Takhtajan Armen. Flowering Plants

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726 Subclass IV. COMMELINIDAE

Roisa MM and VL Scatena. 2003. Floral anatomy of Eriocaulon

elichrysoides and Syngonanthus caulescens (Eriocaulaceae).

Flora 198: 188–199.

Rosa MM and VL Scatena. 2007. Floral anatomy of

Paepalanthoideae (Eriocaulaceae, Poales) and their nectarif-

erous structures. Ann. Bot. 99: 131–139.

Saarela JM, HS Rai, JA Doyle, PK Endress, S Mathews,

AD Marchant, B Briggs, and SW Graham. 2007.

Hydatellaceae identiﬁ ed as a new branch near the base of the

angiosperm phylogenetic tree. Nature 446: 312–315.

Sajo MG. 1992. Estudos morfoanatomicos em orgais foliares de

Xyris L. (Xyridaceae). Bol. Bot. Univ. São Paulo 13: 67–86.

Sajo MG and PJ Rudall. 1999. Systematic vegetative anatomy

and ensiform leaf development in Xyris (Xyridaceae). Bot. J.

Linn. Soc. 130: 171–182.

Scatena VL and F Bauman. 2001. Embryology and seed devel-

opment of Paepalanthus sect. Actinocephalus (Koern.)

Ruhland (Eriocaulaceae). Plant Biol. 3: 341–350.

Scatena VL, AM Giulietti, EL Borba, and C van den Berg. 2005.

Anatomy of Brazilian Eriocaulaceae: correlation with tax-

onomy and habitat using multivatiate analyses. Plant Syst.

Evol. 253: 1–22.

Splett S, W Barthlott, T Stützel, and MAG Barros. 1993. Leaf

anatomy of Brazilian Eriocaulaceae and its diagnostic sig-

niﬁ cance. Flora 188: 399–411.

Stevenson DW. 1983. Systematic implications of the ﬂ oral mor-

phology of the Mayacaceae. Am. J. Bot. 70(5, pt.2): 32.

Stevenson DW. 1998. Mayacaceae. In: K Kubitzki, ed. The

families and genera of vascular plants, vol. 4, pp. 294–296.

Springer, Berlin/Heidelberg/New York.

Stevenson DW, M Colella and B Boom. 1998. Rapateaceae. In:

K Kubitzki, ed. The families and genera of vascular

plants, vol. 4, pp. 415–424. Springer, Berlin/Heidelberg/

New York.

Stevenson DW, JI Davis, JV Freudenstein, CR Hardy,

MP Simmonds, and CD Specht. 2000. A phylogenetic

analysis of the monocotyledons based on morphological and

molecular character sets, with comments on the placement of

Acorus and Hydatellaceae. In: KL Wilson and DA Morrison,

eds. Monocots: systematics and evolution, pp.17–24. CSIRO,

Collingwood.

Steyermark JA. 1984. Flora of the Venezuelan Guyana, part 1.

Ann. Missouri Bot. Gard. 71: 297–340.

Stützel T. 1984. Blüten- und inﬂ oreszenzmorphologische

Untersuchungen zur Systematik der Eriocaulaceae. Diss.

Bot. 71: 1–108.

Stützel T. 1985. Die epipetalen Drüsen der Gattung Eriocaulon

(Eriocaulaceae). Beitr. Biol. Pﬂ anz. 60: 271–276.

Stützel T. 1988. Untersuchungen zur Wurzelanatomie der

Eriocaulaceen. Flora 180: 223–239.

Stützel T. 1990. “Appendices” am Gynoeceum der Xyridaceen.

Beitr. Biol. Pﬂ anzen 65: 275–299.

Stützel T. 1998. Eriocaulaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 4, pp. 197–207. Springer,

Berlin/Heidelberg/New York.

Stützel T and N Gansser. 1995. Floral morphology of North

American Eriocaulaceae and its taxonomic implications.

Feddes Repert 106: 495–502.

Stützel T and F Weberling. 1982. Untersuchungen über

Verzweigung und Inﬂ oreszenzaufbau von Eriocaulaceen.

Flora 172: 105–112.

Suessenguth K and R Beyerle. 1936. Über die Xyridaceengat-

tung Abolboda Humb. et Bonpl. Bot. Jahrb. Syst. 67(2):

132–142.

Thanikaimoni G. 1965. Contribution to the pollen morphology

of Eriocaulaceae. Pollen Spores 7: 181–191.

Thieret JW. 1975. The Mayacaceae in the southeastern United

States. J. Arnold Arbor. 56: 248–255.

Thorsch JA and VI Cheadle. 1996. Vessels in Eriocaulaceae.

IAWA J. 17: 183–204.

Tiemann A. 1985. Untersuchungen zur Embryologie,

Blütenmorphologie, und Systematik der Rapateaceen und

der Xyridaceen-Gattung Abolboda (Monocotyledoneae).

Diss. Bot. 82: 1–202.

Tomlinson PB. 1964. Notes on the anatomy of Triceratella

(Commelinaceae). Kirkia 4: 207–212.

Uphof JCT. 1924. The physiological anatomy of Mayaca ﬂ uvia-

tilis. Ann. Bot. 38: 389–393.

Venturelli M and F Bouman. 1988. Development of ovule and

seed in Rapateaceae. Bot. J. Linn. Soc. 97: 267–294.

Venturelli M and F Bouman. 1986. Embryology and seed devel-

opment in Mayaca ﬂ uviatilis (Mayacaceae). Acta Bot. Neerl.

35: 497–516.

Vyshenskaya TD. 1985a. Rapateaceae, Xyridaceae. In: A

Takhtajan, ed. Comparative seed anatomy, vol. 1, pp. 167–169.

Nauka, Leningrad (in Russian).

Vyshenskaya TD. 1985b. Mayacaceae, Eriocaulaceae. In: A

Takhtajan, ed. Comparative seed anatomy, vol. 1, pp.

177–183. Nauka, Leningrad (in Russian).

Vyshenskaya TD. 1985c. Hydatellaceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol. 1, pp. 190–191. Nauka,

Leningrad (in Russian).

Weinzieher S. 1914. Beiträge zur Entwicklungsgeschichte von

Xyris indica L. Flora 106: 393–432.

Order 27. DASYPOGONALES

Mainly xeromorphic, rhizomatous perennial herbs or

shrubby or arborescent plants sometimes with second-

ary growth. Vessels in root, with simple or scalariform

(Baxteria, Kingia) perforations. Raphides present or

(Calectasiaceae) absent. Sieve-element plastids of

P2c-type (Lomandraceae). Leaves small to large, alter-

nate, leathery or modiﬁ ed into spines, linear or strap-

shaped, more or less V- or U-shaped in transection,

often distichous or crowded in a rosette, rarely scat-

tered along the stem, usually with broad, often sheath-

ing base that persists after lamina falls or is burnt off.

