Takhtajan Armen. Flowering Plants

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604 Subclass I. ALISMATIDAE

Cook CDK. 1998b. Hydrocharitaceae. In: K Kubitzki, ed. The

families and genera of vascular plants vol 4, pp. 234–248.

Springer, Berlin/Heidelberg/New York.

Govindappa DA and TRB Najdu. 1956. The embryo sac and

endosperm Blyxa oryzetorum Hook. f. J. Indian Bot. Soc. 35:

417–422.

Guo YH and SQ Huang. 1999. Evolution of pollination system

and characters of stigmas in Najadales. Acta Phytotax Sinica

37(2): 131–136.

Haynes RR. 1977. The Najadaceae in the Southeastern United

states. J. Arnold Arbor. 58: 161–170.

Haynes RR. 1979. Revision of north and central American Najas

(Najadaceae). SIDA 8: 34–56.

Haynes RR and LB Holm-Nielsen. 2001. The genera of

Hydrocharitaceae in the southeastern United States. Harvard

Pap. Bot. 5: 201–275.

Haynes RR, LB Holm-Nielsen, and DH Les. 1998. Najadaceae.

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

plants, vol 4, pp. 301–306. Springer, Berlin/Heidelberg/

New York.

Islam AS. 1950. A contribution to the life history of Ottelia

alismoides Pers. J. Indian Bot. Soc. 29: 79–91.

Kaul RB. 1965. Development and vasculature of the androecium

in the Butomaceae. Am. J. Bot. 52: 624 (Abstract).

Kaul RB. 1968. Floral morphology and phylogeny in the

Hydrocharitaceae. Phytomorphology 18: 13–35.

Kaul RB. 1969. Morphology and development of the ﬂ owers of

Boottia cordata, Ottelia alismoides, and their synthetic

hybrid (Hydrocharitaceae). Am. J. Bot. 58: 951–959.

Kaul RB. 1970. Evolution and adaptation of inﬂ orescences in

the Hydrocharitaceae. Am. J. Bot. 57: 708–715.

Kaul RB. 1976. Conduplicate and specialized carpels in the

Alismatales. Am. J. Bot. 63: 175–182.

Kun S, QF Wang, JK Chen. 1997. Micromorphological charac-

ters of seed coats of Chinese Najadaceae and their systematic

signiﬁ cance. Acta Phytotax. Sinica 35: 521–526.

Lakshmanan KK. 1961. Embryological studies in the

Hydrocharitaceae: I. Blyxa octandra Planch. J. Madras Univ.

31B: 133–142.

Lakshmanan KK. 1963. Embryological studies in the

Hydrocharitaceae: II. Halophila ovata Gaudich. J. Indian

Bot. Soc. 42: 15–18.

Lakshmanan KK. 1965. Embryological studies in the

Hydrocharitaceae: III. Nechamandra alternifolia. Phyton

(Buenos Aires) 20: 49–58. IV. Post-fertilization development

in the Hydrilla verticillata Royle. Phyton (Buenos Aires)

22: 13–14.

Les DH, DK Garvin, and CF Wimpee. 1993. Phylogenetic stud-

ies in the monocot subclass Alismatidae: evidence for a

reappraisal of the aquatic order Najadales. Mol. Phylogenet.

Evol. 2: 304–314.

Les DH, ML Moody, and SWL Jacobs. 2005. Phylogeny and

systematics of Aponogeton (Aponogetonaceae): the

Australian species. Syst. Bot. 30: 503–519.

Les DH, ML Moodly, and CL Soros. 2006. A reappraisal of phy-

logenetic relationships in the monocotyledon family

Hydrocharitaceae (Alismatidae). In: JT Columbus, EA Friar,

JM Porter, LM Prince, and MG Simpson, eds. Monocots:

comparative biology and evolution (excluding Poales),

pp. 211–230. Rancho Santa Ana Botanical Garden,

Claremont (Aliso 22: 211–230).

Lowden RM. 1986. Taxonomy of the genus Najas L. (Najadaceae)

in the neotropics. Aquat. Bot. 24: 147–187.

Magnus P. 1894. Uber die Gattung Najas. Ber. Deutsch. Bot.

Ges. 12: 214–224.

Miki S. 1937. The origin of Najas and Potamogeton. Bot. Mag.

(Tokyo) 51: 472–480.

Naidoo Y, JR Lawton, AD Barnabas, and J Goetzee. 1990.

Ultrastructure and cytochemistry of squamulae intravagina-

les of the marine angiosperm, Halophila ovalis. South Afr.

Tydskr. Plantk. 56: 546–553.

Pettitt JM. 1980. Reproduction in sea grasses: nature of the pol-

len and receptive surface of the stigma in the Hydrocharitaceae.

Ann. Bot. 45: 257–271.

Pettitt JM. 1981. Reproduction in sea grasses: pollen develop-

ment in Thalassia hemprichii, Halophila stipulacea, and

Thalassodendron ciliatum. Ann. Bot. 48: 609–622.

Posluszny U and R Sattler. 1976. Floral development of Najas

ﬂ exilis. Canad. J. Bot. 54: 1140–1151.

Rangasamy K. 1941. A morphological study of the ﬂ ower of

Blyxa echinosperma

Hook. f. J. Indian Bot. Soc. 20:

123–133.

Rao YS. 1953. Karyosystematic studies of Helobiales:

I. Butomaceae. Proc. Natl. Inst. Sci. India 19: 563–581.

Rendle AE. 1899. A systematic revision of the genus Najas.

Trans. Linn. Soc., Bot., ser. 2, 5: 379–444.

Roper RB. 1952. The embryo sac of Butomus umbellatus L.

Phytomorphology 2: 61–74.

Sane YK. 1939. A contribution to the embryology of the

Aponogetonaceae. J. Indian Bot. Soc. 18: 79–91.

Sattler R and V Singh. 1978. Floral development of Hydrocleis

nymphoides. Canad. J. Bot. 51: 2455–2458.

Scribailo RW and U Posluszny. 1985. Floral development of

Hydrocharis morsus-ranae (Hydrocharitaceae). Am. J. Bot.

72: 1678–1589.

Shaffer-Fehre M. 1991a. The endotegmen tuberculae: An account

of little-known structures from the seed coat of the Hydro-

charitoideae (Hydrocharitaceae) and Najas (Najadaceae).

Bot. J. Linn. Soc. 107: 169–188.

Shaffer-Fehre M. 1991b. The position of Najas within the

subclass Alismatidae (Monocotyledones) in the light of new

evidence from seed coat structures in the Hydrocharitoideae

(Hydrocharitales). Bot. J. Linn. Soc. 107: 189–209.

Singh V. 1965. Morphological and anatomical studies in

Helobiae: III. Vascular anatomy of the node and ﬂ ower of

Najadaceae. Proc. Indian Acad. Sci. 61B: 98–108.

Singh V. 1966. Morphological and anatomical studies in

Helobiae: VII. Vascular anatomy of the ﬂ ower of Butomus

umbellatus Linn. Proc. Indian Acad. Sci. 63B: 313–320.

Singh V and R Sattler. 1974. Floral development of Butomus

umbellatus. Canad. J. Bot. 52: 223–230.

