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Nepenthes rajah

2007 Schools Wikipedia Selection. Related subjects: Plants

                             iNepenthes rajah
   Large lower pitcher of Nepenthes rajah. Mount Kinabalu, Borneo.
   Large lower pitcher of Nepenthes rajah. Mount Kinabalu, Borneo.

                             Conservation status

   Endangered (EN)
                         Scientific classification

   Kingdom:  Plantae
   Division: Magnoliophyta
   Class:    Magnoliopsida
   Order:    Caryophyllales
   Family:   Nepenthaceae
   Genus:    Nepenthes
   Species:  N. rajah

                                Binomial name

   Nepenthes rajah
   Hook.f., 1859
   Borneo, showing natural range of Nepenths rajah highlighted in green.
   Borneo, showing natural range of Nepenths rajah highlighted in green.

                                  Synonyms

     * Nepenthes rajah
       auct. non Hook.f.: A. Slack, 1986
       [= Nepenthes × alisaputrana]
     * Nepenthes rajah
       auct. non Hook.f.: G.Cheers, 1992
       [= Nepenthes × kinabaluensis]

   Nepenthes rajah ( IPA pronunciation: [ni-pen'thēz ra'ja], named after
   the White Rajah of Sarawak) is an insectivorous pitcher plant species
   of the Nepenthaceae family. It has a very localised distribution, being
   restricted to Mount Kinabalu and neighbouring Mount Tambuyukon in
   Sabah, Malaysian Borneo. N. rajah is famous for the giant urn-shaped
   traps it produces, which can grow up to 35 cm high and 18 cm wide.
   These are capable of holding 3.5 litres of water and in excess of 2.5
   litres of digestive fluid, making them probably the largest in the
   genus by volume. Joseph Dalton Hooker described N. rajah in 1859,
   calling it "one of the most striking vegetable productions hither-to
   discovered".

Etymology

   James Brooke
   Enlarge
   James Brooke

   Joseph Dalton Hooker described N. rajah in 1859, naming it in honour of
   Sir James Brooke, the first White Rajah of Sarawak. In the past, the
   latin name was written as Nepenthes Rajah, since it derives from a
   proper noun. However, this capitalisation is considered incorrect
   today. Rajah Brooke's Pitcher Plant is an accurate, but seldom-used
   common name. N. rajah is also sometimes called the Giant Malaysian
   Pitcher Plant 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
   referred to as the "King of Nepenthes".

Plant characteristics

   Mature plant growing among sedges
   Enlarge
   Mature plant growing among sedges

   Nepenthes rajah, like virtually all species in the genus, is a
   scrambling vine. The stem usually grows along the ground, but will
   attempt to climb whenever it comes into contact with an object that can
   support it. The stem is relatively thick (≤30 mm) and may reach up to 6
   m in length, although it rarely exceeds 3 m. N. rajah does not produce
   runners as some other species in the genus, but older plants are known
   to form basal offshoots. This is especially common in plants from
   tissue culture, where offshoots may form at a young age.

Leaves

   Leaves are produced at regular intervals along the stem. They are
   connected to the stem by sheathed structures known as petioles. A long,
   narrow tendril emanates from the end of each leaf. At the tip of the
   tendril is a small bud which, when physiologically activated, develops
   into a functioning trap. Hence, the pitchers are modified leaves and
   not specialised flowers as is often believed. The green structure most
   similar to a normal leaf is specifically known as the lamina or leaf
   blade.
   Characteristic peltate leaf attachment of N. rajah
   Enlarge
   Characteristic peltate leaf attachment of N. rajah

   The leaves of N. rajah are very 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. Three to five longitudinal veins run along
   each side of the lamina and pennate (branching) veins run towards the
   margin. The lamina is oblong to lanceolate-shaped, ≤80 cm long and
   ≤15 cm wide. The tendrils are inserted ≤5 cm below the leaf apex and
   reach a length of approximately 50 cm.

Pitchers

   All Nepenthes pitchers share several basic characteristics. Traps
   consist of the main pitcher cup, which is covered by an operculum or
   lid that prevents rainwater from entering the pitcher and displacing or
   diluting its contents. A reflexed ring of hardened tissue that
   surrounds the entrance to the pitcher is usually present and is known
   as the peristome. A pair of fringed wings runs down the front of every
   trap and these presumably serve to guide terrestrial insects into the
   pitcher's mouth. Accordingly, the wings are greatly reduced or
   completely lacking in aerial pitchers, for which flying insects
   constitute the majority of prey items.
   Terrestrial pitcher
   Enlarge
   Terrestrial pitcher

   N. rajah produces two distinct types of traps. "Lower" or "terrestrial"
   pitchers are the most common. These are very large, richly coloured and
   ovoid in shape. Exceptional specimens may be up to 40 cm high and
   capable of holding 3.5 litres of water and in excess of 2.5 litres of
   digestive fluid, although most do not exceed 200 ml. The lower pitchers
   of N. rajah are probably the largest in the genus by volume, rivaled
   only by those of N. merrilliana, N. truncata and the giant form of N.
   rafflesiana. These traps rest on the ground and are often reclined,
   leaning against surrounding objects for support. They are usually red
   to purple on the outside, whilst the inside surfaces are lime green to
   purple. This gives a striking contrast to all other parts of the plant,
   which are yellow-green. The lower pitchers of N. rajah are unmistakable
   and for this reason it is easy to distinguish it from all other Bornean
   Nepenthes species.

