Takhtajan Armen. Flowering Plants

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736 Subclass IV. COMMELINIDAE

Radhakrishnaiah M, G Nageshwar, and LL Narayana. 1984.

Chemotaxonomy of Pandanus and Typha. Curr. Sci. 53:

759–760.

Rendle A. 1953. The classiﬁ cation of ﬂ owering plants. Vol. 1.

Gymnosperms and monocotyledons, 2nd ed. Cambridge

University Press, Cambridge

Rowlatt U. 1992. Architecture of the leaf of the greater reed

mace, Typha latifolia L. Bot. J. Linn. Soc. 110: 161–170.

Savile DB0. 1979. Fungi as aids in higher plant classiﬁ cation.

Bot. Rev. 45: 377–503.

St. John H. 1941. Teratological Typha. Rhodora 43: 85–91.

Thieret JW. 1982. The Sparganiaceae in the southeastern United

States. J. Arnold Arbor. 63: 341–355.

Wagner P. 1977. Vessel types of monocotyledons: a survey. Bot

Not. 130: 383–402.

Zubkova IG and LK Shabes. 1985. Typhaceae. In: A Takhtajan,

ed. Comparative seed anatomy, vol. 1, pp. 262–263. Nauka,

Leningrad (in Russian).

Order 30. RESTIONALES

Perennial or less often annual herbs, sometimes lianous,

with solid or ﬁ stulose stems. Silica bodies often pres-

ent. Oxalate raphides mostly present (absent in most

Restionaceae and Centrolepidaceae). Vessels in roots

and stems, with simple or scalariform perforations.

Leaves alternate, commonly distichous, entire, with

open or closed sheath and narrow, parallel-veined

lamina that is sometimes reduced or suppressed.

Stomata paracytic, of graminoid type. Flowers small

and inconspicuous, in various kinds of inﬂ orescences,

often in spikelets, bisexual or more often unisexual,

mostly 3-merous. Perianth small, membranous, in two

cycles or in one cycle, sometimes lacking. Stamens 6

in 2 cycles or 3(4) in 1 cycle, sometimes only 2 or

even 1; anthers basiﬁ xed to dorsiﬁ xed, tetrasporangiate

or less often disporangiate, latrorse or introrse, open-

ing longitudinally. Tapetum secretory. Microsporo-

genesis simultaneous (Flagellariaceae) or more often

successive. Pollen grains 3-celled or less often 2-celled,

tectate-columellate, 1-porate, often with annulate

graminoid pore. Gynoecium commonly of three united

carpels, less often two carpels; in Centrolepidaceae of

one carpel; stylodia more or less connate into style;

ovary superior, 3-locular or by reduction of one or two

carpels 2-locular or even 1-locular, with one pendulous

ovule per locule. Ovules orthotropous, bitegmic,

crassinucellate (Flagellariaceae) or more often tenui-

nucellate, without parietal cell. Female gametophyte

of Polygonum- or Allium-type. Endosperm nuclear.

Fruits loculicidal capsules or nuts, in Flagellariaceae

drupaceous. Seeds small, globose or ovoid, with or

without operculum; seed coat formed by both integu-

ments; endosperm copious, starchy, with aleurone

layer; embryo small, lens-shaped or obovate, weakly

differentiated. Bound p-coumaric, ferulic and diferulic

acids present. n = 6–13, 18, 19.

Related to the Typhales and Juncales and have a

common origin from a commelinalean ancestors

(Linder and Kellogg 1995).

Key to Families

1 Anthers disporangiate or rarely (Restionaceae

p.p.) tetrasporangiate.

2 Stamens three; gynoecium generally of 3 carpels.

Usually xeromorphic perennial herbs, tufted or

with an elongate rhizome with overlapping scales,

glabrous or densely woolly with multicellular

hairs. Stem with a continuous cylinder of scleren-

chyma ring inside the chlorenchyma. Silica bod-

ies present or absent; plants accumulated starch

exclusively ‘pteridophyte type’. Vessels in stem

with scalariform, or with simple and/or scalari-

form, or with simple perforations; in the rhizomes

with exclusively simple perforations and in roots

with scalariform perforations. Stems with solid or

spongy, or hollow internodes. Leaves alternate,

mostly distichous, usually reduced to an open

sheath with scarcely any lamina, usually not ligu-

late, stipulate (in the form of membranous lobes at

the tops of the sheath margins), or estipulate.

Stomata, when present, paracytic. Flowers in spi-

cate or paniculate inﬂ orescences, aggregated into

spikelets subtended by a single bract or less often

solitary and mostly are also subtended by bracte-

oles, unisexual, dioecious or rarely monoecious;

in some species of Lepyrodia and in one species

of Restio bisexual ﬂ owers may occur. Perianth

segments six in two series, rarely four in two

cycles or only three or even absent. Stamens oppo-

site the inner perianth segments; ﬁ laments some-

times connate into a column; anthers dorsiﬁ xed,

unilocular, introrse or latrorse, disponagiate or tet-

rasporangiate, appendaged or unappendages.

Pollen grains 2-celled or 3-celled, scrobiculate,

pores with annulus or without annulus. Staminodia

present or absent. Pistillodia may be present in the

male ﬂ owers. Stylodia free or connate below, with

elongate, papillate, or shortly plumose stigma on

their inner sides; ovary 3- to 1-locular, sessile to

Superorder POANAE 737

stipitate. Hypostase present. Ovules 1–3, tenui-

nucellate, rarely (Alexgeorgia) crassinucellate.

Large starch bodies are surrounding polar nuclei.

Female gametophyte of Polygonum-type. Fruits

1–3-locular loculicidal capsules or small nuts.