Singh V and R Sattler. 1977. Floral development of Aponogeton

natans and A. undulatus. Canad. J. Bot. 55: 1106–1120.

Soros CL and DH Les. 2002. Phylogenetic relationships in the

Alismataceae. In Botany 2002: Botany in the Curriculum.

Abstracts, p. 152. Madison, WI.

Stant MY. 1967. Anatomy of the Butomaceae. Bot. J. Linn. Soc.

60: 31–60.

Sun K, Q-F Wang, and J-K Chen. 1997. Micromorphological

characters of seed coats of Chinese Najadaceae and their sys-

tematic signiﬁ cance. Acta Phytotax. Sinica 35: 521–526 (in

Chinese with English summary).

Superorder ALISMATANAE 605

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).

Sun K, J-K Chen, and Z-Y Zhang. 2002. Studies on pollen mor-

phology of Aponogetonaceae. Bull. Bot. Res. (China) 22:

33–36.

Swamy BGL and KK Lakshmanan. 1962. Contributions to the

embryology of the Najadaceae. J. Indian Bot. Soc. 41:

247–267.

Tanaka N, H Setoguchi, and J Murata. 1997. Phylogeny of the

family Hydrocharitaceae inferred from rbcL and matK gene

sequence data. J. Plant Res. 110: 329–337.

Tanaka N, K Uehara, and J Murata. 2004. Correlation between

pollen morphology and pollination mechanisms in the

Hydrocharitaceae. J. Plant Res. 117: 265–276.

Terekhin EC. 1985. Hydrocharitaceae. In: A Takhtajan, ed.

Comparative seed anatomy vol 1, pp. 38–43. Nauka,

Leningrad (in Russian).

Thanikaimoni G. 1985. Palynology and phylogeny. In: HWE

van Bruggen, ed. Monograph of the genus Aponogeton

(Aponogetonaceae). Bibl. Bot. 137: 11–14.

Tomlinson PB. 1969, 1972. On the morphology and anatomy of

turtle grass, Thalassia testudinum (Hydrocharitaceae): II.

Anatomy and development of the root in relation to function.

IV. Leaf anatomy and development. Bull. Marine Sci. 19 (I):

57–71, 1969; 22(1): 75–93, 1972.

Troll W. 1931. Beitrage zur Morphologic des Gynaeceums: I.

Über das Gynaeceum der Hydrocharitaceen. Planta 14:

1–18.

Vijayaraghavan MR and T Kapoor. 1985. Embryogenesis in

Najas marina L.: Structural and histochemical approach.

Aquatic Bot. 22: 45–60.

Order 3. ALISMATALES

Perennial or rarely annual emergent aquatic or marsh

herbs with strongly aerenchymatous and commonly

creeping rhizomes. Roots ﬁ brous. Plants usually with

schizogenous secretory canals that are lined with an

epithelium. Vessels only in roots, with scalariform, or

simple, or scalariform and simple perforations. Sieve-

element plastids of P2c-type (Behnke 2002). Leaves

alternate, more or less crowded near the tip of the rhi-

zome or stolen, usually somewhat expanded and sheath-

ing at the base, sometimes differentiated into lamina

and petiole. Venation arcuate or parallel. Stomata para-

cytic or (some Alismataceae) tetracytic. Flowers in

various kinds of inﬂ orescences or solitary, bisexual or

unisexual, usually actinomorphic, spirocyclic or cyclic,

3-merous, with double or rarely simple perianth, some-

times naked. Stamens 6–9 to many, rarely three; ﬁ la-

ments free; anthers basiﬁ xed or versatile, extrorse or

latrorse, tetrasporangiate, opening longitudinally.

Tapetum plasmodial. Microsporogenesis successive.

Pollen grains tectate-columellate, 3-celled, 2-pantopo-

rate (up to 29 pores) or sometimes inaperturate, spinose.

Gynoecium of three, six, nine, or more (up to 15–20)

free or basally connate carpels; stylodia terminal or (as

in many Alismataceae) gynobasic, with slightly decur-

rent or apical stigma; each carpel with more or less

numerous ovules with laminar-diffuse placentation or

in each carpel one or several basal or subbasal ovules.

Ovules anatropous or amphitropous, bitegmic, weakly

crassinucellate, sometimes with weakly developed

endothelium, without a parietal cell cut off from the pri-

mary archesporial cell. Female gametophyte of Allium-

type. Endosperm helobial or (in some Alismataceae)

nuclear. Fruits multi-follicles or achenes. Seeds hor-

shoe-shaped, testal, without endosperm or only with

vestigial layer (Alismataceae).

Close to the Hydrocharitales. In Engler’s system the

Limnocharitoideae are even included in the Butomaceae

s.l. However, they differ from the Butomaceae s. str. in

anatomy (secretory canals), pollen morphology,

embryology, and seed shape, “whereas in all these fea-

tures the Limnocharitaceae approach very closely the

Alismataceae” (Dahlgren et al. 1985: 301).

Key to Families

1 Each carpel with numerous ovules scattered over its

inner surface. Perennial aquatic herbs rooted in the

substrate or free-ﬂ oating; rhizome with endoder-

mis. Vessels with more or less oblique, scalariform

and simple perforations. Leaves spiral to distichous

( spirodistichous), petiolate, the laminas orbicular to

lanceolate, the base cordate to attenuate; axillary

scales present. Stomata paracytic. Flowers terminal,

solitary or aggregated in pseudoumbels, bracteate,

actinomorphic, bisexual. Periant segments six, free,

in two cycles. Stamens 3, or 6, or 7–100; anthers

extrorse or latrorse. Carpels 3, or 5–9, or 12–20,

conduplicate and distally unsealed, each with a

short, terminal stylodium ending in a shortly

decurrent stigma or the stigma sessile. Ovary supe-

rior, ovules 12–100 per carpel, anatropous to

campylotropous, pseudocrassinucellate. Lateral

surface of the carpels nectariferous toward the

base. Fruits follicular, aggregated in a head. Seeds

with curved, horseshoe shaped or bent; without

endosperm. Flavone and phenolic sulphates and

tannins present. n = 7, 8, 10, chromosomes large.

. . . . . . . . . . . . . . . . . . . . . . 1. limnocharitaceae.

606 Subclass I. ALISMATIDAE

1 Each carpel with solitary or rarely several ventral-

basal ovules. Perennial or rarely annual, rhizoma-

tous herbs, generally rooted in the substrate. Vessels

only in roots, mostly with simple perforations.

Leaves submerged and emergent, alternate, distic-

hous, spirodistichuous or spirally arranged, sessile

or petiolate; the laminas linear, lanceolate, ovate to

rhomboid, the base attenuate, truncate, cordate,

sagittate, or hastate; axillary scales present. Stomata

paracytic or rarely tetracytic. Flowers bisexual or

unisexual, in terminal inﬂ orescences, bracteate.

Perianth segments six, free, in two cycles, white,

red or pink. Stamens 3–6 in one cycle, or 18–100

(to many) in many cycles; anthers extrorse.

Gynoecium of three carpels or of 6–100 (or more);

carpels closed, with a terminal, lateral, or gynobasic

stylodium and often with slightly decurrent stigma.

Ovules 1–2, rarely more per carpel, ascending, ana-

tropous or amphitropous, weakly crassinucellate.

