Takhtajan Armen. Flowering Plants

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614 Subclass I. ALISMATIDAE

Kuo J, K Aioi, and H Iizumi. 1988. Comparative leaf structure

and its functional signiﬁ cance in Phyllospadix iwatensis

Makino and Phyllospadix japonicus Makino (Zosteraceae).

Aquatic Bot. 30: 169–187.

Kuo J, K Seto, T.Nasu, H Iizumi, and K Aioi. 1989. Notes on

Archaeozostera in relation to the Zosteraceae. Aquatic Bot.

34: 317–328.

Kuo J, RW Ridge, and S Lewis. 1990a. Leaf internal morphol-

ogy and ultrastructure of Zostera moelleri Irmisch ex

Aschers.: a comparative study of intertidal and subtidal

forms. Aquatic Bot. 36: 217–236.

Kuo J, K Aioi, and H Iizumi. 1990b. Chromosome numbers and

their systematic implications in Australian marine angio-

sperms: The Posidoniaceae. Plant Syst. Evol. 171:

199–204.

Lakshmanan KK. 1965. Note on the endosperm formation in

Zannichellia palustris L. Phyton 22: 13–14.

Larkum AWD, AJ McComb, and SA Shepherd, eds. 1989.

Biology of sea grasses: a treatise on the biology of sea

grasses with special reference to the Australian region.

Elsevier, Amsterdam.

Larsen K. 1966. Cytotaxonomical note on Lilaea. Bot. Not. 119:

496–497.

Les DH and RR Haynes. 1996. Coleogeton (Potamogetonaceae),

a new genus of pondweeds. Novon 6: 389–391.

Les DH and DJ Sheridan. 1990a. Hagstrom’s concept of phylo-

genetic relationships in Potamogeton L. (Potamogetonaceae).

Taxon 39: 41–58.

Les DH and DJ Sheridan. 1990b. Biochemical heterophylly and

ﬂ avonoid evolution in North American Potamogeton

(Potamogetonaceae). Am. J. Bot. 77: 453–465.

Les DH, ML Moody, SWL Jacobs, and RJ Bayer. 2002.

Systematics of Seagrasses (Zosteraceae) in Australia and

New Zealand. Syst. Bot. 27: 468–484.

Lieu SM. 1979. Organogenesis in Triglochin striata. Canad.

J. Bot. 57: 1418–1438.

Lindqvist C, J de Laet, RR Haynes, L Aagesen, BR Keener,

and VA Albert. 2006. Molecular phylogenetics of an

aquatic plant lineage, Potamogetonaceae. Cladistics 22:

568–588.

Lupnitz D. 1969. Histogenese and Anatomie von Primarwurzeln

und sprossburtigen Wurzeln einer Potamogetonaceae. Beitr.

Biol. Pﬂ anz. 46 : 247–313.

McConchie CA, SC Ducker, and RB Knox. 1982a Biology of

Australian seagrasses: ﬂ oral development and morphol-

ogy in Amphibolis (Cymodoceaceae). Aust. J. Bot. 30:

251–264.

McConchie CA, RB Knox, and SC Ducker. 1982b. Pollen wall

structure and cytochemistry in the seagrass Amphibolis grif-

ﬁ thii (Cymodoceaceae). Ann Bot. 50: 792–732.

McMillan C. 1983. Seed germination in Halodule wrightii and

Syringodium ﬁ liforme from Texas and the US Virgin Islands,

Aquatic Bot. 15: 217–220.

McMillan C and LH Bragg. 1987. Comparison of fruits of

Syringodium (Cymodoceaceae) from Texas, the US Virgin

Islands and the Philippines. Aquatic Bot. 28: 97–100.

Muenscher WC. 1936. The germination of seeds of Potamogeton

(pondweeds). Mich. Bot. 23: 35–38.

Murbeck S. 1902. Über die Embryologie von Ruppia rostellata

Koch. Koninkl. Sven. Vetensk. Ak. Handl. 36(5): 1–21.

Nikiticheva ZI and OB Proskurina. 1992. Embryology of

Scheuchzeria palustris (Scheuchzeriaceae). Bot. Zhurn. 77:

3–18 (in Russian with English summary).

Pettitt JM and AC Jenny. 1975. Pollen in hydrophilous angio-

sperms. Micron 5: 377–405.

Plisco MA. 1985. Scheuchzeriaceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol 1, pp. 47–49. Nauka,

Leningrad (in Russian).

Posluszny U. 1981. Unicarpellate ﬂ oral development in

Potamogeton zosteriformis. Canad. J. Bot. 59: 495–504.

Posluszny U. 1983. Re-evaluation of certain key relationships in

the Alismatidae: ﬂ oral organogenesis of Scheuchzeria palus-

tris. Am. J. Bot. 70: 925–933.

Posluszny U and R Sattler. 1973. Floral development of

Potamogeton densus. Canad. J. Bot. 51: 647–656.

Posluszny U and R Sattler. 1974a. Floral development of

Potamogeton richardsonii. Am. J. Bot. 61: 209–216.

Posluszny U and R Sattler. 1974b. Floral development of Ruppia

maritima var. maritima. Canad. J. Bot. 52: 1607–1612.

Posluszny U and R Sattler. 1976. Floral development of

Zannichellia palustris. Canad. J. Bot. 54: 651–662.

Posluszny U and PB Tomlinson. 1977. Morphology and devel-

opment of ﬂ oral shoots and organs in certain Zannichelliaceae.

Bot. J. Linn. Soc. 75: 21–46.

Posluszny U, WA Charlton, and DK Jain. 1986. Morphology and

development of the reproductive shoots of Lilaea scilloides

(Poir.) Hauman (Alismatidae). Bot. J. Linn. Soc. 92: 323–342.

Reinecke P. 1964. A contribution to the morphology of

Zannichellia aschersoniana Graebn. South Afr. J. Bot. 30:

93–101.

Roth I. 1961. Histogenese der Laubblatter von Zostera nana.

Bot. Jahrb. Syst. 80: 500–507.

Sattler R. 1965. Perianth development of Potamogeton richard-

sonii. Am. J. Bot. 52: 35–41.

Schneider EL and S Carlquist. 1997. Origins and nature of

vessels in monocotyledons. 2. Juncaginaceae and

Scheuchzeriaceae. Nord. J. Bot. 17: 397–401.

Schwantz G. 1967. Untersuchungen zur postmeiotischen

Mikrosporogenese. I. Morphogenese des Ruppia-Pollens.

Pollen et Spores 9: 9–48.

Singh V. 1964. Morphological and anatomical studies in

Helobiae: I. Vegetative anatomy of some members of

Potamogetonaceae. Proc. Indian Acad. Sci. 60B: 214–231.

Singh V. 1965. Morphological and anatomical studies in Helobiae.

V. Vascular anatomy of the ﬂ ower of Lilaea scilloides (Poir.)

Hamm. Proc. Indian Acad Sci. B, 61: 316–535.

Soros-Pottruff C and U Posluszny. 1994. Developmental mor-

phology of reproductive structures of Phyllospadix

(Zosteraceae). Int. J. Plant Sci. 155: 405–420.

Soros-Pottruff C and U Posluszny. 1995. Developmental

morphology of reproductive structures of Zostera and a

reconsideration of Heterozostera (Zosteraceae). Int. J. Plant

Sci. 156: 143–158.

Sorsa P. 1988. Pollen morphology of Potamogeton and

Groenlandia (Potamogetonaceae) and its taxonomic signiﬁ -

cance. Ann. Bot. Fenn. 25: 179–199.

Soueges R. 1943. Embryogenie des Scheuchzeriacées:

Developpement de 1’embryon chez le Triglochin mariti-

mum L. Compt. Rend. Hebd. Seances Acad. Sci. 216:

746–748.

Superorder ARANAE 615

Stenar H. 1935. Embryologische Beobachtungen ﬁ ber

Scheuchzeria palustris L. Bot. Not. 1935: 78–86.