The mesophyll containing calcium oxalate crystals or

without calcium oxalate crystals. Stomata anomocytic

or (Baxteria) paracytic/tetracytic. Flowers in various

kinds of inﬂ orescences or solitary, bisexual or dioe-

cious, actinomorphic, 3-merous, bracteate. Perianth

segments six in two cycles, mostly dry and gluma-

ceous, similar, more or less free or united basally or

Superorder COMMELIANAE 727

(Calectasia) into a tube, persistent. Stamens six in two

cycles, usually attached to base of perianth segments;

anthers tetrasporangiate, basiﬁ xed or (Dasypogon)

dorsiﬁ xed, opening longitudinally or by apical pores,

introrse or rarely extrorse; the endothecial thickenings

spiral, or girdling (Lomandra). Pollen grains mostly

1-colpate and microreticulate or psilate or (Baxteria)

the exine is divided into geometrically deﬁ ned areas, a

nearly unique type among ﬂ owering plants (Chanda

and Ghosh 1976; Clifford et al. 1998). Gynoecium of

three united carpels; stylodia more or less completely

connate; ovary superior or inferior, 3-locular or

1-locular (Dasypogon), with 1 ovule per carpel, with

or without septal nectaries. Ovules anatropous or rarely

campylotropous, bitegmic, crassinucellate. Female

gametophyte of Polygonum-type. Endosperm nuclear.

Fruits dehiscent (capsules or capsular-indehiscent) or

more often indehiscent, enclosed in persistent perianth.

Seeds more or less globose or oblong, with pale yellow

testa; seed coat formed predominantly by the outer

integument; embryo lens-shaped and broad; endosperm

copious, without starch, with a mealy, very thin-walled

parenchyma (at least in Dasypogon). Testa without

phytomelan. Cell wall-bound ferulic acid is present in

all genera (Rudall and Caddick 1994); rarely (only in

one species of Dasypogon) is present chelidonic acid;

in Lomandra detected arthroquinones; n = 7, 8, 9.

A rather isolated group, which, according to Clifford

et al. (1998), is possibly near the Rapateaceae; proba-

bly belong to the Commelinanae.

Key to Families

1 Anthers opening longitudinally.

2 Fruits capsules.

3 Fruits explosively dehiscent capsules; dehis-

cence septifragal, the valves also splitting into

two valves, each half with the inner wall form-

ing a plate held under tension and springing

upward to eject the seed. Perennial stemless

herbs with short, thick rhizomes and thick

roots. Vessels in roots, with scalariform perfo-

rations. Leaves crowded, linear, pungent, with

sheathing bases. Flowers large, 7–8 cm long,

solitary, terminal, sessile, surrounded by sev-

eral bracts, scented like rotten meat. Perianth

segments free, almost similar, indurated, per-

sistent. Stamens six, attached to the bases of

the perianth; anthers apiculate, introrse. Pollen

grains of unique pantocolpate type: the surface

divided by the apertures into geometrically

deﬁ ned areas; the aperture membrane formed

by coarse pieces, the surface pattern of which

is essentially the same as that of the interaper-

turate areas; it seems as if the ends of the colpi

of pantocolpate grains are united to make a

continuous aperture that encloses several

zones of geometrical pattern; ornamentation

microreticulate (Chanda and Ghosh 1976:

538). Ovary 3-locular, with one erect ovule

per locule; style subulate, more or less as long

as perianth, with small 3-lobed stigma. Seeds

globose. n = 7.. . . . . . . . . . 1. baxteriaceae.

3 Fruits loculicidal capsules, ovoid or trigonous.

Perennial tufted herbs with generally short rhi-

zomes and wiry roots or (Lomandra) herbs or

shrubs with thick, woody rhizomes with sec-

ondary thickening and ﬁ brous roots. Stems

elongated (erect or procumbent) or very short.

Vessels only in roots with simple and scalari-

form perforations or (Acanthocarpus) also in

leaves, with scalariform perforations. Leaves

alternate, scleromorphic, perennial or

(Xerolirion) deciduous, generally distichous

or spirodistichuous, sheathing at base, linear,

sometimes prickly or scarious. Stomata gener-

ally anomocytic, although paracytic and tetra-

cytic in some Lomandra species. Flowers in

usually contracted cymose or racemose inﬂ o-

rescences or solitary, subtended by single to

several bracts, bisexual or (Lomandra and

Xerolirion) unisexual and dioecious, usually

pedicillate, pedicels articulated or (Xerolirion)

not articulated. Perianth segments free or

shortly united basally, ﬂ eshy, petaloid or

sometimes chartaceous, persistent. Stamens

six in two cycles, more or less inserted on the

perianth segments; anthers dorsiﬁ xed. Seeds

testal; present napthoquinones (Lomandra);

n = 7, 8. . . . . . . . . . . . . . .2. lomandraceae.

2 Fruits indehiscent, within persistent perianth.

Perennial herbs or secondary arborescent plants

with short rhizome or stout caudex of ﬁ brous pith

surrounded by persistent packed leaf bases.

Roots thick or ﬁ brous. Stems sometimes (Kingia)

with a cover of aerial roots penetrating the per-

sistent leaf bases. Vessels in roots, with scalari-

form (Kingia) or simple (Dasypogon)

perforations. Leaves linear, imbricate at base,

728 Subclass IV. COMMELINIDAE

sheathing. Inﬂ orescence a globose head on ter-

minal peduncle bearing scattered bracts

(Dasypogon) or axillary headlike, their pedun-

cles covered by imbricate sheathing bracts.

Flowers small, sessile; perianth segments narrow

and dry; outer segments free or (Dasypogon)

united above the middle and thickened; inner

segments free or almost free. Stamens inserted

on perianth; anthers dorsiﬁ xed, versatile

(Dasypogon) or basiﬁ xed, introrse (Kingia).

Pollen grains 1-colpate, ﬁ nely reticulate or punc-

tate. Gynoecium of three carpels; style subulate,

stigma small; ovary sessile, 3- locular, with one

ovule per locule; style long, with punctiform

stigma. Septal nectaries present as shallow basal

grooves. n = 7.. . . . . . . . . .3. dasypogonaceae.

1 Anthers opening by two apical pores. Xeromor-

phic, rhizomatous perennial herbs, shrublets or

shrubs, with wiry roots. Branches stiff, covered by

old leaf sheaths or leaves. Vessels in roots, with

simple perforations. Leaves alternate, small, linear

or lanceolate, simple, entire, pungent, the base

indurated and closely sheathing stem. Flowers

bisexual, solitary, terminal on the ends of short

branchlets, surrounded by many scarious, small

imbricate leaves. Perianth segments connate into a

short tube, the lobes spreading, stifﬂ y scarious,

lilac-blue or purple, variously pubescent outside.

Stamens six inserted at the throat of perianth lobes,

all equal to markedly unequal, free of one another.

Anthers basiﬁ xed, terminally dehiscing via pores

to dehiscing via short slits. Pollen grains 1- colpate,

reticulate. Gynoecium of three carpels; style long,

slender, attenuate from the ovary, much longer

than the ovary. Ovary 1-locular, with three basal,

anatropous ovules; style ﬁ liform, with punctiform

stigma. Septal nectaries absent. Fruits nuts,

enclosed in persistent faded perianth. Seeds

elongate, with thin membranous testa, n = 9.. . . .

4. calectasiaceae.

1. BAXTERIACEAE

Takhtajan 1996. 1/1. Southwestern Australia.

