Overview

Brief Summary

Nepenthes is a genus of tropical pitcher plants, ranging from Southeast Asia, its center of diversity, westward into the Seychelles and eastern Madagascar and south to Australia. Nepenthes is the most diverse group of carnivorous plants to have evolved sophisticated pitcher traps and the only genus in the family Nepenthaceae. As in other carnivorous plants, the adaptive value of carnivory is thought to lie in the acquisition of nitrogen in nitrogen-poor environments.

Pitchers grow on tendrils extending from the midribs of leaves and trap prey passively, collecting pools of water into which prey are lured (with bright colors and nectar secreted on the pitcher rim), drowned, and digested with no energetic active movement on the part of the plant. In some species a single plant may grow some pitchers that lie recumbent on the substrate and others that hang suspended in mid-air; this results in leaf dimorphism, in which ground pitchers are different in shape, size, and appearance from aerial pitchers. Though most prey are small nectarivorous insects, the largest Nepenthes species may produce pitchers capable of trapping small vertebrates such as lizards, rodents, and birds. A few species have evolved modifications of the prototypical pitcher morphology and behavior to collect leaf litter (N. ampullaria) or vertebrate droppings (N. lorii, N. rajah, N. macrophylla, and N. rafflesiana).

Phytotelmata, the pools collected in Nepenthes pitchers, provide unique habitats that can support not only opportunistic species but entire faunal communities. These unusual and specialized communities are analogous to the phytotelmata collected by tree holes and New World bromeliads. Not surprisingly, Nepenthes species potentially form many mutualistic and commensalistic interactions with animals in their native ranges. The small organisms associated with Nepenthes traps are known as nepenthephiles, and may consist of both opportunistic and specialized inhabitants; the nature of their relationships with other nepenthephiles and with the plant, and the costs and benefits for all participants is often unclear. The water itself contains a community of protozoa, invertebrates, and even tadpoles that feed on excess prey, undigested prey remains, or each other, and may aid the plant in digestion.

The small mutualistic ant Camponotus (Colobopsis) schmitzi lives in the hollowed-out leaf tendrils of N. calcarata, enhancing the plant's capture ability by protecting developing traps from herbivores, maintaining the slippery interior of the pitchers, and even swimming into the pools to subdue large prey. In exchange, the ants are provided with living space, prey, and nectar. Coprophagous (dung-eating) species lure small arboreal mammals with copious nectar or shelter; while feeding or roosting, the animals defecate or urinate into the pitcher. Small frogs, land crabs, and spiders may also take advantage of the insects and shelter afforded by the pitcher. Some Nepenthes species are known as 'monkey cups' because monkeys are known to drink the water in the pitchers.

Human interest in Nepenthes ranges from the utilitarian to the aesthetic. Its unique carnivorous habit and unusual and varied forms made the genus an object of fascination by early naturalists and a fashionable but difficult plant to rear in captivity, and the culture and study of Nepenthes by an active community of enthusiasts continues. The highly slippery wax surfaces of the pitcher interior have also inspired modern attempts at engineering similar materials. Today many species, some of which are already scarce and occur only in a few localities, are threatened with extinction due to habitat loss and deterioration.

  • Slack, A. 1979. Carnivorous Plants. Cambridge: The MIT Press,. 74-87. Print.
  • Hewitt-Cooper, Nigel, Nigel. "A case of bird capture by a cultivated specimen of the hybrid Nepenthes X mixta." Carnivorous Plant Newsletter. 41.1 (2012): 31. Web. 20 May. 2013. .
  • "Killer plant" BBC News. British Broadcasting Corporation, 5 Aug 2011. Web. 20 May 2013. .
  • Phillipps, A. " A Second Record of Rats as Prey in Nepenthes rajah." Carnivorous Plant Newsletter. 17.2 (1988): 55. Web. .
  • Greenwood, M., Charles Clarke, Ch'ien C. Lee, et al. "A Unique Resource Mutualism between the Giant Bornean Pitcher Plant, Nepenthes rajah, and Members of a Small Mammal Community." PLOS ONE. 6.6 (2011): n. page. Web. .
  • Pavlovic, Andrej, Ľudmila Slovakova, and Jiri Santrucek. "Nutritional benefit from leaf litter utilization in the pitcher plant Nepenthes ampullaria." Plant, Cell & Environment. 34.11 (2011): 1865-1873. Web. 20 May. 2013. .
  • Clarke, Charles, Jonathan A. Moran, and Lijin Chin. "Mutualism between tree shrews and pitcher plants." Plant Signaling & Behavior. 5.10 (2010): 1187-1189. Web. 20 May. 2013. .
  • Moran, Jonathan A., Charles Clarke, Melinda Greenwood, et al. "Tuning of color contrast signals to visual sensitivity maxima of tree shrews by three Bornean highland Nepenthes species." Plant Signaling & Behavior. 7.10 (2012): 1267-1270. Web. .
  • Rembold, Katja, Eberhard Fischer, Boris F. Striffler, et al. "Crab spider association with the Malagasy pitcher plant Nepenthes madagascariensis." African Journal of Ecology. 51.1 (2012): 188-191. Web. .
  • Grafe, T. Ulmar, Caroline R. Schöner, Gerald Kerth, et al. "A novel resource–service mutualism between bats and pitcher plants." Biology Letters. 7.3 (2011): 436-439. Web. 20 May. 2013. .
  • Thornham, Daniel G., Joanna M. Smith, T. Ulmar Grafe, et al. "Setting the trap: cleaning behaviour of Camponotus schmitzi ants increases long-term capture efficiency of their pitcher plant host, Nepenthes bicalcarata." Functional Ecology. 26.1 (2012): 11-19. Web. .
  • Moran, Jonathan A., Gilles Le Moguedec, David J. Marshall, et al. "A Carnivorous Plant Fed by Its Ant Symbiont: A Unique Multi-Faceted Nutritional Mutualism." PLOS ONE. 7.5 (2012): n. page. Web. 20 May. 2013. .
  • Wong, Tak-Sing, Sung Hoon Kang, Sindy K. Y. Tang, et al. "Bioinspired self-repairing slippery surfaces with pressure-stable omniphobicity." Nature. 477.7365 (2011): 443-447. Web. 20 May. 2013. .
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Physical Description

Type Information

Isolectotype for Nepenthes junghuhnii Macfarl. ex Ridl.
Catalog Number: US 1371178
Collection: Smithsonian Institution, National Museum of Natural History, Department of Botany
Verification Degree: ; Original publication and alleged type specimen examined
Preparation: Pressed specimen
Collector(s): H. Robinson & C. Kloss
Year Collected: 1914
Locality: Lorinchi Peak., Sumatra Island, Indonesia, Asia-Tropical
Elevation (m): 2652 to 2652
  • Isolectotype: Ridley, H. N. 1917. J. Fed. Malay States Mus. 8 (4): 79.; Cheek, M. & Jebb, M. 1997. Blumea. 42: 80.
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Evolution and Systematics

Functional Adaptations

Functional adaptation

Pitchers prevent fungal growth: pitcher plant
 

The pitchers of pitcher plants prevent fungal growth using napthoquinones.

   
  "Nepenthes spp. are carnivorous plants that have developed insect  capturing traps, evolved by specific modification of the leaf  tips, and are able to utilize insect degradation products as  nutritional precursors. A chitin-induced antifungal ability,  based on the production and secretion to the trap liquid of droserone  and 5-O-methyldroserone, is described here. Such specific secretion  uniquely occurred when chitin injection was used as the  eliciting agent and probably reflects a certain kind of defence  mechanism that has been evolved for protecting the carnivory-based  provision of nutritional precursors. The pitcher liquid  containing droserone and 5-O-methyldroserone at 3:1 or  4:1 molar ratio, as well as the purified naphthoquinones,  exerted an antifungal effect on a wide range of plant and  human fungal pathogens. When tested against Candida  and Aspergillus spp., the concentrations required for  achieving inhibitory and fungicidal effects were significantly  lower than those causing cytotoxicity in cells of the human  embryonic kidney cell line, 293T. These naturally secreted  1,4-naphthoquinone derivatives, that are assumed to act via  semiquinone enhancement of free radical production, may offer  a new lead to develop alternative antifungal drugs with  reduced selectable pressure for potentially evolved resistance." (Eilenberg et al. 2010:911)

Note: According to the paper, the exact mode of function of napthoquinones is unknown.

  Learn more about this functional adaptation.
  • Eilenberg H; Pnini-Cohen S; Rahamim Y; Sionov E; Segal E; Carmeli S; Zilberstein A. 2010. Induced production of antifungal naphthoquinones in the pitchers of the carnivorous plant Nepenthes khasiana. Journal of Experimental Botany. 61(3): 911-922.
  • 2010. From carnivorous plants to the medicine cabinet? Anti-fungal agents in pitcher plants investigated. Science Daily [Internet],
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Functional adaptation

Relationship provides nutrients: pitcher plant
 

The large pitcher of Nepenthes pitcher plants gathers nitrogen from tree shrews through a mutualistic relationship.

     
 

"Botanists have discovered that the giant montane pitcher plant of  Borneo has a pitcher the exact same size as a tree shrew's body.

 

"But  it is not this big to swallow up mammals such as tree shrews or rats.

 

"Instead,  the pitcher uses tasty nectar to attract tree shrews, then ensures its  pitcher is big enough to collect the feeding mammal's droppings

 

"That suggests these supposedly 'meat-eating' plants have evolved a  mutualistic relationship with tree shrews.

 

"The tree shrews get  nectar, a valuable food source, and in return, the plants get to catch  and absorb the tree shrew's faeces which likely supplies the majority of  nitrogen required by the plant." (Walker 2010)


  Learn more about this functional adaptation.
  • Chin L; Moran JA; Clarke C. 2010. Trap geometry in three giant montane pitcher plant species from Borneo is a function of tree shrew body size. New Phytologist. 186(2): 461-470.
  • Walker M. 2010. Giant meat-eating plants prefer to eat tree shrew poo. BBC Earth News [Internet],
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Functional adaptation

Walls keep insect feet from sticking: pitcherplants
 

The internal walls of pitcherplants prevent insects from escaping by clogging their feet with a flaky, waxy substance and being rough.

