Takhtajan Armen. Flowering Plants

Подождите немного. Документ загружается.

Superorder ROSANAE 303

obdiplostemonous cycles, less often isomer-

ous and alternate with the petals (Crassuloideae);

ﬁ laments free or very rarely connate below, in

sympetalous ﬂ owers borne on the corolla tube

(the antepetalous ones a little higher than the

antesepalous ones); anthers latrorse or more or

less introrse, more or less basiﬁ xed. Pollen

grains 2-celled, usually 3-colporate. Carpels as

many as the sepals or petals, free or connate

only at the base, rarely (Diamorpha, Pagella)

connate nearly to the middle, each tapering into

a short or elongate stylodium; near the base of

each carpel a small nectariferous scale borne

externally, which is larger and petaloid in

Monanthes and some species of Sedum. Ovules

usually numerous in each carpel, rarely few or

sometimes only one, on submarginal placentas,

crassinucellate. Endosperm cellular, some-

times with chalazal haustoria. Fruits mostly

many-seeded multifollicles which dehisce

xerochastically, rarely (Diamorpha) syncarpous

multifollicle with the carpels opening dorsally.

Seeds very small; “seed coat is 4-layered: the

exotestal cells have thickened outer wall,

the inner exotegmic cell layer is pigmented,

and the two middle layers are completely

crushed” (Thiede and Eggle 2007); embryo

long, straight; endosperm copious to scanty.

Plants often with pyridine alkaloids, ﬂ avones,

acylated ﬂ avonol glycosides; red pigment com-

mon, even in roots; n = 4–8 with a reduction to

7 in Crassula and n = 6, 7 in the Leucosedum

clade (Mort et al. 2001). . . . 4. crassulaceae.

5 Gynoecium usually syncarpous, rarely apocar-

pous or semiapocarpous.

7 Fruits juicy berries crowned by persistent calyx.

Seeds usually numerous, rather small, with a

funicular aril investing the seed or reduced to

crenulate ﬂ eshy placental ridge; exotestal cells

palisade, mucilaginous, endotestal cells crystal-

liferous, radial and inner walls ligniﬁ ed; embryo

small, short; endosperm oily, with slightly thick-

ened walls. Flowers small, in racemes or solitary,

bisexual or unisexual, actinomorphic. Sepals

(4)5, basally connate, imbricate or subvalvate,

sometimes petaloid, persistent; calyx tube adnate

to the ovary. Petals (or staminodia) ﬁ ve, small,

squamiform, obovate or subulate, adnate to the

calyx tube. Stamens ﬁ ve, alternating with the

petals. Pollen grains periporate. Gynoecium of

two carpels with free or more or less connate

stylodia and simple stigma. Ovary inferior,

1-locular, with few to numerous ovules on lat-

eral placentas. Endosperm cellular, helobial, or

nuclear. Leaves alternate, simple, variously

lobed, with palmate venation, estipulate but

often dilated at base; hydathodes associated with

the apiculate marginal teeth. Stomata anamo-

cytic. Nodes trilacunar. Vessels with scalariform

perforations or very rarely also with a few sim-

ple perforations; lateral pitting mostly alternate.

Fibers often with bordered pits. Rays heteroge-

neous. Axial parenchyma absent or in very rare

bands. Erect, arching, trailing, or prostrate

shrubs, often with conspicuous 3-forked or sim-

ple nodal spines, and smaller, internodal bristles.

Present ﬂ avonoids and tannins, proanthocyani-

dins, myricetin glycosides and some acids; n = 8,

rarely 16. . . . . . . . . . . . . . 8. grossulariaceae.

7 Fruits mostly capsular, sometimes multifollicles

or syncarpous multifollicles.

8 Vessels usually with simple perforations.

Fibers, when present, small, with bordered

pits. Nodes usually trilacunar or less often

multilacunar (Astilboides, Astilbe, Bergenia,

Mukdenia, Rodgersia, Darmera). Perennial or

rarely annual or biennial, often rhizomatous

herbs; hairs uni- to multiseriate with multicel-

lular glandular head; young stem with separate

bundles (pseudosiphonostele). Leaves gener-

ally basal, usually alternate, rarely opposite,

simple, entire or lobed and pinnately or pal-

mately veined, or pinnately or palmately com-

pound or decompound, estipulate. Stomata

usually anomocytic. Flowers in various kinds

of cymose or racemose inﬂ orescences or rarely

solitary, bisexual or rarely unisexual (as in

Tanakaea), actinomorphic or rarely zygomor-

phic. Perianth usually 5-merous, less often

3-merous, forming a more or less developed

ﬂ oral tube, which is free or variously adnate to

the base of the ovary. Sepals imbricate or val-

vate. Petals 5(-10), imbricate or valvate, rarely

wanting (as in Chrysosplenium and Bensoniella

oregona). Stamens as many as petals and alter-

nate with them or twice that number, in which

case there is a stamen opposite each perianth;

sometimes eight, four or even (Tolmiea) three

304 Subclass VI. ROSIDAE

stamens; anthers slightly introrse or latrorse,

mostly basiﬁ xed, sometimes slightly dorsi-

ﬁ xed, versatile. Pollen grains 2-celled, 3-col-

pate, 3-colporate or 6–9-porate. Gynoecium

mostly syncarpous, rarely more or less apocar-

pous (Astilbe spp. and Darmera), of 2–5

(mostly only 2) carpels, usually with more

or less free, apical stylodia; stigmas dorsal,

papillate; the carpels sometimes more or less

open ventrally in the free distal regions (as

in Huchera, Tiarella, Tolmiea, Mitella,

Bensoniella, and Tellima); ovary superior to

inferior; ovules 9–30 in several rows, pendu-

lous or ascending, anatropous, bitegmic or

rarely (Saxifraga section Micranthes, Darmera)

unitegmic; micropyle zig-zag or not zig-zag.

Endosperm cellular, helobial, or nuclear and in

some members develops various kinds of

haustoria. Fruits mostly septicidal capsules,

sometimes multifollicles or syncarpous multi-

follicles. Seeds small, smooth, exotestal cells

with outer wall more or less thickened; embryo

straight, minute to medium sized, surrounded

by copious, oily endosperm. Present ﬂ avonols

(quercetin, kaempferol, myricetin), sometimes

proanthocyanidins (cyanidin and delphinidin),

n = 7–17, mostly 7.. . . . . . 7. saxifragaceae.

8 Vessels usually with scalariform perforations.

9 Ovary 5-locular. Much branched shrubs.

Conical to peltate glandular hairs present.

Vessels with simple to scalariform perfora-

tions. Nodes trilacunar. Leaves alternate,

dentate, shining, glandular, and glutinous-

resinous above, softly pubescent below;

stipules minute, subulate, deciduous. Stomata

anomocytic. Flowers in few-ﬂ owered,

corymbose cymes, bisexual, actinomorphic,

5-merous. Calyx tube turbinate, adnate to

ovary, lobes triangular-subulate, erect, val-

vate. Petals imbricate, conspicuous, at length

reﬂ exed, pubescent, white. Stamens ten in

two cycles, inserted with the petals, erect,

the ﬁ ve outer fertile, with broad ﬁ laments

denticulate at apex and ovoid, dorsiﬁ xed,

mucronate, introrse anthers. Pollen grains

3-colporate, with complete tectum and com-

plex endoaperture. Gynoecium of ﬁ ve car-

pels; style shortly 5-lobed, with short and

radiate stigmas, separating more with age;

ovary inferior, with 4–6 ascending axile

ovules per locule. Fruits few-seeded, sep-

ticidal, woody capsules crowned by the

erect sepals and reﬂ exed petals. Seeds

with cartilaginous testa, attenuated at either

end; embryo elongate, surrounded by copi-

ous, ﬂ eshy endosperm. Plants contain

C-glycosyl ﬂ avones and 3-O-glycosides of

quercetin.. . . . . . . 9. pterostemonaceae.