Seeds endotegmic, with copious, mealy, starchy

endosperm capped by minute, lenticular or obo-

vate, weakly differentiated embryo. Contain ﬂ a-

vonols (kaempferol, quercetin, and myricetin),

glycoﬂ avones, and proanthcyanidins. n = 7, 8, 11,

12, 13, 16, 20.. . . . . . . . . . . . . .3. restionaceae.

2 Stamen one; gynoecium of one carpel. Small,

tufted, annual or perennial and rhizomatous

herbs of wet habitats. Raphides and silica bodies

lacking. Multicellular, branched or unbranched

ﬁ lamentous hairs sometimes present. Roots usu-

ally without differentiated pericycle. Stems

solid. Vessels with scalariform perforations in

root, stems and leaves. Leaves alternate, basal or

less often imbricately inserted along the stem,

linear, lanceolate or setaceous, basally with dis-

tinct, membranous, open sheath. Flowers min-

ute, in highly reduced, terminal, compound,

spikelike or headlike inﬂ orescences subtended

by one to several leafy bracts and bearing two to

several distichous, glumelike bracts; each bract

encloses a group of male ﬂ owers or one to sev-

eral female ﬂ owers, or, most frequently, one or

more bisexual synanthia; each synanthium is a

spikeletlike cyme of one or two male ﬂ owers

and one to several superposed or collateral

female ﬂ owers and often has one to three small,

hyaline bracts or bracteoles. Filaments ﬁ liform,

glabrous; anthers 1-locular, dorsiﬁ xed, versatile.

Microsporogenesis successive. Pollen grains

3-celled, tectate-granular, 1-porate (ulcerate),

with poorly deﬁ ned, aperturate margins, verru-

cate, tectum perforated by small punctae (scro-

biculi). Stylodium ﬁ liform, with elongate,

ventrally decurrent, papillate stigma; ovary

superior, with one apical, pendulous ovule.

Female gametophyte of Polygonum-type. Fruits

small, membranous, 1-seeded, indehiscent

(Aphelia) or more often dehiscing longitudinally

on a dorsal side, which gives an appearance of

paired valves in the fruiting pseudanthia of

Gaimardia and Centrolepis (Cooke 1998); col-

lateral dehiscing fruits are generally united into

a capsulelike collective fruit. Seeds small; seed

coat rather thin, membranous, formed mainly by

the inner epidermis of the inner integument, the

outer integument being very thin; endosperm

copious, mealy, rich in compound starch grains,

without aleurone layer; embryo minute, weakly

differentiated, globose, obconical, or bell-like,

peripheral, lying on the micropylar side.

n = 10–13. . . . . . . . . . . .6. centrolepidaceae.

1 Anthers tetrasporangiate.

3 Perennial, high climbing lianas with solid stem

arising from sympodial rhizomes, supported by

means of long, coiled tendrils terminating the

leaves and representing the extension of the

thickened midrib. Leaf blades larger than sheaths

and ending in a tendril. Silica bodies present.

Scattered secretory cells present in leaf and stem.

Vessels in leaves and stems, with scalariform and

simple perforations. Leaves alternate, circinately

enrolled in bud; lamina lanceolate, constricted

at base; leaf sheaths closed. Flowers in more or

less branched terminal panicles, bisexual or

very rarely unisexual, bracteate. Perianth seg-

ments ﬁ ve, in two cycles, free. Stamens six in

two cycles, occasionally reduced to staminodia;

ﬁ laments very short; anthers sagittate basiﬁ xed,

latrorse. Microsporogenesis simultaneous. Pollen

grains 2-celled. Cynoecium of three carpels;

styles three, sometimes connate basally, free or

partially joined. Ovary 2-locular, ovules one per

locule, pendulous. Female gametophyte of

Allium-type. Fruits ﬂ eshy, red or black drupes.

Seeds endotestal; embryo minute, lenticular;

endosperm copious, starchy. Flavonoles (kae-

mpferol) present; alkaloids present or absent;

n = 19. . . . . . . . . . . . . . . . 1. flagellariaceae.

3 Reed-like, coarse, unbranched, erect herbs to 5 m

tall. Stems terete, hollow, arising from a short

sympodial rhizomes, without scabrid margins,

unbranched below the inﬂ orescence. Silica bod-

ies and calcium oxalate crystals present. Secretory

cells absent. Vessels in all vegetative organs, with

scalariform and simple perforations. Leaves large

(the lamina up to 1 m long), alternate, distichous,

ﬂ at, sessile, with tubular, open, sheathing bases,

the mouths of sheaths with diminutive ligules,

the lamina abruptly narrowed at base and articu-

lated to the sheath, long, conspicuously plicate,

grasslike, bearing short prickly hairs and

branched ﬁ lamentous hairs. Flowers in long,

738 Subclass IV. COMMELINIDAE

much branched terminal panicles, bisexual, brac-

teate and have caducous bracteoles. Perianth seg-

ments six, in two cycles, free to joined, sepaloid

(bractlike), sometimes the outer members much

larger, persistent. Stamens six; anthers basiﬁ xed

(saﬁ ttate) latrorse. Pollen grains 3-celled. Ovary

3-locular; styler branches or sessile stigmatic

areas three; stigmas covered with short papillae.

Ovules one per locule, pendulous. Fruits drupa-

ceous, red, yellow or black. Seeds with copious,

mealy, starchy endosperm capped by minute,

lenticular, undifferentiated, discoid embryo.

n = 18. . . . . . . . . . . . . . . . . . 2. joinvilleaceae.