   
  "The most elaborate of all these leaves turned into pit-fall traps are developed by the plants that are known, simply and without qualification, as pitcher plants…Their trapping strategy is the same as the trumpet pitchers. They entice insects with fragrant nectar. The walls of their traps are made even more treacherous by a flaky waxy surface that peels off and clogs the feet of insects so that they lose all chance of adhesion. As their victims tumble into the water and start to struggle to save themselves, the disturbance stimulates glands in the pitcher walls which start to discharge a digestive acid. This is so powerful that a fly will be reduced to a hollow shell within days and a midge will disappear entirely within hours. The whole device is so effective that these pitchers can trap not just small insects, but cockroaches, centipedes and scorpions. The rajah is said to be able to consume mice." (Attenborough 1995:77-78)

"[Abstract] Pitcher plants of the genus Nepenthes efficiently trap and retain insect prey in highly specialized leaves. Besides a slippery peristome which inhibits adhesion of insects they employ epicuticular wax crystals on the inner walls of the conductive zone of the pitchers to hamper insect attachment by adhesive devices. It has been proposed that the detachment of individual crystals and the resulting contamination of adhesive organs is responsible for capturing insects. However, our results provide evidence in favour of a different mechanism, mainly based on the stability and the roughness of the waxy surface. First, we were unable to detect a large quantity of crystal fragments on the pads of insects detached from mature pitcher surfaces of Nepenthes alata. Second, investigation of the pitcher surface by focused ion beam treatment showed that the wax crystals form a compact 3D structure. Third, atomic force microscopy of the platelet-shaped crystals revealed that the crystals are mechanically stable, rendering crystal detachment by insect pads unlikely. Fourth, the surface profile parameters of the wax layer showed striking similarities to those of polishing paper with low grain size. By measuring friction forces of insects on this artificial surface we demonstrate that microscopic roughness alone is sufficient to minimize insect attachment. A theoretical model shows that surface roughness within a certain length scale will prevent adhesion by being too rough for adhesive pads but not rough enough for claws." [from Conclusion] In conclusion we cannot rule out the possibility that to a certain degree breaking of extensively protruding single wax crystals may occur and consequently be found as residues attached to the tarsi. Presumably this would increase the anti-adhesive effect of the surface of N. alata. Furthermore, breaking of crystals for catching prey may occur in different species of pitcher plants or even individuals of N. alata under certain circumstances or environmental conditions. However, the results presented clearly indicate that the structural properties of the mechanically very stabile epicuticular wax alone are sufficient to prevent insect adhesion to the conductive zone of N. alata. (Scholz et al. 2010:1115,1125)


  Learn more about this functional adaptation.
  • Attenborough, D. 1995. The Private Life of Plants: A Natural History of Plant Behavior. London: BBC Books. 320 p.
  • Scholz I; Bückins M; Dolge L; Erlinghagen T; Weth A; Hischen F; Mayer J; Hoffmann S; Riederer M; Riedel M; Baumgartner W. 2010. Slippery surfaces of pitcher plants: Nepenthes wax crystals minimize insect attachment via microscopic surface roughness. Journal of Experimental Biology. 213(7): 1115-1125.
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Wikipedia

Nepenthes junghuhnii

Nepenthes junghuhnii (/nɨˈpɛnθz jʊŋˈhni./; after Friedrich Franz Wilhelm Junghuhn, who collected it between 1840 and 1842) is a tropical pitcher plant native to Sumatra. This species has been the source of much confusion since its discovery. The taxon originally named N. junghuhnii by John Muirhead Macfarlane has never been formally published (Henry Nicholas Ridley's N. junghuhnii is a heterotypic synonym of N. singalana).[1][2] In 1994, taxonomist Jan Schlauer described N. junghuhnii as a "rather dubious species based on insufficient specimens".[3] Nepenthes junghuhnii sensu Macfarlane has not been relocated in the wild since the collection of the type specimen.[1] It is characterised by strongly petiolate leaves and appears to be most closely related to N. bongso and N. spathulata;[1] Schlauer considers it a possible synonym of the former.[2]

Herbarium specimens informally named N. junghuhnii by Macfarlane were collected by Junghuhn in the Batak region of North Sumatra, near Lake Toba.[1] The specimens of N. singalana that Ridley used to describe N. junghuhnii originated from a collection by Harold Ernest Robinson and Cecil Boden Kloss on Mount Kerinci, in the Sumatran province of Jambi.[1]

In his 1928 monograph, "The Nepenthaceae of the Netherlands Indies", B. H. Danser wrote of this taxon:[4]

N. Junghuhnii is based upon plants, collected by Junghuhn in Sumatra, named as such by Macfarlane in the Kew Herbarium and seen by Ridley, and with which the latter author identified specimens from G. Kerintji. I have not seen any of these plants and therefore I am not quite certain about them; however, in H. B. there are other plants, collected by Junghuhn in Sumatra and named N. Junghuhnii by Macfarlane, and it seems evident, that these plants may be identical with those in the Kew Herbarium. About the specimens from G. Kerintji, which, according to Ridley, are "undoubtedly the same species", I dare, of course, not say the same. However, the two numbers in H. B. named by Macfarlane seem to be not wholly identical. The first of the above-mentioned plants is a form, certainly intermediate between N. sanguinea and N. singalana. The coarse stems and the large leaves are those of N. sanguinea, the pitchers are smaller than usually are those of N. sanguinea, their shape approaches that of the pitchers of N. singalana and the inflorescences are quite like those of the latter species. The second plant mentioned above is perhaps a piece of an identical plant, though it shows other characters. It is a fragment of the lower portion of a climbing stem; this stem and its leaves are like those of the first mentioned plant, but the pitchers differ from those of N. sanguinea only by the toothed interior margin of the peristome. When the latter character had been absent, I should not have hesitated to record N. sanguinea as occurring in Sumatra; also the provenance of the two mentioned plants from probably the same habitat has kept me from doing this. Of course it is possible, that both plants of Junghuhn represent a new species, but since the material at hand shows only characters intermediate between two species already known and between many related species intermediate forms have been found, I prefer to distinguish also these plants as such. It is very improbable that it is a hybrid, since pure N. sanguinea has not been found in Sumatra, and from the region where Junghuhn collected his plants N. singalana too has not been recorded. As is obvious from a letter written by Macfarlane and extant in the Leiden Herbarium this author intended to make a description of N. Junghuhnii for Koorders to be published in his Plantae Junghuhnianae lneditae, but apparently it had not come to that. The publication of Ridley, not written in Latin, is invalid from a nomenclatorial point of view.

Danser treated N. junghuhnii as a possible natural hybrid between N. singalana and N. sanguinea.[4] The latter species is now known to be absent from Sumatra.[1]

References[edit]

  1. ^ a b c d e f Clarke, C.M. 2001. Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu.
  2. ^ a b Schlauer, J. N.d. Nepenthes junghuhnii. Carnivorous Plant Database.
  3. ^ Schlauer, J. 1994. NEPENTHES comments for Flora Malesiana. Carnivorous Plant Mailing List, September 9, 1994.
  4. ^ a b Danser, B.H. 1928. 40. Nepenthes sanguinea LINDL.. In: The Nepenthaceae of the Netherlands Indies. Bulletin du Jardin Botanique de Buitenzorg, Série III, 9(3-4): 249-438.
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Nepenthes mirabilis

"Nepenthes phyllamphora" redirects here. For other uses, see Nepenthes phyllamphora (disambiguation).

Nepenthes mirabilis (/nɨˈpɛnθz mɪˈræbɨlɪs/; from Latin: mirabilis = wonderful), or the Common Swamp Pitcher-Plant,[11] is a carnivorous plant species. By far the most widespread of all Nepenthes, its range covers continental Southeast Asia and all major islands of the Malay Archipelago (minus the Lesser Sunda Islands and northern Philippines), stretching from China in the north to Australia in the south.[12] The species exhibits great variability throughout its range. One of the more notable varieties, N. mirabilis var. echinostoma, is endemic to Brunei and Sarawak and possesses an extremely wide peristome.[13]

The conservation status of N. mirabilis is listed as Least Concern on the 2006 IUCN Red List of Threatened Species.[14] In Hong Kong, it is a protected species under Forestry Regulations Cap. 96A.

According to Matthew Jebb and Martin Cheek, the pitchers of N. mirabilis are used as toy phallocrypts in New Guinea.

Distribution[edit]

Nepenthes mirabilis has by far the widest distribution of any Nepenthes species and is known from the following countries and regions: Australia (Cape York Peninsula), Borneo, Cambodia, Caroline Islands (Palau and Yap), China (Guangdong Province, Hainan,[15][16] Hong Kong, and Macau), D'Entrecasteaux Islands, Java, Laos, Louisiade Archipelago, Maluku Islands, Myanmar, New Guinea, Peninsular Malaysia, Philippines (Dinagat and Mindanao), Sulawesi, Sumatra, Thailand, and Vietnam.[12][17][18][19][20][21][22][23] It has also been recorded from many smaller islands, including Babi, Bangka, Banyak Islands, Batu Islands, Bengkalis, Enggano, Ko Lanta, Ko Tarutao, Langkawi, Mendol, Mentawai Islands (North Pagai, Siberut, Sipura, and South Pagai), Meranti Islands (Padang, Rangsang, and Tebing Tinggi), Nias, Penang, Phuket, Riau Islands (Lingga Islands and Riau Archipelago), Rupat,[24] Tawi-Tawi,[25] and possibly Wowoni.[26]

Taxonomy[edit]

Nepenthes mirabilis is closely related to N. rowanae and N. tenax, the only two Nepenthes species endemic to Australia.

Differences between N. mirabilis and N. rowanae (Clarke & Kruger, 2005)
CharacterN. mirabilisN. rowanae
Morphology of leaf bladeAcute to roundedContracted towards the apex, then continuing along the tendril as a narrow, acute, extension
Insertion of tendril to leaf bladeSimplePeltate
Pitcher wingsSimple, bearing multicellular fringe elementsOften flattened at front, forming a T-shape in XS, multicellular fringe elements often present
Leaf blade textureUsually chartaceousStrongly coriaceous
Leaf blade attachment to stemSimple, or rarely decurrent for ⅓ the length of the internodeDecurrent for at least ½ the length of the internode, usually more
Gland density in lower portion of pitcher1600-2500 / cm²Approximately 3600 / cm²
Position of pitcher hip in upper pitchersMid-way, to lower halfUpper quarter
Position of pitcher hip in lower pitchersLower third to quarterImmediately beneath peristome

In his Carnivorous Plant Database, taxonomist Jan Schlauer treats N. kongkandana as a synonym of N. mirabilis.[27]

Infraspecific taxa[edit]

Across its range, N. mirabilis exhibits great variability in terms of pitcher morphology and colour, and it has the most synonyms of all Nepenthes species.[28] The following forms and varieties of N. mirabilis have been described. With the exception of N. mirabilis var. echinostoma and N. mirabilis var. globosa, these taxa are not considered valid today.

