Nepenthes bicalcarata

Nepenthes bicalcarata
Intermediate pitcher of N. bicalcarata
Conservation status
Vulnerable (IUCN 2.3)
Scientific classification
Kingdom:	Plantae
(unranked):	Angiosperms
(unranked):	Eudicots
(unranked):	Core eudicots
Order:	Caryophyllales
Family:	Nepenthaceae
Genus:	Nepenthes
Species:	N. bicalcarata
Binomial name
Nepenthes bicalcarata Hook.f. (1873)[1]
Synonyms
Nepenthes dyak S.Moore (1880)[2]

Nepenthes bicalcarata (pronounced /nɨˈpɛnθiːz baɪˌkælkəˈrɑːtə/ or /bɪˌkælkəˈreɪtə/, Latin: bi "two", calcaratus "spur"), also known as the Fanged Pitcher-Plant,^[3] is a tropical pitcher plant endemic to northwestern Borneo.

Botanical history

Illustration of N. bicalcarata from Macfarlane's 1908 monograph

Nepenthes bicalcarata was formally described by Joseph Dalton Hooker in his 1873 monograph, "Nepenthaceae", based on specimens collected by Hugh Low and Odoardo Beccari near Lawas River in Borneo.^[1] The type specimen, Low s.n., is deposited at the Royal Botanic Gardens, Kew.^[4] Seven years later, Spencer Le Marchant Moore described Nepenthes dyak, based on a specimen collected by Johannes Elias Teijsmann, Teijsmann 10962, from Kapuas River near Sintang in western Borneo.^[2][5] This specimen is also held at the Royal Botanic Gardens, Kew^[4] and a duplicate is deposited at the National Herbarium of the Netherlands in Leiden.^[6] Nepenthes dyak was later mentioned several more times in the botanical literature,^[7][8] but is now considered conspecific with N. bicalcarta.^[9]

Nepenthes bicalcarata was introduced to Europe in 1879 by British explorer Frederick William Burbidge, who collected plants for the famous Veitch Nursery. These were cultivated to larger size and distributed in 1881.^[3]

During this time, interest in Nepenthes had reached its peak. A note in The Gardeners' Chronicle of 1881 mentions the Veitch Nursery's N. bicalcarata as follows:^[10]

"Then there is N. bicalcarata, a most robust habited kind with sturdy foliage and bag-like pitchers provided with a vicious-looking rat-trap-like apparatus in its lid which renders it very distinct from its neighbours."

Several years after its introduction, N. bicalcarata was still very much a horticultural rarity. In Veitch's catalogue for 1889, N. bicalcarata was priced at £3.3s per plant, while the famous giant-pitchered N. northiana and N. rajah were selling for £2.2s.^[3]

Description

Nepenthes bicalcarata plants are the largest in the genus, climbing up to 20 m into the forest canopy. The cylindrical stem is thicker than that of any other Nepenthes species, measuring up to 3.5 cm in diameter. Internodes are up to 40 cm long.^[9][11]

Climbing plant with upper pitchers

The leaves of N. bicalcarata are petiolate and coriaceous in texture. The lamina is obovate-lanceolate in form and also reaches huge dimensions, growing to 80 cm in length and 12 cm in width.^[12] It is slightly decurrent on the stem, forming two narrow wings. The lamina has indistinct longitudinal veins and numerous pennate veins. Tendrils may be up to 60 cm long and 8 mm wide. They are hollow and swollen near the pitcher.^[9]

Lower pitchers

Although most parts of the plant are very large, the pitchers themselves do not rival those of species such as N. rajah. Nevertheless, they may have a volume of over one litre and grow up to 25 cm high and 16 cm wide. A pair of prominent fringed wings (≤15 mm wide) run down the front of lower pitchers. These are usually reduced to ribs in aerial pitchers. The peristome (≤20 mm wide) is characteristically flattened and curved inwards. It bears small but distinct teeth. The two sharp spines for which the species is famous are present on the underside of the pitcher lid, and may be 3 cm long. They are derived from the uppermost 10–12 peristome ribs.^[13] The pitcher lid or operculum is reniform to cordate and has no appendages. An unbranched spur (≤15 mm long) is inserted at the base of the lid.^[9]