Baxteria.

A rather isolated family characterized by very large

solitary ﬂ owers, unique type of multiaperturate pollen

grains, and septifragal capsules.

2. LOMANDRACEAE

Lotsy 1911. 5/80 Australia, New Guinea, New Caledonia.

Lomandra, Romnalda, Chamaexeros, Xerolirion,

Acanthocarpus.

Closely related to the Dasypogonaceae.

3. DASYPOGONACEAE

Dumortier 1829 (including Kingiaceae Endlicher

1838). 2/4. Southwestern Australia.

Dasypogon, Kingia.

4. CALECTASIACEAE

Endlicher 1836. 1/3. Southwestern and southeastern

Australia.

Calectasia.

Probably closely related to the Dasypogonaceae

(both are characterized by large chalazosperm, see

Rudall [1994]), but differ from them markedly in

anthers opening by apical pores, and solitary ﬂ owers on

the ends of short branchlets.

Bibliography

Anway JC. 1969. The evolution and taxonomy of Calectasia cta-

nea R. Br. (Xanthorrhoeaceae) in terms of its present day vari-

ation and cytogenetics. Austral. J. Bot. 17: 147–159.

Barrett RL and Dixon KW. 2001. A revision of the genus

Calectasia (Calectasiaceae) with eight new species

described from south-west Western Australia. Nuytsia 13:

411–448.

Bedford DJ, AT Lee, TD MacFarlane, RJF Henderson, and

AS George. 1986. Xanthorrhoeaceae. Fl. Austral. 46:

88–171.

Briggs BG. 1986. Chromosome numbers in Lomandra

(Dasypogonaceae). Telopea 2: 741–744.

Chanda S and K Ghosh. 1976. Pollen morphology and its evolu-

tionary signiﬁ cance in Xanthorrhoeaceae. In: IK Ferguson

and J Muller, eds. The evolutionary signiﬁ cance of the exine,

pp. 527–559. Linn. Soc. Symposium, No. 1. London/

New York.

Chanda S, B Lugardon, and G Thanikaimoni. 1978. On the

ultrastructure of pollen apertures in Calectasia R.Br.

(Xanthorrhoeaceae). Pollen Spores 20: 351–356.

Clifford HT, GJ Keighery, and JG Conran. 1998. Dasypogonaceae.

In: K Kubitzki, ed. The families and genera of vascular

plants, vol. 4, pp. 190–194. Springer, Berlin/Heidelberg/

New York.

Superorder JUNCANAE 729

Conran JG. 1998e. Lomandraceae. In: K Kubitzki, ed. The fami-

lies and genera of vascular plants, vol. 3, pp. 354–365.

Springer, Berlin/Heidelberg/New York.

Fahn A. 1954. The anatomical structure of the Xanthorrhoeaceae

Dumort. Bot. J. Linn. Soc. 55: 158–184.

Keighery GJ. 1983. Balistochory (explosive seed dispersal) in

Baxteria R. Br. (Xanthorrhoeaceae). W. Austral. Nat. 15:

163–166.

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.

Rudall P. 1994. The ovule and embryo sac in Xanthorrhoeaceae

sensu lato. Flora 189: 335–351.

Rudall P and MW Chase. 1996. Systematics of Xanth-

orrhoeaceae sensu lato, evidence for polyphyly. Thelopea 6,

629–647.

Superorder JUNCANAE

Order 28. JUNCALES

Perennial, often rhizomatous herbs or less often

annuals, rarely shrubs or lianas, mostly of wet habitats,

rarely aquatic, very rarely (Prionium) rather large

shrubby plants with hard persistent trunk (homologous

to the rhizome of other genera) without secondary

growth. Rhizomes underground, sympodial, some-

times tuberous, starchy. Stems commonly trigonous,

rarely compressed, very rarely articulated, mostly

solid, but sometimes hollow or transversely septate.

Raphides absent, but silica bodies commonly present.

Vessels usually in all vegetative organs, with scalari-

form or/and simple perforations. Leaves alternate, tris-

tichous or more rarely distichous or polystichous, basal

or cauline, usually sheathing at base, sheats usually

closed, sessile or rarely pseudopetiolate, mostly linear,

parallel-veined, rarely reduced or absent; ligule pres-

ent or absent. Stomata paracytic or sometimes tetra-

cytic. Flowers small and inconspicuous, sessile in axils

of spirally arranged or distichous bracts, forming

spikes or spikelets that are sometimes solitary and ter-

minal, but usually arranged in secondary inﬂ ores-

cences, bisexual or often unisexual (monoecious or

very rarely dioecious). Perianth segments six (rarely 4)

in two cycles, free, glumaceous or herbaceous, equal

or subequal, sometimes replaced by bristles or numer-

ous hairs or lacking; perianth in some genera (such as

Oreobolus) of six scales in two cycles but usually

strongly reduced into bristles or hairs, or wanting.

Stamens mostly three (1 medio-anterior, 2 lateral), less

often reduced to two or one, rarely six or more, in two

cycles, the inner cycle sometimes lacking; ﬁ laments

generally free from each other; anthers tetra-

sporangiate, basiﬁ xed, introrse, opening longitudinally,

often apiculate. Tapetum secretory. Microsporogenesis

simultaneous. Pollen grains 3-celled, tectate-

columellate, 1-porate or less often up to 7-porate, in

monads (Mapanioideae) (Tarasevich 1986) or more

often borne in cryptotetrads (the 4 microspores being

retained within a common wall, but three of them soon

degenerate so that at maturity pollen grain appears to

be single), or (in Juncaceae and Thurneraceae) usually

in tetrahedral tetrads that are surrounded by the com-

mon tetrad wall consisting of sporopollenin, atectate,

with one reduced, poroid, distal aperture. Gynoecium

of three or less often two, rarely four, united carpels;

stylodia free or more often connate into short, linear

style, often swollen at base, with three, two, or very

rarely eight ﬁ liform, linear, or lanceolate stigmatic

branches. Ovary superior, 3-locular or less often

1- locular, with one to many basal ovules per carpels.

Ovules anatropous, bitegmic (outer integument very

thin), crassinucellate, with parietal cell. Female game-

tophyte of Polygonum-type. Endosperm helobial or

(Cyperaceae) nuclear. Fruits commonly trigonous,

lenticular or globose nuts, rarely drupelike, or three to

many-seeded loculicidal capsules or sometimes cir-

cumscissile. Seeds small, free from the pericarp; seed

coat testal-tegmic, thin and delicate, consists of two lay-

ers formed by outer epidermis of the outer integument

and inner integument of the inner integument; embryo

small, basal, usually more or less differentiated, sur-

rounded by abundant endosperm containing lipids,

aleurone, and simple starch grains. Aleurones, ﬂ avonoid

sulphates, ﬂ avone C-glycosides, kestose and isokestose

storage oligosaccharides present. Chromosomes often

with diffuse centromere. n = 3-13 +.

Related to the Commelinanae, and possibly to the

Xyridales.