Nectaries generally borne at the basal margins of

the carpels. Fruits achenes or rarely, as in

Damasonium, basally dehiscent follicles. Seeds

with starch; embryo curved (horsesho-shaped),

achlorophyllous, endosperm lacking; testa without

phytomela, usually membranous, sometimes thinly

leather, then brown. Flavone and phenolic sulphates

and tannins present; Alkaloids and proanthocyani-

dins (cyanidin), ﬂ avonols (kaempferol and querce-

tin) present or absent; n = (5-)7–11(-13).

. . . . . . . . . . . . . . . . . . . . . . . . . . 2. alismataceae.

1. LIMNOCHARITACEAE

Takhtajan ex Cronquist 1981. 3/12. Tropical and sub-

tropical regions of the Old and New World.

Limnocharis, Hydrocleys, Butomopsis.

The most archaic member of the family is

Limnocharis (Takhtajan 1966; Haynes and Holm-

Nielsen 1992), which is also the least specialized genus

in the order Alismatales.

2. ALISMATACEAE

Ventenat 1799 (including Damasoniaceae Nakai 1943).

12/80–90. Subcosmopolitans.

Damasonium (including Machaerocarpus),

Baldellia, Alisma, Luronium, Ranalisma, Echinodorus

(? including Helianthium), Caldesia, Limnophyton,

Astonia, Sagittaria, Wiesneria, Burnatia.

Very closely related to the Limnocharitaceae. The

genus Ranalisma forms a connecting link between the

Alismataceae and Limnocharitaceae (Posluszny and

Charlton 1993).

Bibliography

Argue CL. 1973. The pollen of Limnocharis ﬂ ava Buch.,

Hydrocleis nymphoides (Willd.) Buch., and Tenogacharis

latifolia (Don) Buch. (Limnocharitaceae). Grana 13:

108–113.

Argue CL. 1976. Pollen studies in the Alismataceae with special

reference to taxonomy. Pollen et Spores 18: 161–201.

Chanda S, S Nilsson, and S Blackmore. 1988. Phylogenetic

trends in the Alismatales with reference to pollen grains.

Grana 27: 257–272.

Charlton WA. 1968, 1973, 1991. Studies in the Alismataceae: I.

Developmental morphology of Echinodorus tenellus. II.

Inﬂ orescences of Alismataceae. IX. Development of the

ﬂ ower of Ranalisma humile. Canad. J. Bot. 46: 1345–1360,

1968; 51: 775–789, 1973; 69: 2790–2796, 1991.

Charlton WA. 2004. Studies in the Alismataceae. XII. Floral

organogenesis in Damasonium alisma and Baldellia ranun-

culoides, and comparisons with Butomus umbellatus. Canad.

J. Bot. 82: 528–539.

Charlton WA and A Ahmed. 1973. Studies in the Alismataceae:

III. Floral anatomy of Ranalisma humile. IV.

Developmental morphology of Ranalisma humile and

comparison with two members of the Butomaceae,

Hydrocleis nymphoides and Butomus umbellatus. Canad.

J. Bot. 51: 891–897, 899–910.

Chen J-M, D Chen, GW Robert, Q-F Wang, and Y-H Guo.

2004a. Evolution of apocarpy in Alismatidae using phyloge-

netic evidence from chloroplast rbcL sequence data. Bot.

Bull. Acad. Sinica 45: 33–40.

Chen J-M, GW Robert, and Q-F Wang. 2004b. Evolution of

aquatic life forms in Alismatidae: phylogenetic estimation

from chloroplast rbcL sequence data. Israel J. Plant Sci. 52:

323–329.

Daumann E. 1964. Zur Morphologic der Blüte von Alisma

plantago-aquatica L. Preslia 36: 226–239.

Forni Martins ER and KP Calligaris. 2002. Chromosomal

studies on neotropical Limnocharitaceae (Alismatales).

Aquatic Bot. 74(1): 33–41.

Harley MM. 1982. Palynological evidence of a close association

between Butomopsis Kunth and Hydrocleys L. C. Rich.

(Limnocharitaceae). In: JJ Symoens, SS Hooper, and

P Compere, eds. Studies on aquatic vascular plants,

pp. 61–65. Botanical Society of Belgium, Brussels.

Haynes RR and LB Holm-Nielsen. 1992. Limnocharitaceae.

Flora Neotropica 56: 1–34.

Haynes RR and LB Holm-Nielsen. 1994. Alismataceae. Flora

Neotropica 64: 1–112.

Haynes RR, DH Les, and LB Holm-Nielsen. 1998a.

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

vascular plants, vol 4, pp 11–18. Springer, Berlin/Heidelberg/

New York.

Superorder ALISMATANAE 607

Haynes RR, DH Les, and LB Holm-Nielsen. 1998b.

Limnocharitaceae. In: K. Kubitzki, ed. The families and

genera of vascular plants vol 4, pp 271–275. Springer, Berlin/

Heidelberg/New York.

Hooper SS and JJ Symoens. 1982. Observations on the family

Limnocharitaceae Takhtajan ex Hooper and Symoens. In:

JJ Symoens, SS Hooper, and P Compere, eds. Studies on

aquatic vascular plants, pp. 56–60. Royal Botanical Society

of Belgium, Brussels.

Johri BM. 1936. The life-history of Butomopsis lanceolata

Kunth. Proc. Indian Acad. Sci. 4B: 139–162.

Johri BM. 1938a. The embryo sac of Hydrocleis nymphoides

Buchen. Beih. Bot. Centralbl. 58A: 165–172.

Johri BM. 1938b. The embryo sac of Limnocharis emarginata L.

New Phytol. 37: 279–285.

Kak AM and S Durani. 1989. Seed morphology of the family

Alismataceae. J. Econ. Taxon, Bot. 13: 501–509.

Kaul RB. 1967a. Development and vasculature of the ﬂ owers

of Lophotocarpus calycinus and Sagittaria latifolia

(Alismataceae). Am. J. Bot. 54: 914–920.

Kaul RB. 1967b. Ontogeny and anatomy of the ﬂ ower of

Limmnocharis ﬂ ava (Butomaceae). Am. J. Bot. 54: 1223–1230.

Kaul RB. 1968. Floral development and vasculature in Hydrocleis

nymphoides (Butomaceae). Am. J. Bot. 55: 236–242.

Kaul RB. 1976. Conduplicate and specialized carpels in the

Alismatales. Am. J. Bot. 63: 175–182.

Kudryashov LV and EI Savich. 1963. Some data on the embry-

ology of Alisma plantago-aquatica L. Bull. Moscow Soc.

Naturalists Div. Biol. 68(4): 50–63 (in Russian).

Leins P and P Stadler. 1973. Entwicklungsgeschichtliche

Untersuchungen am Androeceum der Alismatales. Oesterr.

Bot. Z. 121: 51–63.

Liu K-M, L-G Lei, and G-W Hu. 2002. Developmental study on

the inﬂ orescence and ﬂ ower of Caldesia grandis Samuel

(Alismataceae). Bot. J. Linn. Soc. 140: 39–47.

Markgraf F. 1936. Blütenbau und Verwandtschaft bei den ein-

fachsten Helobiae. Ber. Deutsch. Bot. Ges. 54: 191–229.

