Sun Kun (孙坤)[1]; [Robert A. DeFilipps[2]]
露兜树科 lou dou shu ke
Terrestrial, evergreen, dioecious trees, shrubs or woody lianas. Stems simple or often bifurcately branched, ringed with persistent annular leaf scars, often producing adventitious prop (stilt) roots; aerial roots present or absent. Vegetative reproduction by suckers present or absent. Leaves simple, numerous, spirally arranged at apex of stems and branches, sessile, linear to lanceolate, keeled (carinate) beneath, glabrous, leathery, often lustrous or glaucous, parallel-veined, with numerous horizontal cross-veins (secondary veins), open-sheathed and amplexicaul at base, the margin and midrib beneath often serrate (spinulose), the apex often long-acuminate. Male inflorescences axillary and terminal, compound, bracteate, comprising several-branched racemes or panicles with crowded flowers on spicate ultimate branches; bracts (spathes) often enlarged at apex, white or colored. Perianth absent. Male flowers sessile (pedicellate in Sararanga), with pistillode (vestigial ovary) sometimes present; stamens numerous, fasciculate, arising on rachises or spadix-branches; filaments smooth (Pandanus) or papillose (Freycinetia), seemingly branched; anthers basifixed, bilocular with 4 pollen sacs, dehiscing by longitudinal slits, connective often apiculate; pollen grains often spinulose. Female inflorescences terminal, solitary or in spikes, racemes or capitula, short, bracteate, with crowded flowers, often pendulous in fruit. Female flowers with or without staminodes; pistils free or aggregated (congested) and appressed (fused) to adjacent pistils forming 1-to many-carpelled phalanges or clusters; ovary superior, 1-to several-loculed; ovules solitary to numerous, anatropous, placentation basal or parietal (marginal); style absent or very short; stigmas 1 or more, subsessile, glandular-papillose. Fruit a berry (Freycinetia) or drupe (Pandanus), when drupaceous a multiple structure (syncarp, cephalium) composed of 2–20 woody, or basally fleshy or fibrous, cylindrical to globose, connate, free or often crowded and clustered carpels, each cluster of carpels termed a “phalange”, with mesocarp fibrous and often pithy or hollow above, fibrous and fleshy below; endocarp membranous or evanescent. Seeds 1-numerous, minute, often fusiform ; testa membranous (Pandanus) or crustose (Freycinetia); endosperm oily and fleshy, or starchy; strophiole originating from raphe sometimes present; embryo basal, minute; germination often epigeal.
Three genera and approximately 800 species: Old World tropics, several species introduced as economic or ornamental plants; 2 genera and 7 species (1 endemic) in China.
Stone, B.C. 1968. Materials for a monograph of Freycinetia Gaud. IV. Subdivision of the genus, with fifteen new sections. Blumea 16(2): 361–372. Stone, B.C. 1983. Pandanacées. In: Leroy, J.-F., ed., Flore du Cambodge du Laos & du Viêt-Nam 20: 3–48. Stone, B.C. 1988. Notes on the genus Pandanus (Pandanaceae) in Tahiti. Bot. Journ. Linn. Soc. (London) 97: 33–48. Zhou Lingyun and Zhong Xiongwen. 1992. Pandanaceae. In: Sun Xiangzhong, ed., Fl. Reipubl. Popularis Sin. 8: 12–23.
1a. Stems present, climbing by aerial roots, often also with long, pendulous aerial roots not utilized in climbing; prop roots absent; male spadices usually 2–5, rarely solitary, simple, sometimes fleshy; connective of stamen not apiculate; filaments papillose; ovary 1-celled; carpels multiovulate; fruit a berry 1. Freycinetia
1b. Stems present or absent;
often with aerial roots or prop roots or both; male spadices solitary,
branched, thin, leathery; connective of stamen apiculate; filaments smooth;
ovary 1-to 12-celled; carpels always
1-ovulate; fruit a drupe divided into phalanges 2. Pandanus
藤露兜树属 teng lou dou shu shu
Scrambling or climbing shrubs, rarely herbs; aerial roots present. Leaves ± densely crowded, linear or lanceolate, basally with membranous, marcescent or caducous sheath-margins (auricles). Inflorescences terminal or lateral, consisting of peduncled simple spadices (usually 2–5), arranged in an umbel or short raceme and initially enclosed by several, crowded, 3-seriate, green or colored, caducous spathes. Flowers unisexual, rarely bisexual; perianth absent. Stamens densely congested on rachis. Ovaries densely congested on rachis, usually surrounded by minute staminodes, 1-celled; ovules numerous, congested on 2 or more parietal placentas; stigmas 2 or more, separate or confluent. Fruit a berry (baccate), with or without a hardened apex. Seeds numerous.