Key to Families

1 Plants without silica bodies. Annual or perennial

rhizomatous or tuberous herbs or rarely shrubs (a

few with woody trunks, esp. Prionium). Plants lack-

ing crystal oxalate raphides, but accumulated starch

exclusively “pteridophyte type”. Vessels in roots,

stems and leaves, with simple, or scalariform and

simple perforations. Leaves evergreen, alternate,

730 Subclass IV. COMMELINIDAE

nearly always tristichous, or rarely (Distichia) dis-

tichous, folded, or rolled, terete or ﬂ at, leathery or

membranous and sometimes reduced to membra-

nous sheaths; sessile, simple; Prionium has leaves

in terminal rosette, linear, tapering, ﬂ at or canalicu-

late, serrate. Stomata paracytic. Inﬂ orescences ter-

minal or pseudolateral, compound, of open cymose

panicles, or often congested and head- or spike-like,

or ﬂ owers sometimes solitary and terminal

(Rostkovia, Marsippospermum) or lateral (Distichia,

Oxychloe and Patosia). Flowers with one or more

spathal bracts, bracteolate or ebracteolate, small,

actinomorphic, bisexual or unisexual. Perianth seg-

ments six in two cycles, or rarely three, similar or

sometimes very different in size, free, glumaceous.

Stamens 6 or rarely 2–3; ﬁ laments ﬁ liform of ﬂ at

and widened at the base; anthers introrse or latrorse,

appendages or unappendaged. Pollen ulcerate with

granular exine, in tetrahedral and cross tetrads.

Ovary 1-locular or 3- locular; styles one or three,

free of partially joined; stigmas two, brushlike.

Ovules 3 (Luzula), or 7–100 per locule, ascending.

Fruits mostly loculicidal capsules, but in Distichia,

Oxychloe and Patosia less well deﬁ ned and some-

times circumscissile (Baslev 1996). Seeds numerous

or only three (Luzula), ovoid or rarely fusiform, glo-

bose or angular, not hispid, usually not pointed at

ends, with (mucilaginous) exotesta and endotegmen;

testa without phytomelan; embryo straight, small;

endosperm starchy. Free ﬂ avones present (querce-

tin); n = 3–42, mostly 6, 12, 18, 24. 2. juncaceae.

1 Plants with silica bodies.

2 Ovary 3-locular. Large, tough herbs with large,

subterranean rhizomes. Silica bodies spheroidal,

present in some of the cells of parenchyma and

epidermis. Vessels in roots, stems and leaves,

mostly with scalariform perforations. Leaves all

basal, alternate, fourranked, or tristichous, ﬂ at or

canaliculate, leathery, sessile, sheathing, simple,

entire; leaf bundles in pairs above each other in a

leaf transaction, the lower (and smaller) bundle of

a pair with the phloem on top ( adaxial), facing the

phloem of the upper bundle. Stomata paracytic,

but sometimes some of them tetracytic.

Inﬂ orescences of one or more dense, racemose,

globose to ellipsoid heads subtended by spread-

ing, leafy bracts on an otherwise leaﬂ ess, bluntly

triangular or quadrangular peduncle. Flowers

bisexual, actinomorphic, small, with short,

swollen, puberulous pedicles. Perianth segments

six, free, similar, brown, spotted, persistent.

Stamens six, in two cycles, adnate to the base of

the perianth segments and free of one another;

anthers introrse. Styles united and very short,

attenuate from the ovary; stigmas three, elongate.

Ovules 1–7 per locule, ascending, with a zig-zag

micropyle, placentation basal. Fruits loculicidal

capsules. Seeds spindle-shaped, hispid, subulate-

pointed at both ends; testa without phytomelan;

embryo small, straight, cylindrical; endosperm

copious, mealy, starchy. . . . . . . 1. thurniaceae.

2 Ovary 1-locular. Small to tall perennial or annual

herbs, rarely dwarf shrubs, or lianas, terrestrial

or helophytic, rarely aquatic; the perennials

tufted, rhizomatous, stoloniferous, tuberous.

Stems with solid internodes to with spongy

internodes, or with hollow internodes. Vessels

with scalariform, or simple, or scalariform and

simple perforations. Sieve-element plastids of

P-type. Leaves all basal or one to few cauline,

rarely all cauline, alternate, distichous or tristic-

hous, rarely spiral; ﬂ at or folded, sheathing,

simple, entire, but commonly with prickle hairs.

Inﬂ orescences terminal, more rarely pseudola-

teral, very rarely reduced to a single spikelet.

Flowers bracteate or ebracteate, bracteolate,

usually small to minute, bisexual or (function-

ally) unisexual, and then spikelets bisexual or

unisexual, rarely dioecious. Perianth segments

vestigial (represented by bristles or hairs) and

than 6, free, sometimes more or less sepaloid, or

absent. Stamens usually 3 (1 medio-anterior, 2

lateral), but sometimes reduced to 1–2 or 4–6,

and even more numerous (to 22); ﬁ laments some-

times strongly elongating after anthesis, rarely

connate; anthers introrse or latrorse, appendaged

or unappendaged. Gynoecium of 2, or 3(-4)

carpels; styles 2–3, free to partially joined; stig-

mas (1-)2, or 3(-15). Ovule solitary, basal, biteg-

mic, outer integument not contributing to the

micropyle. Fruits achene-like or rarely drupa-

ceous. Seeds with thin testa (phytomelan lack-

ing), raphe and chalaza usually conspicuous;

embryo small, basal, surrounded by abundant

mealy or oily endosperm. Alkaloids and proantho-

cyanidins present or absent; when present – cyani-

din and delphinidin; ﬂ avonols present (quercetin)

or absent; n = 5 or more. . . . . . . . 3. cyperaceae

Superorder JUNCANAE 731

1. THURNIACEAE

Engler 1907. 1/3. Tropical South America (Guyana

Highlands and Brazil).

Thurnia.

Close related to the Juncaceae, but vessels mostly

with scalariform perforations and silica bodies

present.

2. JUNCACEAE

A.L de Jussieu 1789 (including Prioniaceae Munro et

Linder 1998). 8/400-440. The genera Juncus and

Luzula distributed mostly in the cold and temperate

regions of the Northern Hemisphere, the rest in the

Southern Hemisphere; Prionium – Cape Province of

South Africa.

Marsippospermum, Rostkovia, Distichia, Patosia,

Oxychloe, Luzula, Juncus, Prionium.

The taxonomic position of Oxychloe (7, Andes) is

uncertain. Some authors prefer to place it in Cyperaceae

(Plunkett et al. 1995; Thorne 2000), but morphological

traits of Oxychloe are typically juncaceous (Drabkova

et al. 2003). The genus Prionium starts apart in many

characters, especially in its woody trunk and leaf anat-

omy (Cutler 1969; Munro and Linder 1997, 1998), but

in ﬂ oral morphology and pollen tetrads is juncaceous.

3. CYPERACEAE

A. L. de Jussieu 1789 (including Caricaceae Burnett

1835, Elynaceae Reichenbach 1828, Kobresiaceae

Gilly 1952, Mapaniaceae Shipunov 2003, Papyraceae

Burnett 1835, Scirpaceae Batsch ex Borkhausen 1797).

108/5300. Cosmopolitan, but mainly in cold and tem-

perate regions.

Classiﬁ cation after P. Goetghebeur (1998).