Nepenthes bicalcarata has a paniculate inflorescence. The peduncle may be up to 40 cm long and the rachis can reach 100 cm in length. Female inflorescences are usually shorter.^[14] Branches on the flower stem are up to 40 mm long and bear up to 15 flowers. Sepals are either obovate or lanceolate and up to 4 mm long.^[9] A study of 120 pollen samples taken from a herbarium specimen (Fosberg 43860, altitude not recorded) found the mean pollen diameter to be 28.9 μm (SE = 0.4; CV = 7.5%).^[15]

Mature plants are virtually glabrous. Caducous hairs are present on the youngest parts of the plant and on the inflorescences.

Nepenthes bicalcarata varies little across its range. Consequently, no infraspecific taxa have been described.^[9]

Ecology and conservation status

Natural habitat of N. bicalcarata

Nepenthes bicalcarata is endemic to Borneo. It is most common in the peat swamp forests of the western coast of the island, which stretch across Sarawak, Sabah, Kalimantan, and Brunei. There it often grows in the shade of the ubiquitous dipterocarp Shorea albida.^[3] Nepenthes bicalcarata also occurs in kerangas forest and has even been recorded from white sand heath forests in Sarawak and East Kalimantan.^[3] The species is often sympatric with N. ampullaria in these habitats.^[9]

Specimens growing in undisturbed peat swamp forest, where sunlight is greatly diffused and high humidity prevails, reach the largest dimensions.^[9] Nepenthes bicalcarata has a shallow root system that only penetrates the top layer of peat and leaf litter, to a depth of about 25 cm. Below this, high concentrations of tannins and alkaloids render the substrate toxic.^[3]

Nepenthes bicalcarata is generally found below 300 m in altitude, although Johannes Gottfried Hallier reported a single collection in 1894 from between 700 and 950 m above sea level.^[16]

The conservation status of N. bicalcarata is listed as Vulnerable on the 2006 IUCN Red List of Threatened Species based on an assessment carried out in 2000.^[17] In 1997, Charles Clarke informally classified the species as Near Threatened based on the IUCN criteria.^[9] This agrees with the conservation status assigned to N. bicalcarata by the World Conservation Monitoring Centre.^[18]

Carnivory

Upper pitcher with prominent thorns

The two thorns that give N. bicalcarata its name are unique to this species and bear some of the largest nectaries in the plant kingdom.^[19][20][21] The purpose of these structures has long been debated among botanists. Frederick William Burbidge suggested that they might serve to deter arboreal mammals such as tarsiers, lorises and monkeys from stealing the contents of the pitchers. In an article published in 1982, Cliff Dodd speculated on the function of the thorns, but did not believe they play a role in prey capture.^[22] Charles Clarke observed that monkeys and tarsiers rip the pitchers open at the sides in order to feed from them, rather than reaching in through the pitcher mouth.^[23] However, he found that the mammals attacked the pitchers of N. bicalcarata less frequently than those of other species, such as N. rafflesiana.^[23] Clarke's observations suggest that the spines likely serve to lure insects into a precarious position over the pitcher mouth, where they may lose their footing and fall into the pitcher fluid, eventually drowning.^[23]

A similar trapping method is employed by the Sumatran endemic N. lingulata, which has a single filiform appendage positioned over the pitcher mouth. However, in that species the appendage is a structure of the lid and not the peristome.^[24]

Together with N. ampullaria and N. ventricosa, N. bicalcarata is unusual in that the glandular region of the pitcher extends almost to the peristome, such that there is little or no conductive waxy zone.^[25][26][27] The waxy zone functions by causing prey to slip and fall into the digestive fluid. A 2004 study found that the peristome of N. bicalcarata plays a very important role in prey capture. When it is dry, the peristome is very ineffective in catching insect prey, but when wetted, the capture rate increases more than three-fold.^[25]