Stewart JG and L Ludenberg. 1980. Microsporocyte growth and

meiosis in Phyllospadix torreyi, a marine monocotyledon.

Am. J. Bot. 67: 949–954.

Sun K, J-K Chen, and Z-Y Zhang. 2001. Pollen morphology of

Najadaceae and Zannichelliaceae. Acta Phytotax. Sinica 39:

31–37 (in Chinese with English summary).

Takaso T and F Bouman. 1984. Ovule ontogeny and seed devel-

opment in Potamogeton natans L. (Potamogetonaceae), with

a note on the campylotropous ovule. Acta Bot. Neerl. 33:

519–533.

Talavera S, P Garcia-Murillo, and J Herrera. 1993. Chromosome

numbers and a new model for karyotype evolution in Ruppia

L. (Ruppiaceae). Aquatic Bot. 45: 1–13.

Taylor ARA. 1957a. Studies of the development of Zostera

marina L.: 1. The embryo and seed. Canad. J. Bot. 35:

477–499.

Taylor ARA. 1957b. Studies of the development of Zostera

marina L.: 2. Germination and seedling development. Canad.

J. Bot. 35: 681–695.

Terekhin ES. 1985. Potamogetonaceae, Ruppiaceae. In:

A Takhtajan, ed. Comparative seed anatomy, vol 1, pp. 51–55.

Nauka, Leningrad (in Russian).

Terechin ES and SI Chubarov. 1991. The embryological and

carpological investigation of Althenia jiliformis

(Zannichelliaceae). Bot. Zhurn. 76: 226–236 (in Russian

with English summary).

Terechin ES and GV Shibakina. 1985. Zosteraceae. In:

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

pp. 62–64. Nauka, Leningrad (in Russian).

Tomlinson PB and U Posluszny. 1976. Generic limits in the

Zannichelliaceae (sensu Dumortier). Taxon 25: 273–279.

Tomlinson PB and U Posluszny. 1978. Aspects of ﬂ oral mor-

phology and development in the sea grass Syringodium

ﬁ liformis (Cymodoceaceae). Bot. Gaz. 139: 333–345.

Tomlinson PB and U Posluszny. 2001. Generic limits in the

seagrass family Zosteraceae. Taxon 50: 429–437.

Tutin TG. 1938. The autecology of Zostera marina in relation to

its wasting disease. New Phytol. 37: 50–71.

Vijayraghavan MR and A Vidya Kumari. 1974, Embryology and

systematic position of Zannichellia palustris L. J. Indian

Bot. Soc. 53: 292–302.

Waycott M and DH Less. 1996. An integrated approach to the

evolutionary study of seagrasses. In: J Kuo, RC Phillios,

DI Walker, and H Kirkman, eds. Seagrass biology.

Proceedings of an International Workshop, Rottnest Island,

Western Australia. 25–29 January 1996, pp. 71–78. Perth.

Weigleb G. 1988. Notes on pondweeds, outline for a mono-

graphical treatment of the genus Potamogeton L. Feddes

Repert. 99: 249–266.

Yamashita T. 1972. Eigenartige Wurzelanlage der Embryos bei

Ruppia maritima L. Beitr. Biol. Pﬂ . 48: 157–170.

Yamashita T. 1973. Über die Embryo- und Wurzelentwicklung

bei Zostera japonicus Aschers. et Graebn. J. Fac. Sci. Univ.

Tokyo III, Bot. 11: 175–193.

Yamashita T. 1976. Über die Pollenbildung bei Halodule

pinifolia und H. uninervis. Beitr. Biol. Pﬂ . 52: 217–226.

Zapata O and C McMillan. 1979. Phenolic acids in seagrasses.

Aquatic Bot. 7: 307–317.

Superorder ARANAE

Order 5. ARALES

Very small to giant herbs with creeping or tuberous

rhizomes or scrambling, slender shrubs, often climbers

with aerial roots and true epiphytes, rarely free-ﬂ oating

aquatics. Roots mycorrhizal, without root hairs. Silica

bodies lacking, but calcium oxalate crystals occur.

Vessels in roots and seldom also in rhizome and in

stem, always with scalariform perforations. Simple or

uniseriate laticifers present in some genera of the

Araceae. Resin ducts sometimes present as well as

rows of mucilage cells. Sieve-element plastids P2c-

type, or rarely (Pistiaceae) S-type. Leaves alternate,

spirally arranged or rarely distichous, unifacial (Acora

ceae) or bifacial, usually with distinct, basally sheath-

ing petiole and expanded, entire to variously cleft or

perforate or even compound lamina, variously parallel-

veined or pinnately or palmately net-veined, usually

developing acropetally (like typical dicotyledons).

Often a row of intravaginal squamules occurs in the

axils of the leaf sheaths or in the axils of the cataphylls.

Stomata paracytic, less often tetracytic, or cyclocytic,

sometimes anomocytic. Flowers small, usually numer-

ous, in terminal, unbranched spike with ﬂ eshy axis, the

spadix (extremely reduced in Lemnaceae) generally

subtended by ovate and colored or rarely green, mostly

rather large spathe (lacking in Acorus, Gymnostachys,

Orontium, and in some Lemnaceae), which encloses

the spadix in the juvenile stage, bisexual or unisexual

(monoecious or rarely dioecious), actinomorphic,

bractless. Perianth of 4 or 6(8) free or connate seg-

ments in two cycles, in unisexual ﬂ owers generally

much reduced or wanting or always wanting. Stamens

(1-)4 or 6, or 8–12, free or sometimes more or less

connate and sometimes forming synandria; ﬁ laments

mostly short and broad; anthers tetrasporangiate or

disporangiate, basiﬁ xed, extrorse or very rarely

(Zamioculcas, Acorus) introrse, opening by apical

pores or slits or longitudinally. Tapetum plasmodial or

(Acoraceae) secretory. Microsporogenesis successive.

Pollen grains 2-celled or 3-celled, exceptionally

variable, tectate-columellate to tectate-granular or

atectate, 1-colpate and boat-shaped to globose and

inaperturate, with various types of ornamentation.

Gynoecium of (1-)3(-15) carpels; style terminal and

616 Subclass I. ALISMATIDAE

generally short and thick to conical, or the stigmas ses-

sile; ovary superior, plurilocular or unilocular, seldom

pseudomonomerous, with one to many ovules per

locule. Ovules pendulous, or horizontal, or ascending,

anatropous, hemitropous, or orthotropous, sometimes

campylotropous, bitegmic, very rarely unitegmic

(Symplocarpus – Barabe, Froget, Chrétien 1987 and

Gymnostachys – Buzgo 2001), crassinucellate or tenui-

nucellate, sometimes with endothelium, with or without

parietal cell. Female gametophyte of Polygonum-type or

rarely (as in most Lemnaceae) of Allium-type.

Endosperm cellular, or helobial, or nuclear, with chala-

zal haustorial process. Fruits usually baccate, more

rarely rather dry and ﬁ rmly carnose or leathery, densely

packed into a cylindric infructescence. Seeds one to

many, sometimes ﬂ eshy, sometimes with operculum;

seed coat formed by both integuments, predominantly

by the inner one; embryo generally large, green, mostly

linear, straight or sometimes curved, usually well dif-

ferentiated, sometimes macropodous (having swollen

hypocotyl or radicle); endosperm often present and then

usually copious, with aleurone, simple starch grains,

and oil or endosperm wanting; perisperm present only

in Pistia (Mercado-Noriel and Mercado 1978;

Vyshenskaya 1985). Plants contain ﬂ avone C-glycosides,

ﬂ avonols and cyanogenic glucoide triglochinin.