3.1 MAPANIOIDEAE

Pollen grains in monads. Spike-lets cymose with one

terminal female ﬂ ower and several axillary male

ﬂ owers. Intraspicular prophylls two, always opposite,

scalelike or rarely saclike. – hypolytreae: Scirpo-

dendron, Hypolytrum, Principina, Mapania, Mapan-

iopsis, Paramapania, Diplasia; chrysitricheae:

Capitularina, Exocarya, Chori zandra, Lepironia,

Hellmuthia, Chrysitrix.

3.2 CYPEROIDEAE

Pollen grains in cryptomonads. Spikelets without

terminal ﬂ ower, never prophyllate, many-ﬂ owered.

Flowers subtended by ﬂ oral scale, generally bisexual. –

scirpeae: Scirpus, Eriophorum, Phylloscirpus, Oreo-

bolopsis, Amphiscirpus, Trichophorum (including

Baeothryon); fuireneae: Fuirena, Pseudoschoenus,

Bolboschoenus, Actinoscirpus, Schoenoplectus; eleo-

charideae: Eleocharis, Egleria, Websteria; abild-

gaardieae: Fimbristylis, Dichostylis, Crosslandia,

Bulbostylis, Abildgaardia, Nemum, Nelmesia; cypereae:

Ficinia, Desmoschoenus, Isolepis, Scirpoides, Oxyca-

ryum, Kyllingiella, Androtrichum, Cyperus, Juncellus,

Mariscus, Torulinum, Courtoisina, Remirea, Sphaero-

cyperus, Ascopholis, Alinula, Ascolepis, Pycreus,

Queen slandiella, Kyllinga, Volkiella, Lipocarpha;

dulichieae: Dulichium, Sumatroscirpus, Blysmus;

schoeneae: Arthrostylis, Actinoschoenus, Tricho-

schoenus, Trachystylis, Rhynchospora, Pleuro stachys,

Schoenus, Gymnoschoenus, Mesomelaena, Ptilothrix,

Cyathochaeta, Oreobolus, Carpha, Trianoptiles, Tetraria,

Cyathocoma, Neesenbeckia, Epischoenus, Costularia,

Gahnia, Morelotia, Reedia, Evandra, Caustis,

Cladium, Rhynchocladium, Macha erina, Lepido sperma,

Tricostularia.

3.3 SCLERIOIDEAE

Spikelets with few glumes, one to few of which each

subtend a strictly unisexual ﬂ ower. A perianth often

present. – cryptangieae: Lagenocarpus, Didymi-

andrum, Everardia, Cephalocarpus; trilepideae:

Trilepis, Afrotrilepis, Coleochloa, Microdracoides;

sclerieae: Scleria; bisboeckelereae: Becquerelia,

Diplacrum, Bisboeckelera, Calyptrocarya.

Insertae sedis: Exochogyne, Koyamaea.

3.4 CARICOIDEAE

Pollen grains in cryptomonads. Spikelets without ter-

minal ﬂ ower, female ﬂ owers subtended by strongly

metamorphosed prophyll, which is bottle-shaped

or spathaceous; ﬂ owers always unisexual, without

perianth. – Kobresia, Schoenoxiphium, Uncinia,

Vesicarex, Carex, Cymophyllus.

Have many similarities with the Juncaceae (see espe-

cially Shah 1967; Simpson 1995). According to Simpson

characters linking Juncaceae and Cyperaceae are the

following: leaves usually tristichous, stomata usually

paracytic, calcium oxalate raphides absent, chromo-

somes holocentric, occurrence of postreductional

732 Subclass IV. COMMELINIDAE

meiosis, perianth segments bracteal, microsporogenesis

simultaneous, developing pollen grains in tetrads, pol-

len grains ulcerate, embryogeny onagrad, presence of

luteolin 5-methyl ether, and parasitism by similar rust

and smut species. However, the Cyperaceae differ from

the Juncaceae in the presence of silica bodies,

Cyperaceae-type simultaneous microsporogenesis,

nuclear endosperm development, pollen in pseudo-

monads, solitary ovule per locule, indehiscent fruits,

presence of yellow aurone pigments and ﬂ avone tricin

(Simpson 1995).

The Cyperaceae could either derived from some

ancient juncalean ancestor or more probably had a

common origin. It is important to note that according

to Cheadle (1955) no species in the Juncaceae has ves-

sels quite primitive enough throughout the plant to per-

mit use of the Juncaceae as the source of the most

primitive members of the Cyperaceae. Besides, the

pollen grains of the Cyperaceae, especially those of

Mapanioideae, are less specialized.

Bibliography

Balslev H. 1996. Juncaceae. Flora Neotropica Monograph.

New York Botanical Garden, New York.

Balslev H. 1998. Juncaceae. In: K Kubitzki, ed. The families and

genera of vascular plants, vol. 4, pp. 252–269. Springer,

Berlin/Heidelberg/New York.

Barnard C. 1958. Floral histogenesis in monocotyledons: III.

The Juncaceae. Austral. J. Bot. 6: 285–298.

Blaser NW. 1941, 1944. Studies in the morphology of the

Cyperaceae: I. Morphology of ﬂ owers: A. Scirpoid genera.

B. Rhynchosporoid genera. II. The prophyll. Am. J. Bot. 28:

542–551, 832–838, 1941; 31: 53–54, 1944.

Bruhl J. 1995. Sedge genera of the world: relationships and a new

classiﬁ cation of the Cyperaceae. Austral. Syst. Bot. 8: 125–305.

Bruhl J, L Watson, and MJ Dallwitz. 1992. Genera of Cyperaceae:

Interactive identiﬁ cation and information retrieval. Taxon

41: 225–234.

Cheadle VI. 1955. The taxonomic use of specialization of ves-

sels in the metaxylem of Gramineae, Cyperaceae, Juncaceae,

and Restionaceae. J. Arnold Arbor. 36: 141–157.

Cheadle VI and H Kosakai. 1972. Vessels in the Cyperaceae.

Bot. Gaz. 133: 214–223.

Cheadle VI and H Kosakai. 1973. Vessels in Juncales:

I. Juncaceae and Thurniaceae. Phytomorphology 23: 80–87.

Cutler DF. 1965. Vegetative anatomy of Thurniaceae. Kew Bull.

19: 431–441.

Cutler DF. 1969. Juncales. In: CR Metcalfe, ed. Anatomy of the

monocotyledons, vol. 4. Clarendon Press, Oxford.

Drábková K, J Kirschner, O Seberg, G Petersen, and C

Vlc

ek.

2003. Phylogeny of the Juncaceae based on rbcL sequences,

with special emphasis on Luzula DC and Juncus L. Plant

Syst. Evol. 240: 133–147.

Dunbar A. 1973. Pollen development in the Eleocharis palustris

group (Cyperaceae): I. Ultrastructure and ontogeny. Bot.

Not. 126: 197–254.

Eiten LT. 1976. Inﬂ orescence units in the Cyperaceae. Ann.

Missouri Bot. Gard. 63: 81–112.

Goetghebeur P. 1986. Genera Cyperacearum. Masters thesis,

University of Ghent.