4 Monoecious plants with male and female

ﬂ owers together in the same spikelet. Stamens

six or four. Perennial evergreen herbs with

slender cylindrical solid photosynthetic stems

arising from rhizomes. Silica bodies (as silica

sand) present in the culm mesophyll. Vessels

in stems and in roots, with scalariform and

simple perforations. Leaves reduced to 1–4

tightly convoluted sheaths up to 10 cm long;

sheath split to the base, usually with a mucro

which may be developed into a small awn.

Flowers in single- or several-ﬂ owered spike-

lets aggregated into terminal conical or cylin-

drical dense spikelike heads, subtended by a

glossy, dark brown or black, rigid, ovate bract

longer than the ﬂ ower, dorsiventrally ﬂ attened.

Perianth of six, unequal, glumaceous seg-

ments. Female ﬂ owers with three staminodes.

Stamens six or four; ﬁ laments free, slender;

anthers basiﬁ xed, tetrasporangiate, latrorse.

Pollen grains annulate and with a plug; intera-

pertural wall smooth, and lacks scrobiculi.

Pistillodia present. Gynoecium with 2-locular

ovary; styles two, free, attenuate from the

ovary, apical; stigmas two; ovules one per loc-

ule, pendulous Female gametophyte tet-

rasporic, most similar to the Drusa-type. Fruits

a 3-locular capsule or a nut. Seeds with minute

embryo and copious endosperm. n = 32, 33

and ca. 24.. . . . . . . . . . 5. ecdeiocoleaceae.

4 Dioecious plants; culms not striate or with

striations formed by narrow ridges of scleren-

chyma not be deep crypts; stamens three.

Perennial herbs with solid or ﬁ stulose stems

arising from short, rhizomes, which covered

with overlapping, glabous scales; stems simple

or branched, sometimes very compressed.

Vessels with scalariform perforations. Silica

bodies lacking, but oxalate crystals occur in

some species. Leaves linear to ﬁ liform, later-

ally ﬂ attened, equitant, with a small ligule at the

transition to the open sheath. Flowers in few to

many ﬂ owers paniculate inﬂ orescence with

each branch subtended by elongated leaﬂ ike

bract, pedicellate or sessile, each with one or

two short bracts. Perianth of six more or less

similar and glumaceous segments in two cycles.

Stamens three; ﬁ laments free connate; anthers

dorsiﬁ xed, tetrasporangiate, latrorse. Pollen

grains 3-celled, scrobiculate, pores annu-

late. Gynoecium of three carpels; style free,

stigmas decurrent ovary 3-locular with solitary

ovule per locule. Female gametophyte of

Polygonum-type. Fruits 1-seeded angular cap-

sules dehiscing at the angles or nutlets. Seeds

with minute, lenticular embryo and copious

mealy, starchy endosperm. Flavonol glycosides

present. n = 11.. . . . . . . . . . 4. anarthriaceae

1. FLAGELLARIACEAE

Dumortier 1829. 1/4. Tropical Africa, Southeast Asia,

Malesia, Micronesia, New Caledonia, northern

Australia, and eastward to Samoa and Niue.

Flagellaria.

2. JOINVILLEACEAE

Tomlinson et A. C. Smith. 1970. 1/2. From western

Malesia eastward to Samoa and Hawaii, but absent

from southern and eastern Indonesia, New Guinea, and

Australia.

Joinvillea.

3. RESTIONACEAE

R. Brown 1810 (including Elegiaceae Raﬁ nesque 1838,

Hopkinsiaceae B. G. Briggs et L. A. S. Johnson 2000;

Lyginiaceae B. G. Briggs et L. A. S. Johnson

2000). 57/520. Tropical and South Africa, Madagascar,

Southeast Asia, Malesia, New Guinea, Australia,

Tasmania, New Zealand, and southern South America;

Superorder POANAE 739

concentrated mostly in southwestern Africa and extra-

tropical regions of Australia and Tasmania.

Lyginia, Hopkinsia, Staberoha, Ischyrolepis, Elegia,

Chondropetalum, Dovea, Askidiosperma, Platycaulos,

Restio, Calopsis, Thamnochortus, Rhodocoma, Cerato-

caryum, Cannomois, Nevillea, Hydrophilus, Antho-

chortus (including Anthochortus), Mastersiella,

Hypodiscus, Willdenowia, Lepyrodia, Sporadanthus,

Calorophus, Winifredia, Empodisma, Coleocarya,

Desmocladus, Harperia, Onychosepalum, Catacolea,

Kulinia, Lepidobolus, Baloskion, Curingalia, Acion,

Saropsis, Alexgeorgia, Chordifex, Dielsia, Eurychorda,

Platychorda, Tremulina, Melanostachya, Cytogonid-

ium, Loxocarya (including Megalotheca), Taraxis,

Tyrbastes, Pseudoloxocarya, Leptocarpus, Hypolaena,

Meeboldina, Stenotalis, Chaetanthus, Dapsilanthus,

Apodasmia, Pseudoloxocarya.

4. ANARTHRIACEAE

D. F. Cutler et Airy Shaw 1965. 1/7. Southwestern

Australia.

Anarthria.

Closely related to the Restionaceae, differing

mainly in the culm anatomy.

5. ECDEIOCOLEACEAE

D. F. Cutler et Airy Shaw 1965. 2/2. Southwestern

Australia.

Ecdeiocolea, Georgeantha.

6. CENTROLEPIDACEAE

Endlicher 1836. 4/35. Southern China (Hainan),

Southeast Asia, Malesia, New Guinea, Australia,

Tasmania, New Zealand, and subantarctic South

America (Tierra del Fuego and Falkland Islands).

Centrolepis (including Pseudalepyrum), Aphelia,

Brizula, Gaimardia.