Mayr F. 1943. Beiträge zur Anatomic der Alismataceen: Die

Blattanatomie von Caldesia parnassifolia (Bassi) Parl. Beih.

Bot. Centralbl. 62: 61–77.

Meyer FJ 1932. Beiträge zur Anatomic der Alismataceen. Beih.

Bot. Centralbl. 49(I): 54–63, 272–91, 309–68; 50 (I): 54–63;

52B: 96–111; 54A: 156–69.

Pichon M. 1946. Sur les Alismatacees et les Butomacees. Notul.

Syst. (Paris) 12: 170–183.

Rogers GK. 1983. The genera of Alismataceae in the southeast-

ern United States. J. Arnold Arbor. 64: 383–420.

Sattler R and V Singh. 1973. Floral development of Hydrocleis

nymphoides. Canad. J. Bot. 51: 2455–2458.

Sattler R and V Singh. 1977. Floral organogenesis of Limnocharis

ﬂ ava. Canad. J. Bot. 55: 1076–1086.

Sattler R and V Singh. 1978. Floral organogenesis of Echinodorus

amazonicus Rataj and ﬂ oral construction of the Alismatales.

Bot. J. Linn. Soc. 77: 141–156.

Singh V. 1966. Morphological and anatomical studies in

Helobiae: VI. Vascular anatomy of the ﬂ ower of Alismaceae.

Proc. Natl. Acad. Sci. India B 36: 329–344.

Singh V and R Sattler. 1972. Floral development of Alisma triv-

iak. Canad. J. Bot. 50: 619–627.

Singh V and R Sattler, 1973. Nonspiral androecium and gynoe-

cium of Sagittaria latifolia. Canad. J. Bot. 51: 1093–1095.

Singh V and R Sattler. 1977. Development of the inﬂ orescence

and ﬂ ower of Sagittaria cuneata. Canad. J. Bot. 55:

1087–1105.

Stant MY. 1964. Anatomy of the Alismataceae. Bot. J. Linn.

Soc. 59: 1–42.

Troll W. 1932. Beiträge zur Morphologic des Gynaeceums: II.

Über das Gynaeceum von

Limnocharis Humb. and Bonpl.

Planta 17: 453–460.

Wilder GJ. 1974. Symmetry and development of Butomus

umbellatus (Butomaceae) and Limnocharis ﬂ ava

(Limnocharitaceae). Am. J. Bot. 61: 379–394.

Wodehouse RP. 1936. Pollen grains in the identiﬁ cation and

classiﬁ cation of plants: VIII. The Alismataceae. Am. J. Bot.

23: 535–539.

Order 4. POTAMOGETONALES

Perennial or rarely annual plants growing in marshes

or other very wet places, or aquatic or sub-aquatic and

mostly freshwater herbs with rhizome rooted in sub-

strate. Roots ﬁ brous. Vessels only in roots and rarely

(Scheuchzeriaceae) in rhizomes, with scalariform per-

forations, or less often vessels absent. Sieve-element

plastids of P2c-type (Potamogetonaceae – Behnke

2002). Leaves basal and cauline, alternate to opposite

or rarely verticillate, mostly linear or capillary, with

well-developed, open sheath, mostly ligulate, with

intravaginal squamules, submerged or the upper ones

ﬂ oating and with expanded, lanceolate to elliptic, ovate

or orbicular lamina and a slender petiole; veins when

more than one parallel or arcuate, often connected by

cross-veins. Stomata present or absent. Tannin cells

present or absent. Flowers small to minute, in axillary

or terminal raceme-like inﬂ orescences, bisexual or uni-

sexual, actinomorphic or less often slightly zygomor-

phic, mostly 6–4-merous. Perianth segments in one or

two cycles or wanting, when free, or adnate to the

ﬁ laments. Stamens 6, 4 or rarely 2–3, or 1. Anthers

nearly sessile, basiﬁ xed, mostly tetrasporangiate,

extrorse, opening longitudinally. Tapetum plasmodial.

Microsporogenesis successive. Pollen grains 2-celled

or more often 3-celled, tectate-columellate, inapertu-

rate, usually reticulate. Gynoecium of (1–3)4–6(-16)

free or only partly united carpels; stigmas sessile or on

short stylodia, sometimes slightly decurrent ventrally.

Ovary superior, ovules 1–2, pendulous from the ventral

margin of the carpel, anatropous or orthotropous, or

ortho-campylotropous, but after fertilization becoming

amphitropous, bitegmic, crassinucellate, with a parietal

608 Subclass I. ALISMATIDAE

cell cut off from the archesporial cell. Female gameto-

phyte of Polygonum-type, or rarely (Zannichelliaceae)

of Allium-type. Endosperm helobial or nuclear. Fruits

indehiscent or rarely dehiscent, follicles, achenes,

drupe-like or achene-like, nutlets. Seeds without

endosperm or endosperm scanty; embryo straight or

slightly curved, sometimes with large hypocotyl. Seeds

exotestal, or coat crushed. Flavone aglycones, ﬂ avone

glycosides, and C-glycosylated ﬂ avones are common;

n = 6, 8–11.

Related to the Alismatales and have a common

origin from the alismatalean ancestor.

Key to Families

1 Vessels with scalariform perforations.

2 Stamens mostly six in two cycles.

3 Fruits follicles. Perennial marsh herbs with

sympodial rhizome covered with persistent

ﬁ brous remains of older leaves and leafy stem.

Adventitious roots inserted below the nodes.

Vessels both in roots and in rhizomes. Leaves

basal and cauline, alternate, distichous, linear,

compressed-terete, parallel-veined, with open

sheathing base, the apex obtuse with a con-

spicuous apical pore; sheath marginally hya-

line, with two delicate auricles, at the base

with a dense axillary row of numerous long

hairs that replace intravaginal squamules, with

well-developed air lacunae in the mesophyll.

Stomata tetracytic. Tannin cells present, and

some calcium oxalate crystals occur in the

mesophyll. Flowers in terminal raceme-like

inﬂ orescence in the axils of more or less bract-

like leaves, bisexual, actinomorphic. Perianth

segments six in two cycles, all similar, rela-

tively small, yellow-green. Stamens free; the

ﬁ laments short, anthers linear, basiﬁ xed, with

a short connective tip. Pollen grains 3-celled,

in dyads, globose, inaperturate, reticulate, the

reticulum is continuous between the two

grains. Gynoecium of three or rarely six car-

pels shortly connate at the base; carpel mar-

gins not completely fused (“The carpel walls

retain an open ventral suture throughout the

early and mid stages of development and even

in late developmental stages the suture does

not fully fuse” – Posluszny 1983); stigmas

sessile, decurrent on the upper part papillate.

Each carpel with two (rarely one) basal- axile,

anatropous ovules. Endosperm helobial. Fruits

of divaricate follicles. Seeds oblong, testal,

with large straight embryo and scanty vesti-

gial endosperm consisting of several layers of

strongly compressed cells. Testa smooth,

thick, derived mainly from the outer integu-

ment, the outer epidermis of the outer integu-

ment transforms into exotesta. The plants are

rich in the cyanogenic glucoside triglochinin.

n = 11.. . . . . . . . . . . . 1. scheuchzeriaceae.