Approximately 175 species: Sri Lanka to SE Asia, China, Australia, New Zealand and Oceania; 1 species in China.
山露兜 shan lou dou
Freycinetia batanensis Martelli; F. formosana f. typica Kimura; F. williamsii sensu auct. mult. chin., non Merrill (1908).
Scandent or scrambling liana; stems procumbent or prostrate, to 10 m or longer. Branches with aerial roots, glabrous, 0.5–5 cm wide. Leaves leathery, elongate, linear-lanceolate, 10–60 (–200) × 0.8–4 cm, the apex acuminate, the base narrowed, sharply antrorsely serrate toward the tip and the base, sometimes entire in upper part, with scattered spines along midvein beneath. Spadices 2–4, racemose, on stout peduncle ca. 5 cm long; spathes yellow. Female peduncles 10–15 × 1.4–1.8 cm; spathes (bracts) to 6 cm long. Multiple fruit (berry) globose, cylindrical or ovoid, 2.5–11 × 1.5–2 cm, composed of numerous dense drupes, the (individual) drupes ovoid or irregularly angled with asymmetrical edges, ca. 0.5–1 × 0.4–1 cm. Seed oblongoid.
Thickets and cliffs near seashores. Taiwan (Taitung: Lan Yu Is. (Botel Tobago I.), Keelung, Taipei) [Philippines (Batan I.), Ryukyu Is.].
Stone (1968) observed that the Philippine taxon F. batanensis is a “mere” variety, “if not in fact synonym”, of F. formosana, and that F. batanensis (Sect. Taiwaniella) has berry non-rostrate whereas F. williamsii Merrill, also from Philippines (Sect. Devriesella) has berry rostrate. Hui-Lin Li in Hui-Lin Li & al., eds., Flora of Taiwan 5: 819 (1978) and other workers recognize F. williamsii as occurring in Taiwan on the basis of its included synonym F. batanensis, whereas we follow Stone (1968) in considering F. batanensis a synonym of F. formosana. According to Fl. Taiwan and Fl. Reipubl. Pop. Sin., F. formosana has leaves 60–90 × 2–3 cm and globose multiple fruits, while F. williamsii is smaller, with leaves 10–20 × 0.8–1.2 cm and ovoid multiple fruits.
露兜树属 lou dou shu shu
Dioecious, evergreen trees or shrubs. Stems simple or branched, erect or prostrate, often with stiltlike, verrucose prop roots and aerial roots, sometimes virtually absent, often suckering. Leaves terminal, sessile, densely arranged in corkscrew spirals, 3-or 4-seriate, simple, linear, often spinose-serrate on margin and on keeled midvein beneath. Inflorescence of large, bracteate heads or spadices; perianth absent. Male inflorescence paniculate with spiciform branches subtended by secondary, usually colored, bracts (spathes), the branches covered with numerous stamens; flowers not individually distinguishable. Female inflorescence of globose to cylindrical clusters or several-carpelled aggregates; flowers not individually distinguishable; carpels 1-ovuled; staminodes absent in female flower; placentation subbasal. Multiple fruit (syncarp) a hard drupaceous capitulum, comprising an aggregation of individual connate, angled, fibrous (pericarp) phalanges (“drupes”); mesocarp sometimes hollow; exocarp fleshy; endocarp woody or bonelike; locules 1 or more; phalanges separating at maturity; stigma persistent, capitate. Seed solitary.
Approximately 600 species: Old World tropics; 6 species (1 endemic) in China.
Pandanus utilis Bory, native to Madagascar and cultivated in China, unlike most other species is both non-suckering and has reddish purplish marginal spines (cf. Fl. RPR 8: 23. 1992; Check List of Hong Kong Plants 297. 2002). P. boninensis Warburg, native to Japan, is also cultivated in China#1b.(loc. cit. Fl. RPR). P. utilis and P. boninensis are cultivated at the Taiwan Forestry Research Institute, as is P. pygmaeus Thouars of the Mascarenes (pigmy plant ca. 40 cm).
1a. Herbs.
2a. Mature plants virtually sterile; leaf-apex with 2 distinct lateral pleats and dense antrorse spines (prickles) ca. 1 mm along the pleats ....................................................................... 1. P. amaryllifolius
2b. Mature plants fertile; leaf-apex not laterally pleated and densely spinose.
