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Nepenthes fusca

January 01, 1970

Nepenthes fusca /nɪˈpɛnθiːz ˈfʌskə/, or the dusky pitcher-plant,[7] is a tropical pitcher plant endemic to Borneo. It is found throughout a wide altitudinal range and is almost always epiphytic in nature, primarily growing in mossy forest.

Nepenthes fusca
An intermediate pitcher of N. fusca
from the Crocker Range
Scientific classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Caryophyllales
Family: Nepenthaceae
Genus: Nepenthes
Species:
N. fusca
Binomial name
Nepenthes fusca
Danser (1928)[2]
Synonyms

The specific epithet fusca is derived from the Latin word fuscus, meaning "dark brown" or "dusky", and refers to the colour of the pitchers.[5][7]

Botanical history edit

The first known collection of N. fusca was made by Frederik Endert on October 12, 1925, from Mount Kemul in East Kalimantan, at an elevation of 1500 m. It was discovered during an expedition to central Borneo by the Forest Research Institute of Bogor (then known as Buitenzorg),[4] on which Endert also made the only known collection of N. mollis.[8][note a] The N. fusca specimen, designated as Endert 3955, includes male floral material and is deposited at Herbarium Bogoriense (BO), the herbarium of the Bogor Botanical Gardens.[2][9] Endert wrote about this pitcher plant in a detailed 1927 account of the expedition,[4] although he misidentified it as N. veitchii.[2][9]

Nepenthes fusca was formally described[b] in 1928 by Dutch botanist B. H. Danser in his seminal monograph "The Nepenthaceae of the Netherlands Indies".[2] Danser based his description solely on Endert 3955, citing no other specimens. He wrote of N. fusca:[2]

This new species is, together with N. Veitchii and N. stenophylla, very nearly related to N. maxima, but can not be confounded with any of these species. According to Endert it grew in the forest on a narrow, stony mountain ridge covered with humus, and was not rare.

 
The type material of N. fusca (Endert 3955) bears pitchers that appear to be intermediate between typical lower and upper forms

Botanist Jan Schlauer has noted differences between the type specimen of N. fusca and Sabah plants referred to this species,[6] even interpreting plants illustrated in Kurata's Nepenthes of Mount Kinabalu as representing N. stenophylla (as distinct from N. fallax).[9][10][c] Matthew Jebb does not consider these differences significant enough to merit distinction at the species level. He suggests that the type specimen consists of intermediate lower and upper pitchers as opposed to true forms of either, making them appear atypical.[6]

Much of this taxonomic uncertainty stems from the fact that N. fusca has not been recollected from the type locality and many similar plants have been lumped under this taxon.[8] Matthew Jebb and Martin Cheek attempted to resolve this confusion in their 1997 monograph by interpreting N. fusca as a widespread and variable species.[11]

Subspecies edit

Two subspecies of N. fusca have been described, neither of which is presently thought to represent the species:

Both were originally coined by J. H. Adam and C. C. Wilcock and subsequently published in Jebb and Cheek's 1997 monograph, "A skeletal revision of Nepenthes (Nepenthaceae)".[11] As these names were published without an adequate description, they are both considered nomina nuda.[9] The former is likely based on Chai 35939, a specimen collected from Mount Apo.[9] Schlauer considers it synonymous with N. fallax,[9] a taxon that is in turn considered conspecific with N. stenophylla by most authors.[12]

Nepenthes fusca subsp. kostermansiana was named from the herbarium material Kostermans 21495, which was collected by André Joseph Guillaume Henri Kostermans on October 25, 1963, at 1000 m altitude along the Kelai River on Mount Nyapa (Njapa), Berau Regency, East Kalimantan.[9][13] It is deposited at the National Herbarium of the Netherlands in Leiden.[9] This taxon was initially thought to fall within the variability of N. fusca, but in 2011 it was described as a distinct species, N. epiphytica, with Kostermans 21495 designated as its holotype.[13]

Nepenthes maxima edit

 
An upper pitcher of N. maxima from Sulawesi

Nepenthes maxima, a species native to Sulawesi, New Guinea, and the Maluku Islands, was once thought to extend to Borneo as well. Some authors even wrote that it was widespread on the island.[5][14] This confusion stemmed from the likeness of N. fusca and N. maxima, and from apparently mislabelled seeds collected by Charles Curtis. Curtis was not meticulous in recording where he located individual plants; although it was originally believed that he collected N. curtisii (now considered synonymous with N. maxima)[15] in Borneo, botanist Charles Clarke points out that he also visited Sulawesi on the same trip, and N. maxima is common there.[6]