3 Fruits achenes. Perennial or rarely annual

plants growing in marshes or other very wet,

often saline, places, sometimes aquatic or sub-

aquatic. Perennial species have short or elon-

gated rhizome or stolone. Roots ﬁ brous or

sometimes distally tuberous. Vessels only in

roots. Leaves mostly or wholly basal, alter-

nate, linear, with open sheathing base; sheath

adnate to the lamina, with two auricles; intrav-

aginal squamules membranous. Stomata

mostly paracytic. Crystals of calcium oxalate

few and tannin cells lacking. Flowers small

and inconspicuous, in terminal, mostly bract-

less spikes or spikelike racemes, bisexual or

unisexual (in Tetroncium dioecious), actino-

morphic or less often slightly zygomorphic,

3-merous, 2-merous, or 1-merous, generally

ebracteate, anemophilous. Perianth mostly of

six free and more or less similar segments in

two cycles (Triglochin) or of four, three, two,

or even one segments in one or two cycles,

rarely wanting (unisexual ﬂ owers of Lilaea).

Stamens mostly six in two cycles, rarely three,

four or eight, or (in Lilaea) even one; ﬁ laments

very short. Pollen grains 2-celled or 3-celled,

in monads, globose, inaperturate, tectate-colu-

mellate, reticulate. Gynoecium of 3, 4–10, or

commonly six carpels, nearly apocarpous

(Cyanogeton) to weakly connate (Tetroncium,

Maundia), but becoming separate in the fruit

stage; in (Lilaea) the gynoecium is either of

only of one carpel (Cronquist 1981) or

pseudomonomerous, initially consisted of

three carpels (Singh 1965), which is probably

supported by its three dorsal vascular bundles;

stigma mostly sessile or on short or rarely very

long stylodium; in each carpel or in each ovary

locule one or two, basal or less often (Maundia)

subapically pendulous, ovules. Ovules anatro-

Superorder ALISMATANAE 609

pous or less often (Maundia) orthotropous.

with a parietal cell cut off from the archespo-

rial cell. Endosperm nuclear. Fruits of achenes,

in Lilaea of a single achene. Seeds with

straight or nearly straight embryo and vesti-

gial endosperm consisting of thin, inconspicu-

ous membrane. Cyanogenic, containing the

cyanogenic glucoside triglochinin. n = 6, 8,

9, 11. . . . . . . . . . . . . . . . . 2. juncaginaceae.

2 Stamens four. Perennial or rarely annual aquatic

and mostly freshwater herbs with creeping rhi-

zome rooted in substrate. Roots ﬁ brous. Vessels

only in roots. Leaves alternate to opposite or rarely

verticillate, mostly linear or capillary, with well-

developed, open sheath, mostly ligulate, with

intravaginal squamules, submerged or (in some

species of Potamogeton) the upper ones ﬂ oating

and with expanded, lanceolate to elliptic, ovate or

orbicular lamina and a slender petiole; veins when

more than one parallel or arcuate, often connected

by cross-veins; leaves stipulate (commonly, the

stipular appendages adhering to the sheath or not),

or estipulate. Axillary scales present. The ﬂ oating

and sometimes also the upper submerged leaves

with paracytic stomata. Silica bodies and calcium

oxalate crystals are lacking. Flowers small to min-

ute, in axillary or terminal, bractless spikes that

are more or less elevated above the water, bisex-

ual, actinomorphic, mostly 4-merous. Perianth

segments four, in one cycle, free, bractlike, short-

clawed, adnate to the ﬁ laments or connectives of

the opposite stamens. Stamens four or rarely 2–3;

anthers nearly sessile. Pollen grains 3-celled,

ellipsoid to spheroid, semitectate, nearly inapertu-

rate, with an inconspicuous vestigial colpus under

which the intine is somewhat thickened.

Gynoecium of (1–3)4(5–16) free or only partly

united carpels; carpels sometimes shortly stipitate;

stigmas sessile or on short stylodia, slightly decur-

rent ventrally. Ovary superior, ovule solitary, pen-

dulous from the ventral margin of the carpel,

orthotropous or ortho-campylotropous, but after

fertilization becoming amphitropous, bitegmic,

crassinucellate, with a parietal cell cut off from the

archesporial cell. Endosperm helobial. Fruits

drupe-like or rarely (Groenlandia) achene-like.

Seeds without endosperm; embryo slightly curved,

with large hypocotyl. Seeds exotestal, or coat

crushed. Present ﬂ avonoids (apigenin, luteolin),

ﬂ avone glycosides, and C-glycosylated. The basic

chromosome number (x) is 7 (Haynes et al. 1998),

n ranging from 7 to 52. .3. potamogetonaceae.

1 Plants without vessels.

4 Flowers bisexual or rarely male by abortion of

gynoecium. Stamens three. Submerged marine

perennial herbs with thick, laterally ﬂ attened,

creeping, monopodially branching rhizome cov-

ered with ﬁ bers of old leaf sheaths and erect lat-

eral stems. All vegetative parts with scattered

tanniniferous cells containing proanthocyanins.

Xylem poorly developed and without vessels.

Leaves alternate, distichous on the rhizome, lin-

ear, parallel-veined or with only a mid-vein,

basally with open, biauriculate, ligulate sheath

that is persistent after the lamina has been shed.

Intravaginal squamules present at the nodes.

Stomata absent. Flowers in long-pedunculate

modiﬁ ed cymose inﬂ orescences subtended by

2–4 reduced leaves with broadened sheaths;

according to Hartog (1970: 123), the inﬂ ores-

cence “has to be regarded as an anthela of rhip-

idia, of which the rhipidia distally become

reduced to a ‘spike’ of ﬂ owers. Such a ‘spike’

has lost its terminal ﬂ owers as well as its bracts;

its cymose characters is still apparent from the

fact that the axis is not straight but proceeds more

or less zigzag.” Individual ﬂ owers small, sessile,

bractless, bisexual or rarely male by abortion of

gynoecium, hydrophilous. Perianth absent;

anthers large, sessile, with a much expanded,

shield-like connective produced beyond the loc-

uli, the latter widely separated. Pollen grains

3-celled, spheroidal when released but soon

become ﬁ lamentous (pollen tubes), inaperturate,

without exine. Gynoecium of one carpel with

sessile, obliquely placed, irregularly lobed, dis-

coid stigma, and one orthotropous ovule pendu-

lous from the carpellary margin. Fruits slightly

ﬂ eshy, with spongy pericarp, freely ﬂ oating when

shed. Seeds with very thin, membranous seed

coat and without endosperm embryo straight,

large consisting for the greater part of a large,

ﬂ eshy hypocotyl; hypocotyl with numerous tan-

nin cells. n = 10, dimorphic. 4. posidoniaceae.

4 Flowers unisexual.

5 Stamens two.

6 Fruits long-stipitate or sessile drupes.

Annual or rarely perennial submersed

610 Subclass I. ALISMATIDAE

glabrous herbs growing in fresh or brackish

waters. Roots ﬁ brous, few, non- septate,

from lower nodes of stems. Stems slender,

branched or unbranched, often dimorphic,

the lower stems rhizomatous, the upper

erect, leafy, the tips not modiﬁ ed into turi-

ons. Leaves alternate to subopposite, linear,

divided into blade and stipular sheath encir-

cling the stem; infravaginal scales present.