9 Ovary 2-locular. Trees and shrubs with sim-

ple, unicellular and glandular hairs; young

stem with separate bundles. Branches with

lamillate pith. Vessels with scalariform per-

forations that have numerous slender bars;

lateral pitting scalariform. Rays uniseriate,

heterocellular. Nodes trilacunar with three

traces. Leaves alternate, dentate (spiny-

dentate in Itea ilicifolia), pinnately veined,

estipulate or with minute, linear stipules.

Stomata paracytic. Flowers small, in ter-

minal or axillary racemes or panicles,

bisexual or polygamous, actinomorphic.

Sepals ﬁ ve, basally connate into a short

turbinate or obconic tube, lobes valvate or

open, persistent. Petals ﬁ ve, valvate, per-

sistent. Stamens ﬁ ve, alternating with the

petals, inserted under the edge of the more

or less developed annular nectary disc;

ﬁ laments subulate; anthers small, oblong

to ovoid, dorsiﬁ xed, introrse. Pollen grains

2-porate, bilateral, ektexine homogeneous.

Gynoecium of two united carpels; stylodia

connate into a 2-grooved style but become

free at maturity although often still united

by capitate stigmas; ovary nearly superior

to more than three-quarters inferior, longi-

tudinally 2-grooved, with usually numerous

ovules on axile placentas. Fruits septicidally

2-valved capsules; fruit valves often

attached by the stigma. Seeds few to many;

exotestal cells with outer walls thickened;

embryo large, curved, surrounded by sparse

ﬂ eshy endosperm. Present C-gycosyl

glavones, an unusually in Saxifragales

(Bohm et al. 1988); n = 11. .10. iteaceae.

1. TETRACARPAEACEAE

Nakai 1943. 1/1. Mountains of Tasmania.

Tetracarpaea.

Superorder ROSANAE 305

Most closely allied with the Penthoraceae,

Crassulaceae, and Saxifragaceae, which is supported by

comparative morphological studies (Hils et al. 1988)

and also rbcL sequence data (Morgan and Soltis 1993).

2. APHANOPETALACEAE

Doweld 2001. 1/2. Southeastern Australia.

Aphanopetalum.

Probably related to Haloragaceae (see Fishbein

et al. 2001).

3. PENTHORACEAE

Rydberg ex Britton 1901. 1/2. Far East of Russia,

Japan, Korea, China, northern Vietnam, and eastern

parts of the United States from Maine to Ontario and

Minnesota and south to Florida and Texas.

Penthorum.

Related to both the Crassulaceae and Saxifragaceae,

differing from them in having vessels with scalariform

perforations, ﬁ bers with bordered pits, in their chemical

makeup (Jay 1970; Soltis and Bohm 1982), and espe-

cially in the presence of an operculum (Nemirovich-

Danchenko 1994a). Grund and Jensen (1981)

declined to place Penthorum in either Crassulaceae or

Saxifragaceae, and according to Haskins and Hayden

(1987) Penthorum is probably best classiﬁ ed in the

monogeneric Penthoraceae. A separate family

Penthoraceae has been accepted by van Tieghem (1898),

Rydberg (1905), Novak (1961, 1972), Airy Shaw (1973),

Stern (1974) and Thorne (2006). The rbcL sequence

data also suggest that Penthorum should not be accom-

modated in either Saxifragaceae or Crassulaceae

although the genus is more closely related to Crassulaceae

than to Saxifragaceae (Morgan and Soltis 1993).

4. CRASSULACEAE

J. Saint-Hilaire 1805 (including Cotyledonaceae

Martynov 1820, Rhodiolaceae Martynov 1820,

Sedaceae Martynov 1820, Sempervivaceae A.L. de

Jussieu 1782, Tillaeaceae Martynov 1820). About

41/1550–1600. Widely distributed, mainly in dry and

warm regions, but centered in tropical and especially

South Africa.

Classiﬁ cation after J.Thiede and U.Eggli (2007).

4.1 SEMPERVIVOIDEAE (including Sedoideae and

Echeverioideae)

Androecium usually obdiplostemonous. Flowers

3-12(-32)-merous, mostly 5-merous. Petals free or

basally connate, rarely corolla sympetalous. – thele-

phieae: Sinocrassula, Kungia, Meteros tachys,

Orostachys, Hylotelephium Hylotelephium; umbi-

liceae: Umbilicus (including Chiasto-phyllum),

Pseudosedum, Rhodiola, Phedimus; semperviveae:

Sempervivum, Petrosedum; aeonieae: Aichryson,

Monanthes, Perrierosedum, Aeonium (including

Greenovia), sedeae: Pistorinia, Rosularia, Prometheum,

Afrovivella, Sedella (including Parvisedum), Dudleya,

Sedum (including Diamorpha, Cremnophyla,

Mucizonia), Villadia, Lenophyllum, Graptopetalum

(including, Tacitus), Thompsonella; echeverieae:

Echeveria, Pachyphytum.

4.2 KALANCHOIDEAE (including Cotyledonoideae)

Androecium obdiplostemonous. Flowers 5-merous

or 4-merous. Petals more or less forming tubes or

campanulate corolla. – Adromischus, Kalanchoe

(including Bryophyllum), Cotyledon, Tylecodon.

4.3 CRASSULOIDEAE

Androecium haplostemonous. Petals free or basally

connate, rarely forming tubular or campanulate corolla. –

Crassula (including Tillaea, Rochea, Dinacria, Pagella),

Hypagophytum.

Close to the Penthoraceae, differing mainly in

succulent leaves, anisocytic stomata, vessels with

simple perforations, and the presence of small nec-

tariferous appendages near the base of each carpel.

5. HALORAGACEAE

R. Brown 1814 (including Cercodiaceae A.-L. de

Jussieu 1817, Myriophyllaceae Schultz-Schultzenstein

1832). 8/145. Mainly in the Southern Hemisphere, par-

ticularly in Australia; Myriophyllum is cosmopolitan

and Proserpinaca is conﬁ ned to the Northern

Hemisphere; Laurembergia is endemic to South Africa.

Haloragis, Haloragodendron, Glischrocaryon,

Meziella, Gonocarpus, Laurembergia, Proserpinaca,

Myriophyllum.

The Haloragaceae have often been considered to be

allied to the Myrtales, particularly to the Onagraceae.

However, they differ from the Myrtales in lacking

internal phloem and vestured pits, in having free

306 Subclass VI. ROSIDAE

stylodia covered by large papillae and cellular

endosperm in some of their members, and seeds with

well- developed endosperm. They have more features

in common with the Saxifragales, particularly free

stylodia, cellular endosperm of some members (always

nuclear in Myrtales), and seeds with copious endosperm

(absent or very scanty in Myrtales). A rather close

relationship between Haloragaceae and Saxifragales,

particularly between Myriophyllum and Penthorum. In

Thorn’s system (2006) the Haloragaceae are including

in the Saxifragales.

6. GUNNERACEAE

Meisner 1842. 1/40–60. Southeastern tropical and

South Africa, Madagascar, Malesia, New Guinea,

Tasmania, New Zealand, Hawaii, America from

Mexico to Tierra del Fuego.

Gunnera.

