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Flora Malesiana Nepenthaceae Flora Malesiana Series I - Seed Plants Volume 15 - 2001 Nepenthaceae Martin Cheek & Matthew Jebb ISBN 90-71236-49-8 All rights reserved © 2001 FoundationFlora Malesiana No the this be in part of material protected by copyright notice may reproduced or utilized any electronic form or by any means, or mechanical, including photocopying, recording, or by any and retrieval without written the information storage system, permission from copyright owner. Abstract Flora Malesiana. Series I, Volume 15 (2001) iv + 1—157, published by the Nationaal Herbarium Nederland, Universiteit Leiden branch, The Netherlands, under the aus- pices of FoundationFlora Malesiana. ISBN 90-71236-49-8 for i.e. the Contains the taxonomicrevision ofone family, Nepenthaceae, Malesia, area covering the countries Indonesia, Malaysia, Brunei Darussalam, Singapore, the Philip- pines, and Papua New Guinea. Martin Cheek & Matthew Jebb, Nepenthaceae, pp. 1—157*. A palaeotropical family of lianas, shrubs and herbs, with a single genus, Nepenthes. three There are 83 species of the family in the Malesian area, including nothospecies and one little known species. Most of the species are cultivated and traded across the value. in world as ornamental plants with curiosity Locally Malesia, some species are used for cooking specialist rice dishes, for medicinal uses or for making rope. habitat and ecol- The introductory part consists of chapters on distribution, fossils, ogy, reproductive biology, morphology and anatomy, pitcher function, cytotaxonomy, and characters. conservation, taxonomy, uses, collecting notes, spot Regional keys to the species are given. These are based largely on vegetative charac- ters. distribution, notes Foreach species full references, synonymy, descriptions, ecology, on diagnostic characters and relationships withother species are presented. Species are arranged alphabetically. This treatment is illustrated with 19 full-page line drawings. Index to scientific plant names of taxa treated in this volume (accepted names and synonyms) on pp. 159—161. Lists of revised families in Flora Malesiana on pp. 163—164. * Desk-editing: C.G.G. Baak; Digital imaging: B.N. Kieft Flora Malesiana,Series I, Volume 15 (2001) 1-157 Nepenthaceae (Martin Cheek, Royal Botanic Gardens Kew, Richmond, Surrey, England, and Matthew Jebb, National Botanic Gardens, Glasnevin, Dublin 9, Ireland) Nepenthaceae Dumort., Anal. fam. pi. (1829) 16; Hook.f. in A.DC., Prodr. 17 (1873) 90; Benth. & Hook.f., Gen. PI. 3 (1880) 1; Macfarl. in Engl., Pflanzenr. 4, 3 (1908) 2; Danser, Bull. Jard. Bot. Buitenzorg III, 9 (1928) 251; Jebb & Cheek, Blumea 42 (1997) 5. The below does not include characters A single genus of c. 87 species. description seen outside Malesia. NEPENTHES Nepenthes L., Sp. Pl. (1753) 955; Jebb & Cheek, Blumea 42 (1997) 5. — Type species: Nepenthes distillatoria L. Bandura Adans., Fam. 2 (1763) 75. — Type species: Nepenthes distillatoria L. Phyllamphora Lour., Fl. Cochin. 2 (1790) 606. — Type species: Nepenthes mirabilis (Lour.) Druce. Anurosperma(Hook.f.) Hallier f., Bot. Centralbl. 39, Abt. 2, 1 (1921) 162. —Type species: Nepenthes pervillei Blume. terrestrial Carnivorous, dioecious, woody or subwoody climbers or subshrubs, or Buds scales. epiphytic. Stems terete or 2—4-angled, or winged. naked, lacking Phyllo- involute taxy spiral, 2/5 or 1/2. Leaves exstipulate; or convolute, marcescent, simple, chartaceous or coriaceous, petiolate or sessile; midrib extended into an unbranched tendril, the distal part of which expands into an elaborate, animal trapping receptacle (pitcher) containing digestive fluid. Pitchers dimorphic. Lower pitchers produced from rosettes or short stems, oftenresting on the ground, usually ovoid or globular, the mouth facing towards the stem, with two ventral fringed wings running from the base to the pitcher rim and with the tendril straight, not coiled. Upper pitchers (usually absent in N. argentii, N. ampullaria, and N. pectinata) usually more elongated and infundibuli- the form(funnel-shaped) than the lower pitchers, the mouth facing away from the stem, coiled. Pitcher wings reduced to ridges and not fimbriate, or absent, and the tendril mouth apical or subapical, rimmed with a ribbed peristome (except N. inermis), the inner edge often toothedand bearing nectar glands, at the rear sometimesraised to form the lid held the mouth in N. a column, supporting lid; usually over (reflexed e.g. dubia), surface with flattened basal filiform lower a laterally appendage, rarely an apical ap- pendage, or appendages absent; nectar glands usually abundant; spur insertedon dorsal surface at junction with lid, entire or variously divided, flattened or terete. Inflores- cence terminal, appearing lateralby subsequent growth, a paniculoid