytic or
cyclocytic. Flowers always unisexual, in compact
infl orescences with the axis often thickening to
form a head or invaginated common receptacle.
Stamens erect or infl exed in bud. Ovary unilocular
or less often bilocular. Fruits drupaceous, some-
times with dehiscent exocarp. Seed coat undistin-
guished; endosperm absent or present; embryo
strait or curved, n = 7, 12–14.. . . . . . 2. moraceae.
1 Ovules subbasal or basal, erect, more or less ortho-
tropous. Ovary always unilocular, with a single sty-
lodium. Plants usually without milky latex. Fruits
dry, nutlike.
4 Trees, shrubs, or woody lianas (sometimes epi-
phytic), often with stilt roots. Laticifers more or
less reduced or absent, but mucilage cells and sacs
often present. Leaves alternate, simple and entire
to often so deeply palmately or radiately incised as
to appear compound, with fused and often fully
amplexicaul stipules. Cystoliths wanting. Flowers
dioecious. Stamens erect in bud (except for
Poikilospermum subgenus Poikilospermum), not
abruptly bending in dehiscence. Fruits nutlike.
Seeds large and without endosperm or small and
with endosperm, x = 7.. . . . . . . 4. cecropiaceae.
4 Herbs or occasionally subshrubs or rarely small,
soft-wooded trees, very rarely lianas, in Urticeae
provided with stinging hairs. Frequently with
mucilage cells, and sometimes also with nonar-
ticulated latex canals, but usually without milky
juice. Leaves alternate or opposite, simple, entire
or 3–5(-7)-lobed, usually with three subequal
veins from the base, with epidermal cells tending
to be mineralized with silica or calcium carbon-
ate; stipules usually present (absent in some
Parietarieae), lateral or often intrapetiolar, often
280 Subclass V. DILLENIIDAE
fused. Stomata anisocytic or paracytic. Cystoliths
usually present in both the stems and leaves, lin-
ear to punctiform. Flowers unisexual or rarely
bisexual. Stamens infl exed in bud, abruptly bend-
ing in dehiscence. Fruits nutlike or seldom drupa-
ceous. Seeds with thin oily or starchy endosperm,
but sometimes without endosperm, n = 6–14.
. . . . . . . . . . . . . . . . . . . . . . . . . . 5. urticaceae.
1. ULMACEAE
Mirbel 1815 (including Celtidaceae Link 1831). 15/
200. Widely distributed in tropical, subtropical, and
temperate regions; Ulmoideae distributed mainly in
temperate regions.
1.1 CELTIDOIDEAE
Leaves with three main veins from the base and pairs of
secondary veins usually less than six, alternate or
(Lozanella) opposite, brochidodromous. Rays essen-
tially heterogeneous (in Gironniera and Trema mark-
edly heterogeneous). Flowers unisexual or polygamous.
Pollen grains usually 2–5 porate (pantoporate in
Ampelocera), ectexine palisade (Ampelocera and
Chaetachme) or granular in the middle part (not recog-
nizable in Lozanella, Parasponia, and Trema); orna-
mentation fi nely scabrate or consisting of spinules or of
densely spaced warts and microechinules. Fruits more
or less globose, drupaceous, and with thick-walled and
well-lignifi ed endocarp, or (Pteroceltis) small, nutlike,
with spherical endocarps whose reticulate surface sculp-
ture is almost identical to that of Celtis, and with an exo-
carp developed into a pair of wings differing from those
of Ulmoideae in lacking venation. Seeds with curved
embryo and scanty or no endosperm (endosperm fl eshy
in Parasponia and Lozanella); cotyledons mostly vari-
ously folded or involute. Produce quebrachitol and gly-
cofl avones (which are absent in Gironniera subgenus
Galumpita) but not lignans, sesquiterpens, and fl avonols
(the latter present in Gironniera subgenus Galumpita).
x = 10, 14. – Ampelocera, Chaetachme, Gironniera,
Aphananthe, Celtis, Pteroceltis, Trema, Parasponia,
Lozanella.
1.2 ULMOIDEAE
Leaves pinnately veined, with seven or more pairs of
secondary veins, the lowest pairs not prominent,
craspedodromous, alternate. Rays homogeneous or
weakly heterogeneous. Flowers mostly bisexual or
polygamous. Pollen grains 4–5-porate, with the exine
distinctly and fi nely granular throughout; ornamenta-
tion rugulate, with small spinules. Fruits dry, usually
more or less compressed (samaroid) or less often sub-
drupaceous (Zelkova and Planera) or small and nutlike
with a rigid endocarp similar to that of Zelkova and a
crestlike wing (Hemiptelea). Seeds with straight
embryo and usually without endosperm; cotyledons
broad, fl at or longitudinally folded. Produce lignans,
sesquiterpenes, and fl avonols, but not quebrachitol and
glycofl avons. x = 14. – Ulmus, Holoptelea, Hemiptelea,
Zelkova, Planera, Phyllostylon.
Celtidoideae were traditionally included in the
Ulmaceae and only recently have some authors sup-
ported their distinctiveness as a separate family
(Grudzinskaya 1967; Chernik 1975, 1980, 1981, 1982;
Zavada 1983; Takahashi 1989). However, some gen-
era, including Ampelocera and Aphananthe, possess
character combinations (including fl oral anatomy) that
seem to bridge the gap between typical celtidoids and
ulmoids (see Chernik 1975; Gianasi 1978; Manchester
1989; Terabayashi 1991; Zhong et al. 1992).