Closely related to the Restionaceae, but differ from

them markedly in very reduced and specialized inﬂ o-

rescence with unisexual ﬂ owers clustered into synan-

thia, minute and perianthless ﬂ owers, male ﬂ owers

usually with one stamen, disporangiate anthers,

“ centrolepidoid” pollen grains, and monocarpellate

gynoecium. Also in most species of all genera except

Gaimardia, they have an unusual type of epidermal

cell arrangement whereby one cell overlaps the end of

the next in a ﬁ le (a unique feature in monocotyledons),

they have no pericycle in the root, and their root hairs

arise in an abnormal fashion from one side of root epi-

dermal cells (Cutler 1969: 96). Already Dahlgren

(1975) proposed a separate order Centrolepidales

although in his later publications he included the

Centrolepidaceae in the Restionales. Linder and

Ferguson (1985: 75) suggested that the Centrolepidaceae

be excluded from the orders Restionales and Poales.

According to Hamann (1975), the Centrolepidaceae

could have a neotenous origin from the Restionaceae.

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Neyland R. 2002. A phylogeny inferred from large-subunit

(26S) ribosomal DNA sequences suggests that the family

Dasypogonaceae is closely aligned with the Restionaceae

allies. Austral. Syst. Bot. 15: 749–754

Superorder POANAE 741

Pin YK and LF Yew. 1975. Serological evidence on the distinctness

of the monocotyledonous families Flagellariaceae, Huanguan-

aceae and Joinvilleaceae. J. Linn. Soc. Bot. 70: 77–81.

Prakash N. 1970. The ﬂ oral development and embryology of

Centrolepis fascicularis. Phytomorphology 19: 285–291.

Ronse Decraene LP, PH Linder and EF Smets. 2001. Floral

ontogenetic evidence in support of the Willdenowia clade of

South African Restionaceae. J. Plant Res. 114: 329–342.

Ronse Decraene LP, HP Linder, and EF Smets. 2002. Ontogeny

and evolution of the ﬂ owers of South African Restionaceae

with special emphasis on the gynoecium. Plant Syst. Evol.

231: 225–258.

Rudall P. 1990. Development of the ovule and megagametophyte

in Ecdeiocolea monostachya. Austral. Syst. Bot. 3:

265–274.

Rudall P and HP Linder. 1988. Megagametophyte and nucellus

in Restionaceae and Flagellariaceae. Am. J. Bot. 75:

1777–1786.

Rudall P, W Stuppy, J Cunniff, EA Kellogg, BC Briggs. 2005.

Evolution of reproductive structures in grasses (Poaceae)

inferred by sister-group comparison with their putative clos-

est living relatives, Ecdeiocoleaceae. Am. J. Bot. 92:

1432–1443.

Sack FD. 1994. Structure of the stomatal complex of the mono-

cot Flagellaria indica. Am. J. Bot. 81: 339–344.

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Smithson E. 1956. Comparative anatomy of the Flagellariaceae.

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Subramanyam K and HS Narayana. 1972. Some aspects of the

ﬂ oral morphology and embryology of Flagellaria indica L.

In: YS Murty, BM Johri, HY Mohan Ram, and TM Varghese,

eds. Advances in plant morphology, pp. 211–217. Sarita

Prakashan, Meerut, India.

Tillich HJ. 1996. Seeds and seedlings in Hanguanaceae and

Flagellariaceae (Monocotyledons). Sendtnera 3: 187–197

Tillich H-J und E Sill. 1999. Systematische Studien zur

Morphologie und Anatomie von Hanguana Blume

(Hanguanaceae) und Flagellaria L. (Flagellariaceae), mit

der Beschreibung einer neuen Art Hanguana bogneri spec.

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Tomlinson PB and AC Smith. 1970. Joinvilleaceae, a new fam-

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Johnson. 1997. Flavonoid evidence and the classiﬁ cation of

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tion of Australian Restionaceae. Phytochemistry 49:

529–552.

Order 31. POALES

Perennial or less often annual or biennial herbs or with

more or less woody stems. Raphides and crystals of

calcium oxalate lacking, but silica bodies present.

Internodes solid or more often hollow. Vessels com-

monly in all vegetative organs, with simple or some-

times scalariform perforations. Sieve-element plastids

of P2c-type (Behnke 2002). Leaves alternate, distic-

hous, differentiated into open or rarely closed sheath

and usually narrow and elongate, but in some tropical

members broad and in Phyllorachis even sagittate, in

many Bambusoideae with more or less distinct petiole

(up to 25 cm in Anomochloa); venation parallel or

rarely (Leptaspis and Pharus) with pinnately disposed

lateral veins; a membranous ligule generally present at

the juncture of the sheath and lamina on the adaxial

side, sometimes reduced or represented by a row of

hairs or even wholly suppressed. Stomata paracytic,

with dumbbell-shaped guard cells. Flowers small, very

reduced, bisexual or sometimes unisexual (monoe-

cious or dioecious), arranged in panicles or sometimes

in spikelike or racemelike inﬂ orescences, which con-

sist of one to to many-ﬂ owered spikelets; spikelets

usually with a pair of subopposite bracts lacking axil-

lary ﬂ owers (subtending glumes) and one to several

ﬂ orets often on more or less zigzag axis (rachilla);

each ﬂ oral axis arises in the axil of a bract (lemma) and

usually bears a two-keeled prophyll (palea), (1)2 or

(Stipa and Bambusoideae) 3 much smaller and incon-

spicuous scales (lodicules) above these, and the

androecium and gynoecium; upper (adaxial) scale

(palea) typically has 2 main veins and interpreted as

2 connate segments of the outer cycle of the ancestral

3-merous perianth (

elakovsky 1889), the third seg-

ment nearly always suppressed; lodicules, which

sometimes, as in Arundinaria, are relatively large and

well developed, apparently represent the inner cycle of

the ancestral perianth (

elakovsky 1889; Butzin 1965).

