Takhtajan Armen. Flowering Plants

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Superorder LILIANAE 675

integument; embryo small to large, straight to curved;

endosperm storing aleurone, starch, and lipids. Usually

produce chelidonic acid steroidal saponins. Alkaloids

lacking.

Probably derived from some Colchicaceae –

Uvularioideae-like ancestor.

Key to Families

1 Leaves well developed.

2 Ovules 2–4(-6) per carpel.

3 Plants not woody. Rhizomatous or tuberous

herbs with ﬁ brous or sometimes swollen and

ﬂ eshy roots. Calcium oxalate raphides present.

Vessels mainly in roots, with scalariform or

reticulately perforated and scalariform perfo-

rations, and rarely in stems with scaraliform

perforations. Leaves basal or cauline, alternate,

opposite or verticillate, sessile or sometimes

petiolate, simple, entire, linear, lanceolate,

oblong, or ovate, sometimes cordate, sheathing

or not sheathing. Stomata anomocytic. Flowers

in terminal or axillary racemes, spikes, pani-

cles, or ﬂ owers solitary, axillary, bisexual,

actinomorphic, mostly 3-merous. Perianth seg-

ments six or four, rarely six or nine, free or

more often connate into an urceolate or cam-

panulate tube. Stamens six or rarely 4 or 8, 10,

12, in two cycles, inserted at the base of peri-

anth segments or in the tube; ﬁ laments free or

rarely (Ophiopogon) basally connate into a

ring, sometimes nearly absent; anthers bas-

iﬁ xed or dorsiﬁ xed, introrse. Pollen grains

almost always 2-celled, 1-colpate or rarely (as

in Aspidistra, Tupistra) inaperturate, tectate-

columellate. Gynoecium of (2)3(4) carpels;

style columnar or ﬁ liform, with lobed or capi-

tate stigma, rarely stigma sessile; ovary supe-

rior or rarely (Peliosanthes) semi-inferior,

3-locular or rarely 4- or 2-locular, with two to

many ovules per locule. Septal nectaries often

present. Ovules 2–4 anatropous, campylotro-

pous, or nearly orthotropous, generally crassi-

nucellate, usually with parietal cell. Female

gametophyte of Allium-type or less often of

Polygonum-type, or Drusa-type, or Scilla-type.

Endosperm nuclear or rarely helobial. Fruits

berries, ﬂ eshy dehiscent capsule (Gonioscypha),

drupe (Tricalustra) or dry, leathery, 1–3-seeded

structures, rupturing in early development and

exposing seeds. Seeds globose to ovoid, of

various colour; the testal layer is almost com-

pletely obliterated during maturation, and a

phytomelan crust is never formed; the tegmic

layer collapses, but the cellular layer is retained.

Endosperm consists of cells with thick pitted

rugose walls and store aleurones and lipids;

embryo usually more than half as long as the

endosperm. Flavonols present (kaempferol and

quercetin) or absent; steroidal sapinins in abun-

dance); n = 9–20. . . . . . 1. convallariaceae.

3 Woody, generally large, arborescent plants

with a stout, simple or sparingly branched

trunk up to a few meters tall, with terminal leaf

rosettes. Secondary thickening anomalous.

Vessels present in roots (with simple perfora-

tions) and in leaves (with scalariform perfora-

tions). Leaves alternate, linear, tough and hard

or leathery, sessile, simple, entire, serrulate, or

(Dasylirion) margins usually armed with

curved spines. Stomata anomocytic, sunken,

with oily contents. Inﬂ orescences panicles,

often large and many-ﬂ owered. Flowers polyg-

amo-dioecious or dioecious, actinomorphic,

articulated on their pedicels, bracteate. Perianth

segments six, all similar, free, scarious. Stamens

six in two whorls, free, inserted at base of peri-

anth segments, vestigial in female ﬂ owers;

anthers dorsiﬁ xed, introrse. Pollen grains 1-col-

pate, ﬁ nely reticulate. Ovary 1–3-locular, with

two anatropous ovules per carpel, or 3–6 in a

single cavity in Dasylirion, reduced in male

ﬂ owers; ovules tenuinucellate. Septal nectaries

present. Style short, with 3-lobed stigma.

Fruits dry, 3-winged samaras, or 3-lobed ﬂ at-

ted capsules, or 3-ribbed nuts. Seeds without

phytomelan, with cylindrical straight embryo.

Flavonols (quercetin) present n = 18 and more

often 19, chromosomes without marked dif-

ference in size. . . . . . . . . . . . 3. nolinaceae.

2 Ovule one per carpel. Usually shrubby or treelike

plants (Dracaena draco – the “Dragon Blood

Tree” up to 15 m or more), sometimes scandent,

or sometimes rhizomatous geophytes

(Sansevieria); secondary thickening anomalous.

Stems marked with transverse leaf scars or plants

stemless or nearly stemless. Vessels in roots

(mainly with simple perforations) and in leaves

(with scalariform perforations). Leaves basal or

676 Subclass II. LILIIDAE

more often crowded on the tops of branches,

sheathing at base, small to very large, linear-

oblong to ovate or obovate, strap-shaped or ensi-

form, sometimes more or less distinctly petiolate,

thin and ﬂ exible to thick and rigid, sometimes

conspicuously succulent. Stomata anomocytic.

Flowers numerous, in compound terminal or

axillary, paniculate, spicate, or capitate racemes,

bisexual, actinomorphic, bracteate and with one

bracteole, articulated on the pedicel. Perianth

segments six in two cycles, equal, connate into a

short or long tube. Stamens six in two cycles,

inserted on the tube; anthers dorsiﬁ xed, versatile,

introrse. Pollen grains 1-colpate. Ovary supe-

rior, with one anatropous ovule per locule; sep-

tal nectaries present; style one, apical. stigma

3-lobed to capitate. Fruits 1–3-seeded, red or

orange berries, but sometimes capsular-indehis-

cent, hard and woody. Seed coat has a three to

four-layered outer integument, the outer epider-

mis of which is thick walled and lacks phy-

tomelan, while the cells of the inner integument

have collapsed and form a reddish brown mem-

brane (Dahlgren et al. 1985); embryo small,

endosperm oily, containing aleurone and lipids.

Flavonols (kaempferol) present (Dracaena);

n = 19–21. . . . . . . . . . . . . . 2. dracaenaceae.