N. mirabilis var. echinostoma[edit]

An upper pitcher of N. mirabilis var. echinostoma

Nepenthes mirabilis var. echinostoma was discovered by Odoardo Beccari in 1865 and described as a species, N. echinostoma, by Joseph Dalton Hooker in 1873.[11] In 1882, Frederick William Burbidge described this unusual variety in The Gardeners' Chronicle as follows:[11][30]

Beccari's singular N. echinostoma (vide Herb. Kew) is a wonderful thing, as yet unintroduced—indeed, I suppose unseen by any save Beccari ! The mouths of the urns remind one of the deflexed teeth of some gigantic moss of the Hypnoid section.

Nepenthes mirabilis var. echinostoma is the only form of this species that occurs in Brunei. It has also been recorded from parts of Sarawak, but appears to be completely absent from Sabah.[11]

N. mirabilis var. globosa[edit]

A lower pitcher of a cultivated N. mirabilis var. globosa
Nepenthes mirabilis pitchers being used to cook rice by the Jarai people of Kon Tum Province, Vietnam

Nepenthes mirabilis var. globosa has been recorded from a single undisclosed Andaman Sea island off Phang Nga and from the Thai mainland near the city of Trang.[12][29][31]

This variety was featured on the cover of the January 2006 issue of the Journal of Insectivorous Plant Society, identified as "Nepenthes sp. from Thailand".[32] The name Nepenthes globosa appeared in print in an article by Shigeo Kurata in the July 2007 issue of the Journal of Insectivorous Plant Society.[8] The same issue also featured an article by Masahiro Tada that referred to the plant as "Nepenthes Viking".[10] Prior to its description by Marcello Catalano in 2010, this taxon was also published under the informal name Nepenthes sp. Phanga Nga in Stewart McPherson's 2009 book, Pitcher Plants of the Old World.[12]

In the horticultural trade, this variety is popularly known as both Nepenthes globosa[12] (/nɨˈpɛnθz ɡlˈbzə/; from Latin: globosus, "globular") and Nepenthes sp. Viking,[12][33] after the resemblance the pitchers bear to the prow of a Viking ship.[34]

Pitcher infauna[edit]

A great number of infaunal organisms have been found in the pitchers of this species. These include the sarcophagid fly Sarcophaga papuensis and the mite Nepenthacarus warreni, which have both been found in Australian populations of the plant.[35][36] Similarly, the mosquitoes Aedes dybasi and Aedes maehleri reside in the pitchers of N. mirabilis on the islands of Palau and Yap, respectively.[37] Both have unusual life histories and morphological traits associated with this habit.[38][39]

The nematode Baujardia mirabilis has been described from N. mirabilis in Thailand. It is not thought to be an accidental; the pitchers of this species appear to be the nematode's natural habitat. The microecosystems in these pitchers were found to be dominated by mosquito larvae, midges, and B. mirabilis. It is speculated that this nematode might have a phoretic relationship with one or more infaunal insect species.[40]

In southern China, tree frogs have been observed in the pitchers of N. mirabilis. The amphibians do not fall prey to the plant, but rather feed on insects that are caught by the pitchers.[41] They are not affected by the acidic digestive juices (which may have a pH as low as 2), likely due to the mucilagenous outer layer of their skin.[42]

The first record of an aquatic fungus living in the pitcher organ of a carnivorous plant came from a specimen of N. mirabilis growing along the Jardine River in Australia. The mycelial fungus was observerd as both free-living in the trap's fluid and attached to chitinous insect remains.[43][44]

The pitchers of N. mirabilis have also been found to harbour a complex community of bacteria.[45]

Natural hybrids[edit]

Nepenthes mirabilis has the greatest number of known natural hybrids of any species in the genus.[12]

Notes[edit]

a.^ Nepenthes mirabilis was first described under the Linnaean taxonomic system as Phyllamphora mirabilis by João de Loureiro in 1790. It was then transferred to the genus Nepenthes under the incorrect combination Nepenthes phyllamphora by Carl Ludwig Willdenow in 1805. Most sources[12][13][48][51][61] attribute the earliest publication of the correct binomial—Nepenthes mirabilis—to George Claridge Druce in 1916,[62] but Jan Schlauer's Carnivorous Plant Database records a much earlier publication by Rafarin in 1869.[63]

References[edit]

  1. ^ Danser, B.H. 1928. 1. Nepenthes alata Blanco. [pp. 258–262] In: The Nepenthaceae of the Netherlands Indies. Bulletin du Jardin Botanique de Buitenzorg, Série III, 9(3–4): 249–438.
  2. ^ Cheek, M. & M. Jebb 2013. Typification and redelimitation of Nepenthes alata with notes on the N. alata group, and N. negros sp. nov. from the Philippines. Nordic Journal of Botany 31(5): 616–622. doi:10.1111/j.1756-1051.2012.00099.x
  3. ^ Clarke, C. & C.C. Lee 2012. A revision of Nepenthes (Nepenthaceae) from Gunung Tahan, Peninsular Malaysia. Gardens' Bulletin Singapore 64(1): 33–49.
  4. ^ Schlauer, J. N.d. Nepenthes alata. Carnivorous Plant Database.
  5. ^ Bailey, F.M. 1881. On a new species of Nepenthes. The Proceedings of the Linnean Society of New South Wales 5: 185–187.
  6. ^ a b (Latin) Hooker, J.D. 1873. Ordo CLXXV bis. Nepenthaceæ. In: A. de Candolle Prodromus Systematis Naturalis Regni Vegetabilis 17: 90–105.
  7. ^ a b c d (Latin) Blume, C.L. 1852. Ord. Nepenthaceae. In: Museum Botanicum Lugduno-Batavum, sive stirpium exoticarum novarum vel minus cognitarum ex vivis aut siccis brevis expositio. Tom. II. Nr. 1. E.J. Brill, Lugduni-Batavorum. pp. 5–10.
  8. ^ a b (Japanese) Kurata, S. 2007. ネペンテス ・ ノート (XIV) — Nepenthes globosa のネーミングについて. Journal of Insectivorous Plant Society 58(3): 76.
  9. ^ Macfarlane, J.M. 1908. Nepenthaceae. In: A. Engler. Das Pflanzenreich IV, III, Heft 36: 1–91.
  10. ^ a b (Japanese) Tada, M. 2007. 日本における Nepenthes Viking の渡来と現状について. Journal of Insectivorous Plant Society 58(3): 75.
  11. ^ a b c d Phillipps, A. & A. Lamb 1996. Pitcher-Plants of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
  12. ^ a b c d e f g h i j k l m n McPherson, S.R. 2009. Pitcher Plants of the Old World. 2 volumes. Redfern Natural History Productions, Poole.
  13. ^ a b c d e f g Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
  14. ^ Clarke et al. (2000). Nepenthes mirabilis. 2006. IUCN Red List of Threatened Species. IUCN 2006. www.iucnredlist.org. Retrieved on 12 May 2006.
  15. ^ Nepenthes mirabilis. Flora of China.
  16. ^ (Chinese) Yang G., Huang S., Xu S. & Gao Y. 2006. 海南岛野生猪笼草资源调查及其营养成分分析. [The investigation and study on the germplasm resources and growth of the wild common Nepenthes in Hainan.] Chinese Agricultural Science Bulletin 22(11): 440–442. Abstract
  17. ^ Slack, A. 1979. Nepenthes mirabilis. In: Carnivorous Plants. Ebury Press, London. pp. 79–80.
  18. ^ McPherson, S.R. & V.B. Amoroso 2011. Field Guide to the Pitcher Plants of the Philippines. Redfern Natural History Productions, Poole.
  19. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Borneo. Redfern Natural History Productions, Poole.
  20. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Sulawesi. Redfern Natural History Productions, Poole.
  21. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Peninsular Malaysia and Indochina. Redfern Natural History Productions, Poole.
  22. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Australia and New Guinea. Redfern Natural History Productions, Poole.
  23. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Sumatra and Java. Redfern Natural History Productions, Poole.
  24. ^ Clarke, C.M. 2001. Appendix C: Distribution Maps. In: Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu. pp. 299–307.
  25. ^ Cheek, M. & M. Jebb 2013. Recircumscription of the Nepenthes alata group (Caryophyllales: Nepenthaceae), in the Philippines, with four new species. European Journal of Taxonomy 69: 1–23. doi:10.5852/ejt.2013.69
  26. ^ (Indonesian) Sunarti, S., A. Hidayat & Rugayah 2008. Keanekaragaman tumbuhan di hutan Pegunungan Waworete, Kecamatan Wawonii Timur, Pulau Wawonii, Sulawesi Tenggara. [Plants diversity at the mountain forest of Waworete, East Wawonii District, Wawonii Island, Southeast Sulawesi.] Biodiversitas 9(3): 194–198.
  27. ^ Schlauer, J. N.d. Nepenthes kongkandana. Carnivorous Plant Database.
  28. ^ Catalano, M. 2009. Nepenthes. In: Growing Carnivores — an Italian perspective. Prague. pp. 50–57.
  29. ^ a b Catalano, M. 2010. Nepenthes mirabilis var. globosa M. Catal. var. nov. PDF In: Nepenthes della Thailandia: Diario di viaggio. Prague. p. 40.
  30. ^ Burbidge, F.W. 1882. Notes on the new Nepenthes. The Gardeners' Chronicle, new series, 17(420): 56.
  31. ^ Mey, F.S. 2014. 'Nepenthes of Indochina', my 2010 ICPS lecture now on Youtube. Strange Fruits: A Garden's Chronicle, February 3, 2014.
  32. ^ (Japanese) Cover of the January 2006 issue of the Journal of Insectivorous Plant Society. Insectivorous Plant Society.
  33. ^ Mey, F.S. 2011. Nepenthes mirabilis var. globosa, a popular plant?. Strange Fruits: A Garden's Chronicle, December 24, 2011.
  34. ^ Guerini, M. 2011. 2010: new species of Carnivorous Plants. PDF Associazione Italiana Piante Carnivore.
  35. ^ Yeates, D.K., H. de Souza Lopes & G.B. Monteith 1989. A commensal sarcophagid (Diptera: Sarcophagidae) in Nepenthes mirabilis (Nepenthaceae) pitchers in Australia. Australian Entomological Magazine 16: 33–39.
  36. ^ Fashing, N.J. 2002. Nepenthacarus, a new genus of Histiostomatidae (Acari: Astigmata) inhabiting the pitchers of Nepenthes mirabilis (Lour.) Druce in Far North Queensland, Australia. PDF (1.64 MB) Australian Journal of Entomology 41(1): 7–17. doi:10.1046/j.1440-6055.2002.00263.x
  37. ^ Sota, T. & M. Mogi 2006. Origin of pitcher plant mosquitoes in Aedes (Stegomyia): a molecular phylogenetic analysis using mitochondrial and nuclear gene sequences. Journal of Medical Entomology 43(5): 795–800. doi:10.1603/0022-2585(2006)43[795:OOPPMI]2.0.CO;2
  38. ^ Bohart, R.M. 1956. Insects of Micronesia. Diptera: Culicidae. PDF Insects Micronesia 12(1): 1–85.
  39. ^ Mogi, M. 2010. Unusual life history traits of Aedes (Stegomyia) mosquitoes (Diptera: Culicidae) inhabiting Nepenthes pitchers. Annals of the Entomological Society of America 103(4): 618–624. doi:10.1603/AN10028
  40. ^ Bert, W., I.T. De Ley, R. Van Driessche, H. Segers & P. De Ley 2003. Baujardia mirabilis gen. n., sp. n. from pitcher plants and its phylogenetic position within Panagrolaimidae (Nematoda: Rhabditida). PDF Nematology 5(3): 405–420. doi:10.1163/156854103769224395
  41. ^ Hua, Y. & H. Li 2005. Food web and fluid in pitchers of Nepenthes mirabilis in Zhuhai, China. PDF Acta Botanica Gallica 152(2): 165–175.
  42. ^ Hua, Y. & L. Kuizheng 2004. The Special Relationship Between Nepenthes and Tree Frogs. PDF Carnivorous Plant Newsletter 33(1): 23–24.
  43. ^ Cribbs, A.B. 1987. An aquatic fungus from pitchers of Nepenthes mirabilis. Queensland Naturalist 28: 72–73.
  44. ^ Schnell, D.E. 1992. Literature Review. Carnivorous Plant Newsletter 21(3): 80–82.
  45. ^ Yogiara, A. Suwanto & M.T. Suhartono 2006. A complex bacterial community living in pitcher plant fluid. Jurnal Mikrobiologi Indonesia 11(1): 9–14.
  46. ^ Lauffenburger, A. 1995. Guide to Nepenthes Hybrids. OmnisTerra.
  47. ^ Kurata, S. & M. Toyoshima 1972. Philippine species of Nepenthes. The Gardens' Bulletin Singapore 26(1): 155–158. Abstract
  48. ^ a b Cheek, M.R. & M.H.P. Jebb 2001. Nepenthaceae. Flora Malesiana 15: 1–157.
  49. ^ Lowrie, A. 1983. Sabah Nepenthes Expeditions 1982 & 1983. PDF (1.25 MB) Carnivorous Plant Newsletter 12(4): 88–95.
  50. ^ a b c d (Italian) Catalano, M. 2010. Nepenthes della Thailandia: Diario di viaggio. Prague.
  51. ^ a b c Clarke, C.M. 2001. Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu.
  52. ^ Fretwell, S. 2013. Back in Borneo to see giant Nepenthes. Part 2: Mt Tambuyukon and Poring. Victorian Carnivorous Plant Society Journal 108: 6–15.
  53. ^ Rischer, H. 1995. Observations on the Nepenthes species of Irian Jaya, Part I: Nepenthes insignis Danser. PDF (461 KB) Carnivorous Plant Newsletter 24(3): 75–77.
  54. ^ Lee, C.C. 2007. Re: The most accessible limestone hill of Bau. Carnivorous Plants in the tropics.
  55. ^ a b c Phillipps, A., A. Lamb & C.C. Lee 2008. Pitcher Plants of Borneo. Second Edition. Natural History Publications (Borneo), Kota Kinabalu.
  56. ^ Clarke, C.M. & R. Kruger 2005. Nepenthes rowanae (Nepenthaceae), a remarkable species from Cape York, Australia. Carnivorous Plant Newsletter 34(2): 36–41.
  57. ^ a b Mey, F.S., L.H. Truong, D.V. Dai & A.S. Robinson 2011. Nepenthes thorelii, an emended description and novel ecological data resulting from its rediscovery in Tay Ninh, Vietnam. In: McPherson, S.R. New Nepenthes: Volume One. Redfern Natural History Productions, Poole. pp. 104–131.
  58. ^ Bednar, B. 1983. Nepenthes mirabilis variation. PDF (111 KB) Carnivorous Plant Newsletter 12(3): 64.
  59. ^ N. thorelii rediscovered. Carnivorous Plants in the tropics.
  60. ^ Mey, F.S. 2012. Virtual Nepenthes herbarium on "Europeana". Strange Fruits: A Garden's Chronicle, December 19, 2012.
  61. ^ Danser, B.H. 1928. The Nepenthaceae of the Netherlands Indies. Bulletin du Jardin Botanique de Buitenzorg, Série III, 9(3–4): 249–438.
  62. ^ Druce, G. 1916. Nepenthes mirabilis. In: Botanical Exchange Club of the British Isles Report 4: 637.
  63. ^ Schlauer, J. N.d. Nepenthes mirabilis. Carnivorous Plant Database.