   Mature plants may also produce "upper" or "aerial" pitchers, which are
   much smaller, funnel-shaped and usually less colourful than the lowers.
   True upper pitchers are seldom seen, as the stems of N. rajah rarely
   attain lengths greater than a few metres before dying off and being
   replaced by off-shoots from the main rootstock.

   Upper and lower pitchers differ significantly in morphology, as they
   are specialised for attracting and capturing different prey. Pitchers
   that do not fall directly into either category are simply known as
   "intermediate" pitchers.

   The peristome of N. rajah has a highly distinctive scalloped edge and
   is greatly expanded, forming an attractive red lip around the trap's
   mouth. A series of raised protrusions, known as ribs, intersect the
   peristome, ending in short, sharp teeth that line its inner margin. Two
   fringed wings run from the tendril attachment to the lower edge of the
   peristome.

   The huge, vaulted lid of N. rajah, the largest in the genus, is another
   distinguishing characteristic of this species. It is ovate to oblong in
   shape and has a distinct keel running down the middle. The spur at the
   back of the lid is approximately 20 mm long and unbranched.

   N. rajah is noted for having very large nectar- secreting glands
   covering its pitchers. These are quite different from those of any
   other Nepenthes and are easily recognisable (see ). The inner surface
   of the pitcher, in particular, is wholly glandular, with 300 to 800
   glands/cm².
   Flowering plant of N. rajah
   Enlarge
   Flowering plant of N. rajah

Flowers

   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 80 cm,
   and sometimes even 120 cm tall. 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
   ≤8 mm long. Like all Nepenthes species, N. rajah is dioecious, which
   means that each plant produces either male or female flowers, but never
   both. Fruits are orange-brown and 10 to 20 mm long (see ).

Other characteristics

   All parts of the plant are covered in long, white hairs when young, but
   mature plants are virtually glabrous (lacking hair). This covering of
   hair is known as the indumentum.

   The colour of herbarium specimens is dark-brown in varying hues (see ).

   Little variation has been observed within natural populations of
   Nepenthes rajah and, consequently, no forms or varieties have been
   described. Furthermore, N. rajah has no published synonyms, unlike many
   other Nepenthes species, which exhibit greater variability.

Carnivory

   Drowned lizard found in a freshly opened pitcher. The animal was pulled
   out of the digestive zone for the photograph.
   Enlarge
   Drowned lizard found in a freshly opened pitcher. The animal was pulled
   out of the digestive zone for the photograph.

   Nepenthes rajah is a carnivorous plant of the pitfall trap variety. It
   is famous for occasionally trapping vertebrates and 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 Spencer
   St. John, who accompanied Hugh Low on two ascents of Mount Kinabalu. In
   1988, Anthea Phillipps and Anthony Lamb confirmed the plausability of
   this record when they managed to observe drowned rats in a large
   pitcher of N. rajah. N. rajah is also known to occasionally trap 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. Insects,
   and particularly ants, comprise the majority of prey in both aerial and
   terrestrial pitchers. Other arthropods, such as centipedes, also fall
   prey to N. rajah.

   N. rafflesiana is the only other Nepenthes species reliably documented
   as having caught mammalian prey in its natural habitat. In Brunei,
   frogs, geckos and skinks have been found in the pitchers of this
   species. The remains of mice have also been reported.

   On September 29, 2006, at the Jardin botanique de Lyon in France, a
   cultivated N. truncata was photographed containing the decomposing
   corpse of a mouse.

Interactions with animals

Pitcher infauna

   Although Nepenthes are most famous for trapping and digesting animals,
   their pitchers also play host to a large number of other organisms
   (known as infauna). These include fly and midge larvae, spiders (most
   notably the crab spider Misumenops nepenthicola), 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.

   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".

Species specific

   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. 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-live protozoa". At present, nothing is known of this species
   with regards to its adult biology, habitat, or medical importance as a
   vector of diseases. The same is true for T. rajah; nothing is known of
   its biology except that adults are not haematophagous.
   Damage caused by pests
   Enlarge
   Damage caused by pests

   Another species, Culex shebbearei, has also been recorded as an
   infaunal organism of N. rajah in the past. The original 1931 record by
   F. W. Edwards 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".

Pests

   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 remove substantial portions of the lamina. Also, monkeys and
   tarsiers are known to occasionally rip pitchers open to feed on their
   contents.

Classification

                                     Regiae Clade

              N. maxima         N. pilosa     N. clipeata
              N. oblanceolata * N. burbidgeae N. truncata
              N. veitchii       N. rajah      N. fusca
              N. ephippiata     N. boschiana  N. stenophylla
              N. klossii        N. mollis     N. lowii

               * Now considered a synonym of N. maxima.

              Distribution of the Regiae

               Distribution of the Regiae, based on Danser (1928).
               Note: it is now known that N. maxima is absent from Borneo.

   Nepenthes rajah is not generally considered to be closely related to
   any other species, due to its unusual pitcher and leaf morphology.
   However, several attempts have been made to deduce natural groupings
   within the Nepenthes genus, in order to show relationships between taxa
   below those at the genus rank, which have grouped N. rajah with other
   species thought to share certain traits with it.