Gunnera is often included in the Haloragaceae

(Bentham and Hooker 1865; de Candolle 1868; Engler

et Prantl 1893; Schindler 1905; Hutchinson 1973;

Cronquist 1981; Haywood 1993), but from that family

it differs in sieve-element plastids, inﬂ orescence,

pollen morphology, in some embryological features,

including Peperomia-type female gametophyte, and

especially the absence of suspensor hausrotium. They

also related to the Saxifragaceae (Huber 1963;

Hegnauer 1969; Dahlgren 1975, 1980, 1983; Takhtajan

1980, 1987, 1997; Behnke 1986; Doyle and Scogin

1988). According to Fuller and Hickey (2005)

Gunneraceae are closely afﬁ nity to the Saxifragaceae,

especially to the genus Chrysosplenium. However,

they differ from the Saxifragaceae in unicellular hairs,

2-merous ﬂ owers, pollen morphology, female gameto-

phyte, drupaceous fruits, and cordate embryo.

Crane and Hoot (1994) and also D. Soltis et al.

(1999, 2003) placed this family as the sister group to

Myrothamnaceae. But this conclusion strongly con-

tradict morphological differences between this

families.

7. SAXIFRAGACEAE

A.L. de Jussieu 1789 (including Chrysospleniaceae

Berchtold et J. Presl 1820, Pectiantiaceae Raﬁ nesque

1837) 33/525–550. Mainly in northern temper-

ate regions with a few species in southern temperate

regions and in tropical mountains.

Cascadia, Saxifragodes, Micranthes, Peltoboykinia,

Chrysosplenium, Darmera, Astilboides, Rodgersia,

Mukdenia, Oresitrophe, Bergenia, Tellima, Litho-

phragma, Bensoniella, Heuchera, Conimitella, Elmera,

Tolmiea, Mitella, Tiarella, Telesonix, Jepsonia, Bolandra,

Sullivantia, Suksdorﬁ a, Hieronymusia, Boykinia, Astilbe,

Saxifragopsis, Leptarrhena, Tanakaea, Saxifraga (includ-

ing, Saxifragella), Saniculiphyllum.

Close to the Crassulaceae, but somewhat more

advanced. The genus Chrysosplenium (5 species) is

rather isolated within the family by its peculiar inﬂ o-

rescence, 4-merous, apetalous ﬂ owers (Bensel and

Palser 1975), micromorphology of seeds (Kaplan and

Strohschneider 1984), and also by its chemical makeup

(Bohm et al. 1977; Bohm and Collins 1979).

8. GROSSULARIACEAE

A. P. de Candolle 1805 (including Ribesiaceae Marquis

1820). 1/150. Temperate regions of Eurasia, northeast-

ern Africa, North and Central America, Paciﬁ c South

America south to Tierra del Fuego.

Ribes (including Grossularia).

Flower morphology of Ribes is very close to that of

Saxifragaceae (Gelius 1967), and in ﬂ oral structure are

strikingly similar to Tiarella (Bensel and Palser 1975).

Xylem anatomy and embryological characters (Huber

1963) and seed structure (Corner 1976) of Ribes is also

similar to that of Saxifragaceae as well as to that of

Crassulaceae. The afﬁ nity of the Grossulariaceae to

the Saxifragaceae is also supported by chemical (Jay

1970), serological (Grund and Jensen 1981), and rbcL

sequence data (Morgan and Soltis 1993).

9. PTEROSTEMONACEAE

Small 1905. 1/3. High mountains of Mexico.

Pterostemon.

Pterostemonaceae are usually associated with

Escalloniaceae, Hydrangeaceae, and related families,

but ovules in Pterostemon are bitegmic and crassinu-

cellate. Evidence from rbcL sequence suggests that

there is close alliance between Pterostemon and Itea.

However, Pterostemon differs markedly from

Itea in

imbricate petals, ten stamens, 3-colporate pollen grains,

Superorder ROSANAE 307

5-locular inferior ovary and a number of other features.

Most probably the Pterostemonaceae belong to the

Saxifragales but occupy a rather isolated position.

10. ITEACEAE

J. Agardh 1858. 1/27. East, Central, and South Africa,

Himalayas, Assam, eastern Asia and Southeast Asia,

West Malesia, and one species in Atlantic North

America from New Jersey to Florida.

Itea (including Choristylis).

According to Bensel and Palser (1975: 675), the ﬂ ow-

ers of Itea appear very “saxifragaceous” in external mor-

phology, and their ﬂ oral vascular patterns are very similar

to those typical of the Saxifragoideae. The rbcL sequence

data also suggest a close relationship between Itea and

Saxifragaceae s. str. and especially between Itea and

Pterostemon (Morgan 1993). On the other hand, chemi-

cal data suggest that Itea differs from Saxifragaceae s.

str. and more closely resembles other taxa, particularly

Brexia. Plouvier (1965) reported the presence of allitol in

Itea and suggested that this may indicate a relationship

with Brexia, which accumulates dulcitol, an isomer of

allitol. Both allitol and dulcitol are not reported in

Saxifragaceae and related families. Also, Itea accumu-

lates only ﬂ avones but no ﬂ avonols that are characteristic

of the Saxifragales (Bohm et al. 1988). Thus, the Iteaceae

is a rather distinct family that needs further study.

Bibliography

Abe K. 1982. Embryological studies in the family Saxifragaceae

(s.l.): I. Development of the ovule and embryo sac in

Saxifraga fortunei var. partita (Maldno) Nakai. Am. J. Bot.

69: 416–420.

Agababian VS. 1960. On the palynosystematics of the family

Iteaceae. Bull. Armenian Acad. Sci., Biol., 13: 99–102 (in

Russian.)

Agababian VS. 1961. Materials toward the palynological study

of the family Saxifragaceae s.l. Izvestia Armenian Acad.

Sci., Biol., 14(2): 45–61 (in Russian).

Agababian VS. 1963. On palynomorphology of the genus Ribes.

Bull. Armenian Acad. Sci. 16 (4): 93–98.

Agababian VS. 1964. Evolution of pollen in the orders

Cunoniales and Saxifragales in relation to some question of

their systematics and phylogeny. Izvestia of Acad. Sci. of the

Armenian SSR. Biol. Sciences 17: 59–72 (in Russian).

Al-Shammary KI 1991. Systematic studies of the Saxifragaceae,

chieﬂ y from the Southern Hemisphere. Ph.D. dissertation,

University of Leicester.

Al-Shammary KI and RJ Gornall. 1994. Trichome anatomy of

the Saxifragaceae s. 1. from the Southern Hemisphere. Bot.

J. Linn. Soc. 114: 99–131.

Bader F and J Walter. 1961. Das Areal der Gattung Gunnera L.

Bot. Jahrb. Syst. 80: 281–293.

Bates JC. 1933. Comparative anatomical research within the

genus Ribes. Univ. Kansas Sci. Bull. 21: 369–398.

Batham E. 1943. Vascular anatomy of New Zealand species of

Gunnera. Trans. Roy. Soc. N. Zealand 73: 209–216.

Bawa SB. 1969a. Embryological studies on the Halora-gidaceae:

II. Laurembergia brevipes Schindl. and a discussion of sys-

tematic considerations. Proc. Natl. Inst. Sci. India 353:

273–290.

Bawa SB. 1969b. Embryological studies on the Halora-gidaceae:

III. Myriophyllum intermedium DC. Beitr. Biol. Pﬂ . 45:

447–464.

Bawa SB. 1970. Haloragaceae. Bull. Indian Natl. Acad. 41:

226–229.

Behnke H-D. 1986. Contributions to the knowledge of sieve-

element plastids in Gunneraceae and allied families. Plant

Syst. Evol. 151: 215–222.

Bensel CR and BF Palser. 1975a. Floral anatomy in the

Saxifragaceae sensu lato: II. Saxifragoideae and Iteoideae.

Am. J. Bot. 62: 661–675.

Bensel CR and BF Palser. 1975b. Floral anatomy in the

Saxifragaceae sensu lato: IV. Baueroideae and conclusions.

Am. J. Bot. 62: 688–694.

Bergman B, C Johansson, and E Söderbäck. 1992. The Nostoc–

Gunnera symbiosis. New Phytologist 122: 379–400.