Goetghebeur P. 1998. Cyperaceae. In: K Kubitzki, ed. The fami-

lies and genera of vascular plants, vol. 4, pp. 141–198.

Springer, Berlin/Heidelberg/New York.

Holttum RE. 1948. The Spikelets in Cyperaceae. Bot. Rev. 14:

525–541.

Kirpes CC, LG Clark, and NR Lersten. 1996. Systematic signiﬁ -

cance of pollen arrangement in microsporangia of Poaceae

and Cyperaceae, review and observations on representative

taxa. Am. J. Bot. 83: 1609–1622.

Kirschner J. 2002a. Species plantarum: Flora of the World,

Part 6. Juncaceae 1: Rostkovia to Luzula. Australian

Biological Resources Study, Canberra.

Kirschner J. 2002b. Species plantarum: Flora of the World,

Part 7. Juncaceae 2: Juncus subgenus Juncus. Australian

Biological Resources Study, Canberra.

Kirschner J. 2002c. Species plantarum: Flora of the World,

Part 8. Juncaceae 3: Juncus subgenus Agathryon. Australian

Biological Resources Study, Canberra.

Kovtonjuk N. 1999. Systematic signiﬁ cance of seed surfaces in

some Juncaceae and Caryophyllaceae. In: MH Kurmann,

AR Hemsley, eds. The evolution of plant architecture,

pp. 367–374. Royal Botanic Gardens, Kew.

Koyama T. 1961. Classiﬁ cation of the family Cyperaceae, part 1.

J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 8: 37–148.

Kristiansen KA, M Cilieborg, L Drábková, T Jørgensenm,

G Petersen, O Seberg. 2005. DNA taxonomy – the riddle of

Oxychloë (Juncaceae). Syst. Bot. 30: 284–289.

Kubitzki K. 1966. Untersuchungen über den Blütenbau von

Oreobolus R. Br. Bot. Jahrb. Syst. 85: 80–87.

Kubitzki K. 1998. Thurniaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 4, pp. 455–457. Springer,

Berlin/Heidelberg/New York.

Kukkonen I. 1984. On the inﬂ orescence in the family Cyperaceae.

Ann. Bot. Fenn. 21: 257–264.

Laurent M. 1904. Recherches sur le développement des Joncées.

Ann. Sci. Nat. Bot. 19: 97–194.

Lye KA. 2000. Achene structure and function of structure in

Cyperaceae. In: KL Wilson and DA Morrison, eds. Mono-

cots: systematic and evolution, pp. 615–628. CSIRO,

Collingwood.

Makde KH. 1982. Pollen development in the Cyperaceae.

J. Indian Bot. Soc. 61: 242–249.

Makde KH and SM Bhuskute. 1984. Trends of specialization in

endosperm of Cyperaceae. Current Science 53: 504–506.

Metcalfe CR. 1971. Cyperaceae. In: CR Metcalfe, ed. Anatomy

of the monocotyledons, vol. 5. Clarendon Press, Oxford.

Michalska A. 1953. Cytological investigations on Luzula. Acta

Soc. Bot. Polon. 22: 169–186.

Muasya AM, JJ Bruhl, DA Simpson, A Culham, and MW Chase.

2000. Suprageneric phylogeny of Cyperaceae: a combined

analysis. In: KL Wilson and DA Morrison, eds. Mono-

Superorder JUNCANAE 733

cots: systematic and evolution, pp. 593–601. CSIRO,

Collingwood.

Muasya AM, DA Simpson, MW Chase, and A Culham. 1998.

An assessment of the suprageneric phylogeny in Cyperaceae

using rbcL DNA sequences. Plant Syst. Evol. 211:

257–271.

Munro SL and HP Linder. 1997. The embryology and system-

atic relationships of Prionium serratum (Juncaceae:

Juncales). Am. J. Bot. 84: 850–860.

Munro SL and HP Linder. 1998. The phylogenetic position of

Prionium (Juncaceae) within the order Juncales based on

morphological and rbcL sequence data. Syst. Bot. 23: 43–55.

Nemirovich-Danchenko EN. 1985. Thurniaceae. In: A Takhtajan,

ed. Comparative seed anatomy, vol. 1, pp. 161–162. Nauka,

Leningrad (in Russian).

Plunkett GM, DE Soltis, PS Soltis and RE Brooks. 1995.

Phylogenetic relationships between Juncaceae and

Cyperaceae, insights from rbcL sequence data. Am. J. Bot.

82: 520–525.

Reihmann D. 1977. Vergleichende Embryologie und system-

atische Stellung der Juncaceae mit neuen Beiträgen zur

Samenentwicklung von Juncus und Luzula. Thesis,

University of Bochum.

Richards J, JJ Bruhl, and KL Wilson. 2005. Understanding ﬂ oral

morphology of mapaniid Cyperaceae: development of repro-

ductive structures in Exocarya scleroides. In: Botany 2005.

Learning from plants (Abstracts), pp. 37–38.

Roalson EH. 2005. Phylogenetic relationships in the Juncaceae

inferred from nuclear ribosomal DNA internal transcribed

spacer sequence data. Int. J. Plant Sci. 166: 397–413.

Schneider M. 1932. Untersuchungen über die Embryobildung

und Entwicklung der Cyperaceen. Beih. Bot. Centralbl. 49:

649–674.

Schultze-Motel W. 1959. Entwicklungsgeschichtliche und ver-

gleichend-morphologische Untersuchungen im Blütenbereich

der Cyperaceae. Bot. Jahrb. Syst. 78: 129–170.

Shah CK. 1967. A taxonomic evaluation of the families

Cyperaceae and Juncaceae. Bull. Nation. Inst. Sci. India 34:

248–256.

Shah CK. 1968. Development of pericarp and seed coat in the

Cyperaceae. Naturaliste Canad. 95: 39–48.

Shah CK. 1974. Morphology and embryology of the family

Cyperaceae. Adv. Plant Morph. 1972: 102–112.

Simpson D. 1995. Relationships within Cyperales. In: Rudall

PJ, PJ Cribb, DF Cutler, and CJ Humphries, eds.

Monocotyledons: systematics and evolution, vol. 2, pp.

497–509. Royal Botanic Gardens, Kew.

Simpson D. 1998. Mapaniaceae – fact or ﬁ ction? In Mono-

cots II, p. 50 (Abstract). Sydney.

Simpson DA, CA Furness, TR Hodkinson, AM Muasya, and

MW Chase. 2003. Phylogenetic relationships in Cyperaceae

subfamily Mapanioideae inferred from pollen and plastid

DNA sequence data. Am. J. Bot. 90: 1971–1086.

Smith DL and JS Faulkner. 1976. The inﬂ orescence of Carex

and related genera. Bot. Rev. 42: 53–81.

Snell RS. 1936. Anatomy of the spikelets and ﬂ owers of Carex,

Kobresia, and Uncinia. Bull. Torrey Bot. Club. 63:

277–295.

Starr JR, RJ Bayer, and BA Ford. 1999. The phylogenetic posi-

tion of Carex section Phyllostachys and its implications for

phylogeny and subgeneric circumscription in Carex

(Cyperaceae). Am. J. Bot. 86: 563–577.