Stamens in many Bambusoideae six in two cycles, but

mostly three, less often one, two, four, or (in some

Bambusoideae) more than six; ﬁ laments elongate, ﬁ li-

form; anthers tetrasporangiate, elongate, basiﬁ xed but

deeply sagittate so as to appear versatile, opening

longitudinally. Tapetum secretory. Microsporogenesis

successive. Pollen grains 3-celled, tectate-columellate

with a thick foot-layer, 1-porate, operculate, annulate,

psilate, spinulose or minutely scabrate. Gynoecium

742 Subclass IV. COMMELINIDAE

pseudomonomerous, consists of two or (Bambusoideae)

three carpels; stylodia free or more or less connate into

a short style; ovary superior, 1-locular, with one ovule

attached to the wall. Ovules anatropous, hemitropous,

campylotropous, or orthotropous, bitegmic or rarely

(e.g., in Melocanna) unitegmic, crassinucellate, with-

out parietal cell. Female gametophyte of Polygonum-

type. Endosperm nuclear. Fruits mostly a caryopsis

with a thin pericarp usually adhering to the seed or

sometimes (Crypsis, Sporobolus, Eleusine) utricle-like

with a free, membranous or gelatinous pericarp, rarely

nutlike (some members of the Bambusoideae) or bac-

cate (Melocanna baccifera). Seeds with copious,

starchy endosperm usually with proteinaceous tissue

and sometimes also oily, rarely (Melocanna and related

genera) lacking; embryo straight, varying in size from

very small (Festuceae and most of temperate tribes) to

as long as the fruit (Spartina, etc.), placed on the abax-

ial side of the fruit, strongly differentiated and more or

less completely enveloped by the edges of cylindrical

coleoptile (the ﬁ rst leaf modiﬁ ed as a soil-penetrating

sheath for the plumule), radicle enclosed in coleorhiza

(interpreted by Brown [1965] as the remainder of the

base of the proembryo) and well-developed and

strongly modiﬁ ed cotyledon (scutellum) appressed to

the endosperm, n = 2–23 +.

A rather distinct order characterized by unique

spikelets, ﬁ ne channels in tectum, and especially by

its highly differentiated and specialized embryo

unique among monocotyledons. The Poales are

closely related to and derived from the Restionales,

most probably from some extinct ancestor of the type

of the living genus Joinvillea. Hallier (1902, 1912)

included grasses in Enantioblastae and placed them

between Flagellariaceae and Restionaceae, and

Wettstein (1908, 1935) derived them from

Enantioblastae. Various morphological, anatomical,

palynological, and embryological studies conﬁ rm the

close afﬁ nity between Poales and Restionales.

According to Campbell and Kellogg (1987) and

Rudall et al. (2005), Joinvilleaceae and Poaceae are

sister groups based on such a strong and in angio-

sperms unique synapomorphy as large, solitary silica

bodies in epidermal short cells, and in reproductive

structures. Besides, there are such common features

as the ligules, distichy, open leaf sheath, hollow inter-

nodes, vessel perforations, 3-celled pollen grains,

exine structure, morphology of integuments, binucle-

ate antipodal cells, seed anatomy, Asterad-type of

embryogenesis. Relationships are especially close

between Joinvillea and archaic grasses, such as

Streptochaeta. I therefore agree with Campbell and

Kellogg (1987: 218) that the synapomorphy uniting

the Poaceae and Joinvilleaceae is more convincing

than that linking the grasses to the Restionaceae.

1. POACEAE

Barnhart 1895 or Gramineae A.L. de Jussieu 1789

(nom. altern.). (including Aegilopaceae Martynov

1820, Agrostidaceae Burnett 1835, Alopecuraceae

Martynov 1820, Andropogonaceae Martynov 1820,

Anomochloaceae Nakai 1943, Arundinaceae

Hochstetter 1850, Arundinellaceae Herter 1940,

Avenaceae Martynov 1820, Bambusaceae Burnett

1835, Chloridaceae Herter 1940, Eragrostidaceae

Herter 1940, Festucaceae Sprengel 1825, Hordeaceae

Burnett 1835, Lepturaceae Herter 1940, Melicaceae

Martynov 1820, Miliaceae Burnett 1835, Narda-

ceae Martynov 1820, Oryzaceae Burnett 1835,

Panicaceae Voight 1845, Pappophoraceae Herter 1940,

Parianaceae Nakai 1943, Phalaridaceae Burnett 1835,

Pharaceae Herter 1940, Saccharaceae Martynov 1820,

Spartinaceae Burnett 1835, Stipaceae Burnett 1835,

Streptochaetaceae Nakai 1943, Triticaceae Hochstetter

1848, Zeaceae Kerner 1891). 670-850/10500-11000.

Cosmopolitan.

1.1 ANOMOCHLOOIDEAE

Perennial, rhizomatous, herbaceous grasses. Leaves

with either a distichous or spiral phyllotaxis; outer

ligule absent; inner ligule a short fringe of cilia or

absent, not membranous; pseudopetioles short to very

long, with dark, turgid swellings at both ends or only at

the summit; blades with parallel venation. Inﬂ orescences

terminal, stalked, spike-like, erect when young, but

arching and drooping at maturity, with complicated

branching patterns; the bracts within the spikelet

equivalents with distichous or spiral phyllotaxis; lodi-

cules absent or replaced by a ring of short brownish

cilia borne on a low membranous ring; stamens 4–6;

ovary glabrous, stigmas 1–3, hispid. Fruit a caryopsis,

large, fusoid to oblong-rectangular, embryo large, the

hilum linear, shallow and inconspicuous; endosperm

mealy. n = 12 or 18 (Clark and Judziewicz 1996). –

streptochaeteae: Streptochaeta; anomochloeae:

Anomochloa.