Bohm BA and UG Bhat. 1985. Flavonoids of Astilbe and

Rodgersia. Biochem. Syst. Ecol. 13: 437–440.

Bohm BA and FW Collins. 1979. Flavonoids of some species of

Chrysosplenium. Biochem. Syst. Ecol. 7: 195–201.

Bohm BA and R Ornduff. 1978. Chemotaxonomic studies in the

Saxifragaceae s.1.: 9. Flavonoids of Jepsonia. Madroño 25:

39–43.

Bohm BA and CK Wilkins. 1978. The ﬂ avonoids of Heuchera

cylindrica. Canad. J. Bot. 56: 1174–1176.

Bohm BA, FW Collins, and R Bose. 1977. Flavonoids of Bergenia,

Francoa, and Parnassia. Biochem. Syst. Ecol. 13: 221–233.

Bohm BA, LS Donevan, and UC Bhat. 1986. Flavonoids of

some species of Bergenia, Francoa, Parnassia, and

Lepuropetalon. Biochem. Syst. Ecol. 14: 75–77.

Bohm BA, G Chalmers, and UG Bhat. 1988. Flavonoids and the

relationship of Itea to the Saxifragaceae. Phytochemistry 27:

2651–2653.Chapman M. 1933. The ovule and embryo sac of

Saxifraga virginiensis. Am. J. Bot. 20: 151–158.

Collins FW, BA Bohm, and CK Wilkins. 1975. Flavonol glyco-

side gallates from Tellima grandiﬂ ora. Phytochemistry 14:

1099–1102.

Cutler DF and M Gregory. 1998. Anatomy of the Dicotyledons,

Saxifragales (sensu Armen Takhtajan 1983), 2nd ed, vol. 4.

Clarendon Press, Oxford.

Dahlgren KVO. 1930. Zur Embryologie der Saxifra-goideen.

Svensk. Bot. Tidskr. 24: 429–448.

Dandy JE. 1927. The genera of Saxifragaceae. In: J Hutchinson,

Contributions towards a phylogenetic classiﬁ cation of ﬂ ow-

ering plants. Kew Bull. Misc. Inform. 6: 100–118.

Danilova MF. 1996. Crassulaceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol. 5, pp. 25–32. Nauka,

St. Petersburg (in Russian).

308 Subclass VI. ROSIDAE

Dawson MI. 1983. Chromosome numbers of three South

American species of Gunnera (Gunneraceae). New Zealand

J. Bot. 21: 457–459.

Dickison WC, MH Hils, TW Lugansky, and WL Stern. 1994.

Comparative anatomy and systematics of woody Saxifragaceae

Aphanopetalum Endl. Bot. J. Linn. Soc. 114: 167–182.

Doyle MF. 1990. The biosystematics of Gunnera (Gunneraceae)

in the Hawaiian Islands. Ph.D. dissertation, Claremont

Graduate School.

Doyle MF and R Scogin. 1988a. A comparative phyto-chemical

proﬁ le of the Gunneraceae. New Zealand J. Bot. 26: 493–496.

Doyle MF and R Scogin. 1988b. Leaf phenolics of Gunnera

mamcata (Gunneraceae). Aliso 12: 77–80.

Eggli U, ed. 2003. Illustrated handbook of succulent plants.

Crassulaceae. Springer, Berlin.

Elst P van der. 1909. Beiträge zur Kenntnis der Samen-auﬂ age

der Saxifragaceae. Phyl. Dissertation, University of

Utrecht.

Endress PK and A Igersheim. 1999. Gynoecium diversity and

systematics of the basal eudicots. Bot. J. Linn. Soc. 130:

305–393.

Favarger C. 1957. Sur deux criteres nouveaux utilisables dans la tax-

inomie des Saxifragacées. Rev. Cytol. Biol. Veg. 18: 125–137.

Ferguson IK and DA Webb. 1970. Pollen morphology in the

genus Saxifraga and its taxonomic signiﬁ cance. Bot. J. Linn.

Soc. 63: 295–311.

Fernandes Herrera D. 1984. Estudio anatomico y morfologico

de Gunnera insignis (Oerst.) A.DC. Revista Biol. Trop. 32:

197–202 (with English summary).

Fishbein M and DE Soltis. 2004. Further resolution of the rapid

radiation of Saxifragales (Angiosperms, Eudicots) supported

by mixed-model Bayesian analysis. Syst. Bot. 29: 883–891.

Fishbein M, C Hibsch-Jetter, C Soltis, DE Soltis, and L Hufford.

2001. Phylogeny of Saxifragales (Angiosperms, Eudicots):

analysis of a rapid ancient radiation. Syst. Biol. 50: 817–847.

Fuller DQ. 1995. Systematics and leaf architecture of the

Gunneraceae. Am. J. Bot. 82(6): 130–131 (Abstract).

Fuller DQ and LJ Hickey. 2005. Systematics and leaf architec-

ture of the Gunneraceae. Bot. Rev. 71: 295–353.

Gäumann E. 1919. Studien über die Entwicklungsg-eschichte

einiger Saxifragales. Rec. Trav. Bot. Neerl. 16: 285–322.

Ge L-P, A-M Lu, and K-Y Pan. 2002. Floral ontogeny in Itea

yunnanensis (Iteaceae). Act. Bot. Sinica 44: 1261–1267.

Gelius L. 1967. Studien zur Entwicklungsgeschichte an Blüten

der Saxifragales sensu lato mit besonderer Be-rücksichtigung

des Androeceums. Bot. Jahrb. Syst. 87: 253–303.

Gontcharova SB. 1999. Ornamentation of the testa of some east-

ern Asian Sedoideae (Crassulaceae). Bull. Nation. Sci. Mus.,

B (Tokyo) 25(4): 131–141.

Gornall RJ 1986. Trichome anatomy and the taxonomy of

Saxifraga (Saxifragaceae). Nord. J. Bot. 6: 257–275.

Gornall RJ. 1987. Foliar crystals in Saxifraga and segregate gen-

era (Saxifragaceae). Nord. J. Bot. 7: 233–238.

Gornall RJ. 1989. Anatomical evidence and the taxonomic position

of Darmera (Saxifragaceae). Bot. J. Linn. Soc. 100: 173–182.

Gornall RJ and BA Bohm. 1980. The use of ﬂ avonoids in the

taxonomy of Boykinia and allies (Saxifragaceae). Canad. J.

Bot. 58: 1768–1779.

Gornall RJ and BA Bohm. 1985. A monograph of Peltoboykinia,

Boykinia, Bolandra, and Sarsdorﬁ a (Saxifragaceae). Bot. J.

Linn. Soc. 90: 1–71.

Grund C and U Jensen. 1981. Systematic relationships of the

Saxifragales revealed by serological characteristics of seed

proteins. Plant Syst. Evol. 137: 1–22.

Ham RCHJ van and H t’Hart. 1998. Phylogenetic relationships

in the Crassulaceae inferred from chloroplast DNA

restriction-site variation. Am. J. Bot. 85: 123–134.

Hamel J. 1953. Contribution á l’étude cytotaxinomique des

Saxifragacées. Rev. Cytol. Biol. Vég. 14: 113–311.

Harmsen L. 1939. Studies in the embryology and cytology of

Saxifraga. Medd. Gronl. 125: 1–15.

Hart H’t, RDHJ van Ham, JF Stevens, ET Elema, H van der Klis,

and TWJ Gadella. 1999. Biosystematic, molecular and phy-

tochemical evidence for the multiple origin of sympetaly in

Eurasian Sedoideae (Crassulaceae). Biochem. Syst. Ecol.

27: 407–426.

Haskins ML and WJ Hayden. 1987. Anatomy and afﬁ nities of

Penthorum. Am. J. Bot. 74: 164–177.