Nineteenth century

   The Nepenthes were first split up in 1873, when Hooker published his
   monograph on the genus. Hooker distinguished N. pervillei from all
   other taxa based on the fact that seeds of this species lacked
   appendages that were found to be present in all other Nepenthes (though
   greatly reduced in N. madagascariensis) and subsequently placed it in
   the monotypic subgenus Anurosperma (Latin: anuro: without nerves,
   sperma: seeds). All other species were subsumed in the second subgenus,
   Eunepenthes (Latin: eu: true; "true" Nepenthes).

   A second attempt to establish a natural subdivision within the genus
   was made in 1895 by Günther von Mannagetta und Lërchenau Beck in his
   Monographische Skizze (German: Monographische: monographic, Skizze:
   sketch). Beck kept the two subgenera created by Hooker, but divided
   Eunepenthes into three subgroups: Retiferae, Apruinosae and Pruinosae.
   N. rajah formed part of the Apruinosae (Latin: pl. of apruinosa: not
   frosted). Most contemporary taxonomists agree that Beck's groupings
   have little, if any, taxonomical value, as they were based on arbitrary
   traits not suitable for use as a basis for classification.

Twentieth century

   Nepenthes taxonomy was once again revised in 1908 by John Muirhead
   Macfarlane in his own monograph. Oddly, Macfarlane did not name the
   groups he distinguished. His revision is often not considered to be a
   natural division of the genus.

   In 1928, B. H. Danser published his monograph, in which he divided
   Nepenthes into six clades, based on observations of herbarium material.
   The clades were: the Vulgatae, Montanae, Nobiles, Regiae, Insignes and
   Urceolatae. Danser placed N. rajah in the Regiae (Latin: pl. of rēgia:
   royal). The Regiae clade as proposed by Danser is shown in the table to
   the right.

   Most of the species in this clade are large plants with petiolate
   leaves, an indumentum of coarse reddish-brown hairs, raceme-like
   inflorescence, and funnel-shaped (infundibulate) upper pitchers. All
   bear a characteristic appendage on the lower surface of the lid near
   the apex. With the exception of N. lowii, the Regiae all have a mostly
   flattened or expanded peristome. The majority of species comprising
   Regiae are endemic to Borneo. Based on current understanding of the
   genus, Regiae appears to reflect the relationships of its members quite
   well, although the same cannot be said for the other clades. Despite
   this, Danser's classification was undoubtedly a great improvement on
   previous attempts.

   The taxonomical work of Danser (1928) was revised by Hermann Harms in
   1936. Harms divided Nepenthes into three subgenera: Anurosperma
   Hooker.f. (1873), Eunepenthes Hooker.f. (1873) and Mesonepenthes Harms
   (1936) (Latin: meso: middle; "middle" Nepenthes). The Nepenthes species
   found in the subgenera Anurosperma and Mesonepenthes differ from those
   in the Vulgatae, where Danser had placed them. Harms included N. rajah
   in the subgenus Eunepenthes together with the great majority of other
   Nepenthes; Anurosperma was a monotypic subgenus, while Mesonepenthes
   contained only three species. He also created an additional clade, the
   Distillatoriae (after N. distillatoria).

Glandular morphology

   In 1976, Shigeo Kurata proposed that glands present in Nepenthes
   pitchers were unique to individual species and could be used to
   distinguish between closely-related taxa or even be used as a basis for
   their classification. Kurata studied two types: the nectar glands of
   the lid and the digestive glands of the inside of the pitcher cup. He
   divided the latter into the "lower", "upper" and "middle" parts (see ).
   Although this novel approach shed additional light on the similarities
   between certain species, it has since been largely abandoned by
   taxonomists and botanists specialising in the genus, in favour of
   classical taxonomical nomenclature based on a description of
   morphological characters.
      Distribution of phenolic compounds and leucoanthocyanins in N. ×
                  alisaputrana, N. burbidgeae and N. rajah
   Taxon

                                      1

                                      2

                                      3

                                      4

                                      5

                                      6

                                      7

                                      8

   Specimen
   N. × alisaputrana

                                      +

                                     ++

                                     3+

                                     3+

                                      +

                                     ++

                                     3+

                                      +

   J2442
   in vitro

                                      +

                                     ++

                                     3+

                                     3+

                                      +

                                     ++

                                      +

                                      +

   N. burbidgeae

                                     3+

                                     ++

                                     3+

                                     3+

                                      -

                                      +

                                      -

                                      -

   J2484
   N. rajah

                                      -

                                      -

                                      +

                                     ±

                                     ++

                                     ++

                                     3+

                                      +

   J2443
   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, J = Jumaat Source: OnLine Journal of Biological Sciences 2
   (9): 623-625, 2002. PDF

Biochemical analysis

   More recently, biochemical analysis has been used as a means to
   determine cladistical relationships between Nepenthes species. In 1975,
   David E. Fairbrothers et al. first suggested a link between chemical
   properties and certain morphological groupings, based on the theory
   that morphologically similar plants produce chemical constituents with
   similar therapeutic effects.

   In 2002, phytochemical screening and analytical chromatography were
   used to study the presence of phenolic compounds and leucoanthocyanins
   in several naturally-occurring hybrids and their putative parental
   species (including N. rajah) from Sabah and Sarawak. 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. rajah and N. burbidgeae are shown
   in the table to the left. A specimen of N. × alisaputrana grown from
   tissue culture ( in vitro) was also tested.