Flowers in 1-few-ﬂ owered capitate axillary

or terminal spike, the ﬁ rst enclosed by two

supopposite foliage leaves, bisexual, acti-

nomorphic, without perianth segments.

Stamens two, sessile with an expanded con-

nective; anthers with two bisporangiate

thecae dehiscing by longitudinal slits, con-

nective broad. Pollen grains elongate-arcu-

ate, inaperturate, intectate at both ends,

with distinctly separated columellae.

Carpels (2)4(-16), free; stigma sessile, pel-

tate, lobed; ovary 1-locular, ovule solitary,

more or less pendulous. Endosperm helo-

bial. Seed coat consists of a thin, 2-layered

testa and a tegmen of which only the outer

epidermis persists; endosperm wanting,

embryo with enlarged hypocotyls. Present

leucoanthocyanins, ﬂ avonols, and

G-glycosyl ﬂ avones, n = 8, 10–12, 20,

dimorphic.. . . . . . . . . . . . . .5. ruppiaceae.

6 Fruits nutlets with stony pericarp or vivipa-

rous. Marine perennial herbs with creeping

monopodially or sympodially branched rhi-

zomes. Roots often branched, with few to

many root hairs. Tannin cells present.

Leaves alternate, distichously arranged,

sheathing base and a ligule between sheath

and blade, linear, ﬂ at or terete, with three to

several veins. Flowers usually solitary and

terminal on a short branch or (Syringodium)

in small cymes. No vestige of perianth

except for a transient ridge that forms below

the stamens in the male ﬂ ower of

Syringodium ﬁ liforme (Tomlinson and

Posluszny 1978). Male ﬂ owers consist of

two united stamens; anthers on a common

ﬁ lament or virtually sessile, fused back to

back, tetrasporangiate; connective with an

apical prolongation (except in Syringodium).

Pollen grains without exine, become ﬁ la-

mentous (up to about 1 mm long). Gynoecium

of two free carpels, each with long, slender,

and simple (Halodule) or short and bibrachi-

ate or tribrachiate stylodium. Endosperm

nuclear. Fruits indehiscent, either with

stony pericarp (Halodule, Cymodocea,

Syringodium) or viviparous (Amphibolis).

Testa absent. Embryo consists either for the

larger part of the plumule with a lateral radi-

cle and a cylindrical hypocotyl or of a long

hypocotyl and a short plumule without

radicle. Sulphated phenolic compounds are

present in all genera; n = 7, 8, 10, 14, 15.

. . . . . . . . . . . . . . . . . . 7. cymodoceaceae.

5 Stamen solitary.

7 Ovule anatropous. Plants of fresh or alka-

line or brackish water. Not saponiferous.

Leaves 1-veined or incompletely 3-veined.

Flowers in small, axillary, usually complex,

cymose inﬂ orescences. Althenia and

Lepilaena have 3-membered perianth in

ﬂ owers of both sexes, while Zannichellia

have tiny, spathelike structure that may rep-

resent a perianth only in the female ﬂ ower.

Male ﬂ owers with a single ﬁ lament with 4,

8, or 12 microsporangia, and probably in

the last two cases represent two or three

fully connate stamens (Cronquist 1981;

Tomlinson 1982; Dahlgren et al. 1985).

Pollen grains inaperturate, subspheroidal or

subprolate, minutely spinose, without exine

or sometimes with thin exine and 1-colpate.

Female ﬂ owers with small, cupule-like or

3-lobed perianth. Gynoecium of (1-)3–4(-

9) free carpels, each with short or long sty-

lodium and symmetrically or asymmetrically

funnel-form stigma, which is ﬁ mbriate in

Lepilaena. Ovule solitary, ventral-pendu-

lous. Endosperm helobial. Fruits drupa-

ceous, with membranous exocarp, ﬂ eshy

mesocarp and stony endocarp, but in

Althenia they are dehiscent. Seeds with

spirally coiled cotyledon, swollen hypo-

cotyl, and very small radicle. Flavone

sulphates present. n = 6–8.

. . . . . . . . . . . . . . . 6. zannichelliaceae.

7 Ovule orthotropous. Submerged perennial,

monoecious or dioecious, marine herbs with

creeping or sometimes tuberous-thickened,

Superorder ALISMATANAE 611

monopodial, or sympodial rhizomes and

slender, often ﬂ attened stems. Tannin cells

absent. Vessels and stomata wanting. Leaves

alternate, usually decussate, linear or ﬁ li-

form, parallel-veined, sheathing at the base,

sheaths open or closed, commonly with

stipulelike margins, ligulate. Inﬂ orescence

terminal or lateral, sympodial, erect, con-

sisting of a panicle of rhipidia, but often

reduced to a single rhipidium; each rhipid-

ium consisting of 2–5 spathes, but some-

times reduced to single one; peduncle of

each spathe partially coalescent with the

axis from which it springs or completely

free; spathe consisting of a sheath and a leaf

blade; spathial sheath ligulate, enclosing a

sessile or stalked spadix on the dorsal side

of which the ﬂ owers are arranged (Hartog

1970). Individual ﬂ owers small, sessile,

bractless, unisexual (monoecious or dioe-

cious), without perianth. Stamen solitary;

anther sessile, extrorse, tetrasporangiate (but

an alternative explanation is that a ﬂ ower

with two widely separated unithecate (bispo-

rangiate) anthers – Tomlinson and Posluszny

2001), opening longitudinally. Pollen grains

2-celled or 3-celled, inaperturate, ﬁ lamen-

tous, up to about 2 mm long, with very thin

simple exine. Gynoecium of one carpel with

a short, bibrachiate stylodium and one api-

cal, pendulous ovule. Ovule orthotropous; a

parietal cell is not cut off, but there are peri-

clinal divisions in the nucellar epidermis.

Endosperm nuclear. Fruits small, with scari-

ous, dehiscent pericarp or less often

(Phyllospadix) with soft exocarp and a hard

ﬁ brous endocarp. Testa derived from the

outer integument. Tegmen degenerates.

Seeds without endosperm, with slightly

curved embryo consisting for the most part

of the hypocotyl and lacking a radicle; n = 6,

9, 10.. . . . . . . . . . . . . . . . . 8. zosteraceae.

1. SCHEUCHZERIACEAE

F. Rudolphi 1830. 1/1. In bogs of cold and temperate

regions of the Northern Hemisphere.

Scheuchzeria.

2. JUNCAGINACEAE

Richard 1808 (including Heterostylaceae Hutchinson

1934; Lilaeaceae Dumortier 1829; Maundiaceae

Nakai 1943) Triglochinaceae Chevalier 1827). 5/17.

Widely distributed in cold and temperate regions of

Northern and Southern Hemispheres; monotypic

genus Cycnogeton, very closely related to Triglochin,

is endemic to Australia and Tasmania, and another

monotypic genus Tetroncium is restricted to the

Holantarctic South America (Strait of Magellan);

Lilaea in mountains of Paciﬁ c America from British

Columbia to Chile and Argentina, and Maundia –

Northeastern Australia, southern Queensland, north-

ern New South Wales.

Triglochin, Cycnogeton, Tetroncium, Lilaea,

Maundia.