thyrse or raceme, of 6-300 flowers, the main axis with indeterminategrowth, the partial inflorescences 1-flowered(racemose) or 2-flowered, less usually 3—40-flowered, bracteate or ebrac- free teate. Perianthimbricate, a single whorl of 4 (also interpreted as two whorls of 2) flowers androecium lack- or basally united, patent, nectariferous tepals. Female with ing, ovary superior, 4-carpellate, incompletely 4-locular locules antitepalous, placenta- tion lamellar, ovules erect, anatropous, bitegmic, crassinucellate, 200-500, stigmas floriferous, sessile, as many as locules. Male inflorescence usually larger and more 2 Flora Malesiana, Ser. I, Vol. 15 (2001) united into anthers perianth as in female; stamens 4—?12, filaments an androphore, tetrasporangiate, in 1-3 dense whorls, united into a subspherical anther head, locules Fruit sometimes opening by longitudinal slits, extrorse; gynoecium lacking. stipitate, a loculicidally dehiscent capsule with 4 valves containing 50-500 seeds. Seeds filiform, 3-25 mm long, slender due to long basal and apical appendages. Testa reduced to an the outer epidermis with thick outer walls to the cells and irregular thickenings on radial and inner walls. Tegmen produced only around the embryonic cavity, crushed. Endosperm starchy or absent. Embryo minute, central, straight or U-shaped in a sub- ellipsoid cavity, cotyledons and hypocotyl well-developed, though minute. Indumen- tum of simple, bifid, fasciculate, stellate, dendritic non-glandular and sunken, sessile or shortly stipitate glandular hairs. Colour of pitchers entirely green, or yellow or orange or white or purple or red, often marked with red streaks or blotches; peristome often glossy red; inflorescence with green, brown or red tepals. 2n = 80. — Fig. 1-19. DISTRIBUTION Genus of c. 87 species, mostly in Malesia, but with c. 8 outlying species in: Mada- Sri Lanka NE India Indochina Solomon Is- gascar (2), Seychelles (1), (1), (1), (6-9), found and lands (1), New Caledonia (1) and Australia (1). Most species are in Borneo from the Sunda Islands of from Sumatra. No species are known Lesser east Java, nor the northern tip of Sumatra. FOSSILS Krutzsch (1985) identifiedas Nepenthes fossil pollen types known since the 1930s from the European Tertiary as the Droseridites echinosporus group. This group had pre- viously been referred to the Droseraceae which also have spinuliferous pollen pre- dominantly in tetrahedraltetrads. Three species are involved. They occur in the Palaeo- cene fromFrance to Ukraine: Nepenthes echinatus (Hunger) Krutzsch, N. echinosporus (R. Potonie) Krutzsch and N. major (Krutzsch) Krutzsch. However, Krutzsch's identi- fications be without since the tetrads of 40 seem not to ambiguity N. major are over microns in diameter, falling outside of the range of those in present-day Nepenthes (24-35 microns after acetolysis according to Basak & Subramanyam (1966)) and in- side of fossil the lower range of that seen in modern Drosera. We know no non-pollen Nepenthes. References: Basak, R.K. & K. Subramanyam, Pollen grains of some species of Nepenthes. Phyto- morphology 16, 3 (1966) 334-338.— Krutzsch, W., Uber Nepenthes-pollen (alias Droseridites p.p.) in europaischen Tertiar. Gleditschia 13 (1985) 1, 89-93. HABITAT AND ECOLOGY most Nepenthes are commonly encountered in disturbed secondary forest, swamp, kerangas or heath forest from sea level to c. 700 m altitude. In such habitats however, only a few species occur, although they can be locally common. Nepenthes ampullaria, N. rafflesiana, and N. gracilis can be found in these habitats in Peninsular Malaysia, Cheek & Jebb Nepenthaceae 3 Sumatra, and Borneo, while in New Guinea N. neoguineensis and N. ampullaria oc- cupy these niches. More locally distributedin such habitats are N. bicalcarata(Borneo), N. treubiana(W New Guinea), and N. sumatrana (WC Sumatra). Nepenthes mirabilis occurs from Indochina to N Australia, sometimes in such habitats, though it also occurs in Eucalyptus savannah and open fresh-water or brackish swamps. The shade of undisturbed high forest is anathema to most Nepenthes. The majority of Nepenthes species are confined to montane habitats between 1500-2500m altitude, usually in open mossy, stunted, ridge-top forest as climbers or epiphytes. A few species reach montane or subalpine grassland, for example N. lamii which reaches 3520 m alti- tude in Irian Jaya - the highest altitude known for the genus. Some species can occur in both lowland and montane habitats, for example N. alata, N. albomarginata, N. eustachya, N. maxima, N. merrilliana, N. neoguineensis, and N. reinwardtiana, although most species occur in one or the other. Limestone and ultramafic substrates often support open scrub or woodland and are habitats for some Nepenthes. limestone are known
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