A 1999 study of pitcher morphology and spectral reflectance characteristics showed that N. bicalcarata has no colour contrast maximum between the peristome and pitcher cup in the ultraviolet waveband, but small maxima of –0.17 at 450 nm (violet), 0.32 at 548 nm (green), and –0.16 at 668 nm (red).^[28] This means that the peristome is less reflective than the pitcher cup in the violet and red bands, but more reflective in the green band. The contrast maxima in the green and blue regions seem to correspond to insect visual sensitivity maxima, while the others do not.^[28] Based on their findings, the authors made the "tentative prediction" that the upper pitchers of N. bicalcarata are less successful at catching anthophilous (flower-visiting) prey than those of sympatric N. rafflesiana.^[28]

Association with ants and pitcher infauna

Nepenthes bicalcarata plays host to an unusual species of ant that makes its nest in the plant's hollow tendrils.^{[9][29][30][31]} Described as Camponotus schmitzi in 1933,^[32] it is a member of the extremely populous and widespread genus of carpenter ants.

Upper pitcher with swollen tendril colonised by Camponotus schmitzi.

This unique animal-plant interaction was noted by Frederick William Burbidge as early as 1880.^[33] In 1904, Odoardo Beccari suggested that the ants feed on insects found on and around the plant, but may fall prey to it themselves.^[34] In 1990, B. Hölldobler and E.O. Wilson proposed that N. bicalcarata and C. schmitzi form a mutually beneficial association.^[35] At the time, however, no experimental data existed to support such a hypothesis. A series of observations and experiments carried out in Brunei by Charles Clarke in 1992 and 1998,^[36][37][38] and by Clarke and Roger Kitching in 1993 and 1995,^[39][40] strongly support the mutualism theory. John Thompson has suggested that N. bicalcarata may be the only plant species that obtains nutrients through both insect capture and ant-hosting habits.^[41]

Camponotus schmitzi ants feed by descending into the pitcher fluid and retrieving arthropods caught by the plant. They seem to ignore smaller insects and only target larger prey items. Hauling food from the pitcher fluid to the peristome—a distance of no more than 5 cm—may take up to 12 hours.^[9] In this way the contents of N. bicalcarata pitchers is controlled such that organic matter does not accumulate to the point of putrefaction, which could lead to the demise of pitcher infauna (which also appear to benefit the plant) and sometimes the pitcher itself.^[9][42] The ants have also been observed to attack newly caught insects and therefore prevent prey escape.^[43] At other times, the ants are very passive, remaining hidden under the inner peristome fold, presumably so as not to dissuade visitation by potential prey species.^[43] This behaviour is in stark contrast to other myrmecophytic ants, which are typically highly territorial.^[43] In order to create a favourable environment for its pitcher inhabitants, it appears that N. bicalcarata actively maintains the pH of the pitcher fluid at a less acidic level than that found in most other Nepenthes species.^[26][43] This might explain the occasional presence of tree frog eggs, which are sensitive to strongly acidic conditions, in the pitchers of this species.^[26]

Research conducted by Dennis and Marlis Merbach has shown that C. schmitzi is also beneficial to N. bicalcarata in that it protects it from pitcher-destroying weevils of the genus Alcidodes.^[44][45]

The ants seem to favour upper pitchers and rarely colonise lower pitchers.^[46] This is likely due to the fact that terrestrial traps are periodically submerged in water during heavy rains. Flooding of the ants' nest chamber could result in the death of the developing eggs, larvae, and pupae.^[21] Camponotus schmitzi nests solely in the tendrils of N. bicalcarata and rarely ventures onto other plants. The species is completely dependent on N. bicalcarata for food and domicile.^[9] Nepenthes bicalcarata, on the other hand, is able to survive and reproduce without the presence of the ants; it is a facultative mutualist. This being the case, there appear to be few mature plants over 2 metres in height not colonised by C. schmitzi.^[9]

A species of mite, Naiadacarus nepenthicola, appears to be restricted to the pitchers of N. bicalcarata. It is thought to feed on decomposing leaves and insects that are caught in the pitchers. Deutonymphs of this mite are dispersed through phoresy on C. schmitzi.^[47]

Natural hybrids

The following natural hybrids involving N. bicalcarata have been recorded.