The Arales are traditionally associated with Arecales

and Cyclanthales mainly because of their small ﬂ ow-

ers arranged in dense, spadiciform inﬂ orescence sub-

tended by large principal bract or spathe. However, the

accumulated data have shown that they are much closer

to the alismatanae and share with them many taxo-

nomically important features (Gow 1913; Jussen 1929;

Huber 1977; Dahlgren and Clifford 1982; Dahlgren et

al. 1985; Tillich 1985; Grayum 1991), including the

presence of intravaginal squamules (but only in few

taxon of the Arales, including some Philodendron spe-

cies), the occurrence of both a spathe and spicate inﬂ o-

rescences, of laticifers, an amoeboid tapetum as well

as of the cyanogenic substance triglochinin in Araceae

and certain alismatanae (Nahrstedt 1975), and rbcL

sequence (Duvall et al. 1993). However, the Arales dif-

fer markedly from the alismatanae in many respects.

Key to Families

1 Plants terrestrial or epiphytic or sometimes more or

less aquatic, but not free-ﬂ oating.

2 Leaves ensiform, anatomically unifacial. Peren-

nial herbs with creeping, rather thick, aromatic,

sympodial rhizomes. Have ethereal oil cells, but

do not possess calcium oxalate crystals in any

form, and no raphids. Vessels in roots and in rhi-

zome, with scalariform perforations. Leaves

tufted, long, linear, acuminate, equitant, parallel-

veined, not narrowed or scarcely narrowed below,

sheathing and surrounding the neck of rhizome

at base. Stomata paracytic. Inﬂ orescences scapi-

ﬂ orous (the scape keeled, with double vascular-

ization, taken to represent adnation of the

peduncles with the sheath of the subtending leaf;

“solitary, lateral spadices, tepering acropetally to

a blunt tipe, covered with tightly packed ﬂ owers,

espatheate (in that the leafy point which termi-

nates the scape above the manifestly lateral spa-

dix seems to represent the blade of a terminal

leaf, rather than a true spathe” – Watson and

Dallwitz 2006). Flowers small to minute, bisex-

ual, 3-merous. Perianth segments six, in two

cycles, free, thick, incurved and truncate at apex.

Stamens six; ﬁ laments linear; anthers tetraspo-

rangiate, opening longitudinally. Tapetum secre-

tory. Pollen grains 1-colpate, boat-shaped,

without foot layer (Tarasevich, personal commu-

nication); the apertural exine is subpsilate

(Grayum 1984). Gynoecium of (2-)3(-4) carpels;

style very short, with small stigma; ovary supe-

rior, oblong, 2–3-locular. Ovules 2–4(-5), pendu-

lous, orthotropous, bitegmic, tenuinucellate.

Placentation axile to apical. Endosperm cellular,

without haustorial processes. Fruits berries with

thin, leathery pericarp. Seeds 1(-5)-9, oblong,

with copious endosperm and single layer of radi-

ally elongated cells of perisperm; embryo

straight, green, starchy, testa thick, without phy-

tomelan. Contain proanthocyanidins. Basic chro-

mosome number n = 12. . . . . . . . 1. acoraceae.

2 Leaves with a more or less expanded, bifacial

blade. Plants often cormous, or rhizomatous, or

tuberous, sometimes climbing (stems twiners or

scrambling, some very large). Laticifers present

(sap clear, milky, or dark and taniniferous), rarely

absent. Leaves sometimes distichous, simple,

basal or cauline, rarely solitary (as in Dracontium),

small to very large, sheathing (the sheath mem-

branous, sometimes deciduous); blades often

oblong, cordate, sagitate to hastate, sometimes

perforated. Flowers small, very numerous, often

fragrant, or malodorous, in a spadix that is sub-

Superorder ARANAE 617

tended by well developed and often colored

spathe; plants usually monoecious, except

Arisaema, which is dioecious). Stamens 1–4 or

6–12, free or united into synandria; anthers

extrorse, tetrasporangiate or bisporangiate, open-

ing by longitudinal or short slits, or by apical

pores; staminodia sometimes present, free or

united into synandria. Gynoecium of 1–8(-15)

carpels, monomerous or syncarpous; ovary supe-

rior, 1–3-locular, rarely plurilocular; ovules one

to many per locule, pendulous, or horizontal, or

ascending, often with funicle, less often sessile.

Fruits indehiscent (usually baccate, drupaceous,

sometimes juice, often colorful, or nuts) or rarely

capsular, capsules splitting irregularly (Watson

and Dallwitz 2006). Seeds one to many;

endosperm copious, the embryo straight, located

in center of endosperm, or embryo curved, when

endosperm lacking; embryo chlorophyllous or

rarely (Arisaema amurense) achlorophyllous.

Usually contain saponins, proanthocyanidins

(commonly, sometimes very abundantly), some-

times ﬂ avonols (kaempferol and quercetin), cin-

namic acids, cyanogenic glucosides, alkaloids

and amines (Mayo et al. 1998); n = from 7 up

to 84. . . . . . . . . . . . . . . . . . . . . . . . . 2. araceae.

1 Free-ﬂ oating or submerged aquatics. Lactifers

wanting. Inﬂ orescence very reduced, consists of

one female ﬂ ower and one or two male ﬂ owers.

3 Ovules tenuinucellate. Seeds with copious

perisperm and scanty endosperm. Seed coat with

very large hilum and an operculum. Fruits with

many seeds. Small, stoloniferous, ﬂ oating herbs,

rarely anchored by their long, feathery, hanging

roots; primary root absent. Branching sympodial,

but internodes remain short. Leaves in rosettes,

very shortly petiolated, entire, obovate to spatu-

late-oblong, truncate to emarginate at the apex,

long-cuneate at base, with ﬂ abellate venation,

covered with short depressed uniseriate hairs,

moving together at night. Stomata anomocytic.

Inﬂ orescence small, inconspicuous, enclosed in

small, short, greenish spathe that is tubular below,

bisexual, with two perianthless ﬂ owers, male

ﬂ ower above the female one. Male ﬂ ower with

4–6 stamens united into a synandrium. Pollen

grains tectate-granular, 1(2)- colpate, boat-

shaped, longitudinally strongly ridged; ridges

(from 20 to 25) formed by the outer layer of the

exine; the inner layer of exine forms beaklike

thickenings. Female ﬂ owers naked, consisting of

1-locular carpel, partly attached to the spathe;

style conic, with discoid stigma; ovary with

numerous, basal- subparietal, orthotropous

ovules. Micropyle formed by the outer integu-

ment. Female gametophyte Polygonum-type.

Fruits with membranous, partly transparent peri-

carp, few-seeded. Seeds oblong or obovoid, testa

ultimately rugose; embryo minute, apical, cunei-

form; n = 7. . . . . . . . . . . . . . . . . . 3. pistiaceae.

3 Ovules crassinucellate. Seeds without perisperm

but with endosperm. Seed coat with operculum,

but hilum is not so large. Small to minute, ﬂ oat-

ing or submerged annual, much reduced aquatic

herbs with thalloid plate-like bodies (fronds);

thallus small to minute, globular, ﬂ at or linear,

contain mucilage cells, with or without raphides.

Roots simple, solitary or several per frond or

lacking. Vascular system much reduced, the

plants lacking both vessels and tracheids, or

sometimes with tracheids in roots. Inﬂ orescence

very reduced, enclosed in membranous spathe or

naked, consists of one female and one or two

male ﬂ owers. Perianth lacking. Stamen solitary;

anthers tetrasporangiate (Lemna) or disporan-

giate (Wolfﬁ a), opening by apical pores or short

or longitudinal slits. Pollen grains 3-celled,

tectate-columellate, 1-porate, echinate.

Gynoecium of one carpel; style short, stigma

funneliform; ovary 1-locular, with 1–7 basal,

anatropous, hemitropous, or orthotropous ovules.

Micropyle formed by the inner integument.

Female gametophyte of Allium-type. Endosperm

cellular. Fruits 1–6-seeded utricles. Seeds

oblong-elliptical, smooth; embryo straight, rather

large, well differentiated, but without primary

root (Lodkina 1985), endosperm present or

absent; n = 5, 8, 10, 11, 21.. . . . .4. lemnaceae.