1 Leaves much reduced.

4 Seeds with phytomelan in testa and collapsed

tegmen. Subshrubby or herbaceous perennials with

erect or scandent stem growing from sympodial

rhizome or rarely a tuber. Phylloclades perennial

or deciduous, fasciculate or solitary, ﬂ at, and leaf-

like or more often slender and needlelike to ﬁ li-

form, apex mucronate, rarely (Asparagus

densiﬂ orus) bearing minute, scaly leaves, in some

species (e.g., Asparagus turkestanicus) phyllo-

clades lacking. Vessels in roots with scalariform or

simple perforations, in stems only with scalariform

perforations. Leaves more or less scalelike, scari-

ous, with a spiny or soft basal spur. Stomata ano-

mocytic. Flowers small and inconspicuous,

solitary or in umbel-like or racemelike

inﬂ orescences, bisexual or unisexual; pedicels

articulate, usually bracteolate at the base. Perianth

segments six, free and spreading or basally con-

nate and then forming a cup or tube. Stamens six,

free from each other, inserted at the base of peri-

anth segments; ﬁ laments ﬁ liform or ﬂ attened,

rarely spurred at base; anthers more or less

basiﬁ xed, sagittate, tetrasporangiate, dehiscing

longitudinally, introrse. Microsporogenesis suc-

cessive. Pollen grains 1-colpate. Gynoecium of

three united carpels; ovary superior, sometimes

slightly stipitate, 3-locular, style short, with

3-lobed or capitate stigma. Septal nectaries pres-

ent. Ovules 2–12 per locule, hemitropous or almost

orthotropous, crassinucellate. Female gametophyte

of Polygonum-type. Fruits often red, smooth, or

wrinkled berry, rarely nut, or (Hemiphylacus) loc-

ulicidal capsules. Seeds black; endosperm with

distinctly pitted cell walls, contains aleurone, lip-

ids and hemicellulose; embryo almost reach in the

length of endosperm, slightly curved. n = 10, 56

(Hemiphylacus).. . . . . . . . . . . . 5. asparagaceae.

4 Seeds without phytomelan.

5 Shrubs or subshrubs, often thicket forming,

or lianas; secondary thickening absent.

Phylloclades lanceolate to ovate, acute or pun-

gent-pointed. In Ruscus vessels only in roots

and with scalariform and simple perforations

in other genera also in stems and with scalari-

form perforations. Leaves reduced to small,

scarious scales; mesophyll sometimes contain-

ing mucilage cells and calcium oxalate crys-

tals. Stomata anomocytic. Phylloclades bear

on upper or lower surface racemelike inﬂ ores-

cence (Ruscus), umbel-like inﬂ orescence on

their margins (Semele), or short terminal

racemes are free from the phylloclades

(Danae). Flowers small, inconspicuous, bisex-

ual or (Ruscus) unisexual (monoecious or dioe-

cious), articulated with their pedicels. Perianth

segments six, free or partly connate, in the lat-

ter case with a ﬂ eshy corona. Stamens six or

(Ruscus) three; ﬁ laments connate into a col-

umn; anthers extrorse. Pollen grains 1-colpate

or (Semele) inaperturate. Stigma sessile or sub-

sessile, simple or more or less lobed. Ovary

superior, 3-locular with two ovules per locule

or (Ruscus) 1-locular with 1–4 ovules, without

septal nectaries. Ovules hemitropous or ortho-

tropous, tenuinucellate and have a large funi-

cle. Female gametophyte of Allium-type. Fruits

1–4-seeded red or orange-red berries. Seeds

relatively large, pale; testa lacks phytomelan

and disintegrates during development, but

tegmen is well developed and consists of thick-

walled cells; endosperm store aleurones, lipids

and hemicellulose; embryo is about ½ or 1/3 as

Superorder LILIANAE 677

long as endosperm. Anthroquinones detected

(Ruscus), present (Danae) kaempferol and

quercetin, steroidal saponins and chelidonic

acid;. n = 20; karyotypes of Danae and Ruscus

are bimodal and consist of six large and 14

small chromosomes.. . . . . . . . . 4. ruscaceae.

5 Herbs with generally globose hypocotyledon-

ary tuber or a proliferation of tubers and sto-

lons, usually with one to several slender

sheathed necks. Leaves basally concentrated,

solitary or several, basally sheathing, linear or

lanceolate to ovate or broadly cordate and then

prostrate, if broad basally constricted into pseu-

dopetiole; leaves often with unique complex,

pubescent, more or less dissected enations from

the upper part of relatively small lamina.

Stomata anomocytic. Inﬂ orescence simple sca-

pose raceme, the scape subtended by a basal

cataphyll. Flowers ebracteolate. Perianth seg-

ments free, two cycles similar or dissimilar.

Stamens six, all fertile; ﬁ laments basally

attached to the perianth segments; anthers dor-

siﬁ xed, epipeltate, elongate, opening longitudi-

nally. Tapetum secretory. Microsporogenesis

successive. Pollen grains 1-colpate. Style ﬁ li-

form, with punctiform stigma; ovary superior,

3-locular, with few to several ovules per locule.

Fruits smooth loculicidal capsules. Seeds testal-

tegmic, oval to comma-shaped, without phy-

tomelan, covered with long, unicellular,

air-ﬁ lled, white, or later rust-colored hairs;

embryo large, narrowly conical to cylindrical,

and ﬁ lls up most of the seed volume; mature

seed endosperm is strongly reduced (consumed

by developing embryo), but nucellar tissue

divides to form a sheath of perispermatic tissue,

which encloses the embryo either completely

or only for the chalazal 2/3; the peripheral layer

of the perisperm stores aleurone and fatty oils.

n = 5–7, 10, 12 and 21 (the basic number

appears to be x = 7, see Dahlgren and van Wyk

1988). . . . . . . . . . . . . . . . . 6. eriospermaceae.

1. CONVALLARIACEAE

Horaninow 1834 (including Aspidistraceae Endlicher

1841; Ophiopogonaceae Endlicher 1841; Peliosan-

thaceae R.A. Salisbury 1866; Polygonataceae R.A.

Salisbury 1866; Tupistraceae Schnitzlein 1846).

18/170. Eurasia (south to Bali and Lombok) and North

and Central America.

1.1 CONVALLARIOIDEAE

Fruits berries. Seeds not berrylike. Ovary superior.

n = 9–20. – polygonateae: Polygonatum, Heteropo-

lygonatum, Disporopsis, Maianthemum, Smilacina; con-

vallarieae: Theropogon, Speirantha, Conval laria,

Reineckea, Rohdea; aspidistreae: Tricalistra, Aspidistra,

Gonioscypha, Tupistra (including Campylandra).

1.2 OPHIOPOGONOIDEAE

Fruits leathery or berrylike capsules soon dehiscent

and deciduous after anthesis exposing seeds. Seeds

berry like, with ﬂ eshy seed coat. Ovary superior (Liriope)

or semi-inferior. n = 18. – ophiopogoneae:

Ophiopogon, Liriope; peliosantheae: Peliosanthes.

Have many similarities with the Colchicaceae –

Uvularioideae.

2. DRACAENACEAE

Salisbury 1866 (including Sansevieriaceae Nakai

1936). 2/100–160. Mainly tropical and subtropical

regions of the Old World with only a few species in

Hawaii, Central America and Cuba.

Dracaena, Sansevieria.

Close to the Convallariaceae, which is supported

both by serotaxonomic (Chupov and Kutiavina 1978)

and cytotaxonomic (Sharma and Chaudhuri 1964) data.

3. NOLINACEAE

Nakai 1943. 4/50. Southwestern United States, Mexico

and northern Central America.

Nolina, Calibanus, Dasylirion, Beaucamea.

Closely related to the Convallariaceae and Dracae-

naceae (Bogler and Simpson 1995, 1996; Bogler 1999).

The rare ﬂ avonol 3-0-methyl-8-C-methyl-quercetin, has

been found in all genera of the Nolina ceae and also

occurs in Xanthorrhoeaceae (Williams et al. 1988).

4. RUSCACEAE

Sprengel 1826. 3/9–11. Macaronesia, western and cen-

tral Europe, Mediterranean, western Asia.

Danae, Ruscus, Semele.

Closely related to the Convallariaceae.