• N. ampullaria × N. bicalcarata^[9]
• (N. ampullaria × N. gracilis) × N. bicalcarata [=N. × trichocarpa × N. bicalcarata]
• N. bicalcarata × N. gracilis [=N. × cantleyi]^[9]
• N. bicalcarata × N. mirabilis^[48] (including N. bicalcarata × N. mirabilis var. echinostoma)^[9]
• N. bicalcarata × N. rafflesiana^[9]
• ? (N. bicalcarata × N. rafflesiana) × N. mirabilis var. echinostoma^[9]

None of these hybrids are known to be colonised by C. schmitzi as is the parent species N. bicalcarata.^[9]

N. ampullaria × N. bicalcarata

Lower pitcher of N. ampullaria × N. bicalcarata (cultivated)

Since N. ampullaria and N. bicalcarata are often sympatric in the wild, it is not surprising that they occasionally hybridise. Several examples of this natural hybrid are known from Brunei, where it grows in peat swamp forest and heath forest.^[9] It was first recorded by Mitsuru Hotta in 1966.^[49]

Pitchers bear the characteristic thorns of N. bicalcarata, although they are greatly reduced in size. The mouth is round and the lid is ovate-cordate in form. This hybrid resembles N. × hookeriana to a certain extent, but differs in having spines on the underside of the lid and more globose upper pitchers with a smaller lid.^[9]

Aerial pitchers of N. ampullaria × N. bicalcarata are usually dominated by characteristics of N. ampullaria. They are often very small and unable to function in a normal manner.^[9]

(N. ampullaria × N. gracilis) × N. bicalcarata

This rare complex hybrid was discovered in the lowland forests of Brunei. It produces small speckled pitchers with reduced thorns under the lid. The plant is intermediate in appearance between N. × trichocarpa and N. bicalcarata.

N. bicalcarata × N. gracilis

Main article: Nepenthes × cantleyi

Lower pitcher of Nepenthes × cantleyi

This hybrid has been named N. × cantleyi after Rob Cantley. 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.^[9]

N. bicalcarata × N. mirabilis var. echinostoma

This natural hybrid is rare and is only known from Brunei, where it involves N. mirabilis var. echinostoma. It closely resembles N. bicalcarata, although it differs from that species in having reduced thorns and a wider peristome. Unlike N. bicalcarata, this hybrid inhabits open areas with sandy substrates.^[9]

N. bicalcarata × N. rafflesiana

This relatively rare natural hybrid is sometimes found growing among populations of N. bicalcarata and N. rafflesiana. It is intermediate in form between its parent species.

(N. bicalcarata × N. rafflesiana) × N. mirabilis var. echinostoma

Charles Clarke reported a single plant displaying characters intermediate in form between N. bicalcarata, N. rafflesiana and N. mirabilis var. echinostoma growing among populations of these species. It is likely a complex hybrid, although this cannot be proven without the use of genetic testing.^[9]

Cultivation and cultivars

Lower pitcher of a plant cultivated in Kuching, Sarawak.

Nepenthes bicalcarata may be cultivated in a similar manner to other members of the genus, though it requires relatively hot and humid conditions.^[22][50] It needs large pots to do well, and temporary halts in growth are often due to becoming root-bound. Re-potting to a larger pot will often result in a rapid size increase. Under satisfactory conditions, this plant will grow quite fast for a Nepenthes, reaching a large size relatively rapidly.