1. ACORACEAE

Martynov 1820. 1/4. Holarctic and paleotropical: tem-

perate Eurasia from Norway to Eastern Asia, Sulawesi,

New Guinea; western and central North America.

Acorus.

Related to the Araceae, but they differ from them in

the absence of raphides, presence of the ethereal cells,

618 Subclass I. ALISMATIDAE

ensiform, unifacial leaves, presence of dermal

perisperm, From the majority of Araceae (except

Gymnostachys) Acoraceae differ also in the absence of

the spathe. From the Araceae as well as from all other

monocotyledons they markedly differ in the DNA

sequences of the plastid rbcL, which according to

Duvall et al. (1993) shows that Acorus “represents the

most ancient surviving lineage of the ancestral mono-

cotyledons.” This hypothesis based mainly on the

genetic constitution of the endosymbiotic organism,

the chloroplast, is most improbable and strongly con-

tradicts all other available data.

2. ARACEAE

A.L. de Jussieu 1763 (including Arisaraceae Raﬁ -

nesque 1838; Caladiaceae Salisbury 1866; Callaceae

Reichenbach ex Bartling 1830; Cryptocorynaceae

J. Agardh 1858; Dracontiaceae Salisbury 1866;

Lentiscaceae Horaninow 1843; Orontiaceae Bartling

1830; Pothoaceae Raﬁ nesque 1838). 103/2500–3000.

Predominantly tropical, only about 10 genera occur in

temperate regions of the Northern Hemisphere.

2.1 GYMNOSTACHYDOIDEAE

Laticifers and spicular cells lacking. Leaves epetio-

late, grasslike, bifacial. Prophylls foliar. Two to six

more or less pendulous spadices are situated in a

cluster on the 1–2 m high, unifacial peduncle. Spathe

lacking. Flowers bisexual and 2-merous. Pollen grains

1-colpate, boat-shaped, stigma without papillae and

possibly dry, very little secretion in the gynoecium,

ovule unitegmic, integument is shorter than the

nucellus (Buzgo 2001), n = 24. – Gymnostachys (1).

Australia.

2.2 ORONTIOIDEAE

Laticifers present (Orontium) or absent. Swamp plants.

Stems stout, erect. Leaves spirally arranged, oblong-

elliptic or cordate, with acrodromous leaf venation.

Flowers bisexual, 2–3-merous. Anthers terminal, open-

ing by longitudinal slits. Ovary inferior, ovules anatro-

pous, n = 13–15. – orontieae: Orontium;

symplocarpeae: Lasichiton, Symplocarpus.

2.3 POTHOIDEAE

Laticifers and spicular cells usually lacking. Leaves

spirally arranged or distichous, often with winged

petiole. Lateral veins of second and third orders

connected by veinlets to form a reticulum. Flowers

usually bisexual, with perianth. Ovules anatropous

to hemitropous, n = (10, 12) 14 (15). – potheae:

Pothos, Pedicellarum, Pothoidium, Anthurium;

monstereae: Holochlamys, Spathiphyllum, Rhodos-

patha, Stenospermation, Scindapsus, Rhaphidophora,

Anadendrum, Monstera, Alloschemone, Epipremnum,

Amydrium, Heteropsis.

2.4 LASIOIDEAE

Laticifers absent or the African genera often have

laticifers or ducts, and biforines or biforine-like raphi-

des. Leaves often sagittate, often deeply lobes and

fenestrate; primary lateral veins of major lamina divi-

sions pinnate to arcuate-parallel, order venation reticu-

late. Flowers bisexual, usually with perianth. Pollen

grains colpate or inaperturate. Endosperm usually

present, forming a thin but distinct layer, n = 13. –

Cyrtosperma, Lasiomorpha, Podolasia, Lasia,

Anophyllum, Urospatha, Anaphyllopsis, Pycnospatha,

Draconitum, Draconitoides.

2.5 CALLOIDEAE

Laticifers present. Leaves distichous, cordate-pinnate,

venation parallel-pinnate; petiole with type three aer-

enchyma. Flowers usually bisexual, without perigone.

Pollen grains 2-colpate. Ovules anatropous. Endosperm

copious, n = 18, 27. – Calla.

2.6 PHILODENDROIDEAE

Lactifers and ducts present in roots, stems, leaves and

inﬂ orescences, or absent. Flowers unisexual, without

perigone. Ovules anatropous or orthotropous. Endos-

perm copious or rarely absent. – philodendreae:

Montrichardia, Anubias, Furtadoa, Philodendron,

Homalomena; zantedeschieae: Zantedeschia,

Callopsis; stylochaetoneae: Stylochaeton; zamio-

culcadeae: Gonatopus, Zamioculcas; aglaone-

mateae: Nephthytis, Anchomanes, Pseudohydrosme,

Aglaonema, Aglaodorum; culcasieae: Culcasia,

Cercestis; spathicarpeae: Bognera, Dieffenbachia,

Spathantheum, Gorgonidium, Synandrospadix, Gearum,

Spathicarpa, Asterostigma, Mangonia, Taccarum.

2.7 SCHISMATOGLOTTIDOIDEAE

Lactifers non-anastomosing, in all organs or absent

from leaves. Inﬂ orescence solitary, rarely two per sym-

podium spathe tube connate, persistent, spadix with

Superorder ARANAE 619

unisexual ﬂ owers with perigone absent. Fruit a berry

or syncarp. Seeds with endosperm, often copious. –

cryptocoryneae: Cryptocoryne, Lagenandra; schis-

matoglottideae: Phymatarum, Schisma-toglottis,

Aridarum, Heteroaridarum, Hottarum, Piptospatha,

Bucephalandra.

2.8 AROIDEAE

Laticifers present, rarely absent, straight or anastomo-

sing. Spathe usually differentiated into lower, convo-

lute tube and upper, gaping blade. Flowers unisexual,

without perianth. Pollen grains inaperturate. Ovules

anatropous or orthotropous. Seeds usually with

endosperm. – thomsonieae Amorphophallus,

Pseudodracontium; caladieae: Hapaline, Syngo-

nium, Xanthosoma, Chlorospatha, Ulearum, Filarum,

Zomicarpella, Caladium, Scaphispatha, Jasarum,

Zomicarpa; arisareae: Ambrosina, Arisarum; pelt-

andreae: Peltandra, Typhonodorum, Colletogyne,

Carlephyton, Arophyton; arisae-mateae: Arisaema,

Pinellia; areae: Typhonium, Sauromatum,

Theriophonum, Biarum, Arum, Eminium, Dracunculus,

Helicodiceros; colocasieae: Ariopsis, Alocasia,

Remusatia, Colocasia, Steudnera, Protarum.

3. PISTIACEAE

Richard ex C. Agardh 1822. 1/1. Tropical and sub-

tropical regions.

Pistia.

Differ from the Araceae in habit, absence of pri-

mary root, leaf, and inﬂ orescence morphology, absence

of laticifers, anomocytic stomata, unique exine struc-

ture (Kuprianova and Tarasevich 1984; Tarasevich

1990), and especially in their perispermous seeds (see

Mercado-Noriel and Mercado 1978; Vyshenskaya

1985) and in sieve-element plastids of S-type.

According to Behnke (1995), the type of plastids

clearly corroborate the isolated position of Pistia.

Lindley (1847, 1853) included Ambrosina together

with the genera comprising the modern family

Lemnaceae in the Pistiaceae, whereas Buscalioni and

Lanza (1935) included in the Pistiaceae only Pistia

and Ambrosina. However, the genus Pistia differs

markedly from Ambrosina and all other members of

the Aroideae in its connate stamens, epetiolate leaves,

the absence of tuberous stock, and secretion ﬁ les, in

addition to the characters mentioned above. The

Pistiaceae originated probably from some ancient

Aroideae-like ancestor.