Matthew Jebb and Martin Cheek resolved this confusion in their 1997 monograph by referring a number of Bornean plants identified as N. maxima to N. fusca, thereby excluding the former from the island.[11]

Nepenthes zakriana edit

In 1996, J. H. Adam and C. C. Wilcock described Nepenthes curtisii subsp. zakriana.[3][d] Ten years later, Adam and Hafiza A. Hamid elevated it to species status as Nepenthes zakriana (/nɪˈpɛnθiːz ˌzɑːkriˈænə/, not /ˌzækriˈænə/).[citation needed] The authors described the taxon as a Sabah endemic growing at elevations of 1200 to 1500 m.[citation needed]

Adam and Hafiza wrote that N. zakriana "consistently differed from Nepenthes fusca by prominent raised midribs, extended beyond apex forming an apical glandular appendages on lower lid surface of both upper and lower pitchers; and basal half portion of the midrib developed in nail-shaped glandular crest".[citation needed] However, a number of authors soon voiced their doubts as to whether N. zakriana merited species status[16] and in Pitcher Plants of Borneo by Anthea Phillipps, Anthony Lamb, and Ch'ien Lee, these features were considered to fall within the natural variability of N. fusca.[8]

Nepenthes sp. A edit

In his 1997 monograph, Nepenthes of Borneo, Charles Clarke lists the undescribed taxon "Nepenthes sp. A", which has been recorded from Gunung Mulu National Park in Sarawak.[6] It bears a close resemblance to N. fusca and may be conspecific with it,[6] although its colouration is unusual for the species.[17] The pitchers of this plant match J. H. Adam and C. C. Wilcock's description[18] of N. faizaliana,[6] but the latter is now known to have a round lid (as opposed to narrowly triangular in N. fusca and "Nepenthes sp. A"), suggesting that these two taxa are not closely related.[6] Clarke proposes that this taxon might fall under Jebb and Cheek's more inclusive concept of N. fusca, but retains it as an undescribed species because it remains poorly known.[6]

"Nepenthes sp. A" was first illustrated in a 1988 article by Anthea Phillipps and Anthony Lamb, where it was considered an undescribed species.[19]

Description edit

 
A young plant from Mesilau, Mount Kinabalu

Nepenthes fusca is a climbing plant. The stem may attain a length of 10 m[8] and is up to 8 mm in diameter. Internodes are circular in cross section and up to 7 cm long.[6]

The leaves of this species are petiolate and coriaceous in texture. The lamina or leaf blade is obovate-oblong in shape and measures up to 15 cm in length by 6 cm in width. Its apex is acute to obtuse and may even be slightly peltate. The base of the lamina is gradually attenuate towards the petiole. The petiole (≤4 cm long)[5] is grooved lengthwise and bears a pair of narrow wings that form a semi-amplexicaul sheath around the stem. Up to 3 longitudinal veins are present on either side of the midrib,[5] although they are indistinct. Pinnate veins are numerous. Tendrils measure up to 5 cm in length.[6]

Rosette and lower pitchers are cylindrical throughout. They usually grow to 20 cm in height by 4 cm in width, although exceptional specimens up to 28 cm have been recorded.[8] A pair of fringed wings (≤5 mm wide) runs down the ventral surface of the pitcher and bears fringe elements measuring up to 10 mm and spaced 6 mm apart.[5] The glandular region is restricted to the lower portion of the pitcher's inner surface. The glands are small, overarched, and occur at a density of 600 to 650 per square centimetre.[2] The waxy zone is reduced.[20] The pitcher mouth is positioned horizontally at the front, becoming elongated into a neck at the rear. The peristome is flattened and expanded (≤12 mm wide), but bears only indistinct teeth (≤0.3 mm long).[5] The inner portion of the peristome accounts for around 51% of its total cross-sectional surface length.[20] The pitcher lid or operculum is very narrowly ovate in shape and has a distinctive basal crest on its lower surface. An unbranched spur measuring up to 10 mm in length is inserted near the base of the lid.[6]

     
Lower pitchers of N. fusca from the Crocker Range (CR) and the area around Mount Kinabalu (MK). From left to right: CR, MK, MK, CR, and CR.