Related to the Scheuchzeriaceae. With the

Scheuchzeriaceae they share such a unique character

as the presence of triglochinin, but markedly differ in

many other respects, including nuclear endosperm and

pollen grains.

3. POTAMOGETONACEAE

Reichenbach 1828 (including Hydrogetonaceae Link

1829). 2/80–100. Subcosmopolitan.

Potamogeton (including Coleogeton), Groenlandia.

Related to the Juncaginaceae.

4. POSIDONIACEAE

Hutchinson 1934. 1/9. Along the coasts of the

Mediterranean, locally on the Atlantic coast of south-

western Europe, and along the coasts of southern,

southwestern and southwestern Australia and Tasmania.

Posidonia.

Molecular data indicate a close related to Ruppiaceae

and Cymodoceaceae.

5. RUPPIACEAE

Horaninow 1834. 1/7–8. Subcosmopolitan.

Ruppia.

Closely related to the Cymodoceaceae.

612 Subclass I. ALISMATIDAE

6. ZANNICHELLIACEAE

Chevalier 1827. 4/12. Subcosmopolitan.

Zannichellia, (including Pseudalthenia) Althenia,

Lepilaena, Vieisia.

Recent molecular studies indicate that Zanni-

chelliaceae are most closely related to Potamo-

getonaceae.

7. CYMODOCEACEAE

N. Taylor 1909. 5/16. Mainly tropical and subtropical

seas with a few species in warm-temperate seas.

Cymodocea, Halodule, Syringodium, Amphibolis,

Thalassodendron.

Related to the Zannichelliaceae, but much more

specialized.

8. ZOSTERACEAE

Dumortier 1829. 3/14–20. Temperate seas of both

hemispheres, with a few species extending into tropi-

cal seas.

Zostera, Heterozostera, Phyllospadix.

A very specialized group of marine alismatids that

probably derived from the juncaginalean ancestor.

Bibliography

Aalto M. 1970. Potamogetonaceae fruits. I. Recent and subfossil

endocarps of the Fennoscandian species. Acta Bot. Fenn. 88:

1–85.

Agrawal JS 1952. The embryology of Lilaea subulata H. B. K.

with a discussion on its systematic position. Phytomorphology

2: 15–29.

Albergoni FG, B Basso, and G Tedesco. 1978. Considerations

sur l’anatomie de Posidonia oceanica (Zosteraceae). Plant

Syst. Evol. 130: 191–210.

Arber A. 1940. Studies in ﬂ ower structure: VI. On the residual

vascular tissue in the apices of reproductive shoots, with

special reference to Lilaea and Amherstia. Ann. Bot. 2(4):

617–627.

Barnabas AD. 1982. Fine structure of the leaf epidermis of

Thalassodendron ciliatum (Forsk.) den Hartog. Aquatic Bot.

12: 41–55.

Barnabas AD. 1983. Composition and ﬁ ne structural features of

longitudinal veins in leaves of Thalassodendron ciliatum.

South Afr. J. Bot. 2: 317–325.

Barnabas AD. 1994. Anatomical, histochemical and ultrastruc-

tural features of the seagrass Phyllospadix scouleri Hook.

Aquatic Bot. 49: 167–182.

Barnabas AD and HJ Arnott. 1987. Zostera capensis Setchell:

root structure in relation to function. Aquatic Bot. 27:

309–322.

Barnabas AD and S Kasavan. 1983. Structural features of the

leaf epidermis of Halodule uninervis. South Afr. J. Bot. 2:

311–316.

Black JM. 1913. The ﬂ owering and fruiting of Pectinella

antarctica (Cymodocea antarctica). Trans. Proc. Roy. Soc.

South Australia 37: 1–5.

Bowes G, SK Rao, GM Estavillo, and JB Reiskind. 2002. C

mechanisms in aquatic angiosperms: Comparisons with

terrestrial C

systems. Funct. Plant Biol. 29: 379–392.

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

(Araceae) and Potamogeton (Potomagetonaceae). In XVI

Int. Bot. Congr. Abstracts, p. 240. Missouri Botanical

Garden, St. Louis, MO.

Buzgo M, DE Soltis, PS Soltis, S Kim, H Ma, BA Hauser,

J Leebens-Mackl, and B Johansen. 2006. Perianth develop-

ment in the basal monocot Triglochin maritime

(Juncginaceae). In: JT Columbus, EA Friar, JM Prince,

MG Simpson, eds. Monocots: comparative biology and

evolution (excluding Poales), pp. 107–125. Clermont (Aliso

22: 107–125).

Cambridge ML and J Kuo. 1982. Morphology, anatomy and his-

tochemistry of the Australian seagrasses genus Posidonia

sinuosa Cambridge & Kuo. Aquatic Bot. 14: 1–14.

Campbell DH. 1897. A morphological study of Nolas and

Zannichellia. Proc. Calif. Acad. Set., ser. 3, 1: 1–71.

Campbell DH. 1898. Development of the ﬂ ower and embryo of

Lilaea subulata H. B. K. Ann. Bot. 12: 1–12.

Campbell GKG. 1936. The anatomy of Potamogeton pectinatus.

Trans. Proc. Bot. Soc. Edinb. 32: 179–186.

Charlton WA. 1981. Features of the inﬂ orescence of Triglochin

maritimum. Canad. J. Bot. 59: 2108–2115.

Chrysler MA. 1907. The structure and relationships in

Potamogetonaceae and allied families. Bot. Gaz. 44:

161–188.

Cook MT. 1908. The development of the embryo sac and embryo

of Potamogeton lucens. Bull. Torrey Bot. Club 35:

209–218.

Cox PA and CJ Humphries. 1993. Hydrophilous pollination and

breeding system evolution in sea grasses: a phylogenetic

approach to the evolutionary ecology of the Cymodoceaceae.

Bot. J. Linn. Soc. 113: 217–226.

Cox PA, PB Tomlinson, and K Nieznanski. 1992. Hydrophilous

pollination and reproductive morphology in the seagrass

Phyllospadix scouleri (Zosteraceae). Plant Syst. Evol. 180:

65–75.

Dahlgren KVO. 1939. Endosperm- und Embryobildung bei

Zostera marina. Bot. Not. 1939: 607–615.

De Cock AWAM. 1978. Germination of the thread like pollen

grains of the seagrass Zostera marina L. Bull. Soc. Bot.

France Act. Bot. 1–2: 145–148.

De Cock AWAM. 1980. Flowering pollination and fruiting in

Zostera marina L. Aquatic Bot. 9: 201–220.

Ducker SC, NJ Foord, and RB Knox. 1977. Biology of Australian

sea grasses: the genus Amphibolis C. Agardh (Cymodoeaceae).

Aust. J. Bot. 25: 67–95.

Superorder ALISMATANAE 613

Ducker SC, JM Pettitt, and RB Knox. 1978. Biology of

Australian sea grasses: Pollen development and submarine

pollination in Amphibolis antarctica and Thalassodendron

ciliatum (Cymodoceaceae). Aust. J. Bot. 26: 265–285.

Gardner RO. 1976. Binucleate pollen in Triglochin L. N. Z. J.

Bot. 14: 115–116.

Graves AH. 1908. The morphology of Ruppia maritima. Conn.

Acad. Arts. Sci. 14: 59–170.

Grönland J. 1851. Beitrag zur Kenntnis der Zostera marina. Bot.

