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Order 125. HYDROSTACHYALES

Submerged fresh-water perennial or annual herbs

with short, thick, tuberlike stems and ﬁ brous roots;

stems, leaves, and adventitious roots provided with

numerous small, scale-like or fringed emergences.

Vascular system much reduced, in the disc-like stem

procambial tissue with annular- and spiral vessels is

present (Erbar and Leins 2004). Leaves alternate,

basal, elongate, entire to two or three times pinnati-

sect, dilated and ligulate at the base, with inconspicu-

ous, membranous, intrapetiolar stipule. Stomata

wanting. Flowers small and inconspicuous, sessile in

the axils of the bracts of the dense spike, unisexual,

dioecious or seldom monoecious, without a perianth.

The male ﬂ ower usually is interpreted as consisting

of a single stamen, which is deeply divided into two

theca (Rauh and Jager-Zurn 1966; Hutchinson 1973;

Cronquist 1981; Takhtajan 1987). However, accord-

ing to Leins and Erbar (1990), it consists of two inde-

pendent disporangiate stamens that can be derived

from originally tetrasporangiate stamens in which the

sporangia of the neighboring thecae have fused api-

cally (as in some Scrophulariaceae, e.g., Verbascum)

and shifted into a lateral position (as in some

Scrophulariaceae, e.g., in Hebenstretia and

Zaluzianskya) microsporangia opening longitudi-

nally. Tapetum secretory. Micro sporogenesis simul-

taneous. Pollen grains in permanent tetrahedral or

hexahedral to rhomboidal tetrads (Erbar and Leins

2004), 2-celled, inaperturate. Gynoecium of two

united carpels arranged in trans versal position and

with two elongate, persistent, subulate stylodia that

are sometimes connate at the base; ovary superior,

zygomorphic, 1-locular, with numerous ovules on

two parietal placentas. Ovules anatropous, unitegmic,

tenuinucellate, with a narrow, elongated nucellus

with a tiny gametophyte located at its apex, with

endothelium. Female gametophyte of Polygonum-

type, Endosperm cellular, with micropylar hausto-

rium. Fruits septicidal capsules with numerous tiny,

exotestal seeds, outer cell walls much thickened,

mucilaginous (contain pectin); endosperm scanty or

lacking. Contain kaempferol, mono- and di-gluco-

sides (Scogin 1992), and caffeic acid (Rønsted et al.

2002), n = 10–12 (Hydrostachys imbricata – Palm

1915).

Probably derived from Scrophulariaceae (see Jäger-

Zürn 1965; Rauh and Jäger-Zürn 1966; Takhtajan

1966; Leins and Erbar 1990) and probably have a com-

mon origin with Plantaginaceae.

Taxonomic position is unknown.

1. HYDROSTACHYACEAE

Engler 1894. 1/22. Madagascar and tropical and South

Africa.

Hydrostachys

588 Subclass VIII. LAMIIDAE

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A. Takhtajan, Flowering Plants, 589

Embryo, when differentiated, always with one

cotyledon. The cotyledon usually with two main vas-

cular bundles. Leaf venation striate or of derived types,

mostly arcuate-striate or longitudinally striate

(parallel), less often palmate-striate or pinnate-striate,

almost always more or less closed at the apex (the

veins emerging from the leaf base usually run together

again at their apices). Leaves usually not clearly

divided into petiole and lamina, less often more or less

differentiated, but in these cases the “petiole” and the

“lamina” are not homologous to those of magnoliop-

sids (are of secondary origin), often with sheathing

base. Leaf traces usually numerous. Prophylls (includ-

ing bracteoles) usually solitary and nearly always

adaxial. Vascular bundles usually without cambium or

rarely with vestigial cambium only. Vascular system of

the stem usually consists of many separate scattered

bundles or sometimes of two or more rings of vascular

bundles, and the axis mostly attains its full diameter

early, after which no increase in thickness takes place;

only in some groups does thickening of the axis occur

by means of division and enlargement of ground paren-

chyma cells (so-called diffuse secondary growth), as in

palms, or by means of special kind of cambium that

arises in the parenchyma outside the primary vascular

system, as in some herbaceous and woody Lilianae.

Sieve-element plastids of P-type with several to numer-

ous cuneate (triangular) crystalloid bodies (lacking in

all magnoliopsids studied except Saruma and Asarum

in Aristolochiaceae). Phloem without parenchyma.