Despite naturally occurring in peat swamp forests, N. bicalcarata has been successfully grown in a completely inorganic substrate consisting of one part each of Seramis clay perls, lava gravel, and Lecaton expanded clay perls.^[51]

Nepenthes bicalcarata has been artificially crossed with N. hirsuta. A particularly robust-pitchered cultivar of this hybrid was named Nepenthes 'Hortulanus Otten', in honour of Karel Otten, former curator of the Botanic Garden in Ghent, Belgium.^[52]

A cultivar of the artificial cross N. bicalcarata × N. × dyeriana was registered in 1988 under the name Nepenthes 'Nina Dodd'. It is named after a relative of Cliff Dodd, who created the hybrid.^[53]

Two further cultivars of manmade hybrids involving N. bicalcarata have been named. These are Nepenthes 'Bella' ((N. ampullaria × N. bicalcarata) × N. truncata) and Nepenthes 'Rapa' (N. ampullaria × N. bicalcarata).^[54] However, both of these names are not established, as they were published without a description.^[4]

References

1. ^ ^{a b} Hooker, J.D. 1873. Nepenthaceae. In: A. de Candolle Prodromus Systematis Naturalis Regni Vegetabilis 17: 90–105.
2. ^ ^{a b} Moore, S.L.M. 1880. Alabastra diversa. The Journal of Botany 18: 1–8.
3. ^ ^{a b c d e f} Phillipps, A. & A. Lamb 1996. Pitcher-Plants of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
4. ^ ^{a b c} Schlauer, J. 2006. Nepenthes bicalcarata. Carnivorous Plant Database.
5. ^ Jebb, M.H.P. & M.R. Cheek 1997. A skeletal revision of Nepenthes (Nepenthaceae). Blumea 42 (1): 1–106.
6. ^ Specimen detail: Teijsmann 10962. Nationaal Herbarium Nederland.
7. ^ Burbidge, F.W. 1882. Notes on the new Nepenthes. The Gardeners' Chronicle 17(1): 56.
8. ^ Beccari, O. 1896. Malesia, III (1886–1890): Rivista delle specie del genere Nepenthes, p. 1–15.
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10. ^ [Anonymous] 1881. Messr's Veitch's Nepenthes House. The Gardeners' Chronicle n. ser. 16: 598.
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12. ^ Danser, B.H. 1928. 4. Nepenthes bicalcarata. In: The Nepenthaceae of the Netherlands Indies. Bulletin du Jardin Botanique de Buitenzorg, Série III, 9(3–4): 249–438.
13. ^ Cheek, M.R. & M.H.P. Jebb 2001. Nepenthaceae. Flora Malesiana 15: 1–157.
14. ^ Macfarlane, J.M. 1908. Nepenthaceae. In: A. Engler Das Pflanzenreich IV, III, Heft 36: 1–91.
15. ^ Adam, J.H. & C.C. Wilcock 1999. Palynological study of Bornean Nepenthes (Nepenthaceae).PDF Pertanika Journal of Tropical Agricultural Science 22(1): 1–7.
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18. ^ Simpson, R.B. 1995. Nepenthes and conservation. Curtis's Botanical Magazine 12: 111–118.
19. ^ Merbach, M.A., G. Zizka, B. Fiala, D. Merbach & U. Maschwitz 1999. Giant nectaries in the peristome thorns of the pitcher plant Nepenthes bicalcarata Hooker f. (Nepenthaceae): anatomy and functional aspects.PDF Ecotropica 5: 45–50.
20. ^ Clarke, C.M. 2001. A Guide to the Pitcher Plants of Sabah. Natural History Publications (Borneo), Kota Kinabalu.
21. ^ ^{a b} Clarke, C.M. & C.C. Lee 2004. Pitcher Plants of Sarawak. Natural History Publications (Borneo), Kota Kinabalu.
22. ^ ^{a b} Dodd, C. 1982. The most dangerous (looking) Nepenthes.PDF (658 KiB) Carnivorous Plant Newsletter 11(3): 64–65, 73, 78.
23. ^ ^{a b c} Clarke, C.[M.] 1993. The possible functions of the thorns of Nepenthes bicalcarata (Hook.f.) pitchers.PDF (194 KiB) Carnivorous Plant Newsletter 22(1–2): 27–28.
24. ^ Lee, C.C., Hernawati & P. Akhriadi 2006. Two new species of Nepenthes (Nepenthaceae) from North Sumatra. Blumea 51(3): 561–568.
25. ^ ^{a b} Bohn, H.F. & W. Federle 2004. Insect aquaplaning: Nepenthes pitcher plants capture prey with the peristome, a fully wettable water-lubricated anisotropic surface.PDF Proceedings of the National Academy of Sciences 101(39): 14138–14143.
26. ^ ^{a b c} Moran, J.A., B.J. Hawkins, B.E. Gowen & S.L. Robbins 2010. Ion fluxes across the pitcher walls of three Bornean Nepenthes pitcher plant species: flux rates and gland distribution patterns reflect nitrogen sequestration strategies. Journal of Experimental Botany 61(5): 1365–1374. doi:10.1093/jxb/erq004
27. ^ Di Giusto, B., M. Guéroult, N. Rowe & L. Gaume 2009. Chapter 7: The Waxy Surface in Nepenthes Pitcher Plants: Variability, Adaptive Significance and Developmental Evolution. In: S.N. Gorb (ed.) Functional Surfaces in Biology: Adhesion Related Phenomena. Volume 2. Springer. pp. 183–204.
