Nepenthes rajah

Joseph Dalton Hooker described Nepenthes rajah in 1859, naming it in honour of James Brooke, the first White Rajah of Sarawak.Kurata 1976, p. 61. 'Rajah Brooke's Pitcher Plant'Phillipps & Lamb 1996, p. 129. is an accurate, but seldom-used common name. N. rajah is sometimes called the 'Giant Malaysian Pitcher Plant'Gibson 1983. or simply 'Giant Pitcher Plant', although the binomial name remains by far the most popular way of referring to this species. The specific epithet rajah means "King" in Malay and this, coupled with the impressive size of its pitchers, has meant that N. rajah is often called the "King of Nepenthes".Steiner 2002, p. 94.

{{see also|List of Nepenthes clades}}

Nepenthes rajah is a scrambling vine. The stem usually grows along the ground, but climbs whenever it comes into contact with a suitable support. The stem is up to {{cvt|30|mm|in}}) thick and may reach up to {{cvt|6|m|ft}} in length, although it rarely exceeds {{cvt|3|m|ft}}.{{rp|10,120}} The leaves are distinctive and reach a large size. They are leathery in texture with a wavy outer margin. The leaves are characteristically peltate, whereby the tendril joins the lamina on the underside, before the apex. The tendrils are inserted within {{cvt|5|cm|in}} below the leaf apex and reach a length of approximately {{cvt|50|cm|in}}.{{rp|120,122}}

The pitcher is a trap consisting of the main pitcher cup, covered by an operculum or lid. A reflexed ring of hardened tissue, the peristome, surrounds the entrance. N. rajah produces two distinct types of trap. "Lower" or "terrestrial" pitchers, the most common, are very large, richly coloured, and ovoid in shape. Exceptional specimens may be more than {{cvt|40|cm|in}} in length and hold {{cvt|3.5|l|gal}} of water and over {{cvt|2.5|l|gal}} of digestive fluid, although most do not exceed {{cvt|200|ml|U.S.oz}}.Clarke 2001b, p. 7. The lower pitchers rest on the ground. They are usually red to purple on the outside, lime green to purple on the inside.Clarke 2001b, p. 26. Mature plants may in addition produce much smaller "upper" or "aerial" pitchers, which are funnel-shaped, and usually more colourful. The tendril attachment is normally at the rear of the pitcher cup. True upper pitchers are rare, as the stems rarely attain lengths greater than a few metres before dying off.Clarke & Kruger 2005.

The peristome has a distinctive scalloped edge and is greatly expanded, forming a red lip around the trap's mouth. A series of raised ribs intersect the peristome, ending in short, sharp teeth that line its inner margin. The inner portion of the peristome accounts for around 80% of its total cross-sectional surface length.{{cite journal |last1=Bauer |first1=U. |last2=Clemente |first2=C. J. |last3=Renner |first3=T. |last4=Federle |first4=W. |title=Form follows function: morphological diversification and alternative trapping strategies in carnivorous Nepenthes pitcher plants |journal=Journal of Evolutionary Biology |date=January 2012 |volume=25 |issue=1 |pages=90–102 |doi=10.1111/j.1420-9101.2011.02406.x|pmid=22023155 }} The huge, vaulted lid is a distinguishing feature of this species. It is ovate to oblong in shape with a distinct keel down the middle, and two prominent lateral veins.{{in lang|de}} Schmid-Hollinger, R. N.d. [http://www.bio-schmidhol.ch/site/index.cfm?id_art=54826&actMenuItemID=25460&vsprache/DE/Kannendeckel__lid_.cfm Kannendeckel (lid)] {{Webarchive|url=https://web.archive.org/web/20131206025936/http://www.bio-schmidhol.ch/site/index.cfm?id_art=54826&actMenuItemID=25460&vsprache%2FDE%2FKannendeckel__lid_.cfm |date=2013-12-06 }}. bio-schmidhol.ch. The spur at the back of the lid is approximately {{cvt|20|mm|in}} long and unbranched.{{rp|122}}

The plant has very large nectar-secreting glands covering its pitchers. The inner surface of the pitcher is wholly glandular, with {{cvt|300 to 800|/cm2|/in2|disp=preunit|glands}}.Danser 1928, 38.

N. rajah seems to flower at any time of the year. Flowers are produced in large numbers on inflorescences that arise from the apex of the main stem. N. rajah produces a very large inflorescence that can be {{cvt|80|cm|in}}, and sometimes even {{cvt|120|cm|in}} tall.{{rp|122}} The individual flowers of N. rajah are produced on partial peduncles (twin stalks) and so the inflorescence is called a raceme (as opposed to a panicle for multi-flowered bunches). The flowers are reported to give off a strong sugary smell and are brownish-yellow in colour. Sepals are elliptic to oblong and up to {{cvt|8|mm|in}} long.{{rp|122}} Like all Nepenthes species, N. rajah is dioecious, which means that individual plants produce flowers of a single sex. Fruits are orange-brown and {{cvt|10 to 20|mm|in}} long . A study of 300 pollen samples taken from a herbarium specimen (J.H.Adam 2443, collected at an altitude of {{cvt|1930–2320|m|ft}}) found the mean pollen diameter to be {{cvt|34.7|μm|in}} (SE = 0.3; CV = 7.0%).Adam, J.H. & C.C. Wilcock 1999. {{cite web|url= http://psasir.upm.edu.my/3779/1/Palynological_Study_of_Bornean_Nepenthes_%28Nepenthaceae%29.pdf |title=Palynological study of Bornean Nepenthes (Nepenthaceae). }} Pertanika Journal of Tropical Agricultural Science 22(1): 1–7.