Superorder POANAE 743

1.2 PHAROIDEAE

Perennial, rhizomatous, monoecious, herbaceous

grasses. Rhizomes sympodial. Culms hollow or solid.

Leaves distichous, the pseudopetiole prominent and

twisted, the blade resupinate; outer ligule absent, the

inner ligule present, membranous; blades with lateral

nerves diverging obliquely from the midnerve and

running straight to the margins. Inﬂ orescence terminal,

open, paniculate, the rachis and branches disarticulating

or not, covered with uncinate macrohairs, bracts outside

of the spikelets absent. Spikelets unisexual, one-

ﬂ owered, mostly in male-female pairs on short branch-

lets, or some female spikelets solitary. Female spikelets

large, short-pedicelled; glumes two, shorter than the

ﬂ oret; lemma indurate, tubular or inﬂ ated, covered

wholly or in part by macrohairs; palea narrow, bicari-

nate; lodicules absent; ovary glabrous, style one, stigmas

three. Fruit a caryopsis, the hilum extending its full

length, embryo small, endosperm mealy. Male spikelets

small, short to long-pedicelled, membranous; glumes

two, shorter that the ﬂ oret; lodicules three or none, if

present then minute, elliptic, glabrous, and nerveless;

stamens 6. n = 12. – Pharus, Leptaspis, Scrotochloa.

1.3 PUELIOIDEAE

Perennial. Calms herbaceous, usually unbranched

above; internodes hollow. Leaves distichous, not

basally aggregated; outer ligule absent (Guaduella) or

present (Puelia); blades pseudopetiolate, large and

broad; sheaths non-auriculate. Synﬂ orescences race-

mose or paniculate, sometimes with small bracts sub-

tending the branches. Spikelets with two glumes,

several ﬂ orets and elongated rachilla internodes, the

1–3 basal ﬂ orets male, the next several ﬂ orets female-

fertile and bisexual, with apical incomplete ﬂ orets

(Guaduella), or the basal 3–6 ﬂ orets male or neuter

with the single apical ﬂ oret female (Puelia), disarticu-

lating above the glumes, and between the ﬂ orets

(Guaduella) or not (Puelia); stamens six, ﬁ laments

free (Guaduella) or monadelphous (Puelia); ovary

glabrous or hairy, an apical appendage present or

not; stigmas 2–3. Caryopsis with a long-linear hilum;

embryo small. n = 12. – puelieae: Puelia; guadu-

elleae: Guaduella.

1.4 BAMBUSOIDEAE

Perennials (rarely annual) rhizomatous herbaceous or

woody. Culms hollow or solid. Leaves almost always

petiolate, articulate-deciduous or less often persistent;

laminas usually lanceolate to lanceovate, very rarely

linear. Spikelets (or spikelets proper of the pseu-

dospikelets) bisexual (Bambuseae) or unisexual

(Olyreae), often with an indeﬁ nite number of ﬂ owers,

not infrequently branched, often with more than two

glumes at the base; lemmas and paleas usually with a

large and indeﬁ nite number (up to 25) of veins,

lodicules usually three, less often two, more than three,

or absent. Stamens usually six, less often three,

sometimes two, four, or more than six (up to 120);

ﬁ laments not or rarely connate into a tube. Stigmas 2

or 3, less often 1 or 4–6, not rarely on a long style.

Caryopsts often with the pericarp thickened in the

upper part, often nutlike or drupaceous. Meristem

under the base of the plumule absent. Chromosomes

small; n = 9–12, seldom 7. – bambuseae: Arthrostyli-

dium, Bambusa, Chusquea, Dinochloa, Merostachys,

Neurolepis, etc.; olyreae: (including buerger-

siochleae and parianeae): Buergersiochloa,

Lithachne, Olyra, Raddia, Eremitis, Pariana, etc.

1.5 EHRHARTOIDEAE

Plants annual or perennial (rhizomatous or stolonifer-

ous), herbaceous to suffrutescent. Culms hollow or

solid. Leaves distichous; abaxial ligule absent; adaxial

ligule a fringed or unfringed membrane, or a fringe of

hairs; blades rarely basally cordate or sagittate

(Phyllorachideae), somewhat broad to usually narrow,

sometimes pseudopetiolate; sheaths sometimes bear-

ing auricles. Inﬂ orescences paniculate or racemose,

bracts outside of the spikelets rarely present. Spikelets

bisexual or unisexual, with glumes 2 (absent in some

Oryzeae), sterile ﬂ orets 0 to 2, and female-fertile

ﬂ oret 1; palea well developed; lodicules 2, membranous

or rarely ﬂ eshy; stamens usually 3 or 6 (sometimes 1, 2,

or 4); ovary glabrous, apical appendage absent; styles 2,

free, fused basally of for their full length (Zizaniopsis),

close; stigmas 2. Caryopsis with the hilum long-linear;

endosperm hard, without lipid, containing compound

starch grains (rarely simple); embryo small, embryonic

leaf usually with overlapping margins (meeting in

Potamophila). n = 12 (10 in Microlaena, 15 in Zizania). –

oryzeae: Leersia, Luziola, Oryza, Zizania, etc.;

phyllorachideae: Humbertochloa, Phyllorachis;

ehrharteae: Ehrharta, Microlaena, etc.

1.6 POOIDEAE

Perennial or annual herbs, very seldom weakly ligne-

ous. Culms hollow (rarely solid). Leaves without a

744 Subclass IV. COMMELINIDAE

petiole, not deciduous, very seldom petiolate; laminas

usually linear, seldom lanceolate or lanceolate.