Ampelocera and
Aphananthe are celtidoid in most
nonchemical characters (except for special leaf vena-
tion of Ampelocera) although they contain fl avonols
(Gianasi 1978). Zelkova and Hemiptelea, long consid-
ered celtidoid, are in fact fl avonol producers (Gianasi
1978) and included in the Ulmaceae sensu stricto by
Grudzinskaya (1967), which is confi rmed also by
wood anatomy (Zhong et al. 1992). Besides, Gironniera
consists of two distinct entities, one of which (subge-
nus Gironniera) is really celtidoid and the other (sub-
genus Galumpita) is in many respects rather different
(chemically ulmoid) and was even separated as a genus
Galumpita Blume. Finally, tropical American Celtis
monoica (a monotypic genus Mirandaceltis, according
to Sharp [1958]) is in fact a fl avonol producer (Gianasi
1978) and as Sharp (1958) indicates, its leaf venation
and drupe size are somewhat intermediate between
Celtis and Zelkova. “The fl avonoid data at this level do
not suggest any particular Ulmoid relative for
Mirandaceltis monoica, but do indeed support Sharp’s
contention that the taxon should be removed from
Celtis sensu stricto,” states Gianasi (1978: 342). All
these data show that the differences between celtidoid
and ulmoid genera are not clear-cut, and therefore I am
reluctant to accept a separate family rank for celti-
doids. At best they are two subfamilies.
Superorder MALVANAE 281
2. MORACEAE
Link 1831 (including Artocarpaceae Dumortier 1829,
Dorsteniaceae Chevalier 1827, Ficaceae Berchtold et
J. Pres 1,96720). 37/1200+. Widely distributed in trop-
ics and subtropics with a few species in temperate
regions of both hemispheres.
moreae: Morus, Broussonetia, Milicia, Maclura,
Trophis (including Olmedia), Streblus, Bleekrodea,
Fatoua; artocarpeae: Artocarpus, Parartocarpus,
Treculia, Prainea, Hullettia, Antiaropsis, Sparattosyce,
Batocarpus, Bagassa, Sorocea, Clarisia, Poulsenia;
castilleae: Perebea, Maquira, Castilla, Helicostylis,
Pseudolmedia, Naucleopsis, Antiaris, Mesogyne;
dorstenieae: Utsetela, Bosqueiopsis, Helianthostylis,
Trymatococcus, Brosimum, Trilepislum, Scyphosyce,
Dorstenia; ficeae: Ficus.
The subdivision of the Moraceae follows Rohwer
(1993). But as Rohwer notes, the family forms such a
closely knit complex that tribes and genera are diffi cult
to delimit. According to Rohwer, the best defi ned
tribes are Castilleae and Ficeae.
Related to the Ulmaceae, especially to the
Celtidoideae.
3. CANNABACEAE
Martynov 1820 (including Lupulaceae Link 1831).
2–3/4. Temperate regions of the Northern Hemisphere.
Humulus (including Humulopsis ?), Cannabis
Cannabaceae are related to Urticaceae and
Moraceae, but differ in the absence of laticifers and the
erect position of stamens in bud (Kubitzki 1993).
2n = 18 + XX or XY, in Humulus lupuls 2n = 20 + XX
or XY, while in Humulus (Humulopsis) scandens
2n = 14 + XX (female) or YXY (male).
4. CECROPIACEAE
C.C. Berg 1978. 6/200. Mainly tropical America, but
also tropical Africa (Myrianthus and Musanga) and Asia
from eastern Himalayas to Malesia (Poikilospermum).
Myrianthus, Pourouma, Cecropia, Musanga,
Coussapoa, Poikilospermum.
Cecropiaceae are intermediate between the
Moraceae, with which they share possession of latic-
ifers, and the Urticaceae, with which they share ortho-
tropous subbasal or basal ovules (Berg 1978).
According to Berg (1989: 215), the delimitation of the
family is doubtful only with respect to the genus
Poikilospermum, which deviates in several features
(e.g., the elongate “urticaceous” cystoliths) from the
group of African and American genera. He states that
anatomical data (Bonsen and ter Welle 1983) also
suggest that the position of Poikilospermum is separate
from the other fi ve genera and closer to the
Urticaceae. According to Friis (1993) the position of
Poikilospermum is still uncertain.
5. URTICACEAE
A.L. de Jussieu 1879 (including Parietariaceae Berchtold
et J. Presl 1820). 49/1000. Widely distributed in tropical
and subtropical regions, but some species occur in tem-
perate and cold regions. The largest concentration of
genera and species is in tropical Asia.
urticeae: Urtica, Hesperocnide, Nanocnide,
Obetia, Laportea, Discocnide, Girardinia, Dendrocnide,
Urera, Gyrotaenia: lecantheae: Elatostema,
Meniscogyne, Procris, Pilea, Achudemia, Sarcopilea,
Lecanthus, Petelotiella;
boehmerieae: Boehmeria,
Archiboemeria, Chamabainia, Pouzolzia, Hyrtanandra,
Gonostegia, Neodistemon, Cypholophus, Sarcochlamys,
Touchar dia, Neraudia, Pipturus, Nothocnide,
Oreocnide, Debregeasia, Astrothalamus, Leucosyke,
Gibbsia, Phenax, Maoutia, Myriocarpa; pariet ar-
ieae: Gesn ouinia, Hemistylus, Parietaria, Soleirolla,
Rousselia; forsskaoleae: Forsskaolea, Droguetia,
Didymodoxa, Australina.
Bibliography
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