File:Kinabalu Mesilau N. rajah upper pitcher plant 2.jpg|Mature plants with both lower and upper pitchers

File:nep rajah peltate.jpg|The peltate leaf attachment

File:Rajahpitcher3.jpg|A terrestrial pitcher

File:Nepenthes rajah 41 cm pitcher.jpg|The {{cvt|41|cm|in}} pitcher found on The Sabah Society's March 2011 trip to Mesilau

File:Kinabalu Mesilau N. rajah upper pitcher 3.jpg|A rare aerial pitcher

File:Nep rajah2 flower.jpg|Plant in flower

{{See also|Carnivorous plant#Pitfall traps}}

File:Rajahlizard.jpg

N. rajah is a carnivorous plant of the pitfall trap variety. It is famous for occasionally trapping vertebrates, even small mammals. There exist at least two records of drowned rats found in N. rajah pitchers. The first observation dates from 1862 and was made by Spenser St. John, who accompanied Hugh Low on two ascents of Mount Kinabalu. In 1988, Anthea Phillipps and Anthony Lamb confirmed the plausibility of this record when they managed to observe drowned rats in a large pitcher of N. rajah. In 2011, a drowned mountain treeshrew (Tupaia montana) in a N. rajah pitcher was reported. N. rajah occasionally traps other small vertebrates, including frogs, lizards and even birds, although these cases probably involve sick animals, or those seeking shelter or water in the pitcher, and certainly do not represent the norm.{{rp|33}} Insects, and particularly ants, comprise the majority of prey in both aerial and terrestrial pitchers.

File:Rattus baluensis visiting Nepenthes rajah.png feeding on nectar from a pitcher of N. rajah]]

Nepenthes rajah has evolved a mutualistic relationship with mountain treeshrews (Tupaia montana) in order to collect their droppings. The inside of the reflexed lid exudes a sweet nectar. The distance from the pitcher mouth to the exudate is the same as the average body length of the mountain treeshrew. As it feeds, the treeshrew defecates, apparently as a method of marking its feeding territory. It is thought that in exchange for providing nectar, the faeces provide the plant with the majority of the nitrogen it requires.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}}Walker, M. 2010. [http://news.bbc.co.uk/earth/hi/earth_news/newsid_8552000/8552157.stm Giant meat-eating plants prefer to eat tree shrew poo]. BBC Earth News, March 10, 2010.Clarke, C., J.A. Moran & L. Chin 2010. Mutualism between tree shrews and pitcher plants: perspectives and avenues for future research. Plant Signaling & Behavior 5(10): 1187–1189. {{doi|10.4161/psb.5.10.12807}}Clarke, C. & J.A. Moran 2011. Incorporating ecological context: a revised protocol for the preservation of Nepenthes pitcher plant specimens (Nepenthaceae). Blumea 56(3): 225–228. {{doi|10.3767/000651911X605781}}Davies, E. 2012. [https://www.bbc.co.uk/nature/20273855 David Attenborough's life lessons]. BBC Nature Features, November 15, 2012. In N. lowii, N. macrophylla and N. ;rajah, the colour of the lower lid surface corresponds to visual sensitivity maxima of the mountain treeshrew in the green and blue wavebands, making the lid underside stand out against adjacent parts of the pitcher.Moran, J.A., C. Clarke, M. Greenwood & L. Chin 2012. Tuning of color contrast signals to visual sensitivity maxima of tree shrews by three Bornean highland Nepenthes species. Plant Signaling & Behavior 7(10): 1267–1270. {{doi|10.4161/psb.21661}} Of the three species, N. rajah shows the tightest 'fit', particularly in the green waveband.

In 2011, it was reported that N. rajah has a similar mutualistic relationship with the summit rat (Rattus baluensis).Wells, K., M.B. Lakim, S. Schulz & M. Ayasse 2011. Pitchers of Nepenthes rajah collect faecal droppings from both diurnal and nocturnal small mammals and emit fruity odour. Journal of Tropical Ecology 27(4): 347–353. {{doi|10.1017/S0266467411000162}}Greenwood, M., C. Clarke, C.C. Lee, A. Gunsalam & R.H. Clarke 2011. A unique resource mutualism between the giant Bornean pitcher plant, Nepenthes rajah, and members of a small mammal community. PLoS ONE 6(6): e21114. {{doi|10.1371/journal.pone.0021114}} Whereas the mountain treeshrew visits pitchers during daylight hours, the summit rat is primarily active at night; this may be an example of resource partitioning. Daily scat deposition rates were found to be similar for both mammalian species.

{{multiple image

| align = center

| footer = Left: Mammal visitation rates to N. rajah pitchers. White bars show visitation by T. montana and black bars by R. baluensis. The horizontal bar below the x-axis indicates photoperiod, white representing daylight hours and black nighttime.

Right: Scat deposition rates to N. rajah pitchers. Black squares show the mean number of T. montana droppings found within pitchers; white squares show the mean number found outside pitchers. Black and white diamonds show the same for R. baluensis.

| image1 = Mammal visitation rates to Nepenthes rajah pitchers.png

| width1 = 357

| image2 = Scat deposition rates to Nepenthes rajah pitchers.png

| width2 = 350

}}

{{Main|Nepenthes infauna}}

File:Culex rajah & Toxorhynchites rajah.PNG

Although Nepenthes are most famous for trapping and digesting animals, their pitchers play host to a large number of other organisms (known as infauna). These include fly and midge larvae, spiders (crab spiders such as Henriksenia labuanica), mites, ants, and even a species of crab, Geosesarma malayanum. The most common and conspicuous predators found in pitchers are mosquito larvae, which consume large numbers of other larvae during their development. Many of these animals are so specialised that they cannot survive anywhere else, and are referred to as nepenthebionts.Beaver 1979, pp. 1–10.