4. LEMNACEAE

Martynov 1820 (including Wolfﬁ aceae Bubani 1902).

4/c.35. Cosmopolitan.

4.1 LEMNOIDEAE

Raphides present. Roots present. Each frond has two

ﬂ attened, budding pouches, one basal and one lateral

on either side of the axis. Inﬂ orescence is developed in

one of the budding pouches and consists of one female

and two male ﬂ owers enclosed by membranous spathe.

Anther bilocular, opening transversely. Female ﬂ ower

is lateral to the male one. Seeds with longitudinal

ribs. – Spirodela (including Landoltia), Lemna

4.2 WOLFFIOIDEAE

Raphides and roots lacking. Each frond has only one,

median budding pouch. Inﬂ orescence is developed in a

dorsal cavity, lacks a spathe and has one female and

one male ﬂ ower. Anthers unilocular, opening apically.

Seeds nearly smooth. – Wolfﬁ a, Wolfﬁ ella (including

Pseudowolfﬁ a and Wolfﬁ opsis).

The Lemnaceae have some similarities with the

Pistiaceac but differ markedly in pollen morphology

(Kuprianova and Tarasevich 1984; Tarasevich 1990),

crassinucellate ovules, the absence of perisperm, as

well as in extremely reduced and modiﬁ ed vegetative

body. Both the Pistiaceae and Lemnaceae evolved

from Araceae – Aroideae. However, Pistia is not a link

between the Araceae and Lemnaceae, but those two

groups are not at all close in molecular data.

Bibliography

Amelunxen VF and G Gronau. 1969. Untersuchungen an den

Gerbstoffzellen der Niederblatter von Acorus calamus L.

Cytobiologie 1: 58–69.

Arber A. 1919. The vegetative morphology of Pistia and the

Lemnaceae. Proc. Roy. Soc. Lond., Ser. B, Biol. Sci. 91:

96–103.

Barabe D and S Forget. 1988. Anatomie des ﬂ eurs fertiles et

steriles de Zamioculcas (Araceae). Bull. Mus. Natl. Hist.

Nat. B Adansonia 10: 411–419.

Barabé D, S Forget, and S Chrétien 1986. Sur les gynécées

pseudo-monomeres: Cas de Symplocarpus. Compte Rendu

Acad. Sci. Paris, sér.3, 302: 429–434.

620 Subclass I. ALISMATIDAE

Barabé D, S Forget, and S Chrétien. 1987. Organogénese de la

ﬂ eur de Symplocarpus foetidus (Araceae). Canad. J. Bot. 65:

446–455.

Barabe D, A Bruneau, F Forest, and C Lacroix. 2002. The cor-

relation between development of atypical bisexual ﬂ owers

and phylogeny in the Aroideae (Araceae). Plant Syst. Evol.

232: 1–19.

Barabé D, C Lacroix, A Bruneau, A Archambault, and

M Gibernau. 2004. Floral development and phylogenetic

position of Schismatoglottis (Araceae). Int. J. Plant Sci. 165:

173–189.

Behnke H-D. 1995. P-type sieve-element plastids and the

ssystematics of the Arales (sensu Cronquist 1988) – with

S-type plastids in Pistia. Plant Syst. Evol. 195: 87–119.

Beppu T and A Takimoto. 1981. Geographical distribution and

cytological variation of Lemna paucicostata Hegelm. Jpn.

Bot. Mag. (Tokyo) 94: 11–20.

Blanc P. 1977. Contribution a l’etude des aracees. II.

Remarques sur la croissance sympodiale chez l’Anthurium

scandens Engl., le Philodendron fenzlii Engl. Et le

Philodendron speciosum Schott. Rev. Gen. Bot. 84:

319–331.

Blanc P. 1980. Observations sur les ﬂ agelles des Araceae.

Adansonia II 20: 325–338.

Blodgett F-H. 1923. The embryo of Lemna. Am. J. Bot. 10:

336–342.

Bogner J. 1979. A critical list of the aroid genera. Aroideana 1:

63–73.

Bogner J. 1987. Morphological variations in aroids. Aroideana

10(2): 4–16.

Bogner J and A Hay. 2000. Schismatoglottideae (Araceae) in

Malesia. II – Aridarum, Bucephalandra, Phymatarum and

Piptospatha. Telopea 9: 179–222.

Bogner J and M Hesse. 2005. Zamioculcadoideae, a new sub-

family of Araceae. Aroideana 28: 3–20.

Bogner J and SJ Mayo. 1998. Acoraceae. In: K Kubitzki, ed.

The families and genera of vascular plants, vol 4, pp. 7–11.

Springer, Berlin/Heidelberg/New York.

Bogner J and DH Nicolson. 1991. A revised classiﬁ cation of

the Araceae with dichotomous keys. Willdenowia 21:

35–50.

Bown D. 1988. Aroids – plants of the Arum family. Century,

London.

Bown D. 2000. Aroids: Plants of the Arum family. Timber Press,

Portland, OR.

Boyce PC. 1993. The genus Arum. Royal Botanic Gardens,

Kew.

Boyce PC and A Hay. 2001. A taxonomic revision of Araceae

tribe Potheae (Pothos, Pothoidium and Pedicellarum) for

Malesia, Australia and the tropical western Paciﬁ c. Telopea

9: 449–571.

Buell MF. 1935. Seed and seeding of Acorus calamus. Bot. Gaz.

96: 758–795.

Buell MF. 1938. Embryology of Acorus calamus. Bot. Gaz. 99:

556–568.

Buscalioni L and D Lanza. 1935. Le basi morfologiche, anato-

miche, teratologiche della nuova famiglia delle Pistiaceae

(Buscalioni e Lanza) rappresentate dai duegeneri Pistia ed

Ambrosinia. Malpighia 34: 103–180.

Buzgo M. 1994. Inﬂ orescence development of Pistia stratiotes

(Araceae). Bot. Jahrb. Syst. 115: 557–570.

Buzgo M. 2001. Flower structure and development of Araceae

compared with alismatids and Acoraceae. Bot. J. Linn. Soc.

136: 393–425.

Buzgo M and PK Endress. 1998. Floral development of Acorus

calamus (Acoraceae) in comparison with other basal mono-

cots. In Monocots II, p. 13 (abstract). Sydney.

Buzgo M and PK Endress. 1999. The gynoecium of Gymnostachys

(Araceae) and Potamogeton (Potomagetonaceae). In XVI

International Botanical Congress: Abstracts, p. 240.

St. Louis, MO.

Buzgo M and PK Endress. 2000. Floral structure and develop-

ment of Acoraceae and its systematic relationships with

basal angiosperms. Int. J. Plant Sci. 161: 23–41.

Campbell DH 1899. Notes on the structure of the embryo sac in

Sparganium and Lysichiton. Bot. Gaz. 27: 153–166.

Campbell DH. 1900. Studies on the Araceae. 1. Ann. Bot. 14:

1–15.

Carlquist S and EL Schneider. 1997. Origins and nature of vessels

in Monocotyledons. 1. Acorus. Int. J. Plant Sci. 158: 51–56.

Carlquist S and EL Schneider 1998. Origin and nature of vessels

in monocotyledons. 5. Araceae subfamily Colocasioideae.

Bot. J. Linn. Soc. 128: 71–86.

Chao Y and JD Palmer. 1999. Multiple acquisitions via horizon-

tal transfer of a group I intron in the mitochondrial coxI gene

during evolution of the Araceae family. Mol. Biol. Evol. 16:

1155–1165.

Chen YY, Li DZ, and H Wang. 2002. Infrageneric phylogeny

and systematic position of the Acoraceae inferred from ITS,

18S and rbcL sequences. Acta Bot. Yunn. 24: 699–706.

Chouteau M, D Barabé, and M Gibernau. 2006. Pollen-ovule

ratios in some Neotropical Araceae and their putative signiﬁ -

cance. Plant Syst. Evol. 257: 147–157.