   Phenolic and ellagic acids were undetected in N. rajah, while
   concentrations of kaempferol were found to be very weak.
   Chromatographic patterns of the N. × alisaputrana samples studied
   showed complementation of its putative parental species.

   Myricetin was found to be absent from all studied taxa. This agrees
   with the findings of previous authors ( Som, 1988; Jay and Lebreton,
   1972) and suggests that the absence of a widely distributed compound
   like myricetin among the Nepenthes examined might provide "additional
   diagnostic information for these six species".

Sequencing

   Several proteins and nucleotides of N. rajah have been either partially
   or completely sequenced. These are as follows:
     * translocated tRNA-Lys (trnK) pseudogene (DQ007139)
     * trnK gene & maturase K (matK) gene (AF315879)
     * trnK gene & maturase K (matK) gene (AF315880)
     * maturase K (AAK56010)
     * maturase K (AAK56011)

Related species?

   Recently, a striking new species of Nepenthes was discovered in the
   Philippines by Andreas Wistuba. Temporarily dubbed N. sp. Palawan 1, it
   bears a close resemblance to N. rajah in terms of pitcher and leaf
   morphology. Due to the geographic distance separating the two species,
   it is unlikely they are in any way closely related. Thus, this case
   might represent an example of convergent evolution, whereby two
   organisms not closely related independently acquire similar
   characteristics while evolving in separate, but comparable, ecosystems.
   N. sp. Palawan 1 is currently being formally described.

History and popularity

   Hugh Low
   Enlarge
   Hugh Low
   Joseph Dalton Hooker
   Enlarge
   Joseph Dalton Hooker

   Due to its size, unusual morphology and striking colouration, N. rajah
   has always been a very popular and highly sought-after insectivorous
   plant. However, despite its popularity amongst pitcher plant
   enthusiasts, it is fair to say that Nepenthes rajah remains a little
   known species outside the field of carnivorous plants. Due to its
   specialised growing requirements, it is not a suitable candidate for a
   houseplant and, as such, is only cultivated by a relatively small
   number of hobbyists and professional growers worldwide. This being the
   case, N. rajah is nonetheless probably the most famous of all pitcher
   plants. Its reputation for producing some of the most magnificent
   pitchers in the genus dates back to the late 18th century.

Early history

   N. rajah was discovered by Hugh Low on or just before 7 March 1851,
   during Low's first ascent of Mount Kinabalu. It was described eight
   years later by Hooker, who named it after James Brooke, the first White
   Rajah of Sarawak. The species was first collected for the Veitch
   Nurseries by Frederick William Burbidge in 1878, during his second trip
   to Borneo. Shortly after being introduced into cultivation in 1881, N.
   rajah proved very popular among wealthy Victorian horticulturalists and
   it was a much sought-after species. A year later, young N. rajah plants
   were displayed at the Royal Horticultural Society's annual show for the
   first time. During this time, interest in Nepenthes had reached its
   peak. The Garden reported that Nepenthes were being propagated by the
   thousands to keep up with the European demand.

   However, 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 located at the National Botanic Gardens at Glasnevin in
   Ireland, however this soon perished also. It would be many years until
   N. rajah was reintroduced into cultivation.

Recent popularity

   In recent years there has been renewed interest in Nepenthes worldwide.
   Much of the plants' current popularity can probably be attributed to
   Shigeo Kurata, whose book Nepenthes of Mount Kinabalu (1976), which
   featured the best colour photography of Nepenthes to date, did much to
   bring attention to these unusual plants.

   Not surprisingly, N. rajah is a relatively well known plant in
   Malaysia, especially its native Sabah. The species is often used to
   promote Sabah, and specifically Kinabalu National Park, as a tourist
   destination, and features prominently on postcards from the region. N.
   rajah has appeared on the covers of several popular Nepenthes
   publications, including Nepenthes of Mount Kinabalu ( Kurata, 1976) and
   Nepenthes of Borneo ( Clarke, 1997), both published in Kota Kinabalu,
   Malaysia. On April 6, 1996, Malaysia issued a series of four postage
   stamps depicting some of its more famous Nepenthes species. Two 30 ¢
   stamps, featuring N. macfarlanei and N. sanguinea, as well as two 50¢
   stamps, depicting N. lowii and N. rajah, were released. The N. rajah
   stamp has been assigned a unique identification number in two popular
   stamp numbering systems. These are Scott #580 and Yvert #600.
   Curiously, the peltate leaf attachment that is so characteristic of
   this species is not shown.