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Nepenthes

For the drug of forgetfulness mentioned in Greek mythology, see Nepenthe.

Nepenthes (/nɨˈpɛnθz/), popularly known as tropical pitcher plants or monkey cups, is a genus of carnivorous plants in the monotypic family Nepenthaceae. The genus comprises roughly 150 species, and numerous natural and many cultivated hybrids. They are mostly liana-forming plants of the Old World tropics, ranging from South China, Indonesia, Malaysia and the Philippines; westward to Madagascar (two species) and the Seychelles (one); southward to Australia (three) and New Caledonia (one); and northward to India (one) and Sri Lanka (one). The greatest diversity occurs on Borneo, Sumatra, and the Philippines, with many endemic species. Many are plants of hot, humid, lowland areas, but the majority are tropical montane plants, receiving warm days but cool to cold, humid nights year round. A few are considered tropical alpine, with cool days and nights near freezing. The name "monkey cups" refers to the fact that monkeys have been observed drinking rainwater from these plants.

Description[edit]

Nepenthes species usually consist of a shallow root system and a prostrate or climbing stem, often several metres long and up to 15 m (49 ft) or more, and usually 1 cm (0.4 in) or less in diameter, although this may be thicker in a few species (e.g. N. bicalcarata). From the stems arise alternate, sword-shaped leaves with entire leaf margins. An extension of the midrib (the tendril), which in some species aids in climbing, protrudes from the tip of the leaf; at the end of the tendril the pitcher forms. The pitcher starts as a small bud and gradually expands to form a globe- or tube-shaped trap.[3]

Basic structure of an upper pitcher

The trap contains a fluid of the plant's own production, which may be watery or syrupy, and is used to drown the prey. Research has shown this fluid contains viscoelastic biopolymers that may be crucial to the retention of insects within the traps of many species. The trapping efficiency of this fluid remains high, even when significantly diluted by water, as inevitably happens in wet conditions.[4]

The lower part of the trap contains glands which absorb nutrients from captured prey. Along the upper inside part of the trap is a slick, waxy coating which makes the escape of its prey nearly impossible. Surrounding the entrance to the trap is a structure called the peristome (the "lip") which is slippery and often quite colorful, attracting prey, but offering an unsure footing. Above the peristome is a lid (the operculum); in many species, this keeps rain from diluting the fluid within the pitcher, the underside of which may contain nectar glands which attract prey.[3]

Nepenthes species usually produce two types of pitchers, known as leaf dimorphism. Appearing near the base of the plant are the large lower traps, which typically sit on the ground. The upper or aerial pitchers are usually smaller, coloured differently, and possess different features from the lower pitchers. These upper pitchers usually form as the plant reaches maturity and the plant grows taller. To keep the plant steady, the upper pitchers often form a loop in the tendril, allowing it to wrap around nearby support. In some species (e.g. N. rafflesiana), different prey may be attracted by the two types of pitchers. This varied morphology also often makes identification of species difficult.[3]

Prey usually consists of insects, but the largest species (e.g. N. rajah and N. rafflesiana) may occasionally catch small vertebrates, such as rats and lizards.[5][6] There are even records of cultivated plants trapping small birds.[7][8] Flowers occur in racemes or more rarely in panicles with male and female flowers on separate plants. They are insect-pollinated, the primary agents being flies (including blow flies, midges, and mosquitoes), moths, wasps, and butterflies.[9] Their smells can range from sweet to musty or fungus-like.[10] Seed is typically produced in a four-sided capsule which may contain 50–500 wind-distributed seeds, consisting of a central embryo and two wings, one on either side (though see N. pervillei).

The genus is cytologically diploid, with all studied species having a chromosome number of 2n=80.[11][12] This high number is thought to reflect paleopolyploidy (likely 8x or 16x).[12][13][14][15]

Distribution and habitat[edit]

Global distribution of Nepenthes

The genus Nepenthes is mostly found within the Malay Archipelago, with the greatest biodiversity found on Borneo, Sumatra, and the Philippines, especially in the Borneo montane rain forests. The full range of the genus includes Madagascar (N. madagascariensis and N. masoalensis), the Seychelles (N. pervillei), Sri Lanka (N. distillatoria), and India (N. khasiana) in the west to Australia (N. mirabilis, N. rowanae, and N. tenax) and New Caledonia (N. vieillardii) in the southeast. Most species are restricted to very small ranges, including some only found on individual mountains. These limited distributions and the inaccessibility of the regions often means some species go decades without being rediscovered in the wild (e.g. N. deaniana, which was rediscovered 100 years after its initial discovery). About 10 species have population distributions larger than a single island or group of smaller islands. Nepenthes mirabilis has the distinction of being the most widely distributed species in the genus, ranging from Indochina and throughout the Malay Archipelago to Australia.[3][16][17]

Because of the nature of the habitats which Nepenthes species occupy, they are often graded as either lowland or highland species, depending on their altitude above sea level, with 1,200 m (3,937 ft) the rough delineation between lowland and highland. Species growing at lower altitudes require continuously warm climates with little difference between day and night temperatures, whereas highland species thrive when they receive warm days and much cooler nights. Nepenthes lamii grows at a higher altitude than any other in the genus, up to 3,520 m (11,549 ft).[3][17]

Most Nepenthes species grow in environments that provide high humidity and precipitation and moderate to high light levels. A few species, including N. ampullaria, prefer the dense, shaded forests, but most other species thrive on the margins of tree/shrub communities or clearings. Some species (e.g. N. mirabilis) have been found growing in clear-cut forest areas, roadsides, and disturbed fields. Other species have adapted to growing in savanna-like grass communities. The soils in which Nepenthes species grow are usually acidic and low in nutrients, being composed of peat, white sand, sandstone, or volcanic soils. Exceptions to these generalities include species that thrive in soils with high heavy metal content (e.g. N. rajah), on sandy beaches in the sea spray zone (e.g. N. albomarginata). Other species grow on inselbergs and as lithophytes, while others, such as N. inermis, can grow as epiphytes with no soil contact.[3]