28. ^ ^{a b c} Moran, J.A., W.E. Booth & J.K. Charles 1999. Aspects of Pitcher Morphology and Spectral Characteristics of Six Bornean Nepenthes Pitcher Plant Species: Implications for Prey Capture.PDF Annals of Botany 83: 521–528.
29. ^ Shelford, R. 1916. A Naturalist in Borneo. T. Fisher Unwin, London.
30. ^ Cresswell, J.E. 2000. Resource input and the community structure of larval infaunas of an eastern tropical pitcher plant Nepenthes bicalcarata. Ecological Entomology 25(3): 362–366. doi:10.1046/j.1365-2311.2000.00267.x
31. ^ Merbach, M.A., G. Zizka, B. Fiala, U. Maschwitz & W.E. Booth 2000. Carnivory and myrmecophytism—a contradiction? Studies on Nepenthes bicalcarata Hook.f. and its ants. Tagungsband gtö 2000 13. Jahrestagung der Deutschen Gesellschaft für Tropenökologie 1–3. March 2000 in Würzburg Lehrstuhl für Tierökologie und Tropenbiologie Universität Würzburg. p. 106.
32. ^ Schuitemaker, J.P. & A. Stärke 1933. Contribution a l'etude de la faune Nepenthicole. Art. III Un noveau Camponotus de Borneo, habitant les tiges cruises de Nepenthes récolté par J. P. Schuitemaker et décrit par A. Stärke, den Dolder. Overdruk uit het Natuurhistorisch Maandblad 22(3): 29–31.
33. ^ Burbidge, F.W. 1880. The Gardens of the Sun: or, a Naturalist's Journal on the Mountains and in the Forests and Swamps of Borneo and the Sulu Archipelago. John Murray, London.
34. ^ Beccari, O. 1904. Wanderings in the Great Forests of Borneo. Archibald and Constable, London.
35. ^ Hölldobler, B. & E.O. Wilson 1990. The Ants. Springer-Verlag, Berlin, Germany.
36. ^ Clarke, C.M. 1992. The ecology of metazoan communities in Nepenthes pitcher plants in Borneo, with special reference to the community of Nepenthes bicalcarata Hook.f. Ph.D. thesis, University of New England, Armidale, New South Wales. 269 pp.
37. ^ Clarke, C.M. 1998. Initial colonisation and prey capture in Nepenthes bicalcarata (Nepenthaceae) pitchers in Brunei. Sandakania 12: 27–36.
38. ^ Clarke, C.M. 1998. The aquatic arthropod community of the pitcher plant, Nepenthes bicalcarata (Nepenthaceae) in Brunei. Sandakania 11: 55–60.
39. ^ Clarke, C.M. & R.L. Kitching 1993. The metazoan food webs from six Bornean Nepenthes species. Ecological Entomology 18(1): 7–16. doi:10.1111/j.1365-2311.1993.tb01074.x
40. ^ Clarke, C.M. & R.L. Kitching 1995. Swimming ants and pitcher plants: a unique ant-plant interaction from Borneo. Journal of Tropical Ecology 11(4): 589–602. doi:10.1017/S0266467400009160
41. ^ Thompson, J.H. 1981. Reverse animal-plant interactions: the evolution of insectivorous and ant-fed plants. Biological Journal of the Linnean Society 16: 147–155.
42. ^ Hansen, E. 2001. Where rocks sing, ants swim, and plants eat animals: finding members of the Nepenthes carnivorous plant family in Borneo. Discover 22(10): 60–68.
43. ^ ^{a b c d} Bonhomme, V., I. Gounand, C. Alaux, E. Jousselin, D. Barthélémy & L. Gaume 2011. The plant-ant Camponotus schmitzi helps its carnivorous host-plant Nepenthes bicalcarata to catch its prey.PDF Journal of Tropical Ecology 27: 15–24. doi:10.1017/S0266467410000532
44. ^ Merbach, M.A., G. Zizka, B. Fiala, D.J. Merbach, W.E. Booth & U. Maschwitz 2007. Why a carnivorous plant cooperates with an ant – selective defense against pitcher-destroying weevils in the myrmecophytic pitcher plant Nepenthes bicalcarata Hook.f..PDF Ecotropica 13: 45–56.
45. ^ Merbach, M.A. & D.J. Merbach 2002. Auf der Suche nach Nahrung in die Todesfalle.PDF (116 KiB) Forschung Frankfurt 2002(3): 74–77.
46. ^ Clarke, C.M. 1997. The effects of pitcher dimorphism on the metazoan community of the carnivorous pitcher plant Nepenthes bicalcarata Hook.f.. Malayan Nature Journal 50: 149–157.
47. ^ Fashing, N.J. & T.H. Chua 2002. Systematics and ecology of Naiadacarus nepenthicola, a new species of Acaridae (Acari : Astigmata) inhabiting the pitchers of Nepenthes bicalcarata Hook. f. in Brunei Darussalam.PDF (2.12 MiB) International Journal of Acarology 28(2): 157–167.
48. ^ McPherson, S.R. 2009. Pitcher Plants of the Old World. 2 volumes. Redfern Natural History Productions, Poole.
49. ^ Hotta, M. 1966. Notes on Bornean Plants. Acta Phytotaxonomica et Geobotanica 22(1–2): 7–9.
50. ^ D'Amato, P. 1998. The Savage Garden — Cultivating Carnivorous Plants. Ten Speed Press, Berkley, California.
51. ^ Rischer, H. 2000. Growing Nepenthes in a Completely Inorganic Substrate. Carnivorous Plant Newsletter 29(2): 50–53.
52. ^ Cook, D., J.H. Hummer, A. Vogel, O. Gluch & C. Clayton 2004. New Cultivars. Carnivorous Plant Newsletter 33(1): 19–22.
53. ^ Robinson, J.T. 1988. New Cultivars. Carnivorous Plant Newsletter 17(1): 20.
54. ^ Fukatsu, Y. 1999. List of Nepenthes Hybrids.