Starr JR, SA Harris, and DA Simpson. 2004. Phylogeny of the

unispicate taxa in Cyperaceae tribe Cariceae I: generic rela-

tionships and evolutionary scenarios. Syst. Bot. 29: 528–544.

Starr JR, Teoh V, E Roalson, AM Muasya, and DA Simpson.

2006. Towards a phylogenetic classiﬁ cation of sedges

(Cyperaceae): chloroplast (rbcL, matK, NdhF) and nuclear

(ADC) data. In: Botany 2006 – Looking to the future, con-

serving the past (Abstracts), pp. 258–259.

Tarasevich VF. 1986. On the position of the family Cyperaceae

in the system of ﬂ owering plants in connection with the

palynological data. In: VN Tikhomirov, ed. Sources of infor-

mation in the phylogenetic systematics of plants, pp. 75–76.

Nauka, Moscow (in Russian).

Tejavathi DH. 1987. Seed development in some members of

Cyperaceae. Beitr. Biol. Pﬂ anz. 62: 43–55.

Thimm U. 1985. Zur Embryologie, Blüten- und Fruchtanatomie

der isolierten Juncales-Gattungen Prionia und Thurnia.

Diploma Thesis Bochum. University of Bochum, Division of

Biology.

Tucker GC. 1987. The genera of Cyperaceae in the SE United

States. J. Arnold Arbor. 68: 361–445.

Van der Veken P. 1965. Contribution a embryographie system-

atique des Cyperaceae-Cyperoideae. Bull. Jard. Bot. Brux.

35: 285–354.

Van Wichelen J, K Camelbeke, P Chaerle, P Goetghebeur, and

S Huysmans. 1999. Comparison of different treatments for

LM and SEM studies and systematic value of pollen grains

in Cyperaceae. Grana 38: 50–58.

Vovk AG. 1985a. Juncaceae. In: A Takhtajan, ed. Comparative

seed anatomy, vol. 1, pp. 158–159. Nauka, Leningrad

(in Russian).

Vovk AG. 1985b. Cyperaceae. In: A Takhtajan, ed. Comparative

seed anatomy, vol. 1, pp. 163–166. Nauka, Leningrad

(in Russian).

Vrijdaghs A, P Caris, P Goetghebeur, and E Smets. 2004. The

bristles of Dulichium (Cyperaceae), a ﬂ oral ontogenetic

study. In: Botany 2004. Alpine diversity: adapted to the

peaks (Abstracts), p. 164.

Vrijdaghs A, P Caris, P Goetghebeur, and E Smets. 2005a. Floral

ontogeny in Scirpus, Eriophorum and Dulichium

(Cyperaceae), with special reference to the perianth. Ann.

Bot. 95: 1199–1209.

Vrijdaghs A, P Goetghebeur, AM Muasya, P Caris, and E Smets.

2005b. Floral ontogeny in Ficinia and Isolepis (Cyperaceae),

with focus on the nature and origin of the gynophore. Ann.

Bot. 96: 1247–1264.

Wilczek E. 1892. Beiträge zur Kenntnis des Baues der Frucht

und des Samens der Cyperaceen. Bot. Centralbl. 51: 129–

138, 193–201, 225–233, 257–265.

Williams CA and JB Harborne. 1975. Luteolin and daphnetin

derivatives in the Juncaceae and their systematic signiﬁ -

cance. Biochem. Syst. Ecol. 3: 181–190.

Wulff HD. 1939. Die Pollenentwicklung der Juncaceen nebst

einer Auswertung der embryologischen Befunde hinsichtlich

einer Verwandtschaft zwischen den Juncaceen und

Cyperaceen. Jahrb. Wiss. Bot. 87: 533–556.

Yen AC and RG Olmstead. 2000. Molecular Systematics of

Cyperaceae Tribe Cariceae based on two chloroplast DNA

734 Subclass IV. COMMELINIDAE

regions: ndhF and trnL intron-intergenic spacer. Syst. Bot.

25: 479–494.

Zhang X, KL Wilson, and JJ Bruhl. 2004. Sympodial structure

of spikelets in the tribe Schoeneae (Cyperaceae). Am. J. Bot.

91: 24–36.

Superorder POANAE

Order 29. TYPHALES

Perennial herbs of shallow-water and wet habitats,

sometimes wholly submerse and ﬂ oating, with creep-

ing, sympodial rhizomes rich in starch (starch grains

pteridophyte-type). Raphide bundles are common, cal-

cium oxalate crystals present. Silica bodies sometimes

present. Cells with mucilage common. Vessels in all

vegetative organs, with scalariform perforations.

Leaves alternate, distichous, with strongly sheathing

base, long, linear, ﬂ at or keeled, with parallel venation,

possess a prominent system of aerenchymatous chan-

nels. Stomata paracytic. Flowers small or minute,

numerous, basically 3-merius or 2-merous, unisexual,

in compound, basally female and apically male race-

mose inﬂ orescences consisting of complex globular

heads or very dense cylindric spikes. Perianth of one

to several inconspicuous, membranous scales or

more or less numerous thin bristles. Stamens 1–8,

mostly 3; ﬁ laments free or often basally connate;

anthers tetrasporangiate, basiﬁ xed, extrorse. Tapetum

plasmodial. Microsporogenesis successive. Pollen

grains 2-celled, free or in tetrads, tectate-columellate,

1- porate (ulcerate), ﬁ nely reticulate. Gynoecium

mostly pseudomonomerous, in Typhaceae seemingly

monomerous while in Sparganiaceae with one more or

less adaxial fertile locule and one abaxial empty

locule, but some have gynoecia with two or three

fertile locules and correspondingly with two or

three stigmatic branches; style short or long with

shortly decurrent stigma; ovary superior, with one

pendulous ovule in the fertile locule (or fertile locules).

Ovules anatropous, apotropous, bitegmic, crassinucel-

late, with parietal cell. Female gametophyte of

Polygonum-type. Endosperm helobial. Fruits small,

dry, indehiscent or dorsally dehiscent. Seeds ﬁ ll up the

fruit locule, with thin, membranous seed coat, straight,

cuneate or fusiform, well-differentiated embryo, copi-

ous, mealy endosperm containing starch, aleurone, and

small quantity of lipids, and thin perisperm containing

starch, aleurone, and lipids (Sparganium) or only

starch (Typha). The unligniﬁ ed cell walls contain

bound ferulate. n = 15.

Taxonomic position is disputable. The order

Typhales is enigmatic, having many automorphic fea-

tures, but few characters which links it clearly to other

orders (Linder and Kellogg 1995). They are associated

either with the commelinids or more traditionally with

the Arales and Pandanales. They resemble the Arales in

starchy rhizome, ﬂ oral morphology of Sparganium, the

abundance of crystal raphides, plasmodial tapetum,

successive microsporogenesis, starchy endosperm

(only in certain Araceae). Seeds of the Typhales have

perisperm, like those of Pistiaceae and Acoraceae.