Ecological relationships[edit]

A drowned lizard found in a freshly opened pitcher of N. rajah

The most obvious interaction between Nepenthes species and their environments, including other organisms, is that of predator and prey. Nepenthes species certainly attract and kill their prey, albeit passively, through active production of attractive colours, sugary nectar, and even sweet scents. From this relationship, the plants primarily gain nitrogen and phosphorus to supplement their nutrient requirements for growth, given these soil nutrients are typically lacking. The most frequent prey is an abundant and diverse group of arthropods, with ants and other insects topping the menu. Other arthropods found frequently include spiders, scorpions, and centipedes, while snails and frogs are more unusual, but not unheard of. The most uncommon prey for Nepenthes species includes rats found in N. rajah. The composition of prey captured depends on many factors, including location, but can incorporate hundreds of individual insects and many different species.[3] While many Nepenthes species are generalists in what they capture, at least one, N. albomarginata, has specialised and almost exclusively traps termites and produces nearly no nectar. Nepenthes albomarginata gains its name from the ring of white trichomes directly beneath the peristome. These trichomes—or "hairs"—are palatable to termites and will attract them to the pitcher. In the course of collecting the edible trichomes, hundreds or thousands of termites will fall into the pitcher.[18][19]

Prey retention by viscoelastic pitcher fluid of N. rafflesiana[4]
The blue bottle fly (Calliphora vomitoria) can escape after landing in water on its ventral surface. 
The same is true if the fly falls in dorsally (wings-first). 
But the viscoelastic properties of N. rafflesiana digestive fluid prevent prey escape, whether the fall is ventral.. 
..or dorsal. (All videos recorded at 500 frames/s) 

Symbioses[edit]

A lower pitcher of N. attenboroughii supporting a large population of mosquito larvae. The upright lid of this species exposes its pitchers to the elements such that they are often completely filled with fluid.[20]

Nepenthes bicalcarata provides space in the hollow tendrils of its upper pitchers for the carpenter ant Camponotus schmitzi to build nests. The ants take larger prey from the pitchers, which may benefit N. bicalcarata by reducing the amount of putrefaction of collected organic matter that could harm the natural community of infaunal species that aid the plant's digestion.[21]

Nepenthes lowii has also formed a dependent relationship, but with vertebrates instead of insects. The pitchers of N. lowii provide a sugary exudate reward on the reflexed pitcher lid (operculum) and a perch for tree shrew species, which have been found eating the exudate and defecating into the pitcher. A 2009 study, which coined the term "tree shrew lavatories", determined between 57 and 100% of the plant's foliar nitrogen uptake comes from the faeces of tree shrews.[22] Another study showed the shape and size of the pitcher orifice of N. lowii exactly match the dimensions of a typical tree shrew (Tupaia montana).[23][24] A similar adaptation was found in N. macrophylla and N. rajah, and is also likely to be present in N. ephippiata.[24]

Infauna[edit]

Main article: Nepenthes infauna

Organisms that spend at least part of their lives within the pitchers of Nepenthes species are often called Nepenthes infauna. The most common infaunal species, often representing the top trophic level of the infaunal ecosystem, are many species of mosquito larvae. Other infaunal species include fly and midge larvae, spiders, mites, ants, and even a species of crab (Geosesarma malayanum). Many of these species specialise to one pitcher plant species and are found nowhere else. These specialists are called nepenthebionts. Others, often associated with but not dependent on Nepenthes species, are called nepenthophiles. Nepenthexenes, on the other hand, are rarely found in the pitchers, but will often appear when putrefaction approaches a certain threshold, attracting fly larvae that would normally not be found in the pitcher infaunal community. The complex ecological relationship between pitcher plants and infauna is not yet fully understood, but the relationship may be mutualistic: the infauna is given shelter, food, or protection, and the plant that harbours the infauna receives expedited breakdown of captured prey, increasing the rate of digestion and keeping harmful bacterial populations repressed.[21][25][26]

Botanical history[edit]

Plukenet's drawing of N. distillatoria from his Almagestum Botanicum of 1696.

The earliest known record of Nepenthes dates back to the 17th century. In 1658, French colonial governor Étienne de Flacourt published a description of a pitcher plant in his seminal work Histoire de la Grande Isle de Madagascar. It reads:[27]

It is a plant growing about 3 feet high which carries at the end of its leaves, which are 7 inches long, a hollow flower or fruit resembling a small vase, with its own lid, a wonderful sight. There are red ones and yellow ones, the yellow being the biggest. The inhabitants of this country are reluctant to pick the flowers, saying that if somebody does pick them in passing, it will not fail to rain that day. As to that, I and all the other Frenchmen did pick them, but it did not rain. After rain these flowers are full of water, each one containing a good half-glass. [translated from French in Pitcher-Plants of Borneo][10]

Flacourt called the plant Amramatico, after a local name. More than a century later, this species was formally described as N. madagascariensis.[28]

The second species to be described was N. distillatoria, the Sri Lankan endemic. In 1677, Danish physician Thomas Bartholin made brief mention of it under the name Miranda herba, Latin for "marvellous herb".[29] Three years later, Dutch merchant Jacob Breyne referred to this species as Bandura zingalensium, after a local name for the plant.[30] Bandura subsequently became the most commonly used name for the tropical pitcher plants, until Linnaeus coined Nepenthes in 1737.[10]

Nepenthes distillatoria was again described in 1683, this time by Swedish physician and naturalist Herman Niklas Grim.[31] Grim called it Planta mirabilis destillatoria or the "miraculous distilling plant", and was the first to clearly illustrate a tropical pitcher plant.[10] Three years later, in 1686, English naturalist John Ray quoted Grim as saying:[32]

The root draws up moisture from the earth which with the help of the sun's rays rises up into the plant itself and then flows down through the stems and nerves of the leaves into the natural utensil to be stored there until used for human needs. [translated from Latin in Pitcher-Plants of Borneo][10]

One of the earliest illustrations of Nepenthes appears in Leonard Plukenet's Almagestum Botanicum of 1696.[33] The plant, called Utricaria vegetabilis zeylanensium, is undoubtedly N. distillatoria.[10]

Cantharifera as illustrated in Rumphius's Herbarium Amboinensis, Volume 5, published in 1747, although probably drawn in the late 17th century. The vine on the right is not a Nepenthes, but a species of Flagellaria.

Around the same time, German botanist Georg Eberhard Rumphius discovered two new Nepenthes species in the Malay Archipelago. Rumphius illustrated the first one, now considered synonymous with N. mirabilis, and gave it the name Cantharifera, meaning "tankard-bearer". The second, referred to as Cantharifera alba, is thought to have been N. maxima. Rumphius described the plants in his most famous work, the six-volume Herbarium Amboinense, a catalogue of the flora of Ambon Island. However, it would not be published until many years after his death.[34]

After going blind in 1670, when the manuscript was only partially complete, Rumphius continued work on Herbarium Amboinensis with the help of clerks and artists. In 1687, with the project nearing completion, at least half of the illustrations were lost in a fire. Persevering, Rumphius and his helpers first completed the book in 1690. However, two years later, the ship carrying the manuscript to the Netherlands was attacked and sunk by the French, forcing them to start over from a copy that had fortunately been retained by Governor-General Johannes Camphuijs. The Herbarium Amboinensis finally arrived in the Netherlands in 1696. Even then, the first volume did not appear until 1741, 39 years after Rumphius's death. By this time, Linnaeus's name Nepenthes had become established.[10]

Illustration of Bandura zeylanica (N. distillatoria) from Burmann's Thesaurus Zeylanicus of 1737

Nepenthes distillatoria was again illustrated in Johannes Burmann's Thesaurus Zeylanicus of 1737. The drawing depicts the end of a flowering stem with pitchers. Burmann refers to the plant as Bandura zeylanica.[35]

The next mention of tropical pitcher plants was made in 1790, when Portuguese priest João de Loureiro described Phyllamphora mirabilis, or the "marvellous urn-shaped leaf", from Vietnam. Despite living in the country for around 35 years, it seems unlikely that Loureiro observed living plants of this species, as he stated the lid is a moving part, actively opening and closing. In his most celebrated work, Flora Cochinchinensis, he writes:[36]

[...] (the) leaf-tip ends in a long hanging tendril, twisted spirally in the middle, from which hangs a sort of vase, oblong, pot-bellied, with a smooth lip with a projecting margin and a lid affixed to one side, which of its own nature freely opens and closes in order to receive the dew and store it. A marvellous work of the Lord! [translated from French in Pitcher-Plants of Borneo][10]

Phyllamphora mirabilis was eventually transferred to the genus Nepenthes by Rafarin in 1869.[37] As such, P. mirabilis is the basionym of this most cosmopolitan of tropical pitcher plant species.[21]

Loureiro's description of a moving lid was repeated by Jean Louis Marie Poiret in 1797. Poiret described two of the four Nepenthes species known at the time: N. madagascariensis and N. distillatoria. He gave the former its current name and called the latter Nepente de l'Inde, or simply "Nepenthes of India", although this species is absent from the mainland. In Jean-Baptiste Lamarck's Encyclopédie Méthodique Botanique, he included the following account:[28]

This urn is hollow, as I have just said, usually full of soft, clear water, and then closed. It opens during the day and more than half the liquid disappears, but this loss is repaired during the night, and the next day the urn is full again and closed by its lid. This is its sustenance, and enough for more than one day because it is always about half-full at the approach of night. [translated from French in Pitcher-Plants of Borneo][10]

The Nepenthes house of the Veitch Nurseries as illustrated in The Gardeners' Chronicle, 1872

With the discovery of new species and Sir Joseph Banks' original introduction of specimens to Europe in 1789, interest in Nepenthes grew throughout the 19th century, culminating in what has been called the "Golden Age of Nepenthes" in the 1880s.[3][10] However, the popularity of the plants dwindled in the early 20th century, before all but disappearing by World War II. This is evidenced by the fact that no new species were described between 1940 and 1966. The revival of global interest in the cultivation and study of Nepenthes is credited to Japanese botanist Shigeo Kurata, whose work in the 1960s and 1970s did much to bring attention to these plants.[38]

Etymology[edit]

The name Nepenthes was first published in 1737 in Carolus Linnaeus's Hortus Cliffortianus.[39] It references a passage in Homer's Odyssey, in which the potion "Nepenthes pharmakon" is given to Helen by an Egyptian queen. "Nepenthe" literally means "without grief" (ne = not, penthos = grief) and, in Greek mythology, is a drug that quells all sorrows with forgetfulness.[10][40] Linnaeus explained:

If this is not Helen's Nepenthes, it certainly will be for all botanists. What botanist would not be filled with admiration if, after a long journey, he should find this wonderful plant. In his astonishment past ills would be forgotten when beholding this admirable work of the Creator! [translated from Latin by Harry Veitch][41]

The plant Linnaeus described was N. distillatoria, a species from Sri Lanka.[10]

Nepenthes was formally published as a generic name in 1753 in Linnaeus's famous Species Plantarum, which established botanical nomenclature as it exists today. Nepenthes distillatoria is the type species of the genus.[42]

Nepenthes from Carolus Linnaeus's Species Plantarum of 1753

The name "monkey cups" was discussed in the May 1964 issue of National Geographic, in which Paul A. Zahl wrote:[43]

The carriers called them "monkey cups," a name I had heard elsewhere in reference to Nepenthes, but the implication that monkeys drink the pitcher fluid seemed farfetched. I later proved it true. In Sarawak I found an orangutan that had been raised as a pet and later freed. As I approached it gingerly in the forest, I offered it a half-full pitcher. To my surprise, the ape accepted it and, with the finesse of a lady at tea, executed a delicate bottoms-up.