Further reading

• 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.
• Adam, J.H. 1997. Prey spectra of Bornean Nepenthes species (Nepenthaceae) in relation to their habitat.PDF Pertanika Journal of Tropical Agricultural Science 20(2–3): 121–134.
• Bourke, G. 2011. The Nepenthes of Mulu National Park. Carniflora Australis 8(1): 20–31.
• Clarke, C.M. 2006. Introduction. In: Danser, B.H. The Nepenthaceae of the Netherlands Indies. Natural History Publications (Borneo), Kota Kinabalu. pp. 1–15.
• Fashing, N.J. 2010. Two novel adaptations for dispersal in the mite family Histiostomatidae (Astigmata).PDF In: M.W. Sabelis & J. Bruin (eds.) Trends in Acarology: Proceedings of the 12th International Congress. Springer Science, Dordrecht. pp. 81–84. doi:10.1007/978-90-481-9837-5
• Fretwell, S. 2010. Twelve days in Borneo – a dream expedition: part 4. Victorian Carnivorous Plant Society Journal 98: 6–13.
• (Czech) Kubeš, V. 2003. "Panel story" o pěstování láčkovky Nepenthes bicalcarata. Trifid 2003(2): 7–10. (page 2, page 3, page 4)
• (Indonesian) Mansur, M. 2001. Koleksi Nepenthes di Herbarium Bogoriense: prospeknya sebagai tanaman hias.PDF In: Prosiding Seminar Hari Cinta Puspa dan Satwa Nasional. Lembaga Ilmu Pengetahuan Indonesia, Bogor. pp. 244–253.
• Siegara, A. & Yogiara 2009. Bacterial community profiles in the fluid of four pitcher plant species (Nepenthes spp.) grown in a nursery. Microbiology Indonesia 3(3): 109–114.
• Thorogood, C. 2010. The Malaysian Nepenthes: Evolutionary and Taxonomic Perspectives. Nova Science Publishers, New York.
• (German) Zizka, G. 1990. Pflanzen und Ameisen. Palmengarten Sonderheft 15: 112–116.
• (German) Zizka, G. 1991. Über die Kannenpflanze Nepenthes bicalcarata J.D. Hooker. Palmengarten 91(3): 39–44.