Ecology

   Mount Kinabalu, Borneo
   Enlarge
   Mount Kinabalu, Borneo

Kinabalu

   Nepenthes rajah has a very localised distribution, being restricted to
   Mount Kinabalu and neighbouring Mount Tambuyukon, both located in
   Kinabalu National Park, Sabah, Malaysian Borneo. Mount Kinabalu is a
   massive granitic dome structure that is geologically young and formed
   from the intrusion and uplift of a granitic batholith. At 4095.2 m, it
   is by far the tallest mountain on the island of Borneo and one of the
   highest peaks in Southeast Asia. The lower slopes of the mountain are
   mainly composed of sandstone and shale, transformed from marine sand
   and mud about 35 million years ago. Intrusive ultramafic ( serpentine)
   rock was uplifted with the core of the batholith and forms a collar
   around the mountain. It is on these ultramafic soils that the flora of
   Mount Kinabalu exhibits the greatest levels of endemicity and many of
   the area's rarest species can be found here.
   Ultramafic outcrops (yellow) in Kinabalu National Park (green)
   Enlarge
   Ultramafic outcrops (yellow) in Kinabalu National Park (green)

Substrate

   N. rajah seems to grow exclusively on serpentine soils containing high
   concentrations of nickel and chromium, which are toxic to many plant
   species. Its tolerance of these, therefore, means that it can grow in
   an ecological niche where it does not face much competition for space
   or nutrients. These soils are also rich in magnesium and are slightly
   alkaline as a result. They often form a relatively thin layer over a
   base of ultramafic rock and are thus known as ultramafic soils.
   Ultramafic soils are thought to cover approximately 16% of Kinabalu
   National Park. These soils have high levels of endemicity in many
   taxonomic groups, not least the Nepenthes. Four species in the genus,
   including N. rajah, can only be found within the boundaries of the
   park.

   N. rajah usually grows in open, grassy clearings on old land slips and
   flat ridge tops, particularly in areas of seeping ground water, where
   the soil is loose and permanently moist. Although these sites can
   receive very high rainfall, excess water drains away quickly,
   preventing the soil from becoming waterlogged. N. rajah can often be
   found growing in grassy undergrowth, especially among sedges.
   Temperature and humidity readings taken along the "Nepenthes rajah
   Nature Trail" at around 11 am during an overcast sky
   Enlarge
   Temperature and humidity readings taken along the "Nepenthes rajah
   Nature Trail" at around 11 am during an overcast sky

Climate

   N. rajah has an altitudinal distribution of 1500–2650 m a.s.l. and is
   thus considered an (ultra) highland or Upper Montane plant. In the
   upper limit of its range, night-time temperatures may approach freezing
   and day-time maximums rarely exceed 25 ℃. Due to the night-time
   temperature drop, relative air humidity increases significantly, rising
   from 65-75% 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. Average annual precipitation in
   this region is around 3000 mm.

Conservation status

Endangered species

   Nepenthes rajah is classified as Endangered (EN – B1+2e) on the IUCN
   Red List of Threatened Species . It is also listed on Schedule I, Part
   II of the Wildlife Conservation Enactment (WCE) 1997 and, perhaps more
   notably, CITES Appendix I, which prohibits international trade in
   plants collected from the wild. However, due to its popularity among
   collectors, many plants have been removed from the wild illegally, even
   though the species' distribution lies entirely within the bounds of
   Kinabalu Park. This led to some populations being severely depleted by
   over-collection in the 1970s and eventually resulted in the species'
   inclusion in CITES Appendix I in 1981. Along with N. khasiana, it is
   one of only two species in the genus to feature on this list.

   This being the case, however, the short-term future of N. rajah seems
   to be relatively secure and it would perhaps be more accurately
   classified as Vulnerable (VU) or, taking into account protected
   populations in National Parks, Lower Risk conservation dependent (LR
   (cd)). This agrees with the conservation status of N. rajah according
   to the World Conservation Monitoring Centre (WCMC), under which it is
   also considered Vulnerable. Furthermore, the species was originally
   treated as Vulnerable (V) by the IUCN prior to the introduction of the
   1994 threat categories.

   Although N. rajah has a restricted distribution and is often quoted as
   a plant in peril, it is not rare in the areas where it does grow and
   most populations are now off-limits to visitors and lie in remote parts
   of Kinabalu National Park. Furthermore, N. rajah has a distinctive leaf
   shape making it difficult to illegally ship abroad even if the pitchers
   are removed, as an informed customs official should be able to identify
   it.

   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, to a large extent, removed the
   incentive for collectors to travel to Sabah to collect the plant
   illegally, and demand for wild-collected plants has fallen considerably
   in recent years.

   Rob Cantley, a prominent conservationist and artificial propagator of
   Nepenthes plants, assesses the current status of plants in the wild as
   follows:

   "This species grows in at least 2 distinct sub-populations, both of
   which are well protected by Sabah National Parks Authority. One of the
   populations grows in an area public access to which is strictly
   prohibited without permit. However, there has been a decline in
   population of mature individuals in the better known and less patrolled
   site. This is largely due to damage to habitat and plants by careless
   visitors rather than organised collection of plants. Nepenthes rajah
   has become common in cultivation in recent years as a result of the
   availability of inexpensive clones from tissue culture. I believe that
   these days commercial collection of this species from the wild is
   negligible."

   This being the case, however, it appears that the genetic variability
   of cultivated N. rajah plants is very small, as all commercially
   available tissue-cultured plants are thought to belong to just four
   clones originating from the Royal Botanic Gardens, Kew in London,
   England.