3a. Leaves linear, ca. 4–5 cm wide ...................................................... 2. P. austrosinensis
3b. Leaves narrowly linear, ca. 1.5 cm wide .................................................. 3. P. fibrosus
1b. Trees or shrubs.
4a. Plants not suckering; leaves often glaucous beneath; phalanges (drupes) 4–12-locular; persistent stigma simple, not bifurcated ............................................................................... 5. P. tectorius
4b. Plants suckering; leaves not glaucous beneath; phalanges (drupes) 1–3(or 4)-locular; persistent stigma simple or bifurcated.
5a. Phalanges (drupes) 2-to 3-locular; persistent stigma not bifurcated ............... 4. P. kaida
5b. Phalanges (drupes) usually 1-locular; persistent stigma bifurcated .......... 6. P. urophyllus
??? xiang lu dou
Pandanus latifolius Hasskarl (1842), non Perrot (1825) & auct. mult.; P. odorus Ridley.
Herbs evergreen. Stems branched; aerial roots present. Sucker shoots of small growth phase with stems slender, to 1–1.6 m × ca. 2–5 cm, decumbent and ascending. Leaves somewhat glaucous beneath, keeled beneath but unarmed, the apex with distinct twin lateral pleats, the margin entire except at apex, there with very few minute prickles <1 mm long; blades 25–75 × 2–5 cm. Flowers unknown, probably never produced in small growth phase. Large growth phase with stems 2–4.5 m × 15 cm, erect, not or only sparsely branched. Leaves broadly linear, apex acute, glaucous beneath, keeled beneath and with twin lateral pleats with prickles same as in small growth phase; blades to 150–220 × 7–9 cm. Female inflorescence unknown in large growth phase. Male inflorescence (evidently exceedingly rare) probably pendent, to 60 cm; spathes ca. 90 cm; spikes oblong spikes, to 35 cm or more, the upper ones much shorter, ca. 9–10 × 2 cm, of numerous crowded, flat staminal phalanges 1.5–2.5 mm wide; stamens mostly 3–6 per phalange, filament 0.5–1.5 × 0.4–0.6 mm; anther oblong, ca. 2.5 × 0.5 mm, apex bluntly convex, without or with a barely discernible apiculum. .
Cultivated. Hainan (Xing Long, Dan Xian) [Indonesia].
Small growth phase, “perpetuated by continual harvesting of its leaves” (Stone, 1978), is cultivated for the musky aromatic leaves used in cooking, e.g., for scenting rice. Hainan [Indonesia, Malaysia, Philippines, West Irian (New Guinea)].
Warburg, O. 1900. Pandanaceae. In: Engler, A., Das Pflanzenreich 3. Heft. (IV. 9): 87, and Backer, C.A. and R.C. Bakhuizen van den Brink. 1968. Flora of Java 3: 205, noted that this species is known only from sterile (vegetative) specimens. A male flowering collection from a “large growth phase” in Ternate (Moluccas) was later described by Stone (1978). The description denoting two growth phases is from: Stone, B.C. 1978. Studies in Malesian Pandanaceae XVII. On the taxonomy of “pandan wangi”: a Pandanus cultivar with scented leaves. Economic Botany 32(3): 285–293.
露兜草 lou dou cao
Pandanus austrosinensis var. longifolius L.Y. Zhou & X.W. Zhong.
Herbs evergreen. Stems underground branched, with numerous adventitious roots, stems terrestrial unbranched. Leaves linear, subleathery, to 5 m × ca. 5 cm, apex acuminate into triangular, with small serrated flagelliform tip, margin with hamate spine, midrib raised abaxially, sparsely spinose. Male inflorescence of spikes to 10 cm long; stamens mostly 6, filaments connate in a fascicle below, free above, ca. 3.2 mm, inserted into peduncles, umbraculiform; anthers linear, ca. 3 mm, basifixed, introrse, 2-celled, longitudinally dehiscing, midrib with carina abaxially and densely spinulose. Carpels numerous, free at the apex, adnate to nearby carpels at the base. Ovary 1-loculed; ovule 1; styles short; stigma entire or furcate. Multiple fruits ellipsoid-orbicular or suborbicular, ca. 10–18 × 5–12 cm, consisting of ca. 250–300 drupes, pericarp of mature fruits becoming fibrous; drupes obconical, 5-to 6-ridged, ca. 3.3 × 1.2 cm; persistent stigma spinose. Fl. Apr–May.