Z. 9: 185–192.

Guo Y-H and CDK Cook. 1990. The ﬂ oral biology of

Groenlandia densa (L.) Fourreau (Potamogetonaceae).

Aquatic Bot. 38: 283–288.

Gupta BL. 1934. Contribution to the life history of Potamogeton

crispus. J. Indian Bot. Soc. 13: 51–65.

Hagstrom ML. 1916. Critical researches on the Potamogeton.

Kongl. Svenska Vetenskapsakad. Handl. 55: 1–281.

Hartog C den. 1970. Sea grasses of the World. Verh. Kon. Ned.

Akad. Wetensch. Afd. Natuurk. Tweede Sect. 59(I): 1–275.

Haynes RR. 1978. The Potamogetonaceae in the southeastern

United States. J. Arnold Arbor. 59: 170–191.

Haynes RR and LB Holm-Nielsen. 1987. The Zannichelliaceae

in the southeastern United States. J. Arnold Arbor. 68:

259–268.

Haynes RR, DH Les, and LB Holm-Nielsen. 1998a.

Juncaginaceae. In: K Kubitzki, ed. The families and genera

of vascular plants, vol 4, 260–263. Springer, Berlin/

Heidelberg/New York.

Haynes RR, DH Les, and LB Holm-Nielsen. 1998b.

Potamogetonaceae. In: K Kubitzki, ed. The families and gen-

era of vascular plants, vol 4, pp. 408–415. Springer, Berlin/

Heidelberg/New York.

Haynes RR, LB Holm-Nielsen, and DH Les. 1998c. Ruppiaceae.

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

plants, vol 4, pp. 445–448. Springer, Berlin/Heidelberg/

New York.

Haynes RR, DH Les, and LB Holm-Nielsen. 1998d.

Scheuchzeriaceae. In: K Kubitzki ed., The families and gen-

era of vascular plants, vol 4, pp. 449–451. Springer, Berlin/

Heidelberg/New York.

Haynes RR, DH Les, and LB Holm-Nielsen. 1998e.

Zannichelliaceae. In: K Kubitzki, ed. The families and gen-

era of vascular plants, vol 4, pp. 470–474. Springer, Berlin/

Heidelberg/New York.

Hegelmaier F. 1870. Über die Entwicklung der Blütenteile von

Potamogeton. Bot. Z. 18: 283–320.

Hofmeister W. 1852. Zur Entwicklungsgeschichte der Zostera.

Bot. Z. 10: 121–131, 137–149, 157–158.

Holferty GM. 1901. Ovule and embryo of Potamogeton natans.

Bot. Gaz. 31: 339–346.

Isaac FM. 1969 (1970). Floral structure and germination in

Cymodocea ciliata. Phytomorphology 19: 44–51.

Jacobs SWL and MA Brock. 1982. A revision of the genus

Ruppia (Potamogetonaceae) in Australia. Aquatic Bot. 14:

325–337.

Kamelina OP. 1990. Potamogetonaceae. In: TB Batygina and

MS Yakovlev, eds. Comparative embryology of ﬂ owering

plants. Monocotyledons. Butomaceae – Lemnaceae, vol 1,

pp 34–39. Nauka, Leningrad (in Russian).

Kamelina OP and ES Terekhin. 1990. Ruppiaceae. In:

TB Batygina and MS Yakovlev, eds. Comparative embryol-

ogy of ﬂ owering plants. Monocotyledons. Butomaceae –

Lemnaceae, vol 1, pp. 39–44. Nauka, Leningrad

(in Russian).

Kato Y, K Aioi, Y Omori, N Takahata, and Y Satta. 2003.

Phylogenetic analyses of Zostera species based on rbcL and

matK sequences: implications for the origin and diversiﬁ ca-

tion of seagrasses in Japanese waters. Genes Genet. Syst. 78:

329–342.

Kay QON. 1971. Floral structure in the marine angiosperms

Cymodocea serrulata and Thalassodendron ciliatum

(Cymodocea ciliata). Bot. J. Linn. Soc. 64: 423–429.

Keighery GJ and DJ Coates. 1981. Chromosome counts

in Posidonia (Posidoniaceae). Plant Syst. Evol. 137:

221–222.

Kirkman H. 1975. Male ﬂ oral structure in the marine angio-

sperm

Cymodocea serrulata (R. Br.) Ascherson and Magnus

(Zannichelliaceae). Bot. J. Linn. Soc. 79: 267–268.

Kuo J. 1978. Morphology, anatomy and histochemistry of

the Australian sea grasses genus Posidonia Konig

(Posidoniaceae). I. Leaf blade and leaf sheath of Posidonia

australis Hook.f. Aquatic Bot. 5: 171–190.

Kuo J. 1983. The nacreous walls of sieve elements in sea grasses.

Am. J. Bot. 70: 159–164.

Kuo J. 1993a. Functional leaf anatomy and ultrastructure in a

marine angiosperm, Syringodium isoetifolium (Aschers.)

Dandy (Cymodoceaceae). Aust. J. Mar. Freshwater Res. 44:

59–73.

Kuo J. 1993b. Root anatomy and rhizosphere ultrastructure in

tropical sea grasses. Aust. J. Mar. Freshwater Res. 44:

75–84.

Kuo J. 2001. Chromosome numbers of the Australian

Zosteraceae. Plant Syst. Ecol. 226(3–4): 155–163.

Kuo J and ML Cambridge. 1978a. Morphology, anatomy, and

histochemistry of the Australian sea grasses of the genus

Posidonia Konig (Posidoniaceae): I. Leaf blade and leaf

sheath of Posidonia australis Hook. f. Aquatic Bot. 5:

163–170.

Kuo J and ML Cambridge. 1978b. Morphology, anatomy, and

histochemistry of the Australian species of the genus

Posidonia Konig (Posidoniaceae): rhizome and root of

Posidonia australis Hook. f. Aquatic Bot. 5: 191–206.

Kuo J and H Kirkman. 1987. Floral and seeding morphology

and anatomy of Thalassodendron pachyrhizum den Hartog

(Cymodoceaceae). Aquatic Bot. 29: 1–17.

Kuo J and H Kirkman. 1990. Anatomy of vipiparous sea grasses

of Amphibolis and Thalassodendron and their nutrient

supply. Bot. Mar. 33: 117–126.

Kuo J and AJ McComb. 1989. Sea grass taxonomy, structure

and development. In: AWD Larkum, AJ McComb,

SA Shephard, eds. Biology of seagrasses. A treatise on the

biology of seagrasses with special reference to the Australian

region, pp. 6–73. Elsevier Science, Amsterdam.

Kuo J and AJ McComb. 1998a. Cymodoceaceae. In: K Kubitzki,

ed. The families and genera of vascular plants, vol 4,

pp 133–140. Springer, Berlin/Heidelberg/New York.

Kuo J and AJ McComb. 1998b. Posidoniaceae. In: K Kubitzki,

ed., The families and genera of vascular plants, vol 4,

pp 404–408. Springer, Berlin/Heidelberg/New York.

Kuo J and AJ McComb. 1998c. Zosteraceae. In: K Kubitzki, ed.

The families and genera of vascular plants, vol 4, pp. 496–502.

Springer, Berlin/Heidelberg/New York.