The complex relationships between these various organisms are not yet fully understood. The question of whether infaunal animals "steal" food from their hosts, or whether they are involved in a mutually beneficial (symbiotic) association has yet to be investigated experimentally and is the source of considerable debate. Clarke suggests that mutualism is a "likely situation", whereby "the infauna receives domicile, protection and food from the plant, while in return, the infauna helps to break down the prey, increase the rate of digestion and keep bacterial numbers low".{{rp|42-43}}

As the size and shape of Nepenthes pitchers vary greatly between species, but little within a given taxon, it is not surprising that many infaunal organisms are specially adapted to life in only the traps of particular species. N. rajah is no exception, and in fact has two mosquito taxa named after it. Culex (Culiciomyia) rajah and Toxorhynchites (Toxorhynchites) rajah were described by Masuhisa Tsukamoto in 1989, based on larvae collected in pitchers of N. rajah on Mount Kinabalu three years earlier.Tsukamoto 1989, p. 216. The two species were found to live in association with larvae of Culex (Lophoceraomyia) jenseni, Uranotaenia (Pseudoficalbia) moultoni and an undescribed taxon, Tripteroides (Rachionotomyia) sp. No. 2. Concerning C. rajah, Tsukamoto noted that the "body surface of most larvae are covered in Vorticella-like protozoa".Tsukamoto 1989, p. 220.

File:Nep rajah27.jpg

Another species, Culex shebbearei, has been recorded as an infaunal organism of N. rajah. The original 1931 record by F. W. EdwardsEdwards 1931, pp. 25–28. is based on a collection by H. M. Pendlebury in 1929 from a plant growing on Mount Kinabalu. However, Tsukamoto notes that in light of new information on these species, "it seems more likely to conclude that the species [C. rajah] is a new species which has been misidentitied as C. shebbearei for a long time, rather than to think that both C. shebbearei and C. rajah n. sp. are living in pitchers of Nepenthes rajah on Mt. Kinabalu".

Not all interactions between Nepenthes and fauna are beneficial to the plant. N. rajah is sometimes attacked by insects which feed on its leaves and damage substantial portions of the lamina. Monkeys and tarsiers occasionally rip pitchers open to feed on their contents.Burbidge 1880.

{{See also|#Timeline}}

Due to its size, unusual morphology and striking coloration, N. rajah is a popular insectivorous plant. However, it remains a little-known species outside the field of carnivorous plants. Due to its specialised growing requirements, it is not a suitable houseplant and, as such, is only cultivated by hobbyists and professional growers. Its reputation for producing some of the most magnificent pitchers in the genus dates back to the late 19th century.Masters, M.T. 1872. [https://www.biodiversitylibrary.org/page/26129152 The cultivated species of Nepenthes.] The Gardeners' Chronicle and Agricultural Gazette 1872(16): 540–542. N. rajah was first collected by Hugh Low on Mount Kinabalu in 1858. It was described the following year by Joseph Dalton Hooker, who named it after James Brooke, the first White Rajah of Sarawak. The description was published in The Transactions of the Linnean Society of London.Hooker 1859.

Spenser St. John wrote an account of his encounter with N. rajah on Mount Kinabalu in his 1862 Life in the Forests of the Far East.St. John 1862, pp. 324, 334. N. rajah was first collected for the Veitch Nurseries by Frederick William Burbidge in 1878, during his second trip to Borneo.Phillipps & Lamb 1996, p. 20. Shortly after being introduced into cultivation in 1881, N. rajah proved very popular among wealthy Victorian horticulturalists and became a much sought-after species. A note in The Gardeners' Chronicle of 1881 mentions the Veitch plant as follows: "N. rajah at present is only a young Rajah, what it will become was lately illustrated in our columns...".[Anonymous] 1881. A year later, young N. rajah plants were displayed at the Royal Horticultural Society's annual show for the first time.Phillipps & Lamb 1996, p. 22. The specimen exhibited at the show by the Veitch Nurseries, the first of this species to be cultivated in Europe, won a first class certificate.Phillipps & Lamb 1996, p. 21. In Veitch's catalogue for 1889, N. rajah was priced at £2.2s per plant.Phillipps & Lamb 1996, p. 18.

Dwindling interest in Nepenthes at the turn of the century saw the demise of the Veitch Nurseries and consequently the loss of several species and hybrids in cultivation, including N. northiana and N. rajah. By 1905, the final N. rajah specimens from the Veitch nurseries were gone, as the cultural requirements of the plants proved too difficult to reproduce. The last surviving N. rajah in cultivation at this time was at the National Botanic Gardens at Glasnevin in Ireland, however this soon perished also.

File:Hugh Low.png|Hugh Low

File:Nepenthes rajah Life in the forests of the Far East.png|In Life in the Forests of the Far East in 1862

File:First Nepenthes rajah cultivated in Europe.jpg|The first N. rajah plant cultivated in Europe, in The Garden, 1882