In Indonesia the plants are often called kantong semar or "Semar's pocket".

Cultivation[edit]

Cultivated Nepenthes rajah and other species

Nepenthes may be cultivated in greenhouses. Easier species include N. alata, N. ventricosa, N. khasiana, and N. sanguinea. These four species are highlanders (N. alata has both lowland and highland forms), some easy lowlander species are N. rafflesiana, N. bicalcarata, N. mirabilis, and N. hirsuta.[44]

Highland forms are those species that grow in habitats generally higher in elevation, and thus exposed to cooler evening temperatures. Lowland forms are those species growing nearer to sea level. Both forms respond best to rainwater (but some tap water works as long as it is flushed monthly with rainwater or water low in dissolved solid and chemicals), bright light (though some species can grow in full sun), a well-drained medium, good air circulation and relatively high humidity, although easier species such as N. alata can adapt to lower humidity environments. Highland species must have night-time cooling to thrive in the long term. Chemical fertilisers are best used at low strength. Occasional feeding with frozen (thawed before use) crickets may be beneficial. Terrarium culture of smaller plants, such as N. bellii, N. × trichocarpa and N. ampullaria, is possible, but most plants will get too large over time.[45][46]

Plants can be propagated by seed, cuttings, and tissue culture. Seeds are usually sown on damp chopped Sphagnum moss, or on sterile plant tissue culture media once they have been properly disinfected. The seeds generally become nonviable soon after harvesting, so seed are not usually the preferred method of propagation. A 1:1 mixture of orchid medium with moss or perlite has been used for germination and culture. Seed may take two months to germinate, and two years or more to yield mature plants. Cuttings may be rooted in damp Sphagnum moss in a plastic bag or tank with high humidity and moderate light. They can begin to root in one to two months and start to form pitchers in about six months. Tissue culture is now used commercially and helps reduce collection of wild plants, as well as making many rare species available to hobbyists at reasonable prices. Nepenthes species are considered threatened or endangered plants and are listed in CITES appendices 1 and 2.[45]

Species[edit]

Around 157 species of Nepenthes are currently recognised as valid. This number is quickly increasing, with several new species being described each year.[38]

Hybrids and cultivars[edit]

The complex man-made hybrid N. ventricosa × (N. lowii × N. macrophylla)

There are many hybrid Nepenthes and numerous named cultivars. Some of the more well-known, artificially produced hybrids and cultivars include:

  • N. × coccinea ((N. rafflesiana × N. ampullaria) × N. mirabilis)
  • N. × mixta (N. northiana × N. maxima)
  • N. 'Emmarene' (N. khasiana × N. ventricosa)
  • N. 'Judith Finn' (N. spathulata × N. veitchii)

See also[edit]

References[edit]

  1. ^ Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III" (PDF). Botanical Journal of the Linnean Society 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x. Retrieved 2013-07-06. 
  2. ^ Cheek, M. & M. Jebb 2013. The Nepenthes micramphora (Nepenthaceae) group, with two new species from Mindanao, Philippines. Phytotaxa 151(1): 25–34. doi:10.11646/phytotaxa.151.1.2
  3. ^ a b c d e f g h Barthlott, W., Porembski, S., Seine, R., and Theisen, I. 2007. The Curious World of Carnivorous Plants. Portland, Oregon: Timber Press.
  4. ^ a b Gaume, L. & Y. Forterre 2007. A viscoelastic deadly fluid in carnivorous pitcher plants. PLoS ONE 2(11): e1185. doi:10.1371/journal.pone.0001185
  5. ^ Phillipps, A. 1988. A second record of rats as prey in Nepenthes rajah. Carnivorous Plant Newsletter 17(2): 55.
  6. ^ Moran, J.A. 1991. The role and mechanism of Nepenthes rafflesiana pitchers as insect traps in Brunei. Ph.D. thesis, University of Aberdeen, Aberdeen, Scotland.
  7. ^ "Killer plant 'eats' great tit at Somerset nursery". BBC News. 5 August 2011. Retrieved 5 August 2011. 
  8. ^ Hewitt-Cooper, N. 2012. A case of bird capture by a cultivated specimen of the hybrid Nepenthes × mixta. Carnivorous Plant Newsletter 41(1): 31–33.
  9. ^ Clarke, C.M. 2001. Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu.
  10. ^ a b c d e f g h i j k l Phillipps, A. & A. Lamb 1996. Pitcher-Plants of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
  11. ^ Lowrey, T.K. 1991. No. 519: Chromosome and isozyme number in the Nepenthaceae. American Journal of Botany 78(6, supplement): 200–201.
  12. ^ a b Heubl, G.R. & A. Wistuba 1997. A cytological study of the genus Nepenthes L. (Nepenthaceae). Sendtnera 4: 169–174.
  13. ^ Meimberg, H. & G. Heubl 2006. Introduction of a nuclear marker for phylogenetic analysis of Nepenthaceae. Plant Biology 8(6): 831–840. doi:10.1055/s-2006-924676
  14. ^ Cytology of Nepenthes. LMU Department für Biologie.
  15. ^ Brittnacher, J. N.d. Evolution -- Nepenthes Phylogeny. International Carnivorous Plant Society.
  16. ^ McPherson, S.R. 2009. Pitcher Plants of the Old World. 2 volumes. Redfern Natural History Productions, Poole.
  17. ^ a b Jebb, M., and Cheek, M. 1997. A skeletal revision of Nepenthes (Nepenthaceae). Blumea, 42: 1-106.
  18. ^ Moran, J.A., M.A. Merbach, N.J. Livingston, C.M. Clarke & W.E. Booth 2001. Termite prey specialization in the pitcher plant Nepenthes albomarginata—evidence from stable isotope analysis. Annals of Botany 88: 307–311. doi:10.1006/anbo.2001.1460
  19. ^ Merbach, M.A., D.J. Merbach, U. Maschwitz, W.E. Booth, B. Fiala & G. Zizka 2002. Mass march of termites into the deadly trap. Nature 415: 36–37. doi:10.1038/415036a
  20. ^ Robinson, A.S., A.S. Fleischmann, S.R. McPherson, V.B. Heinrich, E.P. Gironella & C.Q. Peña 2009. A spectacular new species of Nepenthes L. (Nepenthaceae) pitcher plant from central Palawan, Philippines. Botanical Journal of the Linnean Society 159(2): 195–202. doi:10.1111/j.1095-8339.2008.00942.x
  21. ^ a b c Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
  22. ^ Clarke, C.M., U. Bauer, C.C. Lee, A.A. Tuen, K. Rembold & J.A. Moran 2009. Tree shrew lavatories: a novel nitrogen sequestration strategy in a tropical pitcher plant. Biology Letters 5(5): 632–635. doi:10.1098/rsbl.2009.0311
  23. ^ Chin, L., J.A. Moran & C. Clarke 2010. Trap geometry in three giant montane pitcher plant species from Borneo is a function of tree shrew body size. New Phytologist 186 (2): 461–470. doi:10.1111/j.1469-8137.2009.03166.x
  24. ^ a b Walker, M. 2010. Giant meat-eating plants prefer to eat tree shrew poo. BBC Earth News, March 10, 2010.
  25. ^ Mogi, M. & H.S. Yong 1992. Aquatic arthropod communities in Nepenthes pitchers: the role of niche differentiation, aggregation, predation and competition in community organization. Oecologia 90: 172–184. doi:10.1007/BF00317174
  26. ^ Beaver, R.A. 1979. Fauna and foodwebs of pitcher plants in west Malaysia. Malayan Nature Journal 33: 1–10.
  27. ^ de Flacourt, É. 1658. Histoire de la Grande Isle de Madagascar.
  28. ^ a b Poiret, J.L.M. 1797. Népente. In: J.B. Lamarck Encyclopédie Méthodique Botanique Vol. 4.
  29. ^ Bartholinus 1677. Miranda herba. Acta Medica et Philosophica Hafniensia 3: 38.
  30. ^ Breyne, J. 1680. Bandura zingalensium etc. Prodromus Fasciculi Rariorum Plantarum 1: 18.
  31. ^ Grimm, H.N. 1683. Planta mirabilis destillatoria. In: Miscellanea curiosa sive Ephemeridum. Med. Phys. Germ. Acad. Nat. Cur. Decuriae 2, ann. prim. p. 363, f. 27.
  32. ^ Ray, J. 1686. Bandura cingalensium etc. Historia Plantarum 1: 721–722.
  33. ^ Plukenet, L. 1696. Utricaria vegetabilis zeylanensium. In: Almagestum Botanicum.
  34. ^ Rumphius, G.E. 1741–1750. Cantharifera. In: Herbarium Amboinense 5, lib. 7, cap. 61, p. 121, t. 59, t. 2.
  35. ^ Burmann, J. 1737. Thesaurus Zeylanicus. Amsterdam.
  36. ^ de Loureiro, J. 1790. Flora Cochinchinensis 2: 606–607.
  37. ^ Schlauer, J. N.d. Nepenthes mirabilis. Carnivorous Plant Database.
  38. ^ a b Clarke, C.M. & C.C. Lee 2004. Pitcher Plants of Sarawak. Natural History Publications (Borneo), Kota Kinabalu.
  39. ^ Linnaeus, C. 1737. Nepenthes. Hortus Cliffortianus. Amsterdam.
  40. ^ Gledhill, D. 2008. The Names of Plants. Fourth Edition. Cambridge University Press, Cambridge.
  41. ^ Veitch, H.J. 1897. Nepenthes. Journal of the Royal Horticultural Society 21(2): 226–262.
  42. ^ Linnaeus, C. 1753. Nepenthes. Species Plantarum 2: 955.
  43. ^ Zahl, P.A. 1964. Malaysia's Giant Flowers and Insect-trapping Plants. National Geographic 125(5): 680–701.
  44. ^ James Pietropaolo; Patricia Ann Pietropaolo (1986). Carnivorous Plants of the World. Timber Press. p. 49. ISBN 978-0-88192-066-6. 
  45. ^ a b Peter D'Amato (2 July 2013). The Savage Garden, Revised: Cultivating Carnivorous Plants. Ten Speed Press. p. 89. ISBN 978-1-60774-411-5. 
  46. ^ Barry A. Rice (2006). Growing carnivorous plants. Timber Press, Incorporated. ISBN 978-0-88192-807-5. 