   However, illegal collection is not the only threat facing plants in the
   wild. The El Niño climatic phenomenon of 1997/98 had a catastrophic
   effect on the Nepenthes species on Mount Kinabalu. The dry period that
   followed severely depleted some natural populations. Forest fires broke
   out in 9 locations in Kinabalu Park, covering a total area of 25 square
   kilometres and generating large amounts of smog. During the El Niño
   period, many plants were temporarily transferred to the park nursery to
   save at least some individuals. These were later replanted in the
   "Nepenthes Garden" in Mesilau (see below). In spite of this, N. rajah
   was one of the less affected species and relatively few plants perished
   as a result. Since then, Ansow Gunsalam has established a nursery close
   to the Mesilau Lodge at the base of Kinabalu Park to protect the
   endangered species of that area, including N. rajah.
   Plant on display at the Kinabalu "Mountain Garden"
   Enlarge
   Plant on display at the Kinabalu "Mountain Garden"

Restricted distribution

   The newly opened Mesilau Nature Resort, which lies near the golf course
   behind the village of Kundasang, is now the only place where regular
   visitors can hope to see this spectacular species in its natural
   habitat. Here, several dozen N. rajah plants grow near the top of a
   steep landslide. Both young and mature plants are present, some with
   sizable pitchers approaching 35 cm in height (see ). Due to their great
   size, several plants are thought to be over 100 years old. Daily guided
   tours are organised to the "Nepenthes Garden" where these plants are
   located. The "Nepenthes rajah Nature Trail", as it is called, is
   subject to a fee and operates daily from 9:00am to 4:00pm. Almost all
   other natural populations of this species occur in remote parts of
   Kinabalu National Park, which are off-limits to tourists. Visitors to
   the park can also see a medium-sized N. rajah plant that is on display
   in the nursery adjoining the "Mountain Garden" at Kinabalu Park
   Headquarters.

   Other known localities of wild N. rajah populations include the Marai
   Parai plateau, Mesilau East River near Mesilau Cave, Upper Kolopis
   River and eastern slope of Mount Tambuyukon.

Natural hybrids

   N. tentaculata × N. rajah
   Enlarge
   N. tentaculata × N. rajah

   Nepenthes rajah is known to hybridise with several other species with
   which it is sympatric. It seems to flower at any time of year and for
   this reason it hybridises relatively easily. Charles Clarke also notes
   that "N. rajah, more than any other species, appears to have been
   successful in having its pollen transported over considerable
   distances. Consequently, a number of putative N. rajah hybrids exist
   without the parent plant growing nearby". However, it appears that the
   limit as to how far pollen can be transported is approximately 10 km.
   Hybrids between N. rajah and all other Nepenthes species on Mount
   Kinabalu (with the exception of N. lowii) have been recorded. Due to
   the slow-growing nature of N. rajah, no hybrids involving it have been
   artificially produced as of yet.

   At present, the following natural hybrids are known:
     * N. × alisaputrana Adam & Wilcock (1992) [= N. burbidgeae × N.
       rajah]
     * N. edwardsiana × N. rajah
     * N. fusca × N. rajah
     * N. × kinabaluensis S. Kurata nom. nud. (1976) [= N. villosa × N.
       rajah]
     * N. macrovulgaris × N. rajah
     * N. stenophylla × N. rajah
     * N. tentaculata × N. rajah

   The "Mountain Garden" of Kinabalu National Park contains a number of
   well-grown Nepenthes, including the rare hybrid N. stenophylla × N.
   rajah. This plant has leaves resembling those of N. stenophylla, but
   the lid and wings are typical of N. rajah. The peristome is strongly
   influenced by N. stenophylla and bristles are present at the border of
   the lid, a unique characteristic of this hybrid. It occurs at an
   altitude of 1500-2600 m.

   Two hybrids of N. rajah have been formally described and given specific
   names: N. × alisaputrana and N. × kinabaluensis. Both are listed on
   CITES Appendix II and the latter is also considered Endangered (EN (D))
   under current IUCN criteria.

Nepenthes × alisaputrana

   N. × alisaputrana (originally published as "Nepenthes ×
   alisaputraiana") is named in honour of Datuk Lamri Ali, Director of
   Sabah Parks. It is only known from a few remote localities within
   Kinabalu National Park where is grows in stunted, open vegetation over
   serpentine soils at around 2000 m above sea level, often amongst
   populations of N. burbidgeae. 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). 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 hyrbid 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, Phillipps and Lamb (1996) gave it the common name Leopard
   pitcher-plant, though this is rarely used. The peristome is green to
   dark red and striped with purple bands. Leaves are often slightly
   peltate. The plant climbs well and aerial pitchers are frequently
   produced. N. × 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 Insect Eating Plants & How To Grow
   Them ( Slack, 1986) as being N. rajah was in fact N. burbidgeae × N.
   rajah.
   Nepenthes × kinabaluensis
   Enlarge
   Nepenthes × kinabaluensis