{{hidden|ta1=left|Click [show] to view a list of early publications, illustrations, and collections of Nepenthes rajah.|Early publications: Transact. Linn. Soc., XXII, p. 421 t. LXXII (1859); MIQ., Ill., p. 8 (1870); HOOK. F., in D.C., Prodr., XVII, p. 95 (1873); MAST., Gard. Chron., 1881, 2, p. 492 (1881); BURB., Gard. Chron., 1882, 1, p. 56 (1882); REG., Gartenfl., XXXII, p. 213, ic. p. 214 (1883); BECC., Mal., III, p. 3 & 8 (1886); WUNSCHM., in ENGL. & PRANTL, Nat. Pflanzenfam., III, 2, p. 260 (1891); STAPF, Transact. Linn. Soc., ser. 2, bot., IV, p. 217 (1894); BECK, Wien. Ill. Gartenz., 1895, p. 142, ic. 1 (1895); MOTT., Dict., III, p. 451 (1896); VEITCH, Journ. Roy. Hort. Soc., XXI, p. 234 (1897); BOERL., Handl., III, 1, p. 54 (1900); HEMSL., Bot. Mag., t. 8017 (1905); Gard. Chron., 1905, 2, p. 241 (1905); MACF., in ENGL., Pflanzenr., IV, 111, p. 46 (1908); in BAIL., Cycl., IV, p. 2129, ic. 2462, 3 (1919); MERR., Bibl. Enum. Born., p. 284 (1921); DANS., Trop. Nat., XVI, p. 202, ic. 7 (1927).

Early illustrations: Transact. Linn. Soc., XXII, t. LXXII (1859) optima; Gard. Chron., 1881, 2, p. 493 (1881) bona, asc. 1; Gartenfl., 1883, p. 214 (1883) bona, asc. 1; Wien. Ill. Gartenfl., 1895, p. 143, ic. 1 (1895) asc. 1; Journ. Roy. Hort. Soc., XXI, p. 228 (1897) optima; Bot. Mag., t. 8017 (1905) optima; BAIL., Cycl., IV, ic. 2462, 3 (1919) asc. 1; Trop. Nat., XVI, p. 203 (1927) asc. 1.

Early collections: North Borneo. Mt. Kinabalu, IX 1913, Herbarium of the Sarawak Museum (material without flowers or fruits); Marai-parai Spur, 1-4 XII 1915, Clemens 11073, Herbarium Bogoriense, the Herbarium of the Buitenzorg Botanic Gardens (male and female material); 1650 m, 1892, Haviland 1812/1852, Herbarium of the Sarawak Museum (male and female material).}}

N. rajah is relatively well known in Malaysia, especially its native Sabah. The species is used to promote Sabah's Kinabalu National Park and features on postcards from the region. N. rajah has appeared on the covers of Nepenthes publications, including Nepenthes of Mount Kinabalu (Kurata, 1976) and Nepenthes of Borneo (Clarke, 1997), both published in Kota Kinabalu. In 1996, Malaysia issued a series of four postage stamps including one of N. rajah.Ellis 2000.{{efn|The stamp has been assigned a unique identification number in two popular stamp numbering systems: Scott #580 and Yvert #600.}} N. rajah was featured in the first episode of Kingdom of Plants 3D, a natural history series presented by David Attenborough.[http://skyatlantic.sky.com/kingdom-of-plants/kingdom-of-plants-episode-1-life-in-the-wet-zone Kingdom of Plants: Episode 1 - Life in the Wet Zone]. Sky Atlantic HD.

N. rajah is restricted to Mount Kinabalu and neighbouring Mount Tambuyukon, both in Kinabalu National Park, Sabah, Malaysian Borneo.{{rp|123}} Mount Kinabalu is a massive granitic dome structure that is geologically young and formed from the intrusion and uplift of a granitic batholith. At {{cvt|4095.2|m|ft}}, it is by far the tallest mountain on the island of Borneo and one of the highest peaks in Southeast Asia.{{Cite web |url=http://www.sabah.gov.my/mocet/ |title=Sabah Ministry of Tourism, Culture and Environment Homepage |access-date=2006-05-02 |archive-url=https://web.archive.org/web/20060423041755/http://www.sabah.gov.my/mocet/ |archive-date=2006-04-23 |url-status=dead }}

N. rajah grows exclusively on serpentine soils with high concentrations of nickel and chromium, toxic to many plant species. This means that it faces less competition for space and nutrients.Adlassnig, Peroutka, Lambers & Lichtscheidl 2005.

File:Mount kinabalu 01.png|Mount Kinabalu, Borneo

File:Kinabalu ultramafic soil.png|Ultramafic outcrops (yellow) in Kinabalu National Park (green)

N. rajah has an altitudinal distribution of {{cvt|1500–2650|m|ft}} above sea level and is thus considered an (ultra) highland or Upper Montane plant.{{Cite web |url=http://homepage.univie.ac.at/christian.puff/AS_Borneo_Kinabalu.htm |title=Vegetation Zones on Mount Kinabalu |access-date=2006-04-14 |archive-url=https://web.archive.org/web/20050915121301/http://homepage.univie.ac.at/christian.puff/AS_Borneo_Kinabalu.htm |archive-date=2005-09-15 |url-status=dead }} In the upper limit of its range, night-time temperatures may approach freezing and day-time maxima rarely exceed {{cvt|25|C|F}}.{{rp|2}} Due to the night-time temperature drop, relative air humidity increases significantly, rising from 65 to 75% to over 95%. Vegetation at this height is very stunted and slow-growing due to the extreme environmental conditions that prevail. Plants are often subjected to fierce winds and driving rain, as well as exposure to intense direct sunlight. The relatively open vegetation of the upper montane forest also experiences greater fluctuations in temperature and humidity compared with lower altitudes. These changes are largely governed by the extent of cloud cover. In the absence of clouds, temperatures rise rapidly, humidity drops, and light levels may be very high. When cloud cover returns, temperatures and light levels fall, while humidity levels increase.{{rp|29}}

N. rajah is an Endangered (EN – B1+2e) on the IUCN Red List of Threatened Species. It is listed on Schedule I, Part II of the Wildlife Conservation Enactment (WCE) 1997[http://www.internationalwildlifelaw.org/Malaysia.html Wildlife Conservation Enactment 1997] and CITES Appendix I,{{cite web |url= http://www.cites.org/eng/cop/08/E-Appendices.pdf |title= APPENDICES I AND II as adopted by the Conference of the Parties |url-status= dead |archive-url= https://web.archive.org/web/20060214042229/http://cites.org/eng/cop/08/E-Appendices.pdf |archive-date= 2006-02-14 }} which prohibits commercial international trade in plants collected from the wild. Many plants have been removed from the wild illegally,Creek, M. 1990. {{cite web|url= http://www.carnivorousplants.org/cpn/articles/CPNv19n3_4p109_112.pdf |title=The conservation of carnivorous plants. }} Carnivorous Plant Newsletter 19(3–4): 109–112. depleting some populations within Kinabalu Park in the 1970s. This led to the CITES listing in 1981.Clarke 2001b, p. 29.