Spikelets with a deﬁ nite or indeﬁ nite number of ﬂ ow-

ers, not branched, provided with two glumes at the

base, seldom without glumes or with only one; lemmas

usually with (1-) 3–9(-11) veins; paleas with 2 veins

forming keels; lodicules two, seldom three, of various

types, or sometimes wanting. Stamens usually three,

very seldom one, two, four, or six; ﬁ laments free.

Stylodia two, very seldom three or one, very rarely

basally connate into a short style. Caryopsis always

with a thin pericarp, this generally adnate to the seed,

more rarely free from it, sometimes saclike.

Meristematic layer present under the coleoptile, as a

result of which the base of the plumule can elongate

during germination and produce adventitious roots.

Chromosomes small or large; n = 2, 4–12 and more. –

brachypodieae: Brachypodium; triticeae: Aegilops,

Agropyron, Elymus, Elytrigia, Eremopyrum, Henrardia,

Hordelymus, Hordeum, Leymus, Psathyrostachys,

Secale, Triticum, etc.; bromeae: Bromus, Boi-

ssiera, etc.; poeae: Agrostis, Aira, Anthoxanthum,

Arrhenatherum, Avena, Briza, Calamagrostis, Cata-

brosa, Cinna, Colpodium, Cutandia, Cynosurus,

Dactylis, Deschampsia, Eremopoa, Festuca, Heli-

ctotrichon, Hierochloe, Holcus, Koeleria, Lolium,

Millum, Polypogon, Puccinellia, Trisetum, Vulpia, etc.;

meliceae: Glyceria, Melica, Lophochlaena, etc;

brylkinieae: Brylkinia; diarrheneae: Diarrhena;

brachyelytreae: Brachyelytrum; ampelodesmeae:

Ampelodesmos; stipeae: Oryzopsis, Piptatherum,

Stipa, etc.; lygeeae: Lygeum; nardeae: Nardus;

phaenospermateae: Phaenosperma.

1.7 ARISTIDOIDEAE

Plants annual or perennial, caespitose, herbaceous,

xerophytic or less commonly mesophytic. Culms solid

or hollow. Leaves distichous. Inﬂ orescences panicu-

late, bracts outside of the spikelets absent. Spikelets

with bisexual ﬂ orets, glumes two, female-fertile ﬂ oret

one, and no rachilla extension, cylindrical or laterally

compressed, disarticulating above the glumes; lemma

with three awns, the awns separate from each other, or

fused below into a twisted column; palea short, less

than half the lemma length; lodicules present or rarely

absent, when present two, free, membranous, glabrous,

heavily vascularized; stamens one to three; ovary

glabrous, apical appendage absent, haustorial syner-

gids absent, styles two, free, close; stigmas two.

Caryopsis with the hilum short or long-linear;

endosperm hard, without lipid, containing compound

starch grains; embryo small (Sartidia) or large

(Aristida, Stipagrostis), epiblast absent, scutellar cleft

present or absent (Sartidia). n = 11, 12. – aristideae:

Aristida, Sartidia, Stipagrostis.

1.8 DANTHONIOIDEAE

Perennial (caespitose, rhizomatous or stoloniferous) or

less commonly annual, herbaceous or rarely suffrutes-

cent plants. Culms solid or very rarely hollow. Leaves

distichous; abaxial ligule usually absent (sometimes

present in Cortaderia, Karroochloa, and Pentaschistis);

adaxial ligule a fringe of hairs or a fringed membrane;

blades relatively narrow, without a speudopetiole;

sheats not auriculate except in Pentameris thuarii.

Inﬂ orescences paniculate or less commonly racemose

or spicate, bracts outside of the spikelets absent.

Spikelets bisexual (but sometimes without bisexual

ﬂ orets in Cortaderia

) or unisexual (Cortaderia,

Lamprothyrsus); glumes 2 and usually equal, female-

fertile ﬂ orets 1–6 (to 20), with apical reduction and a

rachilla extension usually present, laterally com-

pressed; lemma lacking uncinate macrohairs, awn sin-

gle and from a sinus; palea well developed, sometimes

relatively short; lodicules 2, free (rarely joined), ﬂ eshy

or rarely with an apical membranous ﬂ ap, glabrous or

ciliate, often with microhairs, sometimes heavily vas-

cularized; stamens 3; ovary glabrous or rarely with

apical hairs (Pentameris), apical appendage absent,

haustorial synergids present, only weakly developed in

a few taxa; styles 2, the bases usually widely separated,

stigmas 2. Caryopsis with the hilum short or long-

linear; endosperm hard, containing compound starch

grains (simple in Prionanthium); embryo large

or small, epiblast absent, scutellar cleft present. n = 6,

7, 9. – danthonieae: Pentaschistis, Pentameris,

Prionanthium, Chionochloa, Cortaderia, Lampro-

thyrsus, Pseudopentameris, Chaetobromus, Ryti-

dosperma, Notodanthonia, Austrodanthonia, Joycea,

Karroochloa, Schismus, Tribolium, Merxmuellera,

Danthonia, Notochloe, Plinthanthesis.

1.9 ARUNDINOIDEAE

Annuals or perennials, tufted, rhizomatous or stolonif-

erous, herbaceous to somewhat woody. Culms hollow

or less commonly solid. Leaves narrow and the ligule is

membranous or cartilaginous or may form a rim of cilia;

microhairs usually present, arm cells sometimes occur,

Superorder POANAE 745

fusoid cells always absent. Inﬂ orescences usually

paniculate, rarely spicate or racemose, bracts outside of

the spikelets absent. Spikelets with bisexual ﬂ orets,

glumes two, female-fertile ﬂ orets one to several, apical

reduction usually present; pales usually well developed;

lodicules two, free (rarely joined at the base), ﬂ eshy,

glabrous or infrequently ciliate; stamens 1–3; ovary gla-

brous, apical appendage absent, haustorial synergids

absent; styles two, usually free, close; stigmas two.