* Forests, streamsides or roadsides. Guangdong, Hainan (Jian Feng Ling), Guangxi.
小露兜 xiao lou dou
Pandanus gressittii B.C. Stone, Journ. Arnold Arb. 43: 348. 1962.
Herbs evergreen, or small shrubs. Stems branched. Leaves narrowly linear, to 62 × ca. 1.5 cm, margin and midrib beneath spinose. Male inflorescences spicate, branched, 2–5 cm long; spathes 3.5–14 × 0.6–2 cm, margin above the middle spinulose or not; stamens 10–16, in filaments bundles 7 mm long, the free filaments at apex of filament bundles ca. 2 mm long; anthers oblongoid, ca. 10 × 7 mm, small cusp of apex connective ca. 1.2 mm long. Female inflorescences capitate, oblongoid, ca. 3 × ca. 1.2 cm; spathes 10–24 × 1–4 cm, margin sparsely spinose or not; carpel 1, 1-loculed; ovule 1, subbasal; stigma unbranched, ligulate, abaxially smooth, adaxially coarse. Multiple fruits ellipsoid or orbicular, ca. 6 × 3 cm, consisting of numerous drupes; drupes obconical, free after ripening, ca. 1.2 × 0.2–0.3 cm; persistent stigma spiny, simple. Fl. Apr–May.
Streamsides in forests, riversides. Hainan, Taiwan [Vietnam].
Stone (1983) considers P. gressittii a synonym of P. fibrosus.
勒古子 le gu zi
Pandanus forceps Martelli (type: Hong Kong).
Shrubs or small trees. Stems branched, 1–3 m tall; aerial roots absent. Leaves linear, ca. 1 (3–40 m × 3–5 cm, the apex acuminate, with long flagelliform tip, margin and midvein spiny. Male inflorescences consisting of several spikes, each spike ca. 10 cm long, spathes in lower part ca. 45 × 4.5 cm; stamens usually 10, few male flowers with more than 20 stamens; filaments adnate beneath, 7 mm long, umbraculiform, free filaments ca. 1 mm long; anthers narrowly ellipsoid, ca. 3 × 0.7 mm, apex with small cusp ca.0.5 mm long. Female inflorescences capitate, paniculate, ca. 4 × 1.7 cm; spathes numerous, 14–20 × 2–3 cm; stigma short, with 2 serrated branches; carpels 2–3 connate in a bundle, 2–3-loculed; ovule 1 per locule. Multiple fruits ellipsoid, consisting of 150 phalanges; phalanges obconical, ca. 3 × 1.7 cm, the projected part 5-angled above; persistent stigmas 2, branched, serrulate. Fl. May–Jun.
Seasides, stream banks, sunny low-lying areas, forests. Guangdong, Hainan, Hong Kong (Tai Tam, Tai Mo Shan), Kowloon, Kwangtung, Nodoa, Ooi Dick, Sunning, White Cloud Mt. [Vietnam].
Stone (1983) considers P. forceps a synonym of P. kaida. Plants sometimes used for hedges or living fences; young shoots edible. Stone (Ceylon. Journ. Sci. (Bio. Sci.) 11 (2): 119. 1975) notes that P. kaida is currently known only in cultivation.
露兜树 lou dou shu
Pandanus fascicularis Lamarck; Pandanus odoratissimus Linnaeus fil. var. sinensis (Warburg) Martelli (1930) [P. odoratissimus var. sinensis (Warburg) Kanehira (1936), nom. superfl.]; P. remotus St. John; P. sinensis (Warburg) Martelli; P. tectorius var. sinensis Warburg (1900).
Tree or shrub 3-c.10 m. Stem erect or ascending, many-branched, non-suckering; prop roots present or absent; numerous aerial roots often present. Leaves linear-ensiform, green, often glaucous beneath, spinose-serrate on margin and midvein beneath, to 180 × 10 cm, abruptly long-acuminate at apex. Male inflorescence to 60 cm, pedunculate, paniculately compound with ultimate spiciform branches; bracts 13–18, narrowly lanceolate, white, 10–60 × 1.5–4 cm, serrate on margin and midvein; spikes pendulous, 8–20 mm; stamens 10 (–25), racemosely fasciculate; filaments connate below, 1–3 mm; anthers linear, 3 × 0.6 mm, connective mucronate. Female inflorescence capitate, solitary, globose to ovoid-ellipsoid, 10–25 cm, with numerous white spathes 15–30 × 1.4–2.5 cm, the margin serrate; ovule 1 per locule. Fruit (syncarp) pendulous, orbicular or oblongoid, to 17 × 15 cm; segments (phalanges, “drupes”) 40–80 per aggregate-head, each phalange comprising c.5–12 fused obconical carpels (= locules, “cells”, drupes) connate below and free above, fragrant, yellow to orange to yellowish brown from base to apex, 4–10 × 1–6.5 cm; carpel-apices distinct, flattened or truncate (angled), tuberculate; persistent stigma subsessile, slightly protruded. Fl. Jan–May, Fr. Oct.