678 Subclass II. LILIIDAE

5. ASPARAGACEAE

A.L. de Jussieu 1789 (including Hemiphylacaceae

Doweld 2005). 2/210. Widely distributed in the Old

World, mainly in arid and subarid regions.

Asparagus (? including Myrsiphyllum and Protas-

paragus), Hemiphylacus (5, Mexico).

According to Rudall et al. (1998) Hemiphylacus

must be included in the Asparagaceae, but from this

family it differs in fused perianth segments and chro-

mosome number (n = 56).

6. ERIOSPERMACEAE

Endlicher 1841. 1/102. Africa south of the Sahara;

with concentration of species in the southwestern Cape

Province.

Eriospermum.

A rather isolated taxon, which afﬁ nity is still obscure.

It shares some synapomorphies with the Cyanastraceae

(Tecophileales), such as nuclear formation of endo-

sperm, chalazosperm, successive microsporogenesis

and large female gametophyte, but sharply differs in

successive microsporogenesis, pollen ultrastructure,

structure of the seed coat, embryo shape. According to

Lu (1985), of the genera with which Eriospermum has

been compared, Bowiea of the Hyacinthaceae may be

the closest, but the differences are so obvious that a very

close relationship is very unlikely. Recent molecular

data support close relationships with the Asparagaceae

(Chase et al. 1995, 2000; Fay et al. 2000).

Bibliography

Arber A. 1924a. Danaë, Ruscus, and Semele: a morphological

study. Ann. Bot. 38: 229–260.

Arber A. 1924b. Myrisphyllum and Asparagus. Ann. Bot. 38:

635–659.

Bjornstad I. 1970. Comparative embryology Aspara-goideae-

Polygonatae. Nytt. Mag. Bot. 17: 169–207.

Baijnath H and P Perry. 1980. Preliminary observations of leaf

surface structures in Eriospermum Jacq. Proc. Electr.

Microsc. Soc. S. Afr. 10: 39–40.

Baker JG. 1875. Revision of the genera and species of

Asparagaceae. J. Linn. Soc. Bot. 14: 508–632.

Baker JG. 1877. Revision of the genera and species Anthericeae

and Eriospermeae. Bot. J. Linn. Soc. 15: 253–363.

Björnstad I. 1970. Comparative embryology of Asparagoideae-

Polygonateae, Liliaceae. Nytt. Mag. Bot. 17: 160–207.

Bogler D. 1998. Nolinaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 3, pp. 392–397. Springer,

Berlin/Heidelberg/New York.

Bogler DJ and BB Simpson. 1995. A chloroplast DNA study of

the Agavaceae. Syst. Bot. 20: 191–205.

Bogler DJ and BB Simpson. 1996. Phylogeny of Agavaceae

based on ITS rDNA sequence variation. Am. J. Bot. 83:

1225–1235.

Bos JJ. 1998. Dracaenaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 3, pp. 238–240. Springer,

Berlin/Heidelberg/New York.

Bugnon F. 1960. Contributions a 1’etude de quelques problemes

d’anatomie vegetale: V. Vascularisation de 1’eperon foliar chez

l’Asparagus sprengeri Regel. Bull. Soc. Bourg. 20: 65–71.

Conran JG. 1987. A phenetic study of the relationships of the

genus Drymophila R. Br. within the reticulate-veined

Liliiﬂ orae. Aust. J. Bot. 35: 283–300.

Conran JG. 1989. Cladistic analysis of some net-veined

Liliiﬂ orae. Plant Syst. Evol. 168: 123–141.

Conran JG and MN Tamura. 1998. Convallariaceae. In:

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

vol. 3, pp. 186–198. Springer, Berlin/Heidelberg/New York.

Cooney-Sovetts C and R Sattler. 1986. Phylloclade development

in the Asparagaceae: an example of homeosis. Bot. J. Linn.

Soc. 94: 327–371.

Cutler DF. 1992. Vegetative anatomy of Ophiopogoneae

(Convallariaceae). Bot. J. Linn. Soc. 110: 385–419.

Dai LK and SY Liang. 1991. Epidermal features of leaves and

their taxonomic signiﬁ cance in subfamily Ophiopogonoideae

(Liliaceae). Acta Phytotax. Sinica 29: 335–346.

Dickson A. 1886. On the occurrence of foliage leaves in Ruscus

(Semele) androgynus with some structural and morphological

observations. Trans. Proc. Bot. Soc. Edinb. 16: 130–149.

Duthie AV. 1940. Contribution to our knowledge of the genus

Eriospermum. Ann. Univ. Stellenbosch, Reeks A, Wiss.-

Natuurk. 18: 1–64.

Eguiarte LE et al. 1994. The systematic status of the Agavaceae

and Nolinaceae and related Asparagales in the monocotyle-

dons: an analysis based on the rbcL gene sequence. Bol. Soc.

Bot. Mex. 54: 35–56.

Gu Z, Q Yang, and K Kondo. 1990. A karyomorphological study

on Disporopsis Hance in China. La Kromosomo 57:

1916–1925.

Hernandez L. 1995. Taxonomic study of the Mexican genus

Hemiphylacus (Hyacinthaceae). Syst. Bot. 20: 546–554.

Hirsch A. 1977. A developmental study of the phylloclades of

Ruscus aculeatus L. Bot. J. Linn. Soc. 74: 355–365.

Hong D-Y and X-Y Zhu. 1990. Report on karyotypes of 6 spe-

cies in 4 genera of Poligonateae from China. Acta Phytotax.

Sinica 28: 185–198 (in Chinese with English summary).

Hong D-Y, LM Ma, and T Chen. 1987. A discussion on the

karyotype and evolution of the tribe Convallarieae s. 1.

(Liliaceae). In: DY Hong, ed. Plant chromosome research.

Nishiki Print Co., Hiroshima.

Jang CG and M Pfosser. 2002. Phylogenetics of Ruscaceae

sensu lato based on plastid rbcL and trnL-F DNA sequences.

Stapﬁ a. 80: 333–348.

Joyeux L. 1928. Valeur morphologique du cladode chez les

Ruscees. Mem. Acad. Royale Belgique, Bruxelles 9: 1–94.

Judd WS. 2003. The genera of Ruscaceae in the Southeastern

United States. Harvard Pap. Bot. 7: 93–149.

Superorder LILIANAE 679

Ko SC, YO Kim, and YS Kirn. 1985. A cytotaxonomical study

of the tribe Ophiopogoneae in Korea. Korean J. Plant Taxon.

15: 111–125.

Kubitzki K and PJ Rudall. 1998. Asparagaceae. In: K Kubitzki,

ed. The families and genera of vascular plants, vol. 3,

pp. 125–129. Springer, Berlin/Heidelberg/New York.

Lazarte JE and BF Falser. 1979. Morphology, vascular anatomy,

and embryology of pistillate and staminate ﬂ owers of

Asparagus ofﬁ cinalis. Am. J. Bot. 66: 753–764.

Lu A-M. 1985. Embryology and probable relationships of

Eriospermum Jacq. (Eriospermaceae). Nord. J. Bot. 5:

229–240.

Lü H-L, S-A Wu, J Yang, and G-Y Rao. 2000. Systematic

study on the tribe Polygonateae (Liliaceae s.l.) with the

evidence from leaf abaxial epidermis and seed coat. Acta

Phytotax. Sinica 38: 30–42 (in Chinese with English

summary).