Besides, Sparganium and Acorus share the same rust

parasite, Uromices sparganii (Parmelee and Savile

1954; Savile 1979). There are also many similarities to

the Pandanales, including vessel characteristics (Wagner

1977), close resemblances between the branching in

Sparganium and that in Pandanus, and striking resem-

blances in the arrangement of ﬂ owers (see Rendle

1953). As Rendle points out, the inﬂ orescence of

Sparganium recalls that of Pandanus on a smaller scale.

According to Müller-Doblies (1969), there is

“die frappierende Ahnlichkeit der kopfehenformigen

Partialinﬂ oreszenzen von Sparganium mit jenen man-

cher Pandani (z. B. Pandanus sparganioides Bak.).”

The afﬁ nity between these two orders is supported also

by cytological data (Sharma 1964; Malick and Sharma

1966). In both of them n = 15. “Moreover, in the mor-

phology of the chromosomes too homogeneity is evi-

dent, all being characterized by very small chromosomes,

with nearly identical types of restrictions. It is very dif-

ﬁ cult to distinguish the three genera on the basis of

their karyotypes as they look very similar to each other”

(Sharma 1964). Therefore Sharma even suggests the

inclusion of all of them under one order Pandanales.

Chemical data also support the afﬁ nity between

Typhales and Pandanales. Both of them have caffeic

and p-coumaric acids (Gibbs 1974), and in both of them

similar ﬂ avonoids, simple phenols, raphides, steroids,

tannins, and triterpenoids (Radhakrishnaiah et al. 1984)

are present. Finally, a close phylogenetic relationship

between Typhales and Pandanales is supported by sero-

logical study (Bergner and Jensen 1989). However,

Typhales are herbaceous, silica bodies sometimes pres-

ent, cells with mucilage absent, leaves not aggregated

at the end of branches, distichous, with prominent sys-

tem of aerenchymatous channels compartimentized by

Superorder POANAE 735

diaphragms, stomata paracytic, endosperm formation

helobial (probably of particular type with small chala-

zal chamber), fruits dry, seed coat very reduced, seeds

with perisperm and always rich in starch, and the unlig-

niﬁ ed cell walls are ﬂ uorescent under UV and contain

bound ferulic acid. In addition, the epicuticular wax is

consistently of Strelitzia type. Dahlgren et al. (1985)

place the Typhales in the vicinity of Pontederiaceae,

Haemodoraceae and Philydraceae, but don’t exclude

the possibility that they also come near the ancestors of

Cyperales or Pandanales. According to Linder and

Kellogg (1995) Typhales are the sister-group to Poales

s. l. I agree with Kubitzki (1998: 459), that a close rela-

tionship to Poales s. l. (Poanae sensu mihi) at present is

the best supported hypothesis.

Key to families

1 Inﬂ orescence axillary, of dense globose units, each

comprising twice-thrice-compound heads; the inﬂ o-

rescence system basally female, apically male.

March or aquatic perennial herbs. Leaves emergent

and ﬂ oating, alternate, distichous, sessile, sheathing

simple, entire. Mesophyll sometimes containing

mucilage cells (with raphides) and calcium oxalate

crystals. Flowers bracteate (female ﬂ owers) or

ebrancheate (male ﬂ owers), small. Floral receptacle

developing an androphore. Perianth of the female

ﬂ owers of (1-)3–4(-6) small segments. Stamens

(1-)3(-6). Gynoecium of 1–2 carpels, monomerous

or syncarpous (and then pseudomonomerous, with

one locule empty). Ovary 2-locular. Fruits 1-seeded,

drupaceous or nucular, hydrochorous or endozoo-

chorous, sessile or nearly so; seeds starchy, epicarp

consists of three layers: ﬁ rm exocarp, spongy meso-

carp and thick and stony endocarp; embryo small,

endosperm oily. . . . . . . . . . . . . 1. sparganiaceae.

1 Inﬂ orescence a terminal, dense, elongate, cylindri-

cal spike. Aquatic herbs; leaves emergent, alternate,

leathery, sessile, sheathing, simple. Silica bodies

present or absent. The mesophyll crystals raphides,

or druses, or solitary-prismatic, or styloids. Perianth

of the female ﬂ owers mostly of numerous capillary

bristles. Stamens (1-)3(-5); anthers appendaged

(via apical projection of the connective). Gynoecium

of one carpel. Fruits anemochorous, a follicle (but

tiny and achene-like before dehiscence), long-stip-

itate, epicarp consists of two layers: thin exocarp of

thin-walled cells and thin endocarp of thick-walled

cells. . . . . . . . . . . . . . . . . . . . . . . . .2. typhaceae.

1. SPARGANIACEAE

Hanin 1811. 1/14. Extratropical regions of the Northern

Hemisphere, mountains of Sumatra and New Guinea,

and southeastern Australia and New Zealand.

Sparganium.

2. TYPHACEAE

A. L. de Jussieu 1789. 1/10-15. Nearly cosmopolitan,

but mainly in Eurasia and North America.

Typha (including Rohrbachia).

Very close to the Sparganiaceae. The differences

between them are clearly related to the switch from

hydrochory to anemochory (Kubitzki 1998).

Bibliography

Asplund I. 1972. Embryological studies in the genus Typha.

Svensk Bot. Tidskr. 66: 1–17.

Bergner I and U Jensen. 1989. Phytoserological contribution to

the systematic placement of the Typhales. Nord. J. Bot.

8: 447–456.

Cook CDK and MS Nicholls. 1986, 1987. A monographic study

of the genus Sparganium (Sparganiaceae). Part I. Subgenus

Xanthosparganium Holmberg. Bot. Helv. 96: 213–267, 1987;

Part II. Subgenus Sparganium. Bot. Helv. 97: 1–44, 1987.

Gibbs RD. 1974. Chemotaxonomy of ﬂ owering plants, pp. 1–4.

McGill-Queen’s University Press, Montreal.

Graef PE. 1955. Ovule and embryo sac development in Typha

latifolia. Am. J. Bot. 42: 806–809.

Haines RW and KA Lye. 1972. Studies in African Cyperaceae:

VII. Panicle morphology and possible relationships in

Sclerieae and Cariceae. Bot. Not. 125: 331–343.

Lee DW and DE Fairbrothers. 1972. Taxonomic placement of

the Typhales within the monocotyledons: preliminary sero-

logical investigation. Taxon 21: 39–44.

Malik R and AK Sharma. 1966. Chromosome studies in Indian

Pandanales. Cytologia 31: 402–410.

Mavrodoev EV. 2001. Rohrbachia, a new genus of the Typhaceae.

Bot. Zhurn. 86, 9: 120–124 (in Russian with English

summary).

Meyer FJ. 1933. Beiträge zur vergleichenden Anatomie der

Typhaceae (Gattung Typha). Beitr. Bot. Centralbl. 51:

335–376.

Müller-Doblies U. 1969. Über die Blütenstande und Blüten

sowie zur Embryologie von Sparganium. Bot. Jahrb. Syst.

89: 459–450.

Müller-Doblies D. 1970. Über die Verwandtschaft von Typha

und Sparganium im Inﬂ oreszenz- und Blütenbau. Bot. Jahrb.

Syst. 89: 451–562.

Parmelee J and BDO Savile. 1954. Life history and relationship

of rusts of Sparganium and Acorus. Mycologia 46:

823–836.