Seashores, sandy beaches; sometimes used as a living fence, or leaves for weaving. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hong Kong, Kowloon, Lantao I., Macao, Taipei, Taiwan, Yunnan [India, Asia, tropical Australia, Polynesia].
According to FRPR 8 (1992), var. sinensis (a variety not accepted in this treatment on the authority of Stone, 1983) occurs on seaside beaches in Guangdong, Guangxi, Hainan and Taiwan, and differs from the typical variety in having leaves narrower, attenuate into a long flagelliform apex, to 15 cm long; ovary (4–)5–6(–7)-loculed; fruits smaller, orbicular, ca. 8 × 8 cm, consisting of 50–60 drupe bundles, each drupe bundle ca. 2.5 × ca. 2 cm. Stone (1983) considered var. sinensis, with phalanges 3.5 × 4 cm, a synonym of P. tectorius, whose spectrum of phalange dimensions is 3–8 × 2.5–4.5 cm. Correct name of this taxon has long been controversial (cf. Martelli, U. 1929. “Pandanus odoratissimus” o “Pandanus tectorius”? Nuovo Giorn. Bot. Ital., n.s. 36: 329–337). Stone (1988) reinstated P. tectorius, reversing his former (Stone, 1983) acceptance of P. odoratissimus. The type specimen of P. remotus St. John, a name synonymized by Stone (1983), is from Hong Kong (Shek-o Bay).
Stone (1988) noted “variegated-leaf mutants of P. tectorius, known in horticulture variously as P. sanderi and P. veitchii…(which can back-mutate to all green leaf forms).” The mutant (cultivar) P. sanderi is cultivated as an ornamental in Hong Kong (Wu Te-lin (ed.-in-chief). 2002. Check List of Hong Kong Plants 297). Leaves of Pandanus sanderi Hort. Sander ex M.T. Masters have longitudinal yellow or golden marginal stripes. P. veitchii Hort. Veitch ex M.T. Masters & T. Moore has white or silvery marginal stripes; it is cultivated at the Taiwan Forestry Research Institute.
分叉露兜 fen cha lou dou
Pandanus furcatus sensu auct. mult. chin. & Warburg (1910), non Roxburgh ex Roxburgh, Hort. Bengal. 71. 1814.
Trees 7–13 m tall. Stems simple or branched at apex; aerial roots present. Leaves clustered at apex of stem; blade linear, leathery, 1–4.2 m × 2–11 cm, apex incurved, flagelliform and triangularly caudate, margin densely serrate with incurved spines, abaxially sparsely spinose along midrib. Male inflorescences consisting of several spikes; spikes golden yellow, terete, 10–15 cm long; spathes ca. 1 m × 10 cm; male flowers numerous; stamens (anthers) 3–14 per bundle, aggregated at the apex of stalked filaments, anthers linear, ca. 5 mm long, apex of connective with long, curved aristiform cusp. Female inflorescence pendulous on peduncle to 80 cm long, capitate, to 80 × 3 cm, with ca. 15 spathes; female flowers with 1 carpel, rarely 2, stigma bifurcate and curved. Fruit solitary or in racemes, to 23 × 12 cm, oblongoid to ellipsoid, reddish brown or orange-red; drupes connate, obconical-pyramidal, exocarp fleshy, sweet; styles bony, spiny, projected part of the apex pyramidal, 1–2-locular; persistent stigma bi-or 3-furcate. Fl. Aug.
Riversides, streamsides of forests, and cultivated as hedges and living fences. Canton, Guangdong, Guangxi, Hong Kong (Tai Po Kan, Tai Mo Shan, Lantau I.), Kowloon, Linchow, Lofausian, Taomoshan, Swatow, Xizang, Yunnan [Sikkim to Indochina; Vietnam].
Leaves used to make raincoats. Roots are medicinal (Fl. RPR), but details are lacking. Maintained as a species distinct from P. furcatus by Stone (1983).