Ma L-M and D-Y Hong. 1990. Pollen morphology and epider-

mal characters of leaves in Convallarieae (s.l.). Acta phyto-

tax. Sinica 28: 228–236.

McPherson MA, MF Fay, MW Chase, and SW Graham. 2004.

Parallel loss of a slowly evolving intron from two closely

related families in Asparagales. Syst. Bot. 29: 296–307.

Nakai T. 1936. Subdivision of Convallariaceae Link. Jpn. J. Bot.

12: 145–150.

Oganezova GH. 2000. Fruit and seed structure of some

Asparagaceae s.l. in connection with the volume and phylog-

eny of the family. Bot. Zhurn. 85(8): 14–31. (in Russian with

English summary).

Perry PL. 1994. A revision of the genus Eriospermum

(Eriospermaceae). Contrib. Bolus Herb. 17: 1–320.

Perry PL and PJ Rudall. 1998. Eriospermaceae. In: K Kubitzki,

ed. The families and genera of vascular plants, vol. 3,

pp. 241–244. Springer, Berlin/Heidelberg/New York.

Pires JC, IJ Maureira, TJ Givnish, KJ Sytsma, O Seberg,

G Petersen, JI Davis, DW Stevenson, PJ Rudall, MF Fay, and

MW Chase. 2004. Phylogeny, genome size, and chromosome

evolution in Asparagaceae. In: JT Columbus, EA Friar,

CW Hamilton, JM Porter, LM Prince, and MG Simpson, eds.

Monocots: Comparative biology and evolution, vol. 1,

pp. 287–304. Rancho Santa Ana Botanical Garden, Claremont.

Rao G-Y and K-Y Pan. 1994. Pollen morphology of the

Polygonateae and its systematic signiﬁ cance. Cathaya 6:

75–91.

Rao RP and A Kaur. 1979. Embryology and systematic position

of Ophiopogon intermedius. Proc. Indian Natl. Sci. Acad.

45: 175–187.

Rudall PJ and DF Cutler. 1995. Asparagales, a reappraisal. In:

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

Monocotyledons: systematics and evolution, pp. 157–168.

Royal Botanic Gardens, Kew.

Rudall PJ and G Campbell. 1998. Flower and pollen structure of

Ruscaceae in relation to Aspidistreae and other

Convallariaceae. Flora 194: 201–214.

Rudall PJ, CA Furness, MW Chase, and FF Michael. 1997.

Microsporogenesis and pollen sulcus type in Asparagales

(Lilianae). Canad. J. Bot. 75: 408–430.

Rudall PJ, EM Engleman, L Hanson, and MW Chase. 1998.

Embryology, cytology and systematics of Hemiphylacus,

Asparagus and Anemarrhena (Asparagales). Plant Syst.

Evol. 211: 181–199.

Rudall PJ, JG Gonran, and MW Chase. 2000. Systematics of

Ruscaceae/Convallariaceae: a combined morphological and

molecular investigation. Bot. J. Linn. Soc. 134: 73–92.

Sarkar AK, N Datta, and U Chatterjee. 1981 (1982). Cytology of

Peliosanthes Andr. (Liliaceae) as an aid to taxonomy.

Caryologia 34: 467–472.

Schlitder J. 1953. Blütenartikulation und Phyllokladien der

Liliaceae organophylogenetisch betrachtet, part 1 and 2.

Feddes Repert. 55: 154–258.

Schlitter J. 1960. Die Asparageenphyllokladien erweisen sich

auch ontogenetisch als Blätter. Bot. Jahrb. Syst. 79: 428–446.

Schulze W. 1982. Beiträge zur Taxonomie der Liliiﬂ oren X.

Asparagaceae. Wiss. Z. Friedrich Schiller Univ. Jena 31:

309–330.

Sharma AK and M Chaudhuri. 1964. Cytological studies as an

aid in assessing the status of Sansevieria, Ophiopogon, and

Curculigo. Nucleus 7: 43–58.

Sharma AK and PC Datta. 1960. Chromosome studies in species

of Dracaena with special reference to their means of specia-

tion. J. Genet. 57: 43–76.

Sharma AK and I Ghosh. 1968. Cytotaxonomy of Dracaena.

J. Biol. Sci. 11: 45–55.

Shinwari ZK, H Kato, R Terauchi, and S Kawano. 1994.

Phylogenetic relationships among genera in the Liliaceae-

Asparagoideae-Polygonatae s. 1. inferred from rbcL gene

sequence data. Plant Syst. Evol. 192: 263–277.

Stützel T, U Reck, and D Müller-Doblies. 1991. Morphologische

Studien zur Systematik der Convallariaceae. In: G Wagenitz,

ed. 10th Symposium Morphologie, Anatomie und Systematik,

p. 74. Göttingen.

Tamura MN. 1993. Biosystematic studies on the genus

Polygonatum (Liliaceae) III. Morphology of staminal ﬁ la-

ments and karyology of eleven Eurasian species. Bot. Jahrb.

Syst. 115: 1–26.

Tamura MN. 1995. A karyological review of the orders

Asparagales and Liliales (Monocotyledoneae). Feddes

Repert. 106: 83–111.

Tamura MN, M Ogisu, and J-M Xu. 1997a.

Heteropolygonatum,

a new genus of the tribe Polygonatae (Convallariaceae). Kew

Bull. 52: 949–956.

Tamura MN, AE Schwarzbach, S Kruse, et al. 1997b.

Biosystematic studies on the genus Polygonatum

(Convallariaceae) IV. Molecular phylogenetic analysis based

on restriction site mapping of the chloroplast gene trnK.

Feddes Repert. 108: 159–168.

Utech FH. 1979. Floral vascular anatomy of the Himalayan

Theropogon pallidus Maxim. (Liliaceae-Convallarieae).

Ann. Carnegie Mus. 48(3): 25–41.

Utech FH and S Kawano. 1976. Floral vascular anatomy of

Convallaria majalis L. and C.keiskei Miq. (Liliaceae –

Convallariinae). Bot. Mag. (Tokyo) 89: 173–182.

Vaikos NP, SK Markandeya, and RM Pai. 1989. Floral anatomy

of the Liliaceae: tribe Convallarieae. Proc. Indian Acad. Sci.

Plant Sci. 99: 91–95.

Velenovsky J. 1903. Zur Deutung der Phyllokladien der

Asparageen. Beih. Bot. Centralbl. 15: 257–268.

Wu S-A, H-L LU, J Yang, G-Y Rao, R-L You, S Ge, and

Y Zhong. 2000. Molecular systematic studies on the tribe

Polygonateae (s.l.) in China based on RFLPs data of PCR-

ampliﬁ ed chloroplast DNA fragments. Acta Phytotax. Sinica

38: 97–110 (in Chinese with English summary).

680 Subclass II. LILIIDAE

Yamashita J and MN Tamura. 2000. Molecular phylogeny of the

Convallariaceae (Asparagales). In: KL Wilson and

DA Morrison, eds. Monocots: systematics and evolution,

pp. 387–400. CSIRO, Collingwood.

Yamashita J and MN Tamura. 2004. Phylogenetic analysis and

chromosome evolution in Convallarieae (Ruscaceae sensu

lato), with some taxonomic treatments. J. Plant Res. 117:

363–370.