External links

• Photographs of N. bicalcarata at the Carnivorous Plant Photofinder

v · d · e

Nepenthes (list)

Species

N. adnata N. alata N. alba N. albomarginata N. ampullaria N. andamana N. angasanensis N. argentii N. aristolochioides N. attenboroughii N. beccariana N. bellii N. benstonei N. bicalcarata N. bokorensis N. bongso N. boschiana N. burbidgeae N. burkei N. campanulata N. ceciliae N. chang

N. chaniana N. clipeata N. copelandii N. danseri N. deaniana N. densiflora N. diatas N. distillatoria N. dubia N. edwardsiana N. ephippiata N. eustachya N. eymae N. faizaliana N. flava N. fusca N. gantungensis N. glabrata N. glandulifera N. gracilis N. gracillima N. gymnamphora

N. hamata N. hamiguitanensis N. hirsuta N. hispida N. holdenii N. hurrelliana N. inermis N. insignis N. izumiae N. jacquelineae N. jamban N. junghuhnii N. kampotiana N. kerrii N. khasiana N. klossii N. kongkandana N. lamii N. lavicola N. leonardoi N. lingulata N. longifolia

N. lowii N. macfarlanei N. macrophylla N. macrovulgaris N. madagascariensis N. mantalingajanensis N. mapuluensis N. masoalensis N. maxima N. merrilliana N. micramphora N. mikei N. mindanaoensis N. mira N. mirabilis N. mollis N. muluensis N. murudensis N. naga N. neoguineensis N. northiana N. ovata

N. palawanensis N. paniculata N. papuana N. peltata N. pervillei N. petiolata N. philippinensis N. pilosa N. pitopangii N. platychila N. pulchra N. rafflesiana N. rajah N. ramispina N. reinwardtiana N. rhombicaulis N. rigidifolia N. rowanae N. sanguinea N. saranganiensis N. sibuyanensis N. singalana

N. smilesii N. spathulata N. spectabilis N. stenophylla N. sumatrana N. suratensis N. surigaoensis N. talangensis N. tenax N. tentaculata N. tenuis N. thai N. thorelii N. tobaica N. tomoriana N. treubiana N. truncata N. veitchii N. ventricosa N. vieillardii N. villosa N. vogelii

Incompletely diagnosed taxa: N. sp. Misool • N. sp. Papua • N. sp. Sulawesi
Possible extinct species: N. echinatus • N. echinosporus • N. major

Natural hybrids

N. × alisaputrana N. × bauensis N. × cantleyi N. × cincta

N. × ferrugineomarginata N. × harryana N. × hookeriana N. × kinabaluensis

N. × kuchingensis N. × merrilliata N. × mirabilata N. × pangulubauensis

N. × pyriformis N. × sarawakiensis N. × sharifah-hapsahii N. × trichocarpa

N. × truncalata N. × trusmadiensis N. × tsangoya N. × ventrata

Related topics

Nepenthes taxonomy • Nepenthes infauna • Nepenthes species by distribution • Nepenthes cultivars • Nepenthes literature
Nepenthesin • Nepenthales