Nepenthes × kinabaluensis

   N. × kinabaluensis is another impressive plant. The pitchers get large
   also, but do not compare to those of N. rajah or N. × alisaputrana. It
   is a well-known natural hybrid of what many consider to be the two most
   spectacular Nepenthes species of Borneo: N. rajah and N. villosa. N. ×
   kinabaluensis can only be found on Mount Kinabalu (hence the name) and
   nearby Mount Tambuyukon, where the two parent species are occur
   sympatrically. More specifically, plants are known from a footpath near
   Paka Cave and several places along an unestablished route on a
   south-east ridge, which lies on the west side of the Upper Kolopis
   River. The only accessible location from which this species is known is
   the Kinabalu summit trail, between Layang-Layang and the helipad, where
   it grows at about 2900 m in a clearing dominated by Dacrydium gibbsiae
   and Leptospermum recurvum trees. N. × kinabaluensis has an altitudinal
   distribution of 2420 to 3030 m. It grows in open areas in cloud forest.
   This species can be distinguished from N. rajah by the presence of
   raised ribs that line the inner edge of the peristome and end with
   elongated teeth. These are more prominent than those found in N. rajah
   and are clue as to the hybrid's parentage (N. villosa has very
   developed peristome ribs). The peristome is coarse and expanded at the
   margin (but not scalloped like that of N. rajah), the lid orbiculate or
   reniformed and almost flat. In general, pitchers are larger than those
   of N. villosa and the tendril joins the apex about 1-2 cm below the
   leaf tip, a feature which is characteristic of N. rajah. In older
   plants, the tendril can be almost woody. N. × kinabaluensis is an
   indumentum of villous hairs covering the pitchers and leaf margins,
   which is approximately intermediate between the parents. Lower pitchers
   have two fringed wings, whereas the upper pitchers usually lack these.
   The colour of the pitcher varies from yellow to scarlet. N. ×
   kinabaluensis seems to produce upper pitchers more readily than either
   of its parents. In all respects N. × kinabaluensis is intermediate
   between the two parent species and it is easy to distinguish from all
   other Nepenthes of Borneo. However, it has been confused once before,
   when the hybrid was labelled as N. rajah in Letts Guide to Carnivorous
   Plants of the World (Cheers, 1992).

   N. × kinabaluensis was first collected near Kambarangoh by Gibbs in
   1910 and later mentioned by Macfarlane as "Nepenthes sp." in 1914.
   Although Macfarlane did not formally name the plant, he noted that
   "[a]ll available morphological details suggest that this is a hybrid
   between N. villosa and N. rajah". It was finally described in 1976 by
   Kurata as N. × kinabaluensis. The name was published in Nepenthes of
   Mount Kinabalu, though the specific epithet "kinabaluensis" is a nomen
   nudum, as it was published with an inadequate description and lacked
   information on the type specimen. The name was subsequently republished
   by Kurata in 1984 and by Adam and Wilcock in 1996.

Hybrid or species?

   It is worth noting that N. × alisaputrana and N. × kinabaluensis are
   often fertile and thus may breed among themselves. Clive A. Stace
   (1980) writes that we may speak of "stabilised hybrids when they have
   developed a distributional, morphological or genetic set of characters
   which is no longer strictly related to that of its parents, ... if the
   hybrid has become an independent, recognisable, self-producing unit, it
   is de facto a separate species". N. hurrelliana and N. murudensis are
   two examples of species that have a putative hybrid origin. N. ×
   alisaputrana and N. × kinabaluensis are sufficiently stabilised that a
   species status has been discussed. Indeed, N. kinabaluensis was
   described as a species by Adam & Wilcock in 1996. However, this
   interpretation does not have much support in the scientific community
   and the name has not been published in any other work since.

   Due to their dioecious nature, a hybrid involving a pair of Nepenthes
   species can represent one of two possible crosses, depending on which
   species was the female and which was the male. When the cross is known,
   the female (or pod) parent is usually referred to first, followed by
   the male (or pollen) parent. This is an important distinction, as the
   hybrid will usually display different morphological features according
   to the type of cross; the pod parent is thought to be dominant in most
   cases and hybrid offspring usually resemble it more than the pollen
   parent. Most wild plants of N. × kinabaluensis, for example, show a
   greater affinity to N. rajah than N. villosa and are thus thought to
   represent the cross N. rajah × N. villosa. However, specimens have been
   found that seem to be more similar to N. villosa, suggesting that they
   might be the reverse cross (see ). The same is true for other hybrids
   involving N. rajah. It is unknown whether these crosses are fertile or
   not and this element of uncertainty only adds to the confusion over the
   distinction between a stable hybrid and a species.

Cultivation

   Nepenthes rajah has always been considered to be one of the more
   difficult Nepenthes species to cultivate. However, in recent years, it
   has become apparent that the plant may not be deserving of its
   reputation.
   Cultivated N. rajah plant with large lower pitcher
   Enlarge
   Cultivated N. rajah plant with large lower pitcher

Environmental factors

   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 ℃, and cool
   nights, with temperatures of about 10 to 15 ℃. Here, it is important to
   note that the temperatures themselves are not vital (when kept within
   reasonable limits), but rather the temperature drop itself; N. rajah
   needs considerably cooler nights, with a drop of 10 ℃ or more being
   preferable. Failure to observe this requirement will almost certainly
   doom the plant in the long term or, at best, limit it to being a small,
   unimpressive specimen.

   In addition, like all Nepenthes, this plant needs a fairly humid
   environment to grow well. Values in the region of 75% R.H. are
   generally considered optimal, with increased humidity at night (~90%
   R.H.). However, N. rajah does tolerate fluctuations in humidity,
   especially when young, provided that the air does not become too dry
   (below 50% R.H.). Humidity can be easily controlled using an ultrasonic
   humidifier in conjunction with a humidistat.
   Cultivated N. rajah plant
   Enlarge
   Cultivated N. rajah plant

   In its natural habitat, N. rajah grows in open areas, where it is
   exposed to direct sunlight - it therefore needs to be provided with a
   significant amount of light in cultivation as well. To meet this need,
   many growers have used metal halide lamps in the 500–1000 watt range,
   with considerable success. The plant should be situated a fair distance
   from the light source, 1 to 2 m is recommended. Depending on location,
   growers can utilise natural sunlight as a source of illumination.
   However, this is only recommended for those living in equatorial
   regions, where light intensity is sufficient to satisfy the needs of
   the plant. A photoperiod of 12 hours is comparable to that experienced
   in nature.