Yang D-Q and K-F Zhu. 1990. Studies on karyotypes of 5 spe-

cies of Rohdea and Tupistra. Acta Phytotax. Sinica 28: 199–

206 (in Chinese with English summary).

Yang Y, H Li, X Liu, and K Katsuhiko. 1990. Karyotype study

on the genus Ophiopogon in Yunnan. Acta Bot. Yunn. Suppl.

3: 94–102.

Yeo PF. 1968. A contribution to the taxonomy of the genus

Ruscus. Notes Roy. Bot. Gard. Edinb. 28: 237–264.

Yeo PF. 1998. Ruscaceae. In: K Kubitzki, ed. The families and

genera of vascular plants, vol. 3, pp. 412–416. Springer,

Berlin/Heidelberg/New York.

Zimmermann MH and PB Tomlinson. 1969. The vascular sys-

tem in the axis of Dracaena fragrans (Agavaceae):

1. Distribution and development of primary strands. J. Arnold

Arbor. 50: 370–383.

Zimmermann MH and PB Tomlinson. 1970. The vascular sys-

tem in the axis of Dracaena fragrans (Agavaceae): 2. Origin

and distribution of secondary tissues. J. Arnold Arbor. 51:

478–491.

Superorder PANDANANAE

Order 16. PANDANALES

More or less arborescent plants, sometimes (Freycinetia)

lianas with clasping, aerial roots, sometimes epiphytic;

prop-roots usually at base of stem. Stems simple or

sympodially branched, woody, but have no secondary

growth or secondary vascular tissue and increase of

trunk diameter is the result of primary thickening growth

alone. Silica absent, but calcium oxalate occurs both as

raphides and other kinds of crystal. Sieve-element plas-

tids of form-P2cf. Vessels in all vegetative organs, in

roots with very primitive scalariform perforations.

Leaves spirally arranged, usually aggregated at the ends

of the branches, tristichous or (Sararanga) tetrastichous,

but the ranks run in well-marked spirals owing to spiral

growth of the stem, usually linear-ensiform, long, usu-

ally rigid, coriaceous, sheathing at base, parallel-veined.

Stomata more or less distinctly tetracytic. Flowers min-

ute, numerous, much reduced, condensed in terminal or

sometime lateral or axillary, usually unisexual inﬂ ores-

cences, subtended by bracts; lower bracts foliaceous,

upper bracts variously coloured. Sometimes fragrant

and slightly sweet; individual ﬂ owers strictly unisexual

(most Pandanus), or with vestiges of the other sex, occa-

sionally bisexual in some species of Freycinetia, with-

out perianth or sometimes (Sara ranga) with perianth-like

cupule. Male ﬂ owers have many stamens (9–300); ﬁ la-

ments ﬂ eshy, free or often connate; anthers basiﬁ xed,

tetrasporangiate, sometimes apiculate. Tapetum plas-

modial or secretory. Microsporogenesis successive.

Pollen grains 2-celled, 1-porate, echinate, psilate, retic-

ulate. Female ﬂ owers free or many connate into clusters.

Gynoecium of one to several more or less united car-

pels; style short or wanting; stigmas reniform or hippo-

crepiform, sometimes elongated ovary superior,

1-locular, with one ovule in Pandanus and Sararanga or

several or numerous in Freycinetia. Ovules anatropous,

bitegmic, crassinucellate, with parietal cell. Female

gametophyte of Polygonum-type or Allium-type.

Endosperm nuclear. Fruits baccate or drupes, in heads.

Seeds rather small, straight or variously curved; seed

coat thin and membranous or (Sararanga) thickened;

embryo small and straight; endosperm copious, oily or

(Freycinetia) starchy. n = 25, 28, 30, 60.

Related to the Cyclanthales.

1. PANDANACEAE

R. Brown 1810 (including Freycinetiaceae Brongniart

ex Le Maout et Decaisne 1868). 4/800–900. Tropical

regions of the Old World, especially in Malesia,

Melanesia, and Madagascar, with a few species in tem-

perate regions (China, Japan, New Zealand).

1.1 PANDANOIDEAE

Trees generally supported basally by aerial roots.

Flowers in repeatedly branched panicles (Sararanga)

or in spikes or racemes of spikes (Pandanus). Ovary

with one ovule. Fruits monodrupes or polydripes

depending on the number of carpels (Pandanus) or

polypyrenoid berries (Sasraranga). Endosperm oily. –

Sararanga, Pandanus.

1.2 FREYCINETIOIDEAE

Woody lianas with clasping roots and linear to ovate or

obovate leaves. Flowers in pseudoumbels (rarely

racemes) of spikes. Ovary multiovulate. Fruits ﬂ eshy

or ligneous berries. Endosperm starchy. – Freycinetia,

Martellidendron.

Superorder PANDANANAE 681

Bibliography

Callmander MW, P Chassot, Küpfer, and PP Lowry. 2003.

Recognition of Martelliodendron, a new genus of

Pandanaceae, and its biogeographic implications. Taxon 52:

747–762.

Cox PA. 1981. Bisexuality in the Pandanaceae: new ﬁ ndings in

the genus Freycinetia. Biotropica 13: 195–198.

Cox PA. 1990. Pollination and the evolution of breeding systems

in Pandanaceae. Ann. Missouri Bot. Gard. 77: 816–840.

Cox PA, K-L Huynh, and BC Stone. 1995. Evolution and sys-

tematics of Pandanaceae. In: PJ Rudall, PJ Cribb, DF Cutler,

and CJ Humphries, eds. Monocotyledons: systematics and

evolution, vol. 2, pp. 663–684. Royal Botanic Gardens, Kew.

Fagerlind F. 1940. Stempelbau und Embryosackentwicklung bei

einigen Pandanazeen. Ann. Jard. Bot. Buitenzorg 49: 55–78 +

Taf. 7–13.

Furness CA and PJ Rudall. 2006. Comparative structure and

development of pollen and tapetum in Pandanales. Int.

J. Plant Sci. 167: 331–348.

Huynh K-L. 1974. La morphologic microscopique et la tax-

onomie du genre Pandanus. Bot. Jahrb. Syst. 94:

190–256.

Huynh K-L. 1991. The ﬂ ower structure in the genus Freycinetia,

Pandanaceae (part 1) – Potential bisexuality in the genus

Freycinetia. Bot. Jahrb. Syst. 112: 295–328.

Huynh K-L. 1992. The ﬂ ower structure in the genus Freycinetia,

Pandanaceae (part 2) – Early differentiation of the sex organs,

especially of the staminodes, and further notes on the anthers.

Bot. Jahrb. Syst. 114: 417–441.

Huynh K-L. 2001. Contribution to the ﬂ ower structure of

Sararanga (Pandanaceae). Bot. J. Linn. Soc. 136: 239–245.

Huynh K-L and PA Cox. 1992. Flower structure and potential

bisexuality in Freycinetia reineckei (Pandanaceae), a species

of the Samoa Islands. Bot. J. Linn. Soc. 110: 235–265.

Jarzen DM. 1983. The fossil pollen record of the Pandanaceae.

Gard. Bull. 36: 163–175.