Hayden WJ and JD Lewandowski. 1997. Gynoecium structure

in Penthorum. Am. J. Bot. 84(6): 201 (Abstract).

Hegnauer R. 1969. Chemical evidence for classiﬁ

cation of some

plant taxa. In: JB Harborne and T. Swain, eds. Perspectives

in phytochemistry, pp. 128–138. Academic Press, London/

New York.

Herr JM. 1954. The development of the ovule and female game-

tophyte in Tiarella cordifolia. Am. J. Bot. 41: 333–338.

Hideux MJ. 1979. Le pollen donnees nouvelles de la microscopie

electronique et de 1’informatique: Structure du sporoderme des

rosidae-saxifragales, etude comparative et dynamique. Paris.

Hideux MJ and IK Ferguson. 1976. The stereostructure of the

exine and its evolutionary signiﬁ cance in Saxifragaceae

sensu lato. In: IK Ferguson and J Muller, eds. The evolution-

ary signiﬁ cance of the exine, pp. 327–377. Linn. Soc.

Symposium, No. 1. London/New York.

Hils MH. 1985. Comparative anatomy and systematics of twelve

woody Australasian genera of the Saxifragaceae. Matthew

Hils, Florida.

Hils MH, WC Dickison, TW Lucansky, and WL Stern. 1988.

Comparative anatomy and systematics of woody

Saxifragaceae; Tetracarpaea. Am. J. Bot. 75: 1687–1700.

Jay M. 1970 (1971). Quelques problmes taxinomiques et phylo-

génétique des Saxifragacées vus á la lumiére de la biochimie

ﬂ avonique. Bull. Mus. Hist. Nat., Paris, sér. 2, 42: 754–775.

Jay M and B Voirin. 1976. Les ﬂ avonoides de deux espéces du

genre Chrysosplenium. Phytochemistry 15: 517–519.

Jensen LCW. 1968. Primary stem vascular patterns in three

subfamilies of the Crassulaceae. Am. J. Bot. 55: 553–563.

Johansson C and B Bergman. 1992. Early events during the

establishment of the Gunnera/Nostoc symbiosis. Planta 188:

403–413.

Johnson LA and DE Soltis. 1994. MatK DNA sequences and

phylogenetic reconstruction in Saxifragaceae sensu stricto.

Syst. Bot. 19: 143–156.

Johnson LA and DE Soltis. 1995. Phylogenetic inference in

Saxifragaceae sensu stricto and Gilia (Polemoniaceae) using

matK sequences. Ann. Missouri Bot. Gard. 82: 149–175.

Juel HO. 1907. Studien über die Entwicklungsgeschichte von

Saxifraga granulata. Nova Acta Reg. Soc. Sci. Uppsal. 4(1):

1–41.

Kapil RN and SB Bawa. 1968. Embryological studies on the

Haloragidaceae: I. Haloragis colensoi Skottsb. Bot. Not.

121: 11–28.

Superorder ROSANAE 309

Kaplan K. 1981. Embryologische pollen- und samen-morpholo-

gische Untersuchungen zur Systematik von Saxifraga

(Saxifragaceae). Bibl. Bot. 134: 1–54.

Kaplan K and M Strohschneider. 1984. Mikromorpho- logische

Untersuchungen an Samenoberﬂ ächen in der Gattung

Chrysosplenium (Saxifragaceae). Bot. Jahrb. Syst. 104:

469–482.

Kellermann WA. 1881. Die Entwicklungsgeschichte der Blüten

von Gunnera chilensis. Zürich.

Klopfer K. 1968, 1970. Beiträge zur ﬂ oralen Morpho-genese

und Histogenese der Saxifragaceae: 1. Die In-ﬂ oreszenz-

Entwicklung von Tellima grandiﬂ ora. 2. Die

Blütenentwicklung einiger Saxifragen-Arten. Flora 157B:

461–476, 1968; 158: 1–21, 1968; 159: 347–365, 1970.

Klopfer K. 1973. Florale Morphogenese und Taxonomie der

Saxifragaceae sensu lato. Feddes Repert. 84: 475–516.

Knapp U. 1994. Skulptur der Samenschale und Gliederung der

Crassulaceae. Bot. Jahrb. Syst. 116: 157–187.

Komar GA. 1967. On the nature of inferior ovary of the

Grossulariaceae. Bot. Zhurn. 52: 1611–1629 (in Russian).

Komar GA. 1970. Development of the ﬂ ower and inﬂ orescence

in some representatives of the family Grossulariaceae. Bot.

Zhurn. 55: 954–971 (in Russian).

Komar GA. 1996. Grossulariaceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol. 5, pp. 60–67. Nauka, St.

Petersburg (in Russian).

Krach JE. 1976. Samenanatomie der Rosiﬂ oren: I. Die Samen

der Saxifragaceae. Bot. Jahrb. Syst. 97: 1–60.

Krach JE. 1977. Seed characters in and afﬁ nities among the

Saxifragineae. Plant Syst. Evol., Suppl., 1: 141–153.

Kubitzki K. 2007a. Aphanopetalaceae. In: K Kubitzki, ed. The

families and genera of vascular plants, vol. 9, pp. 29–30.

Sprinegr. Berlin/Heidelberg/New York.

Kubitzki K. 2007b. Haloragaceae. In: K Kubitzki, ed. The fami-

lies and genera of vascular plants, vol. 9, pp. 184–190.

Springer, Berlin/Heidelberg/New York.

Kubitzki K. 2007c. Iteaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 9, pp. 202–204. Springer,

Berlin/Heidelberg/New York.

Kubitzki K. 2007d. Pterostemonaceae. In: K Kubitzki, ed. The

families and genera of vascular plants, vol. 9, pp. 405–406.

Springer, Berlin/Heidelberg/New York.

Kubitzki K. 2007e. Tetracarpaceae. In: K Kubitzki, ed. The

families and genera of vascular plants, vol. 9, pp. 456–457.

Springer, Berlin/Heidelberg/New York.

Kulbaeva B Zh. 1992a. Seed anatomy in representatives of

Saxifragaceae family. Bot. Zhurn. 77(3): 36–49 (in Russian

with English summary).

Kulbaeva B Zh. 1992b. The seed surface structure in representa-

tives of the family Saxifragaceae. Bot. Zhurn. 77(4): 61–68

(in Russian with English summary).

Kulbaeva B Zh. 1992c. Seed surface in representatives of the

family Saxifragaceae: Typology and signiﬁ cance for system-

atics and phylogeny. Bot. Zhurn. 77(8): 98–105 (in Russian

with English summary).

Kulbaeva B Zh. 1996. Saxifragaceae. In: A. Takhtajan,

ed. Comparative seed anatomy, vol. 5, pp. 34–60. Nauka,

St. Petersburg (in Russian).

Lobova TA. 2000. Pterostemonaceae. In: A Takhtajan,

ed. Comparative seed anatomy, vol. 6, pp. 296. Nauka,

St. Petersburg (in Russian).

Lowrey TK and ER Robinson. 1988. The interaction of gynomo-

noecy, dichogamy, and wind-pollination in Gunnera

perpensa L. (Gunneraceae) in South Africa. Monograph

Syst. Bot. 25: 237–246.

MacGaughey V. 1917. Gunnera petaloidea Gaud., a remarkable

plant of the Hawaiian Islands. Am. J. Bot. 4: 33–39.

Mandrik VY and LV Golyshkin. 1973. Embryological studies of

some species of the family Crassulaceae. Bot. Zhurn. 58:

263–272 (in Russian).

Mauritzon J. 1933. Studien über die Embryologie der Familien

Crassulaceae und Saxifragaceae. Ph.D. dissertation,

University of Lund.