The recent advent of artificial tissue culture, or more specifically in vitro, technology in Europe and the United States has meant that plants can be produced in large numbers and sold at relatively low prices (~US$20–$30 in the case of N. rajah). In vitro propagation refers to production of whole plants from cell cultures derived from explants (generally seeds). This technology has largely removed the incentive for collectors to travel to Sabah to collect the plant illegally, and demand for wild-collected plants has fallen.{{rp|172}} The conservationist Rob Cantley assesses the current status of plants in the wild as due mainly to habitat damage rather than actual theft of plants.{{Cite web |url=http://www.cites.org/common/com/ac/19/E19-10-1.doc |title=Nineteenth meeting of the Animals Committee. Geneva (Switzerland), 18–21 August 2003. |access-date=2006-05-15 |archive-url=https://web.archive.org/web/20060928183812/http://www.cites.org/common/com/ac/19/E19-10-1.doc |archive-date=2006-09-28 |url-status=dead }} The plants in the wild are further threatened by climate phenomena including El Niño; the 1997/98 event and subsequent drought had a catastrophic effect on the Nepenthes on Mount Kinabalu.{{cite book |last1=Clarke |first1=Charles |title=Nepenthes of Sumatra and Peninsular Malaysia |date=2001 |publisher=Natural History Publications (Borneo) |location=Kota Kinabalu |isbn=983-812-050-2}}{{rp|236}}

File:Nepenthes rajah mountain garden.jpg

The Mesilau Nature Resort near the village of Kundasang is now the only place where visitors can hope to see this species in its natural habitat.Clarke 2001b, p. 38. Daily guided tours are organised. Almost all other populations are in remote parts of Kinabalu National Park, off-limits to tourists.

Other localities include the Marai Parai plateau, Mesilau East River near Mesilau Cave, the Upper Kolopis River, and the eastern slope of Mount Tambuyukon.Kurata 1976, p. 64–65. On Pig Hill, N. rajah grows at 1950–2320 mAdam, J.H., C.C. Wilcock & M.D. Swaine 1992. {{cite web |url= http://myais.fsktm.um.edu.my/8918/1/10.pdf |title= The ecology and distribution of Bornean Nepenthes. |url-status= dead |archive-url= https://web.archive.org/web/20110722233248/http://myais.fsktm.um.edu.my/8918/1/10.pdf |archive-date= 2011-07-22 }} Journal of Tropical Forest Science 5(1): 13–25. and is sympatric with N. burbidgeae, N. tentaculata, and the natural hybrid N. × alisaputrana.Thong, J. 2006. {{cite web |url= http://www.vcps.au.com/journals/82_Dec_2006Public.pdf |title= Travels around North Borneo – Part 2. |access-date= 2011-07-04 |archive-url= https://web.archive.org/web/20110707185406/http://www.vcps.au.com/journals/82_Dec_2006Public.pdf |archive-date= 2011-07-07 |url-status= dead }} Victorian Carnivorous Plant Society Journal 82: 6–12.

{{main|List of Nepenthes natural hybrids}}

File:Nep rajah hybrid.jpg

N. rajah hybridises with all the other Nepenthes species with which it is sympatric. Charles Clarke notes that the pollen of N. rajah can be transported as much as {{cvt|10|km|mi}}.{{rp|143}}[http://lhnn.proboards.com/index.cgi?action=display&board=pforum&thread=3061&page=1 A rare find: N. rajah nat. hybrid]. Flora Nepenthaceae.

{{See also|Nepenthes#Cultivation}}

File:2005-12-18 N rajah 034.jpg

N. rajah is a montane species or "highlander", growing at altitudes ranging from 1500 to 2650 m. As such, it requires warm days, with temperatures ranging (ideally) from approximately 25 to 30 °C,[https://web.archive.org/web/20020811093531/http://www.cpzine.com/article.aspx?cid=13&y=2002&m=6&d=21 On the Cultivation of Nepenthes rajah] and cool nights, with temperatures of about 10 to 15 °C. It needs a fairly humid environment to grow well. Values in the region of 75% R.H. have been recommended. In its natural habitat, N. rajah grows in open areas, exposed to direct sunlight. In the tropics, growers can utilise natural sunlight as a source of illumination. In temperate zones, a photoperiod of 12 hours, comparable to that of the tropics, is recommended. The plant produces an extensive root system, requiring a wide pot. The species can tolerate hard water though purified water is recommended by specialists.D'Amato 1998, p. 7. From trials carried out by a commercial nursery,micronutrient solutions appear to be beneficial. Actual fertilizers (containing NPK) were found to be damaging.{{Cite web |url=http://borneoexotics.com/aspx/guides.aspx |title=Nepenthes Cultivation and Growing Guides |access-date=2006-01-22 |archive-url=https://web.archive.org/web/20070721034030/http://www.borneoexotics.com/aspx/guides.aspx |archive-date=2007-07-21 |url-status=dead }}