Nambudiri EM and WD Tidwell. 1978. On probable afﬁ nities of

Viracarpon Sahni from the Deccan Intertrappean ﬂ ora of

India. Paleontographica 166: 30–43.

North CA and AJ Willis. 1971. Contributions to the anatomy of

Sararanga (Pandanaceae). Bot. J. Linn. Soc. 64: 411–421.

Pijl L van der. 1956. Remarks on pollination by bats in

Freycinetia, Duabanga, and Haplophragma, and on chi-

ropterophily in general. Acta Bot. Neerl. 5: 135–144.

Poppendieck H-H. 1987. Monoecy and sex changes in

Freycinetia (Pandanaceae). Ann. Missouri Bot. Gard. 74:

314–320.

Stone BC. 1968a. Morphological studies in Pandanaceae:

I. Staminodia and pistillodia of Pandanus and their hypo-

thetical signiﬁ cance. Phytomorphology 18: 498–509.

Stone BC. 1968b. Materials for a monograph of Freyci-netia

Gaud.: IV. Subdivision of the genus, with ﬁ fteen new sec-

tions. Blumea 16: 361–372.

Stone BC. 1970. Materials for a monograph of Freycinetia

Gaud. (Pandanaceae): V. Singapore, Malaya, and Thailand.

Gardens’ Bulletin. Straits Settlements 25: 189–207.

Stone BC. 1972a. Materials for a monograph of Freyci-netia

Gaud. (Pandanaceae): XV. The Sumatran species. Federat.

Mus. J., n.s. 15: 203–207.

Stone BC. 1972b. A reconsideration of the evolutionary status of

the family Pandanaceae and its signiﬁ cance in monocotyle-

don phylogeny. Quart. Rev. Biol. 47: 34–45.

Stone BC. 1990. New evidence for the reconciliation of ﬂ oral

organization in Pandanaceae with normal angiosperm pat-

terns. In: P Baas, K Kalkman, and R Geesink, eds. The plant

diversity of Malesia, pp. 33–35. Kluwer, Dordrecht.

Stone BC, K-L Huynh, and H-H Poppendieck. 1998.

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

vascular plants, vol. 3, pp. 397–404. Springer, Berlin/

Heidelberg/New York.

Tomlinson PB. 1965. A study of the stomatal structure in

Pandanaceae. Pac. Sci. 19: 38–54.

Vaughan RE and PO Wiehe. 1953. The genus Pandanus in the

Mascarene Islands. Bot. J. Linn. Soc. 55: 1–32.

Zimmermann MH, PB Tomlinson, and J LeClaire. 1974.

Vascular construction and development in the stems of cer-

tain Pandanaceae. Bot. J. Linn. Soc. 68: 21–41.

Order 17. CYCLANTHALES

Palm-like terrestrial or sometimes epiphytic perennial

herbs or less often herbaceous vines or lianas. Stem

very short to long and slender, rhizomatous to aerial,

usually more or less ligniﬁ ed. Commonly with raphide

sacs and sometimes also with other types of calcium

oxalate crystals; silica bodies lacking. Watery or milky

juice present in all organs. Mucilage canals occur in the

vegetative parts of all genera except Cyclanthus; in this

genus there are laticiferous vessels. Sieve-element

plastids of form-P2c. Vessels in roots and leaves, with

scalariform perforations. Leaves large, spiral, spirodis-

tichous, or orthodistichous, sheathing at base, mostly

petiolate; lamina very deeply bilobed to biﬁ d or more

rarely ﬂ abelliform-divided or entire, sometimes plicate,

with parallel or parallel-pinnate venation and with

cross-veins. Stomata tetracytic. Flowers small, in

axillary or terminal spadices, sometimes screwlike,

subtended by 2–4 (to 11) conspicuous, lanceolate-

ovate, variously coloured bracts, the spathes, which

enclose the spadix when young; individual ﬂ owers

much reduced, unisexual, densely crowded, the female

ones more or less embedded in axis, male and female

ﬂ owers in same spadix. Male ﬂ owers without perianth

or with symmetrical or asymmetrical perianth; perianth

segments vary in shape and may have an adaxial secre-

tory glandule. Stamens mostly numerous; ﬁ laments

basally more or less connate and sometimes adnate to

the perianth; in most genera swollen below, possessing

a basal bulb, anthers basiﬁ xed, tetrasporangiate,

682 Subclass II. LILIIDAE

usually opening longitudinally, sometimes with an api-

cal secretory glandule. Tapetum secretory.

Microsporogenesis successive. Pollen grains 2-celled,

usually boat-shaped, tectate with a thick foot-layer and

no endexine, 1-colpate or less often 1–2-porate or

(Dianthoveus and Evodianthus) inaperturate, usually

foveolate. Female ﬂ owers with simple perianth consist-

ing of four free or connate segments, four opposite sta-

minodia and gynoecium of four united carpels

alternating with staminodia; stylodia free or basally

connate into a short style; often stylodia are very short

or wanting and then four stigmas nearly sessile; ovary

superior, 1-locular, with numerous parietal, subapical

or apical ovules. Ovules anatropous, bitegmic, weakly

crassinucellate or almost tenuinucellate, with parietal

cell. Female gametophyte of Polygonum-type.

Endosperm helobial. Fruits many-seeded, baccate,

often coalescent into multiple fruit. Seeds with gener-

ally straight, linear, small to medium-sized embryo and

copious endosperm containing oil, aleurone, often

hemicellulose, Dicranopygium rich in starch, which is

also present in embryo, n = 9, 15, 16.

This order has features, especially in the plicate

leaves, that are similar to those of the palms (Dahlgren

et al. 1985), but it differs sufﬁ ciently in predominantly

herbaceous habit, absence of silica, morphology of

inﬂ orescence, constant syncarpous gynoecium,

absence of septal nectaries, constant successive

microsporogenesis, helobial endosperm, starch-rich

endosperm and embryo in Dicranopygium, as well as

in the epicuticular wax without crystalloids. Probably

related to the Pandanaceae. According to Harling

(1958), Freycinetia is probably the nearest to the

Cyclanthales. However, the order differs sufﬁ ciently

from the Pandanales in secretory tapetum, helobial

endosperm, more differentiated and plicate leaves, the

non spiral growth of the stem.

1. CYCLANTHACEAE

Poiteau ex A. Richard 1824. 12/235. Tropical America,

West Indies.

1.1 CARLUDOVICOIDEAE

Inﬂ orescence with ﬂ owers in spirally arranged groups.

Placentas four or one. Latex absent. Leaves apically

biﬁ d, fanlike, or entire, plicate. Fruits baccate, syncar-

pous or not. n = 9, 15, 16. – Carludovica, Dianthoveus,

Evodianthus, Aplundia, Thoracocarpus, Schulte-

siophytum, Dicranopygium, Ludovia, Sphaeradenia,

Stelestylis, Chorigyne.

1.2 CYCLANTHOIDEAE

Male and female ﬂ owers in separate, alternating cycles

or ﬂ at spirals. Placentas numerous. Latex present.

Leaves deeply cleft with a forked main rib, non plicate.

Fruits dry, syncarpous. n = 9. – Cyclanthus.

Bibliography

Dahlgren RMT. 1982. Cyclanthaceae. Monocot Newslett. 2:

7–32.

Eriksson R. 1989. Chorigyne, a new genus of the Cyclanthaceae

from Central America. Nord. J. Bot. 9: 31–45.