Mayuzumi S and H Ohba. 2004. The phylogenetic position of

Eastern Asian Sedoideae (Crassulaceae) inferred from

chloroplast and nuclear DNA sequences. Syst. Bot. 29:

587–598.

Messinger W, K Hummer and A Liston. 1999. Ribes

(Grossulariaceae) phylogeny as indicated by restriction site

polymorphisms of PCR-ampliﬁ ed chloroplast DNA. Plant

Syst. Evol. 217: 185–195.

Miehe H. 1924. Entwicklungsgeschichtliche Untersu-chungen

der Algensymbiose bei Gunnera macrophylla Bl. Flora 17:

1–15.

Modilewski J. 1908. Zur Embryobildung von Gunnera chilensis.

Ber. Deutsch. Bot. Ges. 26a: 550–556.

Moody ML and DH Les. 2001. Phylogenetic relationships in

Haloragaceae emphasizing the aquatic genus Myriophyllum.

In Botany 2001: Plants and People, p. 128. Albuquerque

(Abstracts).

Moreau F. 1971. Apport des caractéres stomatiques á la taxino-

mie et á la phylogénie des Saxifragées. Bull. Soc. Bot. France

118: 381–427.

Morf E. 1950. Vergleichend-morphologische Untersu-chungen

am Gynoeceum der Saxifragaceen. Ber. Schweiz. Bot. Ges.

60: 516–590.

Morgan DR and DE Soltis. 1993. Phylogenetic relationships

among members of Saxifragaceae sensulato based on

rbcL sequence data. Ann. Missouri Bot. Gard. 80:

631–660.

Mort ME, DE Soltis, PS Soltis, J Franciso-Ortega, and A Santos-

Guerra. 2001. Phylogenetic relationships and evolution of

Crassulaceae inferred from matK sequence data. Am. J. Bot.

88: 76–91.

Mort ME, DE Soltis, PS Soltis, J Francisco Ortega, and A Santos

Guerra. 2002. Phylogenetics and evolution of the

Macaronesian clade of Crassulaceae inferred from nuclear

and chloroplast sequence data. Syst. Bot. 27: 271–288.

Nagaraj M and BHM Nijalingappa. 1967. Embryological stud-

ies in Myriophyllum intermedium DC. Proc. Indian Acad.

Sci. 65B: 210–220.

Nagaraj M and BHM Nijalingappa. 1974. Embryological stud-

ies in Laurembergia hirsuta. Bot. Gaz. 135: 19–28.

Nemirovich-Danchenko EN. 1994a. Seed structure in Penthorum

sedoides and P. chinense (Penthoraceae). Bot. Zhurn. 79 (6):

56–64 (in Russian with English summary).

Nemirovich-Danchenko EN. 1994b. Morphology and anatomy

of the seeds of Iteaceae. Bot. Zhurn. 79(9): 83–87 (in Russian

with English summary).

Nemirovich-Danchenko EN. 1996a. Penthoraceae, Crassulaceae.

In: A Takhtajan, ed. Comparative seed anatomy, vol. 5,

pp. 22–32. Nauka, St. Petersburg (in Russian).

310 Subclass VI. ROSIDAE

Nemirovich-Danchenko EN. 1996b. Iteaceae. In: A Takhtajan,

ed. Comparative seed anatomy, vol. 5, pp. 67–72. Nauka, St.

Petersburg (in Russian).

Nemirovich-Danchenko EN. 2000. Tetracarpaceae. In: A

Takhtajan, ed. Comparative seed anatomy, vol. 6, pp. 232–

233. Nauka, St. Petersburg (in Russian).

Nicholls KW, BA Bohm, and EF Wells. 1986. The ﬂ avonoids of

Mitella, Bensoniella, and Conimitella (Saxifragaceae).

Canad. J. Bot. 64: 525–530.

Nikiticheva ZI. 1985. Haloragaceae. In: MS Yakovlev, ed.

Comparative embryology of ﬂ owering plants: Brunelliaceae-

Tremandraceae, pp. 116–120. Nauka, Leningrad (in

Russian).

Orchard AE. 1975. Taxonomic revisions in the family

Haloragaceae: I. The genera Haloragis, Haloragodendron,

Glischrocaryon, Meziella, and Gonocarpus. Bull. Auckland

Inst. and Mus. 10: 1–299.

Orchard AE. 1990. Gunneraceae. Flora of Australia 18: 85–87.

Canberra.

Pacheco P, DJ Crawford, TF Stuessy, and MO Silva. 1993.

Flavonoid chemistry and evolution of Gunnera (Gunneraceae)

of the Juan Fernandez Islands, Chile. Paciﬁ c Sci. 45:

389–399.

Pastre A and A Pons. 1973. Quelques aspects de la systematique

des Saxifragacees a la lumiere des donnees de la palynolo-

gie. Pollen Spores 15: 117–133.

Planchon JE. 1854. Afﬁ nités et synonymic de quelques genres

nouveaux. Ann. Sci. Nat. Bot. ser. 4, 2: 256–266.

Plouvier V. 1965. Etudes chimiotaxinomiques sur les

Saxifragacees. Bull. Soc. Bot. France, Mem., 1965:

150–161.

Praglowski J. 1970. The pollen morphology of the Haloragaceae

with reference to taxonomy. Grana 10: 159–239.

Ramamonjiarisoa BA. 1980. Comparative anatomy and system-

atics of African and Malagasy wood Saxifragaceae s.l. Ph.D.

dissertation. University of Massachusetts. Amherst, MA.

Reinke J. 1873. Untersuchungen über die Morphologie der

Vegetationsorgane von Gunnera. In: Morphologische

Abhandlungen, S. 47–122. Leipzig.

Rocen T. 1928. Beiträge zur Embryologie der Crassulaceen.

Svensk Bot. Tidskr. 22: 368–376.

Roeland CHJ Van Ham and H ‘t Hart. 1998. Phylogenetic rela-

tionships in the Crassulaceae inferred from chloroplast DNA

restriction-site variation. Am. J. Bot. 85: 123–134.

Rylova TB. 1989. Morphological features of pollen in some

fossil and extant species of Itea (Iteaceae). Bot. Zhurn. 74:

694–699 (in Russian with English summary).

Savile DBO. 1953. Splash-cup dispersal mechanism in

Chrysosplenium and Mitella. Science 117: 250–251.

Savile DBO. 1975. Evolution and biogeography of Saxifragaceae

with guidance from their rust parasites. Ann. Missouri Bot.

Gard. 62: 354–361.

Saxena NP. 1964. Studies in the family Saxifragaceae: I.

A contribution to the morphology and embryology of

Saxifraga diversifolia Wall. Proc. Indian Acad. Sci. B 60:

38–51.

Scherwin PA and RL Wilbur. 1971. The contributions of ﬂ oral

anatomy to the generic placement of Diamorpha smallii and

Sedum pusillum. J. Elisha Mitchell Sci. Soc. 87: 103–114.

Schnegg H. 1902. Beiträge zur Kenntnis der Gattung Gunnera.

Flora 90: 161–208.

Senters AE and DE Soltis. 2003. Phylogenetic relationships in

Ribes (Grossulariaceae) inferred from ITS sequence data.

Taxon 52: 51–66.

Silvester WB and DR Smith. 1969. Nitrogen ﬁ xation by

Gunnera-Nostoc symbiosis. Nature 224: 1231.

Skottsberg C. 1929. Bemerkungen über die Morphologic von

Gunnera macrophylla Bl. Meddelanden fran Göteborgs bot.

Träd. V: 115–126.

Soltis DE. 1980. Karyotypic relationships among species of

Boykinia, Heuchera, Mitella, Sullivantia, Tiarella, and

Tolmiea (Saxifragaceae). Syst. Bot. 5: 17–29.