Caryopsis with the hilum short or long-linear; endosperm

hard, without lipid, containing compound starch grains;

embryo large or small. n = 6, 9, 12. – arundineae:

Arundo, Amphipogon, Dregeochloa, Hakonechloa,

Molinia, Phragmites, Triodia, etc.

1.10 MICRAIROIDEAE

Annual or perennial, terrestrial and rarely aquatic, rhi-

zomatous, matforming, sometimes decumbent herbs.

Culms solid or hollow. Leaves distichous or sometimes

spirally arranged; abaxial ligule occasionally present

as a line of hairs, adaxial ligule a fringe of hairs.

Inﬂ orescences panicles, racemes or spikes, terminal or

axillary. Spikelets bisexual, with two glumes (one in

Sphaerocaryum); glumes tardily or rarely early decid-

uous. Pales well developed, lodicules two or some-

times absent, cuneate, free, ﬂ eshy. Stamens 2–3; ovary

usually glabrous, apical appendage absent; styles 2,

free, stigmas 2. Fruits caryopsis with the hilum short

linear to long linear; endosperm hard, containin simple

starch grains; embryo small; n = 10. – eriachneae:

Eriachne, Pheidochloa; isachneae: Coelachne,

Heteranthoecia, Isachne, Limnopoa, Sphaerocaryum;

micraireae: Micraira.

1.11 CHLORIDOIDEAE

Annual or perennial herbs, tufted or rhizomatous (stolo-

niferous) and sometimes robust (rarely woody). Culms

solid or hollow. Leaves mostly narrow and the ligule is

hair-fringed or reduced to a rim of hairs. Silica bodies

are mostly saddle-shaped and microhairs are always

present. Distal cell usually inﬂ ated and sometimes

glandular. Mesophyll lacks arm cells and fusoid cells

and has a Kranz type PS anatomy. Inﬂ orescences

paniculate, paniculate with spicate branches, racemose,

or spicate, bracts outside of the spikelets absent.

Spikelets bisexual or sometimes unisexual (if so the

plants dioecious or monoecious), with one to several

ﬂ orets and the rachilla usually prolonged; pales well

developed, lodicules two or absent, ﬂ eshy, glabrous.

Stamens 1–3, stigmas 2; ovary glabrous, apical

appendage absent, haustorial synergids absent. Fruits

caryopsis or utricle, hilum short; endosperm hard, with-

out lipid, containing simple or compound starch grains;

embryo large or rarely small. n = 7, 8, 9 or 10. – cyno-

donteae (chlorideae): Bouteloua, Soderstroimia,

Grifﬁ thsochloa, Pringleochloa, Buchloe, Cathestecum,

Cyclostachya, Opizia, Pentarrhaphis, Melanocenchris,

Schaffnerella, Lintonia, Pommereulla, Pleuraphis,

Chloris, Arachne, Afrotrichloris, Apochiton, Crypsis,

Cynodon, Diplachne, Enteropogon, Eustachys,

Trichloris, Astrebla, Brachyachne, Cypholepis,

Eleusine, Tetrapogon, Ochthochloa, Saugetia, Tragus,

Aegopogon, Decaryella, Dignathia, Farrago, Hilaria,

Lepturus, Zoysia, Perotis, Lophiolepis, Lycurus,

Tetrachaete, etc.; eragrostideae: Coelochyrum,

Eleusine, Eragrostis, Acamptoclados, Ectrosia, Harpa-

chne, Heterachne, Panichloa, Steirachne, Viguierella,

Entoplocamia, Desmostachya, Myrios tachya,

Tetrachne, Distichlis, Jouvea, Psilolemma, Crypsis,

Urochondra, Allolepis, Dasyochloa, Erion euron,

Munroa, Dinebra, Trichoneura, Tridens, Aeluropus,

Muhlenbergia, Tripogon, Oropetium, Craspedorhachis,

Kengia, Bewsia, Ctenium, Sporobolus, Monodia,

Plectrachne, Fingerhuthia, Uniola, Neo stapﬁ a,

Orcuttia, Pappophorum, Spartina, etc.;

1.12 CENTOTHECOIDEAE

Annuals or rhizomatous or /and stoloniferous

perennials, herbaceous or reedlike. Calms solid or hol-

low, Leaves typically petiolate with relatively broad tes-

sellate blades; ligules membranous or ciliate, or

membranous with ciliate margins. Inﬂ orescences race-

miform or paniculiform; bracts outside of the spikelets

absent. Spikelets bisexual or unisexual, often com-

pressed laterally, (1-)2 to many-ﬂ owered with reduction

either below or above the fertile ﬂ orets; lemma lacking

uncinate macrohairs, if awned, the awn single; palea

usually well developed, sometimes relatively short; lod-

icules 2 or 0, more or less cuneate; stamens (1-)2–3;

stigmas 2; ovary glabrous, apical appendage absent.

Caryopsis with the hilum basal, usually ﬂ attened later-

ally; endosperm hard, without lipid, containing simple

or compound starch grains; embryo small or large.

n = 12 (Tenório 1976; Clayton 1978, Soderstrom 1981). –

centotheceae: Centotheca, Zeugites, Bromuniola,

Calderonella, Chasmanthium, Chevalierella, Lopha-

therum, Megastachya, Orthoclada; thysanolaeneae:

Thysanolaena.