Usually without clearly differentiated bark and pith.

The primary root is usually ephemeral, dries out early

in the growth of the plant, and is replaced by an adven-

titious root system that develops from the stem or (as

in grasses) directly from the hypocotyl. Ontogenetically

root cap and root epidermis are of different origin.

Usually herbs, but often secondarily arborescent plants

(primary woody plants are absent among the monocots).

Flowers usually 3-merous, sometimes 4- or 2-merous,

very rarely 5-merous. Nectaries predominantly septal.

Pollen grains mostly 1-colpate (sulcate) or of derived

types, often 1-porate.

The Liliopsida most probably originated from some

very ancient vesselless herbaceous member of Magno-

liopsida that had atactostelic vascular system, P-type

sieve-element plastids, 3-merous ﬂ owers, apocarpous

gynoecium with laminar-diffuse (scattered) placenta-

tion, bitegmic and crassinucellate ovules (with parietal

tissue between the female gametophyte) and the nucel-

lar epidermis, and primitive 2-celled and 1-colpate

pollen grains.

Unfortunately there is no convincing dicotylidonous

sister group to the monocotyledons. According to some

authors the nearest group are nymphaeids. Some of the

relatively most archaic monocots have some similari-

ties with the nymphaeids (Hallier 1905; Schaffner 1929,

1934; Eber 1934; Takhtajan 1954, 1959, 1969, 1987;

Kaul 1967; Cronquist 1968, 1981, 1988). As long ago

as 1905, Hallier suggested that the Nymphaeaceae (s.1.)

were the “ancestors of the whole division of monocoty-

ledons” though later (1912) he changed his opinion.

According to Arber (1920: 309), the Nymphaeaceae

“descended from a stock closely related to that which

gave rise to the monocotyledons.” Similar ideas have

also been expressed by some other botanists.

The nymphaeoids and some archaic monocots do

indeed have some important characters in common.

In the morphology of their gynoecia the families

Butomaceae and Limnocharitaceae resemble the

Cabombaceae, and in their laminar-diffuse placenta-

tion they recall the Nymphaeaceae. There are also

some other important similarities, including atacto-

stelic vascular cylinder and especially root ontogeny

(see Voronin 1964) as well as the development of

Class LILIOPSIDA

(MONOCOTYLEDONS)

590 Class LILIOPSIDA

female gametophytes, stomatal patterns, seed anatomy,

and the arrangement of the ﬁ rst leaves (prophylls) on

lateral axes. However, the sieve-element plastids of the

nymphaeids are of S-type, and they are too specialized

to be considered the ancestors of monocots. It is much

more probable that they evolved from some remote

common ancestor that was already more or less adapted

to a relatively wet (but not yet aquatic) habitat.

Henslow (1911) considered the distinctive features of

monocots the result of the primary adaptation to an

aquatic habitat while Jeffrey (1917: 415) in his classical

“Anatomy of Woody Plants” suggested an aquatic or

amphibious way of life might have led to the loss of

cambial activity. Henslow’s hypothesis has been criti-

cized by Sargant (1903, 1904), who concluded that many

of the characteristic features of the monocots may be

easier explained as having arisen as a result of adaptation

to a geophilous habit. But apparently nearer to the truth

was Parkin (1923: 59), who suggested “the golden mean”

between the two hypotheses. He writes: “Respecting the

relative merits of an aquatic or geophilous ancestry of

monocotyledons, the two views may be somewhat rec-

onciled by regarding the earliest ones as neither mark-

edly aquatic nor extremely geophilous—in fact, marsh

plants with stout rhizomes. Some of their descendants

have become completely hydrophytic, others sharply

geophytic, while others have returned to the arborescent

habit by fresh means.” Apparently the ancient common

ancestor of both the Nymphaeidae and monocots was a

hygrophilous or perhaps even amphibious geophyte in

which geophyly arose under wet terrestrial conditions –

most probably under the forest canopy or in the forest

margin. But as is well known, underground storage is

usually a response to a resting season, and geophytes are

abundant and highly diversiﬁ ed in areas with a pro-

nounced resting season (see Bews 1927). Therefore,

they could originate in a climate having a marked dry

season (Sargant 1903; Stebbins 1974).

The class Liliopsida includes 4 subclasses, 31

orders, 120 families, more than 3,000 genera, and

about 65,000 species.

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