Soltis DE. 1984a. Karyotypic relationships among Elmera,

Heuchera, and Tellima

(Saxifragaceae). Syst. Bot. 9: 6–11.

Soltis DE. 1984b. Karyotypes and relationships of species of

Jepsonia (Saxifragaceae). Syst. Bot. 9: 137–141.

Soltis DE. 1984c. Karyotypes of Leptarrhena and Tana-kaea

(Saxifragaceae). Canad. J. Bot. 62: 671–673.

Soltis DE. 1986. Karyotypic relationships among Astil-boides,

Bergenia, Darmera, and Mukdenia and their implications for

subtribal boundaries in Saxifrageae (Saxifragaceae). Canad.

J. Bot. 64: 586–588.

Soltis DE. 1988. Karyotypes of Bensoniella, Conimitella,

Lithophragma, and Mitella, and relationships in

Saxifragaceae. Syst. Bot. 13: 64–72.

Soltis DE. 2007. Saxifragaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 9, pp. 418–435. Springer,

Berlin/Heidelberg/New York.

Soltis DE and BA Bohm. 1982. Flavonoids of Penthorum

sedoides. Biochem. Syst. Ecol. 10: 221–224.

Soltis DE and L Hufford. 2002. Ovary position diversity in

Saxifragaceae: clarifying the homology of epigny. Int. J.

Plant Sci. 163(2): 277–293.

Soltis DE and PS Soltis. 1997. Phylogenetic relationships in

Saxifragaceae sensu lato: a comparison of topologies based

on 18S rDNA and rbcL sequences. Am. J. Bot. 84:

504–522.

Soltis DE, PS Soltis, and KD Bothel. 1990. Chloroplast DNA

evidence for the origins of the monotypic Bensoniella and

Conimitella (Saxifragaceae). Syst. Bot. 15: 349–362.

Soltis DE, PS Soltis, MT Clegg, and M Durbin. 1990. rbcL

sequence divergence and phylogenetic relationships in

Saxifragaceae sensu lato. Proc. Nad. Acad. Sci. USA 87:

4640–4644.

Soltis DE, A Crable, D Morgan, PS Soltis, and R Kuzoff. 1993.

Molecular systematics of Saxifragaceae sensu stricto. Am. J.

Bot. 80: 1056–1081.

Soltis DE, RK Kuzoff, E Conti, R Gornall, and K Ferguson. 1996.

matk and rbcL gene sequence data indicate that Saxifraga

(Saxifragaceae) is polyphyletic. Am. J. Bot. 83: 371–382.

Soltis DE, RK Kuzoff, ME Mort, M Zanis, M Fishbein,

L Hufforl, J Koontz, and MK Arroyo. 2001a. Elucidating

deep-level phylogenetic relationships in Saxifragaceae using

sequences for six chloroplastic and nuclear DNA regions.

Ann. Missouri Bot. Gard. 88: 669–693.

Soltis DE, M Tago-Nakazawa, Q-Y Xiang, S Kawano, J Murata,

M Wakabayashi, and C Hibsch-Jetter. 2001b. Phylogenetic

relationships and evolution in Chrysosplenium

(Saxifragaceae) based on matK sequence data. Am. J. Bot.

88: 883–893.

Soltis DE, AE Senters, MJ Zanis, S Kim, JD Thompson,

PS Soltis, LP Ronse Decraene, PK Endress, and JS Farris.

Superorder ROSANAE 311

2003. Gunnerales are sister to other core eudicots: implica-

tions for the evolution of pentamery. Am. J. Bot. 90:

461–470.

Spongberg SA. 1978. The genera of Crassulaceae in the south-

eastern United States. J. Arnold Arbor. 59: 197–248.

Stern WL. 1974. Saxifragales. In: Encyclopaedia Britannica,

15th ed., vol. 16, pp. 291–302. Chicago.

Stern WL, EM Sweitzer, and RE Phipps. 1970. Comparative

anatomy and systematics of woody Saxifragaceae: Ribes. In:

NKB Robson, DF Cutler, and M Gregory, eds. New research

in plant anatomy, pp. 215–237. Bot. J. Linn. Soc. 63 (Suppl.).

London/New York.

Steyn EMA, PJ Robbertse, and AE van Wyk. 1987. Floral devel-

opment in Greyia ﬂ anaganii with notes on inﬂ orescence ini-

tiation and sympodial branching. S. Afr. Tydskr. Plantk. 53:

194–201.

Stolt KAH. 1928. Die Embryologie von Myriophyllum alterni-

ﬂ orum DC. Svensk. Bot. Tidskr. 22: 305–319.

Subramanyam K. 1962. Embryology in relation to systematic

botany with particular reference to the Crassulaceae. In:

Plant embryology: A symposium, pp. 94–112. New Delhi.

Taneyama M and S Yoshida. 1978. Studies on C-glycosides in

higher plants: I. Occurrence of bergenin in Saxifragaceae.

Bot. Mag. (Tokyo) 91: 109–112.

Thiede J. 2007. Penthoraceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 9, pp. 292–296. Springer,

Berlin/Heidelberg/New York.

Thiede J and U Eggli. 2007. Crassulaceae. In: K Kubitzki, ed.

The families and genera of vascular plants, vol. 9, pp. 83–118.

Springer, Berlin/Heidelberg/New York.

Tillson AH. 1940. The ﬂ oral anatomy of the Kalan-choideae.

Am. J. Bot. 27: 595–600.

Towata EM. 1985a. Mucilage glands and cyanobacterial coloni-

zation in Gunnera kaalensis (Haloragaceae). Bot. Gaz. 146:

56–62.

Towata EM. 1985b. Morphometric and cytochemical ultrastruc-

tural analyses of the Gunnera kaalensis/Nostoc symbiosis.

Bot. Gaz. 146: 293–301.

Uhl CH. 1963. Chromosomes and phylogeny of the Crassulaceae.

Cact. Succ. J. (Los Angeles) 35: 80–84.

Van der Meijden R, and N Caspers. 1971. Haloragaceae. In:

CGGJ van Steenis, ed. Flora Malesiana, ser.1, 7: 239–263.

Noordhoff, Groningen.

Wakabayashi M. 1970. On the afﬁ nity in Saxifragaceae s. lato

with special reference to the pollen morphology. Acta

Phytotax. Geobot. 24: 128–145 (in Japanese, with English

summary).

Wanntorp L, J Praglowski, and E Grafström. 2004. New insights

into the pollen morphology of the genus Gunnera

(Gunneraceae). Grana 43: 15–21.

Wanntorp L and LP Ronse Decraene. 2005. The Gunnera ﬂ ower:

Key to eudicot diversiﬁ cation or response to pollination

mode? Int. J. Plant Sci. 166: 945–953.

Wanntorp L and H-E Wanntorp. 2003. The biogeography of

Gunnera L.: Vicariance and dispersal. J. Biogeogr. 30:

979–987.

Wanntorp L, H-E Wanntorp, B Oxelman, and M Källersjö.

2001. Phylogeny of Gunnera. Plant Syst. Evol. 226:

85–107.

Wanntorp L, H-E Wanntorp, and M Källersjä. 2002. Phylogenetic

relationships of Gunnera based on nuclear ribosomal DNA

ITS region, rbcL and rps16 intron sequences. Syst. Bot. 27:

512–521.

Wanntorp L, H-E Wanntorp, and R Rutishauser. 2003. On the

homology of the scales in Gunnera (Gunneraceae). Bot. J.

Linn. Soc. 142: 301–308.

Warming E. 1909. Saxifragaceae: 1. Morphology and biology.

Medd. Gronl. 36: 169–236.

Watari S. 1930. Anatomical studies on the leaves of some saxi-

fragaceous plants, with special reference to the vascular

system. J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 5: 195–316.