File:Kinabalu water fall N. rajah 11.JPG

Several stories have been told about the species. One such example is the famous legend that N. rajah grows exclusively in the spray zones of waterfalls, on ultramafic soils. Although it is true that they will grow in such places, N. rajah is certainly not found solely in the spray zones of waterfalls. It is likely that the misconception was popularised by Shigeo Kurata's 1976 book Nepenthes of Mount Kinabalu, in which he states that "N. rajah is rather fond of wet places like swamps or the surroundings of a waterfall". Some N. rajah plants do indeed grow near waterfalls (as noted by Hugo Steiner, 2002) "providing quite a humid microclimate". Another myth is that it occasionally catches small monkeys and other large animals in its pitchers. Such tales can probably be explained as rodents being mistaken for other species.D'Amato 1998, XV.

{{notelist}}

{{Reflist|24em}}

{{Refbegin|33em}}

• [Anonymous] 1881. [https://www.biodiversitylibrary.org/page/26126548 Messrs. Veitch's Nepenthes-house.] The Gardeners' Chronicle, new series, 16(410): 598–599.
• [Anonymous] 1883. [https://www.biodiversitylibrary.org/page/26130371 Mr. A. E. Ratcliff's Nepenthes]. The Gardeners' Chronicle 20(497): 18–19.
• [Anonymous] 1884. [https://www.biodiversitylibrary.org/page/26109896 Nepenthes Rajah.] The Gardeners' Chronicle, new series, 21(524): 52.
• [Anonymous] 1884. [https://www.biodiversitylibrary.org/page/26109960 Nepenthes Rajah.] The Gardeners' Chronicle, new series, 21(526): 116.
• [Anonymous] 1887. [https://www.biodiversitylibrary.org/page/26088044 Nepenthes at Messrs. Veitch's, Chelsea.] The Gardeners' Chronicle, series 3, 2(41): 438.
• [Anonymous] 2006. [http://www.spiegel.de/wissenschaft/erde/0,1518,440010,00.html Pflanze verdaut Maus.] Spiegel Online 29 September 2006.
• "T. B." 1881. [https://www.biodiversitylibrary.org/page/26126521 Home correspondence. Nepenthes Rajah.] The Gardeners' Chronicle, new series, 16(409): 571.
• Adam, J. H. & C. C. Wilcock 1992. A new natural hybrid of Nepenthes from Mt. Kinabalu (Sabah). Reinwardtia 11: 35–40.
• Adam, J. H. 1997. {{cite web |url= http://psasir.upm.edu.my/3641/1/Prey_Spectra_of_Bomean_Nepenthes_Species_%28Nepenthaceae%29_in.pdf |title= Prey spectra of Bornean Nepenthes species (Nepenthaceae) in relation to their habitat. }} Pertanika Journal of Tropical Agricultural Science 20(2–3): 121–134.
• Adam, J. H. & C. C. Wilcock 1998 ['1996']. Pitcher plants of Mt. Kinabalu in Sabah. The Sarawak Museum Journal 50(71): 145–171.
• Adlassnig, W., M. Peroutka, H. Lambers & I. K. Lichtscheidl 2005. [https://doi.org/10.1007%2F1-4020-4099-7_6 The roots of carnivorous plants.] Root Physiology: from Gene to Function 4: 127–140. {{ISBN|978-1-4020-4098-6}} (print) {{ISBN|978-1-4020-4099-3}} (online) {{doi|10.1007/1-4020-4099-7_6}}
• Arx, B., J. Schlauer & M. Groves 2001. [http://www.kew.org.uk/conservation/CITES_Checklists/CITESCarnivorousPlantChecklist.pdf CITES Carnivorous Plant Checklist.]{{Dead link|date=September 2018 |bot=InternetArchiveBot |fix-attempted=yes }} The Trustees of the Royal Botanic Gardens Kew. 99 pp. {{ISBN|1-84246-035-8}}
• Beaman, J.H. & C. Anderson 2004. The Plants of Mount Kinabalu: 5. Dicotyledon Families Magnoliaceae to Winteraceae. Natural History Publications (Borneo), Kota Kinabalu.
• Beaver, R. A. 1979. Fauna and food webs of pitcher plants in West Malaysia. The Malayan Nature Journal 33 (1): 1–10.
• Beck, G. 1895. Die Gattung Nepenthes. Wiener Illustrierte Gartenzeitung.
• Burbidge, F. W. 1880. The Gardens of the Sun. Murray, London. 364 pp.
• Burbidge, F.W. 1882. [https://www.biodiversitylibrary.org/page/25598125 Notes on the new Nepenthes.] The Gardeners' Chronicle, new series, 17(420): 56.
• Camilleri, T. 1998. Carnivorous Plants. Kangaroo Press, Roseville, New South Wales, Australia.
• Chan, S. 2005. [http://www.cuti.com.my/news.php?op=show_article&aid=102 Pitcher plant paradise]. The Star, August 27, 2005.
• Cheers, G. 1992. Letts Guide to Carnivorous Plants of the World. Letts of London House, Parkgate Road, London SW11 4NQ. x + 174 pp. {{ISBN|1-85238-124-8}}
• Clarke, C. M. 2001b. A Guide to the Pitcher Plants of Sabah. Natural History Publications (Borneo), Kota Kinabalu. iv + 40 pp. {{ISBN|983-812-015-4}}
• Clarke, C. 2013. [http://flashstream1.jcu.edu.au/html/tess-13-seminar-sep25.html What Can Tree Shrews Tell Us about the Effects of Climate Change on Pitcher Plants?] {{Webarchive|url=https://web.archive.org/web/20160305062128/https://flashstream1.jcu.edu.au/html/tess-13-seminar-sep25.html |date=2016-03-05 }} [video] TESS seminars, 25 September 2013.
• Clarke, C. M. & R. Kruger 2005. [http://www.carnivorousplants.org/cpn/Species/v34n2p36_41.html Nepenthes rowanae (Nepenthaceae), a remarkable species from Cape York, Australia.] Carnivorous Plant Newsletter 34 (2): 36–41.
• {{cite iucn|author= Clarke, C.|author2= Cantley, R.|author3= Nerz, J. |author4= Rischer, H.|author5= Witsuba, A.|year= 2000|page= e.T39690A10251581 |title= Nepenthes rajah|access-date= 4 February 2020}}
• Corner, E.J.H. 1996. Pitcher-plants (Nepenthes). In: K.M. Wong & A. Phillipps (eds.) [https://books.google.com/books?id=oiMIAQAAMAAJ Kinabalu: Summit of Borneo. A Revised and Expanded Edition.] The Sabah Society, Kota Kinabalu. pp. 115–121. {{ISBN|9679994740}}.
• D'Amato, P. 1998. The Savage Garden: Cultivating Carnivorous Plants. Ten Speed Press, Berkeley, California. xxii + 314 pp. {{ISBN|0-89815-915-6}}
• Danser, B. H. 1928. [http://www.omnisterra.com/botany/cp/pictures/nepenthe/dansermg/dans49.htm 38. Nepenthes Rajah.] In: The Nepenthaceae of the Netherlands Indies. Bulletin de Jardin de Botanique, Buitenzorg, Série III, 9 (3–4): 249–438.
• Douglas, J. 1884. [https://www.biodiversitylibrary.org/page/26109995 Home correspondence. Nepenthes Rajah.] The Gardeners' Chronicle, new series, 21(527): 151.
• Edwards, F. W. 1931. Mosquitoes breeding in plant pitchers. Natural History Magazine 3: 25–28.