Crawford DJ, E Landolt, DH Les, and RT Kimb. 2001. Allozyme

studies in Lemnaceae: variation and relationships in Lemna

sections Alatae and Biformes. Taxon 50: 987–999.

Crawford DJ, E Landolt, DH Les, and RT Kimball. 2006.

Speciation in duckweeds (Lemnaceae): phylogenetic and

ecological inferences. In: JT Columbus, EA Friar, JM Porter,

LM Prince, and MG Simpson, eds. Monocots: comparative

biology and evolution (excluding Poales), pp. 231–242,

Rancho Santa Ana Botanical Garden, Claremont.

Croat TC. 1990. A comparison of aroid classiﬁ cation systems.

Aroideana 13: 44–63.

Croat TC. 1998. History and current status of systematic research

with Araceae. Aroideana. 21: 26–145.

Daubs EN. 1965. A monograph of Lemnaceae. Illinois Biological

Monographs 34.

Den Hartog C and F van der Plas. 1970. A synopsis of the

Lemnaceae. Blumea 18: 355–368.

Duvall MR. 2001. An anatomical study of anther development

in Acorus L.: phylogenetic implications. Plant Syst. Evol.

228: 143–152.

Duvall MR, MT Clegg, MW Chase, WD Clark, WJ Kress,

HG Hims, LE Eguiarte, JF Smith, BS Gaut, EA Zimmer, and

GH Learn, Jr. 1993a. Phylogenetic hypotheses for the mono-

cotyledons constructed from rbcL sequence data. Ann.

Missouri Bot. Gard. 80: 607–619.

Duvall MR, GH Leaen, Jr., LE Eguiarte, and MT Clegg. 1993b.

Phylogenetic analysis of rbcL sequences identiﬁ es Acorus

calamus as the primal extant monocotyledon. Proc. Natl.

Acad. Sci. USA 90: 4641–4644.

Superorder ARANAE 621

Engler A. 1876. Vergleichende Untersuchungen über die mor-

phologischen Verhaltnisse der Araceae: I. Naturliches

System der Araceae. Nova Acta Acad. Caes. Leop.-Carol.

German. Nat. Cur. 39: 133–155.

Engler A. 1884. Beitrage zur Kenntnis der Araceae: V. 1, 2. Über

den Entwicklungsgang in der Familie der Araceen und über

die Blutenmorphologie derselben. Bot. Jahrb. Syst. 5:

141–188, 287–336.

Erti PO. 1932. Vergleichende Untersuchungen über die

Entwicklung der Blattnervatur der Araceen. Flora 126:

115–248.

Evstatieva LN, MN Todorova, IV Ognyanov, and LV Kuleva.

1996. Chemical composition of the essential oil in Acorus

calamus L. (Araceae). Fitologija (Bulgaria) 48: 19–23.

Eyde RH, DH Nicolson, and P Sherwin. 1967. A survey of ﬂ oral

anatomy in Araceae. Am. J. Bot. 54: 478–497.

Fox MG and JC French. 1988. Systematic occurrence of sterols

in latex of Araceae: Colocasioideae. Am. J. Bot. 75: 132–137.

French JC. 1986a. Patterns of stamen vascularity in the Araceae.

Am. J. Bot. 73: 434–449.

French JC. 1986b. Ovular vasculature in Araceae. Bot. Gaz.

147: 478–495.

French JC. 1986c. Patterns of endothecial wall thickenings in

Araceae: subfamilies Colocasioideae, Aroideae, and

Pistioideae. Bot. Gaz. 147: 166–179.

French JC. 1987a. Systematic occurrence of a sclerotic hypo-

dermis in roots of Araceae. Am. J. Bot. 74: 891–903.

French JC. 1987b. Structure of ovular and placental trichomes of

Araceae. Bot. Gaz. 148: 198–208.

French JC. 1987c. Systematic survey of resin canals in roots of

Araceae. Bot. Gaz. 148: 360–371.

French JC. 1988. Systematic occurrence of anastomosing latic-

ifers in Araceae. Bot. Gaz. 149: 71–81.

French JC. 1997. Vegetative anatomy. In: SJ Mayo, J Bogner,

and PC Boyce. The genera of Araceae, pp. 9–24. Royal

Botanic Gardens, Kew.

French JC and CT Kessler. 1989. Molecular systematics of the

Araceae: are Acorus and Gymnostachys aroids? Am. J. Bot.

76(Suppl.): 242.

French JC and PB Tomlinson. 1981a. Vascular patterns in stems

of Araceae: subfamily Pothoideae. Am. J. Bot. 68: 713–729.

French JC and PB Tomlinson. 1981b. Vascular patterns in stems

of Araceae: subfamily Monsteroideae. Am. J. Bot. 68:

1115–1129.

French JC and PB Tomlinson. 1981c. Vascular patterns in stems

of Araceae: subfamilies Calloideae and Lasioideae. Bot.

Gaz. 142: 366–381.

French JC and PB Tomlinson. 1981d. Vascular patterns in stems

of Araceae: subfamily Philodendroideae. Bot. Gaz. 142:

550–563.

French JC and PB Tomlinson. 1983. Vascular patterns in stems

of Araceae: subfamilies Calocasioideae, Aroideae and

Pistioideae. Am. J. Bot. 70: 756–771.

Gonçalves EG, Élder AS Paiva, and MA Nadruz Coelho. 2004.

A preliminary survey of petiolar collenchyma in the Araceae.

Ann. Missouri Bot. Gard. 91: 473–484.

Goremykin VM, B Holland, KI Hirsch-Ernst, and FH Hellwig.

2005. Analysis of Acorus calamus genome and its phylo-

genetic implications. Mol. Biol. Evol. 22: 1813–1222.

Gow JE. 1913. Phylogeny of the Araceae. Proc. Iowa Acad. Sci.

20: 161–168.

Govaerts R and DG Frodin. 2002. World checklist and bib-

liography of Araceae (and Acoraceae). Royal Botanic

Gardens, Kew.

Grayum MH. 1984. Palynology and phylogeny of the

Araceae. Ph.D. dissertation, University of Massachusetts,

Amherst, MA.

Grayum MH. 1985. Evolutionary and ecological signiﬁ cance of

starch storage in pollen of the Araceae. Am. J. Bot. 72:

1565–1577.

Grayum MH. 1986. Phylogenetic implications of pollen nuclear

number in the Araceae. Plant Syst. Evol. 151: 145–161.

Grayum MH. 1987. A summary of evidence and arguments

supporting the removal of Acorus from the Araceae. Taxon

36: 723–729.

Grayum MH. 1990. Evolution and phylogeny of the Araceae.

Ann. Missouri Bot. Gard. 77: 628–697.

Grayum MH. 1991. Systematic embryology of the Araceae. Bot.

Rev. 57: 167–203.

Grayum MN. 1992. Comparative external pollen ultrastructure

of the Araceae and putatively related taxa. Monogr. Syst.

Bot. Missouri Bot. Gard. 43: 1–167.

Grob GB, B Gravendeel, MCM Eurlings, and WLA Hetterscheld.

2002. Phylogeny of the tribe Thomsonieae (Araceae) based

on chloroplast matK and trnL intron sequences. Syst. Bot.

27: 453–467.

Gupta BL. 1935. Studies on the development of the pollen grain

and embryo sac of Wolfﬁ a arrhiza. Curr. Sci. 4: 104–105.

Haccius B and KK Lakshmanan. 1966. Vergleichende

Untersuchung der Entwicklung von Kotyledon und

Sproﬁ scheitel bei Pistia stratiotes und Lemna gibba: Ein

Beitrag zum Problem der sogenannten terminalen

Blattorgane. Beitr. Biol. Pﬂ . 42: 425–443.

Hartog C den and F von der Plas. 1970. A synopsis of the

Lemnaceae. Blumea 18: 355–368.