Further reading[edit]

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Nepenthes sp. Luzon

Nepenthes sp. Luzon is an undescribed tropical pitcher plant known from a single peak in northern Luzon, the Philippines, where it grows at 1220 m above sea level.[1] It shows similarities to N. burkei and N. ventricosa.[1] The species was originally discovered in April 2002 by ornithologist Herman Nuytemans[1] and was only relocated in the wild just over 10 years later.[2]

References[edit]

  1. ^ a b c McPherson, S.R. 2011. Discovery of an incompletely diagnosed pitcher plant from Luzon. In: New Nepenthes: Volume One. Redfern Natural History Productions, Poole. pp. 457–459.
  2. ^ Suarez, W. 2013. The rediscovery of Nepenthes sp. Luzon. Carnivorous Plants in the tropics, April 18, 2013.


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Nepenthes sp. Anipahan

Nepenthes sp. Anipahan is an undescribed tropical pitcher plant known only from Mount Anipahan in central Palawan, the Philippines, where it grows in upper montane forest at 1200–1400 m above sea level.[1][2][3] It is very closely allied to N. leonardoi and may be conspecific with it.[1][2]

References[edit]

  1. ^ a b McPherson, S.R. 2011. The Discovery of Nepenthes sp. 'Anipahan'. In: New Nepenthes: Volume One. Redfern Natural History Productions, Poole. pp. 330–345.
  2. ^ a b McPherson, S.R. 2011. Comparison of the highland Palaweño Nepenthes. In: New Nepenthes: Volume One. Redfern Natural History Productions, Poole. pp. 364–381.
  3. ^ Exploration of Mount Anipahan and Mount Kiamo. [video] Redfern Natural History Productions.


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Nepenthes × pangulubauensis

Nepenthes × pangulubauensis (/nɨˈpɛnθz pæŋˌɡlbˈɛnsɪs/; from Mount Pangulubau) is a natural hybrid between N. mikei and N. gymnamphora (or N. xiphioides, depending on whether or not it is considered a distinct species). It is endemic to the Indonesian island of Sumatra.[1]

References[edit]

  1. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Sumatra and Java. Redfern Natural History Productions, Poole.


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Nepenthes × ferrugineomarginata

Nepenthes × ferrugineomarginata (/nɨˈpɛnθz ˌfɛrʉˈɪni.ɵˌmɑrɨˈnɑːtə/; from Latin: ferrugineus = rust coloured, marginatus = margined) is a natural hybrid between N. albomarginata and N. reinwardtiana.[1] Its natural range covers the islands Borneo and Sumatra.[2] The type specimen was collected by Shigeo Kurata in Kenukat, West Kalimantan, in 1981. Kurata described the hybrid the following year.

N. × ferrugineomarginata has the longest binomial name of any plant in the genus.

References[edit]

  1. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Borneo. Redfern Natural History Productions, Poole.
  2. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Sumatra and Java. Redfern Natural History Productions, Poole.


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Nepenthes × hookeriana

Nepenthes × hookeriana (/nɨˈpɛnθz ˌhʊkəriˈɑːnə/; after Joseph Dalton Hooker), or Hooker's Pitcher-Plant,[5] is a common natural hybrid involving N. ampullaria and N. rafflesiana. It was originally described as a species.

It is a relatively common natural hybrid found in lowland conditions throughout Peninsular Malaysia, Borneo, Singapore and the Indonesian island of Sumatra.[6][7][8] Like its parental species, the hybrid is generally found in recently disturbed clearings.

Infraspecific taxa[edit source | edit]

  • Nepenthes hookeri var. elongata Hort.Veitch ex Wilson (1877)[9] sphalm.typogr.
  • Nepenthes hookeriana f. elongata (Hort.Veitch ex Wilson) Divers (1879)

References[edit source | edit]

  1. ^ von Arx, B., J. Schlauer & M. Groves 2001. CITES Carnivorous Plant Checklist. PDF The Cromwell Press, United Kingdom.
  2. ^ Masters, M.T. 1881. New garden plants. Nepenthes Hookeriana. The Gardeners' Chronicle, new series, 16(417): 812–813.
  3. ^ Hooker, J.D. 1873. Nepenthaceae. In: A. de Candolle Prodromus Systematis Naturalis Regni Vegetabilis 17: 90–105.
  4. ^ Schlauer, J. N.d. Nepenthes hookeriana. Carnivorous Plant Database.
  5. ^ Phillipps, A. & A. Lamb 1996. Pitcher-Plants of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
  6. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Borneo. Redfern Natural History Productions, Poole.
  7. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Peninsular Malaysia and Indochina. Redfern Natural History Productions, Poole.
  8. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Sumatra and Java. Redfern Natural History Productions, Poole.
  9. ^ [Anonymous] 1877. Reports of Societies. Royal Horticultural. The Gardeners' Chronicle 8(197): 441.

Further reading[edit source | edit]

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Nepenthes × alisaputrana

Nepenthes × alisaputrana (/nɨˈpɛnθz ˈɑːlɨsəpʊˈtrɑːnə/ preferably, or /əˈlɪsəpʊˈtrɑːnə/, after Datuk Lamri Ali), or the Leopard Pitcher-Plant,[1] is a hybrid of two well-known Nepenthes pitcher plant species: N. burbidgeae and N. rajah. The plant is confined to Mount Kinabalu in Sabah, Borneo.[2]

N. × alisaputrana (right) is often sympatric with N. rajah (left)

Nepenthes × alisaputrana was described in 1992 by J. H. Adam and C. C. Wilcock and is named in honour of Datuk Lamri Ali, a former Director of Sabah Parks.[3] It is only known from a few remote localities within Kinabalu National Park, where it grows in stunted, open vegetation over serpentine soils at around 2000 m above sea level, often amongst populations of N. burbidgeae.[4] It grows alongside both parent species on Pig Hill,[5] where it is found at 1930–1950 m.[6]

This plant is notable for combining the best characters of both parent species, not least the size of its pitchers, which rival those of N. rajah in volume (≤35 cm high, ≤20 cm wide).[4] The other hybrids involving N. rajah do not exhibit such impressive proportions. The pitchers of N. × alisaputrana can be distinguished from those of N. burbidgeae by a broader peristome, larger lid and simply by their sheer size. The hybrid differs from its other parent, N. rajah, by its lid structure, indumentum of short, brown hairs, narrower and more cylindrical peristome, and pitcher colour, which is usually yellow-green with red or brown flecking. For this reason, Anthea Phillipps and Anthony Lamb gave it the common name "Leopard Pitcher-Plant".[1] The peristome is green to dark red and striped with purple bands. Leaves are often slightly peltate. The hybrid is a strong climber and frequently produces upper pitchers.[2]

Nepenthes × alisaputrana more closely resembles N. rajah than N. burbidgeae, but it is difficult to confuse this plant with either. However, this mistake has previously been made on at least one occasion; a pitcher illustrated in Adrian Slack's Insect-Eating Plants and How to Grow Them as being N. rajah[7] is in fact N. burbidgeae × N. rajah.[2]

Distribution of phenolic compounds and leucoanthocyanins in
N. burbidgeae, N. rajah, and N. × alisaputrana
[8]
Taxon
1
2
3
4
5
6
7
 8 
Specimen
N. burbidgeae
3+
++
3+
3+
-
+
-
-
Jumaat 2484
N. rajah
-
-
+
±
++
++
3+
+
Jumaat 2443
N. × alisaputrana
+
++
3+
3+
+
++
3+
+
Jumaat 2442
N. × alisaputrana (in vitro)
+
++
3+
3+
+
++
+
+
Key: 1: Phenolic acid, 2: Ellagic acid, 3: Quercetin, 4: Kaempferol, 5: Luteolin, 6: 'Unknown Flavonoid 1', 7: 'Unknown Flavonoid 3', 8: Cyanidin

±: very weak spot, +: weak spot, ++: strong spot, 3+: very strong spot, -: absent

In 2002, phytochemical screening and analytical chromatography were used to study the presence of phenolic compounds and leucoanthocyanins in N. × alisaputrana and its putative parent species.[8] The research was based on leaf material from nine dry herbarium specimens. Eight spots containing phenolic acids, flavonols, flavones, leucoanthocyanins and 'unknown flavonoid' 1 and 3 were identified from chromatographic profiles. The distributions of these in the hybrid N. × alisaputrana and its putative parental species N. burbidgeae and N. rajah are shown in the table to the left. A specimen of N. × alisaputrana grown from tissue culture (in vitro) was also tested.[8]

Luteolin, cyanidin and 'Unknown Flavonoid 3' were undetected in N. burbidgeae, while concentrations of 'Unknown Flavonoid 1' were found to be weak. Chromatographic patterns of the N. × alisaputrana samples studied showed complementation of its putative parental species.[8]

Myricetin was found to be absent from all studied taxa. This agrees with the findings of previous authors[9][10] and suggests that the absence of a widely distributed compound like myricetin among the Nepenthes examined might provide additional diagnostic information for these taxa.[8]

References[edit]

  1. ^ a b Phillipps, A. & A. Lamb 1996. Pitcher-Plants of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
  2. ^ a b c Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
  3. ^ Adam, J.H. & C.C. Wilcock 1992. A new natural hybrid of Nepenthes from Mt. Kinabalu (Sabah). Reinwardtia 11: 35–40.
  4. ^ a b Clarke, C.M. 2001. A Guide to the Pitcher Plants of Sabah. Natural History Publications (Borneo), Kota Kinabalu.
  5. ^ Thong, J. 2006. Travels around North Borneo – Part 2. PDF Victorian Carnivorous Plant Society Journal 82: 6–12.
  6. ^ Adam, J.H., C.C. Wilcock & M.D. Swaine 1992. The ecology and distribution of Bornean Nepenthes. PDF Journal of Tropical Forest Science 5(1): 13–25.
  7. ^ Slack, A. 1986. Insect-Eating Plants and How to Grow Them. Alphabooks, Dorset, UK.
  8. ^ a b c d e Adam, J. H., R. Omar & C. C. Wilcock 2002. Phytochemical Screening of Flavonoids in Three Hybrids of Nepenthes (Nepenthaceae) and their Putative Parental Species from Sarawak and Sabah. PDF OnLine Journal of Biological Sciences 2(9): 623–625. doi:10.3923/jbs.2002.623.625
  9. ^ Jay, M. & P. Lebreton 1972. Chemotaxonomic research on vascular plants. The flavonoids of Sarraceniaceae, Nepenthaceae, Droseraceae and Cephlotaceae, a critical study of the order Sarraceniales. Naturaliste Canadien 99: 607–613.
  10. ^ Som, R.M. 1988. Systematic studies on Nepenthes species and hybrids in the Malay Peninsula. Ph.D. thesis, Fakulti Sains Hayat, Universiti Kebangsaan Malaysia, UKM Bangi, Selangor Darul Ehsan.
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Nepenthes × tsangoya

Nepenthes × tsangoya (/nɨˈpɛnθz sæŋˈɡɔɪ.ə/; after Peter Tsang) is a tropical pitcher plant. It reportedly represents the complex natural hybrid (N. alata × N. merrilliana) × N. mirabilis.