• Ellis, R. 2000. [http://www.carnivorousplants.org/cpn/samples/Cons293Stamps.htm Carnivores on Stamps and Currency]. Carnivorous Plant Newsletter 29 (3): 90–92.
• Fairbrothers, D. E., J. J. Mabry, R. L. Scogin & B. L. Turner 1975. Annals of the Missouri Botanical Garden 62: 765–800.
• Fretwell, S. 2013. Back in Borneo for giant Nepenthes. Part 1: Mesilau Nature Reserve, Ranau. Victorian Carnivorous Plant Society Journal 107: 6–13.
• Gibson, T. C. 1983. {{cite web |url= http://www.carnivorousplants.org/cpn/articles/CPNv12n4p82_84.pdf |title= On the Cultivation of the Giant Malaysian Pitcherplant (Nepenthes rajah). }}Carnivorous Plant Newsletter 12 (4): 82–84.
• Gibson, T. C. 1988. {{cite web |url= http://www.carnivorousplants.org/cpn/articles/CPNv17n3p84.pdf |title= A Further Note on Nepenthes rajah Cultivation. }}Carnivorous Plant Newsletter 17 (3): 84.
• Hemsley, W. B. 1905. Nepenthes Rajah. Curtis's Botanical Magazine, CXXXI (4th series, I) t. 8017.
• Hooker, J. D. 1859. [https://www.biodiversitylibrary.org/page/12906432 XXXV. On the origin and development of the pitchers of Nepenthes, with an account of some new Bornean plants of that genus]. The Transactions of the Linnean Society of London 22(4): 415–424. {{doi|10.1111/j.1096-3642.1856.tb00113.x}}
• 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.
• Jebb, M. H. P. & M. R. Cheek 1997. A skeletal revision of Nepenthes (Nepenthaceae). Blumea 42 (1): 1–106.
• Johnson, N. 2013. The history of the genus Nepenthes at Kew. Planta Carnivora 35(1): 6–19.
• Kaul, R. B. 1982. Floral and Fruit Morphology of Nepenthes lowii and N. villosa, Montane Carnivores of Borneo. American Journal of Botany 69 (5): 793–803.
• Kurata, S. 1976. Nepenthes of Mount Kinabalu. Sabah National Parks Publications No. 2, Sabah National Parks Trustees, Kota Kinabalu. 80 pp.
• Kurata, S. 1984. Journal of the Insectivorous Plant Society (Japan) 35: 65.
• Malouf, P. 1995. {{cite web |url= http://carnivorousplants.org/cpn/articles/CPNv24n3p64_69.pdf |title= A visit to Kinabalu Park.}} Carnivorous Plant Newsletter 24(3): 64–69.
• Macfarlane, J. M. 1908. Nepenthaceae. A. Engler, Das Pflanzenreich IV, 111, Heft. 36: 1–91.
• Macfarlane, J.M. 1911. [https://www.biodiversitylibrary.org/page/34938989 New species of Nepenthes.] Contributions from the Botanical Laboratory of the University of Pennsylvania 3(3): 207–210. ([https://www.biodiversitylibrary.org/page/34939103 plates I–II])
• Macfarlane, J.M. 1914. Nepenthes sp.. Biological Journal of the Linnean Society, XLII.
• Masters, M.T. 1881. [https://www.biodiversitylibrary.org/page/26126442 New garden plants. Nepenthes Rajah, Hook. f.]. The Gardeners' Chronicle, new series, 16(407): 492–493.
• McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Borneo. Redfern Natural History Productions, Poole.
• {{in lang|de}} Meimberg, H. 2002. {{cite web |url= http://edoc.ub.uni-muenchen.de/1078/1/Meimberg_Harald.pdf |title= Molekular-systematische Untersuchungen an den Familien Nepenthaceae und Ancistrocladaceae sowie verwandter Taxa aus der Unterklasse Caryophyllidae s. l.. }} Ph.D. thesis, Ludwig Maximilian University of Munich, Munich.
• 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}}
• Mey, F.S. 2014. [http://carnivorousockhom.blogspot.fr/2014/02/joined-lecture-on-carnivorous-plants-of.html Joined lecture on carnivorous plants of Borneo with Stewart McPherson.] Strange Fruits: A Garden's Chronicle, February 21, 2014.
• 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.
• Nelson, E. C. The waxing of glorious rajah. Kew magazine [May 1991] 8 (2): 81–89.
• Nerz, J. & A. Wistuba 2007. Nepenthes mantalingajanensis (Nepenthaceae), eine bemerkenswerte neue Spezies aus Palawan (Philippinen). Das Taublatt 55(3): 17–25.
• {{in lang|ja}} Oikawa, T. 1992. Nepenthes Rajah Hook.f.. In: {{Nihongo||無憂草 – Nepenthes|Muyū kusa – Nepenthes}}. [The Grief Vanishing.] Parco Co., Japan. pp. 8–9.
• Phillipps, A. 1988. {{cite web |url= http://www.carnivorousplants.org/cpn/articles/CPNv17n2p55.pdf |title= A Second Record of Rats as Prey in Nepenthes rajah. }} Carnivorous Plant Newsletter 17 (2): 55.
• Phillipps, A. & A. Lamb 1996. Pitcher Plants of Borneo. Natural History Publications (Borneo), Kota Kinabalu. x + 171 pp. {{ISBN|983-812-009-X}}
• Reginald 1883. Nepenthes Rajah J. D. Hooker. Gartenflora, XXXII, p. 213.
• Saint-John, S. 1862. Life in the Forests of the Far East; or, Travels in northern Borneo. 2 volumes. London: Smith, Elder & Co. (reprinted by Oxford University Press, 1974)
• Simpson, R. 1991. Plants in peril, 15: Nepenthes rajah. Kew magazine [May 1991] 8 (2): 89–94.
• Slack, A. 1979. Nepenthes rajah. In: Carnivorous Plants. Ebury Press, London. p. 85.
• Slack, A. 1986. Insect-Eating Plants and How to Grow Them. Alphabooks, Dorset, UK. 172 pp. {{ISBN|0-906670-42-X}} (hardback) {{ISBN|0-906670-35-7}} (paperback)
• 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.
• Stace, C.A. 1980. Plant Taxonomy and Biosystematics. Arnold, London. {{ISBN|0-521-42785-1}}
• Steiner, H. 2002. Borneo: Its Mountains and Lowlands with their Pitcher Plants. Toihaan Publishing Company, Kota Kinabalu. viii + 136 pp. {{ISBN|983-40421-1-6}}
• Thorogood, C. 2010. The Malaysian Nepenthes: Evolutionary and Taxonomic Perspectives. Nova Science Publishers, New York.
• Tsukamoto, M. 1989. {{cite web |url= http://www.mosquitocatalog.org/files/pdfs/133700-0.pdf |title= Two New Mosquito Species from a Pitcher Plant of Mt. Kinabalu, Sabah, Malaysia: Culex rajah and Toxorhynchites rajah (Diptera: Culicidae) |access-date= 2009-12-01 |archive-url= https://web.archive.org/web/20110723204219/http://www.mosquitocatalog.org/files/pdfs/133700-0.pdf |archive-date= 2011-07-23 |url-status= dead }}. Japanese Journal of Tropical Medicine and Hygiene, 17 (3): 215–228.
• Yeo, J. 1996. A trip to Kinabalu Park. Bulletin of the Australian Carnivorous Plant Society, Inc. 15(4): 4–5.