Hay A. 1992. Tribal and subtribal delimitation and circumscrip-

tion of the genera of Araceae tribe Lasieae. Ann. Missouri

Bot. Gard. 79: 184–205.

Hesse M. 2002. The uniquely designed pollen aperture in

Lasioideae (Araceae). Aroideana. 25: 51–59.

Hesse, M. 2006a. Pollen wall ultrastructure in Araceae and

Lemnaceae in relation to molecular classiﬁ cations. In:

JT Columbus, EA Friar, JM Porter, LM Prince, and

MG Simpson, eds. Monocots: comparative biology and evo-

lution (excluding Poales), pp. 204–208. Rancho Santa Ana

Botanical Garden, Claremont.

Hesse M. 2006b. Reasons and consequences of the lack of

sporopollenin ektexine in Aroideae (Araceae). Flora 201:

421–428.

Hesse M, M Weber, and H Halbritter. 1998. Pollen wall stratiﬁ -

cation: its possible role in Araceae systematics. In Monocots

II, p. 28. Sydney.

Hesse M, J Bogner, H Halbritter, and M Weber. 2001. Palynology

of the perigoniate Aroideae: Zamioculcas, Gonatopus and

Stylochaeton (Araceae). Grana 40: 26–34.

Hotta M. 1971. Study of the family Araceae – general remarks.

Jpn. J. Bot. 20: 269–310.

Jussen FJ. 1929. Die Haploidgeneration der Araceen und ihre

Verwertung fur die Systematik. Bot. Jahrb. Syst. 62: 155–283.

Il’ina GM. 1990. Lemnaceae. In: TB Batygina and MS Yakovlev,

eds. Comparative embryology of ﬂ owering plants.

Monocotyledons, pp. 279–286. Nauka, Leningrad (in Russian).

622 Subclass I. ALISMATIDAE

Ivanova IE. 1973. On the systematics of the family Lemnaceae.

Bot. Zhurn. 58: 1413–1428 (in Russian).

Kaplan DR. 1970. Comparative foliar histogenesis of Acorus

calamus and its bearing on the phyllode theory of monocoty-

ledonous leaves. Am. J. Bot. 57: 331–361.

Kaplan DR. 1973. Comparative developmental analysis of het-

eroblastic leaf series of axillary shoots of Acorus calamus L.

Cellule 69: 253–290.

Keating RC. 2000. Collenchyma in Araceae: trends and relation

to classiﬁ cation. Bot. J. Linn. Soc. 134: 203–214.

Keating RC. 2002. Leaf anatomical characters and their value in

understanding morphoclines in the Araceae. Bot. Rev. 68:

510–523.

Keating RC. 2003a. Acoraceae and Araceae. In M Gregory and

D Cutler, eds. The anatomy of the Monocotyledons, vol. 9,

pp. 1–327. Oxford University Press, Oxford.

Keating RC. 2003b. Leaf anatomical characters and their value

in understanding morphoclines in the Araceae. Bot. Rev. 68:

510–523.

Keating RC. 2004a. Vegetative anatomical data and its relation-

ship to a revised classiﬁ cation of the genera of Araceae. Ann.

Missouri Bot. Gard. 91: 485–494.

Keating RC. 2004b. Systematic occurrence of raphide crystals in

Araceae. Ann. Missouri Bot. Gard. 91: 495–504.

Korobova SN and NA Zhinkina. 1990. Araceae. In: TB Batygina

and MS Yakovlev, eds. Comparative embryology of ﬂ ower-

ing plants. Monocotyledons, pp. 275–279. Nauka, Leningrad

(in Russian).

Kozhevnikov DA. 1878. On the history of development of ﬂ ower

in the fam. Araceae. Nauka, Moscow (in Russian).

Kulkarni AR, D Dosi, and VM Manoj. 1990. Fruit and seed

structure in Araceae. Proc. Indian Acad. Sci. 100B:

61–70.

Kuprianova LA and VF Tarasevich. 1984. The ultra-structure of

the surface of pollen grain wall in some genera of the family

Lemnaceae and the related genera of the family Araceae.

Bot. Zhurn. 69: 1656–1661 (in Russian with English

summary).

Landolt E. 1986. The family of Lemnaceae – a monographic

study. Vol. 1: Morphology; karyology; ecology; geographic

distribution; systematic position; nomenclature; descrip-

tions. Veroff. Geobot. Inst. Eidg. Tech. Hochsch. Stift. Rubel

Zuer. 71: 566.

Landolt E. 1998. Lemnaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol 4, pp. 264–270. Springer,

Berlin/Heidelberg/New York.

Landolt E and R Kandeler. 1987. The family of Lemnaceae – a

monographic study. Vol. 2: Phytochemistry; physiology;

application; bibliography. Veroff. Geobot. Inst. Eidg. Tech.

Hochsch. Stift Rubel Zuer. 95: 638.

Lawalree A. 1952. L’embryologie des Lemnaceae. Observations

sur Lemna minor. Cellule 54: 305–326.

Lawalree A. 1961. La polinisation de Lemna minor L. Nat. Belg.

42: 164–165.

Lemon GD and U Posluszny. 2000a. Shoot development and

evolution in Pistia stratiotes (Araceae). Int. J. Plant Sci. 161:

721–732.

Lemon GD and U Posluszny. 2000b. Comparative shoot devel-

opment and evolution in the Lemnaceae. Int. J. Plant Sci.

161: 733–748.

Les DH and DJ Crawford. 1999. Landoltia (Lemnaceae) a new

genus of duckweeds. Novon 9: 530–533.

Les DH, E Landolt, and DJ Crawford. 1994. Molecular system-

atics of the Lemnaceae. Am. J. Bot. 81: 168–169.

Les DH, E Landolt, and DJ Crawford. 1997. Systematics of

Lemnaceae (duckweeds), inferences from micromolecular

and morphological data. Plant Syst. Evol. 204: 161–177.

Les DH, DJ Crawford, E Landolt, JD Gabel, and RT Kimball.

2002. Phylogeny and systematics of Lemnaceae, the duck-

weed family. Syst. Bot. 27: 221–240.

Lodkina MM. 1985. Lemnaceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol 1, pp. 275–280. Nauka,

Leningrad (in Russian).

Maheshwari SC. 1954. The embryology of Wolfﬁ a.

Phytomorphology 4: 355–365.

Maheshwari SC. 1956. The endosperm and embryo of Lemna and

systematic position of Lemnaceae. Phytomorphology 6: 51–55.

Maheshwari SC. 1958. Spirodela polyrrhiza: the link between

the aroids and the duckweeds. Nature 181: 1745–1756.

Maheshwari SC and RN Kapil. 1963a. Morphological and

embryological studies on the Lemnaceae. I. The ﬂ oral struc-

ture and gametophytes of Lemna paucicostata. Am. J. Bot.

50: 677–686.

Maheshwari SC and RN Kapil. 1963b. Morphological and embry-

ological studies on the Lemnaceae. II. The endosperm and

embryo of Lemna paucicostata. Am. J. Bot. 50: 907–914.

Maheshwari SC and PP Khanna. 1956. The embryology of

Arisaema wallichianum Hook. f. and the systematic position

of the Araceae. Phytomorphology 6: 379–388.

Maheshwari SC and N Maheshwari. 1963. The female gameto-

phyte, endosperm and embryo of Spirodela polyrrhiza

. Beitr.

Biol. Pﬂ anz. 39: 179–188.

Mayo SJ, J Bogner, and PC Boyce. 1995. The Arales. In:

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

Monocotyledons: systematics and evolution, pp. 277–286.

Royal Botanic Gardens, Kew, London.

Mayo SJ, J Bogner, and PC Boyce. 1997. The genera of Araceae.

Royal Botanic Gardens, Kew.