Eriksson R. 1993. Systematics of the Cyclanthaceae, especially

Sphaeradenia and Chorigyne. Goteborg.

Eriksson R. 1994. Phylogeny of the Cyclanthaceae. Plant Syst.

Evol. 190: 31–47.

Furness CA and PJ Rudall. 2006. Comparative structure and

development of pollen and tapetum in Pandanales. Int.

J. Plant Sci. 167: 331–348.

French CH, K Klancy, and PB Tomlinson. 1983. Vascular pat-

terns in stems of the Cyclanthaceae. Am. J. Bot. 70:

1386–1400.

Gottsberger G. 1991. Pollination of some species if the

Carludovicoideae, and remarks on the origin and evolution

of the Cyclanthaceae. Bot. Jahrb. Syst. 113: 221–235.

Hammel BE and GJ Wilder. 1989. Dianthoveus: A new genus of

Cyclanthaceae. Ann. Missouri Bot. Gard. 76: 112–123.

Harling G. 1946. Studien über den Blütenbau und die

Embryologie der Familie Cyclanthaceae. Svensk Bot. Tidskr.

40: 257–272.

Harling G. 1958. Monograph of the Cyclanthaceae. Acta Horti

Berg. 18: 1–428.

Harling G, GL Wilder, and R Eriksson. 1998. Cyclanthaceae.

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

plants, vol. 3, pp. 202–215. Springer, Berlin/Heidelberg/

New York.

Surange KR. 1949. A contribution to the morphology and anat-

omy of the Cyclanthaceae. Trans. Natl. Inst. Sci. India,

Calcutta 3(4): 159–209.

Tomlinson PB and GJ Wilder. 1984. Systematic anatomy of

Cyclanthaceae (Monocotyledoneae) – an overview. Bot.

Gaz. 145: 535–549.

Wilder GJ. 1976. Structure and development of leaves of

Carludovica palmata (Cyclanthaceae) with reference to

other Cyclanthaceae and Palmae. Am. J. Bot. 63:

1237–1256.

Wilder GJ. 1981a. Structure and development of Cyclanthus

bipartitus Poit. (Cyclanthaceae) with reference to other

Cyclanthaceae. II. Adult leaf. Bot. Gaz. 142: 96–114.

Wilder GJ. 1981b. Morphology of adult leaves in the

Cyclanthaceae (Monocotyledoneae). Bot. Gaz. 142:

564–588.

Superorder PANDANANAE 683

Wilder GJ. 1984. Anatomy of noncostal portions of lamina in

the Cyclanthaceae (Monocotyledoneae), part 5. Bot. Mus.

Leaﬂ . 30: 103–133.

Wilder GJ. 1985. Anatomy of noncostal portions of lamina in

the Cyclanthaceae (Monocotyledoneae), parts 1–4. Bot. Gaz.

146: 82–105, 213–231, 375–394, 545–563.

Wilder GJ. 1986. Anatomy of ﬁ rst-order roots in the

Cyclanthaceae (Monocotyledoneae), parts 1 and 2. Canad.

J. Bot. 64: 2622–2644, 2848–2864.

Wilder GJ. 1987. Contributions to taxonomy and morphology of

Schultesiophytum chorianthum Harl. and Dicranopygium

mirabile Harl. (Cyclanthaceae). Opera Bot. 92: 277–291.

Wilder GJ. 1988. Inﬂ orescence position as a taxonomic charac-

ter in the Cyclanthaceae. Bot. Gaz. 149: 110–115.

Wilder GJ. 1989. Morphology of Dianthoveus cremnophilus

(Cyclanthaceae). Canad. J. Bot. 67: 2450–2464.

Wilder GJ. 1992a. Comparative morphology and anatomy of

absorbing roots and anchoring roots in three species of

Cyclanthaceae (Monocotyledoneae). Canad. J. Bot. 70: 38–48.

Wilder GJ. 1992b. Orthodistichous and dorsiventral symmetry

on adult shoots of Cyclanthus bipartitus (Cyclanthaceae,

Monocotyledoneae). Canad. J. Bot. 70: 1388–1400.

Wilder GJ and DH Harris. 1981. Laticifers in Cyclanthus bipar-

titus Poit. (Cyclanthaceae). Bot. Gaz. 143: 84–93.

Order 18. TRIURIDALES

Usually perennial, commonly small, achlorophyllous,

mycotrophic herbs with slender system of underground

rhizomes often covered with scales. Raphides and silica

bodies lacking. Roots ﬁ liform, with a cortex consisting

of l–3 layers of parenchyma cells containing mycor-

rhizal hyphae. Stems usually slender, erect. Vascular

system weakly developed, with the vascular bundles in

a single ring. Vessels lacking. Sieve-element plastids of

P2c-type (Behnke 2002). Leaves reduced to alternate

scales, have a single vascular strand. Stomata present or

absent. Flowers very small, in terminal racemes (rarely

reduced to a solitary ﬂ ower), bracteate, bisexual (spp. of

Sciaphila) or more often unisexual, monoecious or dioe-

cious), actinomorphic. Perianth of 3–6 (-10) segments

in a single or two cycles; segments mostly more or less

basally connate, often reﬂ exed and may form a starlike

conﬁ guration, often conspicuously caudate apically,

persistent. Stamens 2–6 (-10), often some staminodial,

ﬁ lamented, sessile or immersed in the conical andro-

phore; anthers tetrasporangiate or less often 2–3-spo-

rangiate, more or less extrorse, dehiscing longitudinally

or more often transversely, typically extrorse, some-

times connective extends into a long, slender, terminal

appendage. Tapetum plasmodial or secretory, with inter-

mediate forms occurring within the Sciaphileae

(Rubsamen-Weustenfeld 1991; Furness et Rudall 2006).

Microsporogenesis successive. Pollen grains 3-celled,

globose, inaperturate, with very thin exine which shows

more or less spiny-gemmate, granulate or verrucate

(Furness and Rudall 2006). Gynoecium of ten to many

free or basally connate carpels, each with one basal

ovule and nearly terminal or more often with more or

less lateral or gynobasic stylodium. Ovules anatropous,

bitegmic, tenuinucellate, without parietal cell, nucellar

cap absent. Female gametophyte of Polygonum-type,

but in Triuris very probably of Fritillaria-type

(Rubsamen-Weustenfeld 1991). Endosperm nuclear.

Fruits of small, thick-walled, achenelike follicles or

rarely of achenes. Seeds small, straight or slightly

curved, with small, undifferentiated embryo consisting

of a globose, embryo proper and a more or less short

suspensor; endosperm copious, containing hemicellu-

lose, protein, oil, and exclusively in the Triurideae starch

that disappears continuously during seed maturation;

seed coat formed by the outer integument only, espe-

cially by its inner layer; middle cuticle very strongly

thickened, directly adjacent to the endosperm tissue

(Rubsamen-Weustenfeld 1991). n = 9, 11, 12 (mostly),

13, 14, 15, 16.

According to Chase et al. (2000), the Triuridaceae

related to the Pandanaceae and Cyclanthaceae.