{{Refend}}

{{Commons}}

{{Wikispecies}}

{{Wikisource|Description of Nepenthes rajah Hook.f. (1859)}}

{{Wikiquote}}

• [https://web.archive.org/web/20060526141837/http://www.arcbc.org.ph/arcbcweb/pdf/vol1no4/focus_pitcher_plant.pdf Focus: Rajah Brooke's Pitcher Plant]
• [http://www.fleischfressendepflanzen.de/db/species.ffp?id=435 Die Karnivoren-Datenbank: Nepenthes rajah] {{in lang|de}}

• [https://web.archive.org/web/20160921065900/http://www.cpzine.com/displayarticle.asp?cid=13&y=2002&m=6&d=21 N. rajah Cultivation Notes]
• [http://www.coloradocarnivorousplantsociety.com/Nepenthes_rajah.htm Colorado Carnivorous Plant Society: Nepenthes rajah]

• [http://www.ipni.org/ipni/idPlantNameSearch.do?id=603791-1 The International Plant Names Index: Nepenthes rajah]
• [http://www.omnisterra.com/botany/cp/pictures/nepenthe/dansermg/dans49.htm Nepenthes rajah entry from Danser's Monograph]
• [http://www.redlist.org/search/details.php?species=39690 IUCN Red List of Threatened Species: Nepenthes rajah]
• [https://www.youtube.com/watch?v=trWzDlRvv1M Video about Nepenthes rajah] from The Private Life of Plants

{{Nepenthes}}

{{Taxonbar|from=Q132973}}

Category:Carnivorous plants of Asia

Category:Coprophagous plants

rajah

Category:Endemic flora of Borneo

Category:Flora of Sabah

Category:Threatened flora of Asia

Category:Plants described in 1859

Category:Articles which contain graphical timelines

Category:Flora of Mount Kinabalu

Category:Flora of the Borneo montane rain forests