Mayo SJ, J Bogner, and PC Boyce. 1998. Araceae. In:

K Kubitzki, ed. Families and genera of vascular plants, vol 4,

pp. 26–74. Springer, Berlin/Heidelberg/New York.

Mayo SJ, L Cabrera, G Salazar, and MW Chase. 2003. Aroids

and their watery beginnings. Ms.

McClure JW and RE Alston. 1966. A chemotaxonomic study of

Lemnaceae. Am. J. Bot. 53: 849–860.

Mercado-Noriel LR and BT Mercado. 1978. Floral anatomy and

seed morphology of water lettuce (Pistia stratiotes). Philipp.

Agric. 61: 281–290.

Mucke M. 1908. Über den Bau und die Entwicklung der Fruchte

und über die Herkunft von Acorus calamus L. Bot. Zeit. 66:

1–123.

Nahrstedt A. 1975. Triglochinin in Araceen. Phytochemistry 14:

2627–2628.

Nicolson DH. 1984. Suprageneric names attributable to Araceae.

Taxon 33: 680–690.

Nicolson DH. 1988. History of Araceae systematics. Aroideana

10: 23–30.

Oganezova GG and NA Barsegyan. 1999. Some peculiarities of

the generative organs of Acorus calamus L. from Armenian

populations. Flora Rastitelnost Rast. Res. Armenia 12: 39–41

(in Russian).

Pan YH, KM Liu, and LG Lei. 2002. Advances in the systemat-

ics of Acorus L. and the re-establishment of Acoraceae. Bull.

Bot. Res. (China). 22: 417–421.

Superorder ARANAE 623

Petersen G. 1989. Cytology and systematics of Araceae. Nord. J.

Bot. 9: 116–166.

Ray TS. 1987. Leaf types in the Araceae. Am. J. Bot. 74:

1359–1372.

Ray TS. 1988. Survey of shoot organization in the Araceae. Am.

J. Bot. 75: 56–84.

Remizova M and D Sokoloff. 2003. Inﬂ orescence and ﬂ oral

morphology in Toﬁ eldia (Toﬁ eldiaceae) compared with

Araceae, Acoraceae and Alismatales s.str. Bot. Jahrb. Syst.

124: 255–271.

Riaz M, S Qamar, and FM Chaudhary. 1995. Chemistry of the

medicinal plants of the genus Acorus (family Araceae).

Hamdard Med. 38(2): 50–62.

Rost LCM. 1979. Biosystematic inversigations with Acorus. 4.

Communication: a synthetic approach to the classiﬁ cation of

the genus. Planta Med. 37: 289–307.

Rostowzew S. 1905. Biology and morphology of duckweeds.

Nauka, Moscow (in Russian).

Rothwell GW, MR Van Atta, HE Ballard Jr, and RA Stockey.

2004. Molecular phylogenetic relationships among

Lemnaceae and Araceae using the chloroplast trnL-trnF

intergenic spacer. Mol. Phylogenet. Evol. 30: 378–385.

Rudall PJ and CA Furness. 1997. Systematics of Acorus: ovule

and anther. Int. J. Plant Sci. 158(5): 640–651.

Schneider EL and S Carlquist. 1998. Origin and nature of ves-

sels in monocotyledons. 4. Araceae subfamily

Philodendroideae. J. Torrey Bot. Soc. 125: 253–260.

Scribailo RW and PB Tomlinson. 1992. Shoot and ﬂ oral devel-

opment in Calla palustris (Araceae-Calloideae). Int. J. Plant

Sci. 153: 1–13.

Seubert E. 1993. Die Samenmerkmale der Araceen und ihre

Bedeutung für die Gliederum der Familie. Koeltz,

Koenigstein.

Seubert E. 1997a. The sclereids of Araceae. Flora 192: 31–37.

Seubert E. 1997b. A comparative study of the seeds of Lasieae

(Araceae). Bot. Jahrb. Syst. 119: 407–426.

Shadowsky AF. 1931. Einige Angaben ﬁ ber die Embryogenie von

Pistia stratiotes L. Ber. Deutsch. Bot. Ges. 49: 350–356.

Silva CJ da. 1981. Observacoes sobre a biologia repro-dutiva

Pistia stratiotes L. (Araceae). Acta Amazonica 11:

487–504.

Soukup A, JL Seago Jr, and O Votrubová. 2005. Developmental

anatomy of the root cortex of the basal Monocotyledon,

Acorus calamus (Acorales, Acoraceae). Ann. Bot. 96:

379–385.

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

MP Simmonds, and CD Specht. 2000. A phylogenetic analy-

sis 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.

Su_XH, GM He, KL Sun, DL Bi, and M Wang. 2004. Study of

comparative anatomy on structure of seedlings of Acorus

tatarinowii and Zantedeschia aethiopica. Acta Bot. Bor.

Occid. Sinica 24(3): 504–509.

Tam S-M, PC Boyce, TM Upson, D Barabé, A Bruneau,

F Forest, and JS Parker. 2004. Intergeneric and infrafamilial

phylogeny of subfamily Monsteroideae (Araceae) revealed

by chloroplast trnL-F sequences. Am. J. Bot. 91: 490–498.

Tarasevich VF. 1989. Pollen grain ultrastructure in the genus

Anthurium (Araceae) in connection with its systematics. Bot.

Zhurn. 74: 314–324 (in Russian with English summary).

Tarasevich VF. 1990. Palynological evidence on the position

of the Lemnaceae family in the system of ﬂ owering

plants. Bot. Zhurn. 75: 959–965 (in Russian with English

summary).

Tillich H-J. 1985. Keimlingsbau und verwandtschaftliche

Bezeihungen der Araceae. Gleditschia 13: 63–73.

Tillich H-J. 2003. Seedling diversity in Araceae and its system-

atic implications. Feddes Repert. 114: 454–487.

Van der Ham RWJM, WLA Hetterscheid, and BJ Van Heuven.

1998. Notes on the genus Amorphophallus (Araceae) – 8

Pollen morphology of Amorphophallus and

Pseudodracontium. Rev. Palaeobot. Palynol. 103: 95–142.

Vyshenskaya TD. 1985. Araceae. In: A Takhtajan, ed. Comparative

seed anatomy, 1: 264–275. Nauka, Leningrad (in Russian).

Wang HZ, YG Chen, and CS Fan. 1998. Review of studies on

chemical constituents and pharmacology in genus Acorus in

China. Acta Bot. Yunn. Suppl. 10: 96–100.

Wang HZ, WL Li, ZJ Gu, and YY Chen. 2001. Cytological

study on Acorus L. in southwestern China, with some cyto-

geographical notes on A. calamus. Acta Bot. Sinica 43:

354–358.

Wang PL and H Li. 1998. Report of pollen morphology of

Araceae. Acta Bot. Yunn. Suppl. 10: 41–42.

Wang W and NX Zhao. 2002. Epidermal characters of leaves in

Araceae. J. Wuhan Bot. Res. 20: 343–349.

Watling JR, SA Robinson, and RS Seymour. 2006. Contribution

of the alternative pathway to respiration during thermo-

genesis in ﬂ owers of the Sacred Lotus. Plant. Physiol. 140:

1367–1373.

Weber M, H Halbritter, and M Hesse. 1999. The basic pollen

wall types in Araceae. Int. J. Plant Sci. 160: 415–423.

Williams NH, JB Harborne, and SJ Mayo. 1981. Anthocyanin

pigments and leaf ﬂ avonoids in the family Araceae.

Phytochemistry 20: 217–234.

Wilson KA. 1960. The genera of the Arales in the southeastern

United States. J. Arnold Arbor. 41: 47–72.

Zennie TM and JW McClure. 1977. The ﬂ avonoid chemistry of

Pistia stratiotes L. and the origin of the Lemnaceae. Aquatic

Bot. 3: 49–54.

Zhu ZY. 1985 Some new taxa of Acorus (Araceae) from Sichuan.

Acta Bot. Bor.-Occid. Sinica 5: 118–121.