Potting and watering

   Pure long-fibre Sphagnum moss is an excellent potting medium, though
   combinations involving any of the following - peat, perlite,
   vermiculite, sand, lava rock, pumice, Osmunda fibre, orchid bark and
   horticultural charcoal - may be used with equal success. The potting
   medium should be well-drained and not too compacted. Moss is useful for
   moisture retention near the roots. The mix should be thoroughly soaked
   in water prior to potting the plant.

   It has been noted that N. rajah produces a very extensive root system
   (for a Nepenthes) and, for this reason, it is recommended that a wide
   pot be used to allow for proper development of the root system. This
   also eliminates the need for frequent re-potting, which can lead to
   transplant shock and the eventual death of the plant.
   Plants fresh from tissue culture
   Enlarge
   Plants fresh from tissue culture

   Purified water should be used for watering purposes, although ' hard
   water' is tolerated. This is done to minimise the build-up of minerals
   and chemicals in the soil. Water purity greater than 100  p.p.m. of
   total dissolved solids is often quoted as ideal. A reverse osmosis unit
   can be used to filter the water or, alternatively, bottled distilled
   water can be purchased. Watering should be done regularly. However,
   plants should not be allowed to sit in water, as this may lead to root
   rot.
   N. rajah plant with Drosera species in foreground
   Enlarge
   N. rajah plant with Drosera species in foreground

Feeding and fertilising

   N. rajah is a carnivorous plant and, as such, supplements nutrients
   gained from the soil with captured prey (espescially insects) to
   alleviate deficiencies in important elements such as nitrogen,
   phosphorus and potassium. Just as in nature, a cultivated plant's
   'diet' may include insects and other prey items, although this is not
   necessary for successful cultivation. Crickets are recommended for
   their size and low cost. These can be purchased online or at specialist
   pet stores. They can simply be dropped into the pitchers by hand or
   placed inside using metal tongs or similar, whether dead or alive.

   From trials carried out by a commercial Nepenthes nursery, it appears
   that micronutrient solutions have "a beneficial effect on plants of
   improved leaf colouration, with no deleterious affects" as far as can
   be seen. However, more research is required to verify these results.
   Actual fertilisers (containing NPK) were, on the other hand, found to
   "cause damage to plants, promote pathogens and have no observable
   benefits". Hence, the use of chemical fertilisers is usually not
   advised.

   Above all, it should be remembered that the primary goal of any growing
   setup is to try to mirror the conditions the plants experience in their
   natural habitat as much as possible.

   It is important to note that N. rajah is a very slow growing Nepenthes.
   Under optimal conditions, N. rajah can reach flowering size within
   10 years of seed germination, although it is thought that it may take
   100 years to reach full size. Theoretically, given climatic conditions
   do not change, N. rajah has an indefinite lifespan.

Common misconceptions

   N. rajah growing on ultramafic soil
   Enlarge
   N. rajah growing on ultramafic soil

   Nepenthes rajah has been a well known and highly sought after species
   for over a century and, as a result, there are many stories woven
   around this plant. One such example is the famous legend that N. rajah
   grows exclusively in the spray zones of waterfalls, on ultramafic
   soils. Although the latter is true, N. rajah is certainly not found
   solely in the spray zones of waterfalls and this statement seems to
   have little basis in fact.

   It is likely this misconception was popularised by S. Kurata's 1976
   book Nepenthes of Mount Kinabalu, in which he erroneously states that
   "N. rajah is rather fond of wet places like swamps or the surroundings
   of a waterfall". With the exception of N. mirabilis, no Nepenthes
   species have been reliably recorded from swamps. Such claims were
   presumably originally the result of attempts to further increase the
   legendary status of the plant.

   This being the case, it does appear, however, that certain N. rajah
   plants do in fact grow in the vicinity of waterfalls (as noted by H.
   Steiner, 2002) "providing quite a humid microclimate", which may indeed
   be the source of this particular misconception.

   Another myth surrounding this species is that it occasionally catches
   small monkeys and other large animals in its pitchers. Such tales have
   persisted for a very long time, but can probably be explained as
   rodents being mistaken for other species. It is interesting to note
   that one common name for Nepenthes plants is ' Monkey Cups'. The name
   refers to the fact that monkeys have been observed drinking rainwater
   from these plants. Thus, in a sense, this mythology may have some basis
   in fact.

Gallery

   N. rajah growing in ultrabasic soil

   Young plants with protomorphic pitchers

   Large terrestrial pitcher

   Plant growing in grassy undergrowth

   Pitcher viewed from above

   A lower pitcher of N. rajah

   Modified leaf: lamina, tendril and pitcher

   Infaunal mosquito larvae in pitcher fluid

   N. rajah plant among sedges

   Three medium-sized plants

   Recumbent lower pitcher

   Lower pitcher of four year old cultivated plant

   Timeline

      Timeline of Nepenthes rajah and its natural hybrids

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