N. × tsangoya was mentioned as a natural hybrid in Guide to Nepenthes Hybrids (1995).[1] The known ranges of the parent species only overlap in Mindanao, the Philippines.

References [edit]

  1. ^ Lauffenburger, A. 1995. Guide to Nepenthes Hybrids. OmnisTerra.


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Nepenthes × mirabilata

Nepenthes × mirabilata (/nɨˈpɛnθz mɪˌræbɪˈlɑːtə/; a blend of mirabilis and alata) is a natural hybrid involving N. alata and N. mirabilis.[1]

N. × mirabilata was mentioned as a natural hybrid in Guide to Nepenthes Hybrids (1995). The hybrid is restricted to Mindanao, the Philippines, the only location where the parent species overlap.[2]

References [edit]

  1. ^ Cheek, M.R. & M.H.P. Jebb 2001. Nepenthaceae. Flora Malesiana 15: 1–157.
  2. ^ Co, L. & W. Suarez 2012. Nepenthaceae. Co's Digital Flora of the Philippines.


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Nepenthes × ventrata

Nepenthes × ventrata (/nɨˈpɛnθz vɛnˈtrɑːtə/; a blend of ventricosa and alata) is a natural hybrid involving N. alata and N. ventricosa. Like its two parent species, it is endemic to the Philippines. The name was originally published in the Carnivorous Plant Newsletter in 1979.[1]

Nepenthes × ventrata is one of the most common tissue cultured Nepenthes plants, although it is often mislabelled as Nepenthes alata.[2] It is relatively easy to grow indoors and is usually the first tropical pitcher plant seen by consumers due to its availability in many garden shops and home centres.

The cultivar N. 'LeeAnn Marie' is a later synonym of N. × ventrata, although the name is not established as it was not validly published.[3]

References [edit]

  1. ^ Fleming, R. 1979. Hybrid Nepenthes. PDF (626 KiB) Carnivorous Plant Newsletter 8(1): 10–12.
  2. ^ Catalano, M. 2009. Nepenthes. In: Growing Carnivores — an Italian perspective. Prague. pp. 50–57.
  3. ^ Schlauer, J. N.d. Nepenthes 'LeeAnn Marie'. Carnivorous Plant Database.

Further reading [edit]


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Nepenthes × merrilliata

Nepenthes × merrilliata (/nɨˈpɛnθz mɛˌrɪliˈɑːtə/; a blend of merrilliana and alata) is a natural hybrid involving N. alata and N. merrilliana. Like its two parent species, it is endemic to the Philippines, but limited by the natural range of N. merrilliana to Samar[2] as well as Mindanao and its offshore islands.

References [edit]

  1. ^ Fukatsu, Y. 1999. The List of Nepenthes species and Hybrids. (PDF version PDF)
  2. ^ Robinson, A. 2012. Nepenthes merrilliana on Samar. Carnivorous Plants in the tropics, June 29, 2012.


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Nepenthes × truncalata

Nepenthes × truncalata (/nɨˈpɛnθz ˌtrʌŋkjəˈlɑːtə/; a blend of truncata and alata) is a natural hybrid involving N. alata and N. truncata. Like its two parent species, it is endemic to the Philippines, but limited in distribution by the natural range of N. truncata on Mindanao.

References [edit]

  1. ^ Bednar, B.L. & O.C. Bramblett 1994. Nepenthes hybrid crosses made by: Bruce Lee Bednar & Orgel Clyde Bramblett. Carnivorous Plant Newsletter 23(1): 4.


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Nepenthes × sarawakiensis

Nepenthes × sarawakiensis (/nɨˈpɛnθz səˌrɑːwɑːkiˈɛnsɨs/, after Sarawak, Borneo) is a natural hybrid involving N. muluensis and N. tentaculata. It is quite a rare plant as one of its parent species, N. muluensis, is only known from several isolated mountains.

References [edit]


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Nepenthes × pyriformis

Nepenthes × pyriformis (/nɨˈpɛnθz ˌpɪrɨˈfɔrmɨs/ or /ˌpaɪərɨˈfɔrmɨs/; from Latin: pyrus = pear, -forma = shaped) is a natural hybrid involving N. inermis and N. talangensis.[6] It is known only from Mount Talang in Sumatra, to which N. talangensis is endemic. Nepenthes talangensis was only described as a distinct species in 1994.[7] Prior to this it was placed within N. bongso and some of the older literature identifies this hybrid as N. bongso × N. inermis.[5]

Nepenthes inermis × N. talangensis has been the subject of taxonomic confusion in the past. In an article published in 1973 on the Nepenthes of Borneo, Singapore, and Sumatra,[4] botanist Shigeo Kurata incorrectly identified specimens of this hybrid as belonging to N. dubia.[8]

In 1997, Matthew Jebb and Martin Cheek published their monograph "A skeletal revision of Nepenthes (Nepenthaceae)", in which they referred to N. dubia plant material from Mount Talang (Kurata s.n. SING).[3] Charles Clarke later identified Kurata s.n. as representing N. inermis × N. talangensis.

The natural hybrid is similar to N. dubia, but can be distinguished on the basis of several stable characters. The hybrid has a wider pitcher lid that is never relfexed beyond 90 degrees and the pitcher cup is not appressed in the lower parts as in N. dubia. In addition, the mouth of N. inermis × N. talangensis is raised towards the back as opposed to being horizontal.[8]

In 2001, Kurata described this hybrid as a new species, N. pyriformis.[2] Clarke rejected this interpretation in his monograph Nepenthes of Sumatra and Peninsular Malaysia, published the same year. Clarke found that the type specimen of N. pyriformis, Kurata & Mikil 4230 NDC, matches the appearance of N. inermis × N. talangensis "in most respects".[8] Following Clarke's interpretation, Kurata excluded N. pyriformis from his list of Nepenthes species the following year.[9]

References [edit]

  1. ^ von Arx, B., J. Schlauer & M. Groves 2001. CITES Carnivorous Plant Checklist. PDF The Cromwell Press, United Kingdom.
  2. ^ a b (Japanese) (English) Kurata, S. 2001. スマトラ島およびミンダナオ島産ウツボカズラの2新種(英文). [Two new species of Nepenthes from Sumatra (Indonesia) and Mindanao (Philippines).] Journal of Insectivorous Plant Society 52(2): 30–34.
  3. ^ a b Jebb, M.H.P. & M.R. Cheek 1997. A skeletal revision of Nepenthes (Nepenthaceae). Blumea 42(1): 1–106.
  4. ^ a b Kurata, S. 1973. Nepenthes from Borneo, Singapore and Sumatra. The Gardens' Bulletin Singapore 26(2): 227–232.
  5. ^ a b Hopkins, M., R. Maulder & B.[R.] Salmon 1990. A real nice trip to Southeast Asia. PDF (1.72 MiB) Carnivorous Plant Newsletter 19(1–2): 19–28.
  6. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Sumatra and Java. Redfern Natural History Productions, Poole.
  7. ^ Nerz, J. & A. Wistuba 1994. Five new taxa of Nepenthes (Nepenthaceae) from North and West Sumatra. Carnivorous Plant Newsletter 23(4): 101–114.
  8. ^ a b c Clarke, C.M. 2001. Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu.
  9. ^ Kurata, S. 2002. Revision trial in recent enumeration of Nepenthes species. PDF Proceedings of the 4th International Carnivorous Plant Conference: 111–116.
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Nepenthes × bauensis

Nepenthes × bauensis (/nɨˈpɛnθz bˈɛnsɪs/; after Bau, Sarawak) is a natural hybrid involving N. gracilis and N. northiana.[1]

N. × bauensis is intermediate in appearance between its two parent species. It displays the clumping habit and vine growth of N. gracilis, but can be distinguished from that species on the basis of its larger leaves and stems. The influence of N. northiana is most obvious in the pitcher morphology. In particular, the peristome is wider than in N. gracilis and has scattered red bands. Pitchers are up to 15 cm high and may be pale green to purplish-red in colour.

Like its parent species, N. × bauensis is a lowland plant that grows at an elevation of around 100 m. It is terrestrial in nature and inhabits swampy areas surrounding the limestone hills to which N. northiana is endemic.

This hybrid appears to be very rare and only a few plants have been found. It is known from a single location. N. × bauensis is one of only three known natural hybrids involving N. northiana, the others being N. × cincta and a cross with N. mirabilis.

References [edit]

  1. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Borneo. Redfern Natural History Productions, Poole.


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Nepenthes × cantleyi

Nepenthes × cantleyi (/nɨˈpɛnθz kæntˈl/; after Rob Cantley) is a natural hybrid involving N. bicalcarata and N. gracilis.[1] It is quite rare outside of Brunei, despite its parent species being more widespread throughout Borneo.[2]

The morphology of the pitchers closely resembles N. bicalacarata, although the peristome is more similar to that of N. gracilis. The characteristic spines of N. bicalcarata are greatly reduced and are only present as small bumps. This hybrid has the growth habit of N. gracilis, with the stem scrambling along the ground. It grows in open, sandy areas. The pitcher fluid is notably acidic like that of N. gracilis, with pH values as low as 1.82 being recorded.[2]

References [edit]

  1. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Borneo. Redfern Natural History Productions, Poole.
  2. ^ a b Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.