Webb DA and RJ Gornall. 1989. Saxifrages of Europe.

Christopher Helm, London.

Webb CJ, WR Sykes, and PJ Garnock-Jones. 1988. Gunneraceae.

Flora of New Zealand IV: 748–750.

Weigend M. 2007. Grossulariaceae. In: K Kubitzki, ed. The

families and genera of vascular plants, vol. 9, pp. 168–176.

Springer, Berlin/Heidelberg/ New York.

Weigend M, T Motley, and O Mohr. 2002. Phylogeny and clas-

siﬁ cation in the genus Ribes (Grossulariaceae) based on

5S-NTS sequences and morphological and anatomical data.

Bot. Jahrb. Syst. 124: 163–182.

Wiggins IL. 1959. Development of the ovule and megagameto-

phyte in Saxifraga hiericifolia. Am. J. Bot. 46: 692–697.

Wilkinson HP. 1994. Leaf and twig anatomy of the

Pterostemonaceae (Engl.) Small: Ecological and systematic

features. Bot. J. Linn. Soc. 115: 115–131.

Wilkinson HP. 1998. Gunneraceae. In: DF Cutler, M Gregory,

eds. Anatomy of the Dicotyledons, 2nd ed. Saxifragales, 4:

260–272. Oxford.

Wilkinson HP. 2000. A revision of the anatomy of Gunneraceae.

Bot. J. Linn. Soc. 134: 233–266.

Wilkinson HP and L Wanntorp. 2007. Gunneraceae. In:

K Kubitzki, ed. The families and genera of vascular plants,

vol. 9, pp. 177–183. Springer, Berlin/Heidelberg/New York.

Wu C-Y and T-C Ku. 1992. A new tribe with a new monotypic

genus of Saxifragaceae (s.1.) from China. Acta Phytotax.

Sinica 30: 193–196.

Yen TK. 1936. Floral development and vascular anatomy of the

fruit of Ribes aureum. Bot. Gaz. 98: 105–120.

Order 82. PODOSTEMALES

Aquatic, annual or perennial herbs living in rushing

water and growing only on rocks and stones in rivers

or cataracts, mostly submerged or with some of the

parts ﬂ oating. Plants with more or less conﬂ uent stems

and leaves, usually more or less thalloid and often

resembling bryophytes, lichens, algae, or unlike any

other plants in appearance. Primary root absent. The

thalloid body usually attached to the substrate by

numerous hairs or by specialized rootlike branches

(haptera). Shoots nearly always arising as endogenous

buds from roots; stems reduced or elongate, simple or

branched, sometimes dimorphic, occasionally only

present when ﬂ owering. Xylem very much reduced,

312 Subclass VI. ROSIDAE

usually represented only by a few tracheids with spiral

or annular thickenings or completely wanting. Sieve

plates recorded only in a few genera. Sieve-element

plastids of S-type and large. Leaves, when discernible,

alternate, entire or more or less dissected, without axil-

lary buds; stipules present or absent. Flowers very

small, solitary, or in cymose inﬂ orescences, bisexual,

actinomorphic or zygomorphic, apetalous, anemophil-

ous, entomophilous, or cleistogamous, subtended or

enclosed by two bracteoles (spathe-like and enclosing

up to 20 ﬂ owers in Podostomoideae). Perianth of 2 or

3(-5) free or more or less connate petaloid sepals or

small, annular scale or completely wanting; tepals (in

Podostemoideae) 2–20, small, or large and 4–6, imbri-

cate. Stamens 1–40 in one to many cycles or in more

reduced ﬂ owers there is only a solitary stamen; ﬁ la-

ments free or more often basally connate; anthers

2-locular, extrorse, opening longitudinally. Tapetum

secretory. Microsporogenesis simultaneous or

(Podostemum) successive. Pollen grains 2-celled, in

monads or (Podostemoideae) in dyads or rarely in tet-

rads, from 3-colpate and 3-colporate to pantoporate,

sometimes inaperturate. Gynoecium of (1)2(3) united

carpels with as many locules and free or less often

more or less basally connate stylodia. Ovary superior,

with numerous or rarely (Farmeria) 2–4 ovules on

thickened axile placentas. Ovules anatropous, biteg-

mic, tenuinucellate. As a result of the disintegration of

the cells of the nucellus lying immediately below the

female gametophyte, the so-called nucellar plasmo-

dium or “pseudoembryo” develops, which serves as

nutrition for the developing embryo. Female gameto-

phyte monosporitc or bisporic, 4-celled or sometimes

5- or 6-celled; it contains an egg, one or two synergids,

and one or two chalazal cells; polar nuclei absent, and

therefore there is no triple fusion. Fruits septicidal cap-

sules (except Farmeria metzgerioides, it has indehis-

cent fruit), with usually numerous seeds. Seeds minute,

exotesta thick-walled, usually mucilaginous, endoteg-

men ligniﬁ ed; endosperm wanting; embryo straight,

large, with two massive cotyledons and large suspen-

sor; the large basal cell of the suspensor gives rise to a

prominent and usually branched haustorium, which

draws nourishment from the external tissues. Present

xanthones, mangostin (in Podostemum ceratophyl-

lum); n = 8, 10, 12, 14, 15, 17, 20.

In spite of its high specialization and very far-

reaching reduction processes, the order Podostemales

reveals clearly expressed links with the Saxifragales

(Warming 1891 and many others), particularly with

the Crassulaceae (Magnus 1913; Mauritzon 1933–

1939; Maheshwari 1945; van Royen 1951;

Subramanyam 1962; Kapil 1970, Les et al. 1997, Ueda

et al. 1997). The presence in both of these families of

such a unique structure as well-developed suspensor-

haustorium is one of many common features in favor

of their afﬁ nity.

1. PODOSTEMACEAE

Kunth 1816 (including Marathraceae Dumortier

1829, Philocrenaceae Bongard 1834, Tristichaceae

J. C. Willis 1915). 49/280–300. Pantropical, especially

in Asia and America, extending into temperate eastern

North America and temperate East Asia.

1.1 WEDDELLINOIDEAE

Young ﬂ owers not enclosed in spathella or cupule.

Stomata anomocytic. Sepals (4)5, imbricate; anthers

5–25; pollen grains in monads, 3-colporate. Capsula

not ribbed, opening by two equal valves. Tegmen thick

walled. – Weddellina.

1.2 TRISTICHOIDEAE

Young ﬂ owers subtended or enclosed by two bracteoles,

which are not spathelike. Inﬂ orescences terminal.

Sepals three, imbricate; anthers (1-)3; pollen grains in

monads, pantoporate. Gynoecium of three carpals;

stigmas three. Capsule strongly ribbed, opening by

three equal valves. – Tristicha (including Malacco-

tristicha), Indotristicha, Dalzellia.

1.3 PODOSTEMOIDEAE

Young ﬂ owers enclosed in spathelike bracteoles (spa-

tella). Inﬂ orescences axillary to leaves or bracts. Sepals

2–20, reduced to small scales or wanting. Anthers

2–40, often free. Pollen grains in monads or dyads,

rarely (Diamantina) in tetrads. Gynoecium of 3–7 car-

pals. Capsule ribbed, opening by two equal or unequal

valves, or rarely indehiscent. – podostemeae: Apinagia,

Castelnavia, Ceratolacis, Cipoia, Crenias, Devillea,

Marathrum, Rhyncholacis, Jenmaniella, Lophogyne,

Hydrobryum, Leiothylax, Diamantina, Cladopus,

Zeylanidium (including Hydrobryopsis), Podostemum,

Farmeria, Macarenia, Oserya, Vanroyenella, Wetts-

teiniola, Angolaea, Dicraeanthus, Endocaulos, Butumia,

Djinga, Ledermanniella, Letestuella, Macropodiella,