1. TRIURIDACEAE

Gardner 1843 (including Lacandoniaceae E. Marttnes

et C.H. Ramos 1989). 10–11/45–50. Tropical regions of

America, Africa, and Asia. The largest genus Sciaphila

(30) is concentrated in the Old World.

sciaphileae: Sciaphila, Seychellaria, Andruris,

Hyalisma, Soridium; triurideae: Triuris (including

Lacandonia), Triuridopsis, Peltophyllum. kupeaeae:

Kupea, Kihansia.

In a variation of Triuris, described as a new genus

Lacandonia, with central stamens and peripheral car-

pels, is probably no more than a very interesting case

of homeotic mutation (Takhtajan 1997).

Bibliography

Ambrose BA, S Espinosa-Matís, S Vázquez-Santana, J Márquez-

Guzmán, and ER Alvarez-Buylla. 2006. Comparative devel-

opmental series of the Mexican triurids support a euanthial

684 Subclass II. LILIIDAE

interpretation for the unusual reproductive axes of Lacan-

donia schismatica (Triuridaceae). Am. J. Bot. 93: 15–35.

Cheek M. 2003. Kupeaeae, a new tribe of Triuridaceae from

Africa. Kew Bull. 58: 939–949.

Cheek M, Williams SA, and Etuge M. 2003. Kupea martinetugei,

a new genus and species of Triuridaceae from western

Cameroon. Kew Bull. 58: 225–228.

Davidse G and SE Martinez. 1990. The chromosome number of

Lacandonia schismatica (Lacandoniaceae). Syst. Bot. 15:

635–637.

Green PS and O Solbrig. 1966. Sciaphila dolichostyla

(Triuridaceae). J. Arnold Arbor. 47: 266–269.

Furness CA and PJ Rudall. 2006. Comparative structure and

development of pollen and tapetum in Pandanales. Int.

J. Plant Sci. 167: 331–348.

Furness CA, PJ Rudall, and A Eastman. 2002. Contribution

of pollen and tapetal characters to the systematics of

Triuridaceae. Plant Syst. Evol. 235: 209–218.

Gandolfo MA, KC Nixon, and WL Crepet. 2002. Triuridaceae

fossil ﬂ owers from the Upper Cretaceous of New Jersey.

Am. J. Bot. 89: 1940–1957.

Igersheim A, M Buzgo, and PK Endress. 2001. Gynoecium

diversity and systematics in basal monocots. Bot. J. Linn.

Soc. 136: 1–65.

Imhof S. 1998. Subterranean structures and mycotrophy of the

achlorophyllous Triuris hyaline Miers (Triuridaceae). Canad.

J. Bot. 76: 2011–2019.

Imhof S. 2003. A dorsiventral mycorrhizal root in the achloro-

phyllous Sciaphila polygyna (Triuridaceae). Mycorrhiza 13:

327–332.

Imhof S. 2004. Morphology and development of the subterra-

nean organs of the achlorophyllous Sciaphila polygyna

(Triuridaceae). Bot. J. Linn. Soc. 146: 295–301.

Maas PJM and T Rubsamen. 1986. Triuridaceae. Flora

Neotropica 40: 1–55. New York.

Maas van de Kramer H. 1995. Triudiﬂ orae – Gardener’s delight?

In: PJ Rudall, PJ Cribb, DF Cutler, and CJ Humphries, eds.

Monocotyledons: systematics and evolution, pp. 287–301.

Royal Botanic Gardens, Kew.

Maas van de Kamer H and PJM Maas. 1994. Triuridopsis, a new

monotypic genus in Triuridaceae. Plant Syst. Evol. 192:

257–262.

Maas van der Kamer H and T Rübsamen-Weustenfeld. 1998.

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

vascular plants, vol. 3, pp. 452–458. Springer, Berlin/

Heidelberg/New York.

Marquez-Guzman J, S Vazquez-Santana, ME Engleman,

A Martinez-Mena, and E Martinez. 1993. Pollen development

and fertilization in Lacandonia schismatica (Lacandonia-

ceae) [Triuridaceae]. Ann. Missouri Bot. Gard. 80: 891–897.

Martinez E and CH Ramos. 1989. Lacandoniaceae (Triuridales):

Una nueva familia de Mexico. Ann. Missouri Bot. Gard. 76:

128–135.

Meerendonk JPM van der. 1984. Triuridaceae. In: CGGJ van

Steenis, ed. Flora Malesiana, ser. 1, 10: 109–121. Dodrecht.

Rübsamen-Weustenfeld T. 1991. Morphologische, embryologis-

che, und systematische Untersuchungen an Triuridaceae.

Bibl. Bot. 149: 1–113.

Rudall P. 2003. Monocot pseudanthis revisited: ﬂ oral structure

of the mycoheterotrophic family Triuridaceae. Int. J. Plant

Sci. 164: S307–S320.

Sahashi N, T Ohmoto, K Uehara, M Ikuse, and C Chuma. 1991.

Pollen morphology of Andruris japonica (Triuridaceae).

Grana 30: 597–600.

Tomlinson PB. 1982. Helobieae (Alismatidae). In: CR Metcalfe,

ed. Anatomy of the monocotyledons, 7: 1–522. Clarendon,

Oxford.

Vergara-Silva F, S Espinosa-Matías, BA Ambrose, S Vázquez-

Santana, A Martínez-Mena, J Márquez-Guzmán, E

Martínez, EM Meyerowitz, and ER Alvarez-Buylla. 2003.

Inside-out ﬂ owers characteristic of Lacandonia schismat-

ica evolved at least before its divergence from a closely

related taxon, Triuris brevistyla. Int. J. Plant Sci. 164:

345–357.

Order 19. VELLOZIALES

Usually arborescent or shrubby perennials of various

sizes (sometimes over 6 m tall in South America) with

simple or sparsely dichasially branched woody stems.

Tanniniferous cells sometimes present. Commonly

secreting resin or gum. Sieve-element plastids of Pf,

-Pcf, -Pcfs, -Pfs, -P2c-types (Behnke et al. 2000).

Vessels in roots with simple perforations and in leaves

with scalariform perforations. Leaves xeromorphic,

tristichous or spirotristichuosly arranged, crowded at

the end of stem or its branches when new, linear, often

pungent-pointed, parallel-veined, entire or often

dentate-spinulose along the margins, sheathing at the

base. Stomata paracytic or sometimes tetracytic.

Adventitious roots alternating with the leaves. Flowers

solitary, bisexual or (Barbaceniopsis) functionally

unisexual, actinomorphic, 3-merous, in terminal,

1-many-ﬂ owered inﬂ orescences. Perianth segments

six, in two cycles, usually similar, petaloid, spreading,

free or usually more or less connate into a short or

long tube; perianth tube often with six appendages

that are free or united into corona. Stamens six in two

cycles or often, in most species of Vellozia, numerous

(up to 76) and in six bundles; anthers tetrasporan-

giate or disporangiate, exceptionally 10-sporangiate

(Xerophyta schnizleiniana), usually linear and long,

basiﬁ xed to dorsiﬁ xed, opening longitudinally, latrorse

to introrse, rarely extrorse. Tapetum mostly secretory.

Microsporogenesis successive. Pollen grains 2- or

3-celled, solitary or (species of Vellozia) in tetrads,

tectate-columellate, 1-colpate or rarely (Vellozia)

inaperturate, ﬁ nely verrucate to reticulate (Furness

and Rudal 2006). Gynoecium of three united carpels;

style slender, generally widening apically into a