Upper pitchers differ markedly in shape, being narrowly infundibular in the lower two-thirds and becoming widely infundibular above. They are similar in size to their lower counterparts, typically measuring up to 18 cm, with some larger forms reaching 26 cm.[8] The tiny digestive glands are overarched and number 1500 to 2000 per square centimetre.[2] The pitcher lid is very narrowly triangular with the margins and apex curved downwards.[8] In aerial pitchers, the wings are reduced to ribs.[6]

     
Upper pitchers of N. fusca from the Crocker Range (CR) and the area around Mount Kinabalu (MK). From left to right: MK, MK, CR, MK, and CR.

Nepenthes fusca produces a compact racemose inflorescence. The peduncle is up to 6 cm long, while the rachis is not known to exceed 10 cm. Partial peduncles are one- or two-flowered, up to 8 mm long, and lack a bract. Sepals are elliptic and up to 4 mm long.[6] A study of 120 pollen samples taken from the type specimen (Endert 3955) found the mean pollen diameter to be 34.8 μm (SE = 0.6; CV = 9.1%).[21]

Developing parts of the plant bear an indumentum of long, brown hairs. However, most of these disappear during the normal course of development, and mature parts only have a sparse covering of short, brown hairs.[6]

Ecology edit

 
Nepenthes fusca, N. reinwardtiana, and N. stenophylla growing in an exposed site along a logging road to Mount Murud in Sarawak

Nepenthes fusca is endemic to Borneo, where its range stretches from Central Kalimantan to northwestern Sabah.[6] The species is confined to north-western Borneo.[1] The species has a wide altitudinal distribution and is typically found at elevations of 1200 to 2500 m above sea level.[6] However, N. fusca has occasionally been reported from lowland hills down to 600 m[22] and in Sarawak it has been found at only 300 m in hill forest.[8]

Nepenthes fusca is most commonly found as an epiphyte in shady mossy forest on ridge tops, where it may grow 10 to 15 m off the ground.[8] This makes it particularly difficult to find and often the only evidence of its presence are dead pitchers that have fallen to the forest floor.[22][23][24] In this respect, it can be considered the "ecological equivalent" of N. bongso from Sumatra.[25] More rarely, N. fusca grows terrestrially in exposed sites near montane forest[6] or along logging roads.[8] It is often sympatric with species such as N. reinwardtiana, N. stenophylla, and N. tentaculata,[26] and natural hybrids with all of these have been recorded.[8][12]

 
Nepenthes fusca from Mount Alab in the Crocker Range

Locations edit

The species has been recorded from numerous mountains across Borneo. It can be found at several sites on Mount Kinabalu,[27][28] including Kambarangoh, the Marai Parai plateau, the mountain's East Ridge, and the Bambangan River at around 1500 m.[5] It also grows on the side of the road connecting the park headquarters and the power station, at an elevation of around 1550 m,[29] despite being frequently cut back as part of road maintenance.[23] This is one of the few places where N. fusca can be easily seen by visitors.[23] These roadside plants were greatly affected by the El Niño climatic phenomenon of 1997 to 1998. The resulting dry period severely depleted the population, such that "almost all the plants were destroyed".[17] Nearby plants from more sheltered locations fared better and appeared to have recovered by the following year.[17] A number of N. fusca plants have also been transplanted to the Mesilau nature trail.[30]

Nepenthes fusca can be viewed along the road leading to the disused Mamut Copper Mine, adjacent to Mount Kinabalu. There it is sympatric with N. macrovulgaris, N. stenophylla, and the natural hybrid N. fusca × N. stenophylla; N. burbidgeae grows a short distance away.[30] The species also occurs on nearby Mount Tambuyukon.[5]

On Mount Trusmadi, N. fusca has been observed growing epiphytically on Eleocarpus trees at an elevation of almost 1800 m.[31] A lone plant growing on a ridge top at 1962 m was found to have a small bush frog (probably Philautus aurantium) in one of its partly dried, 15 cm-long pitchers.[32] Scattered plants have also been recorded from a rocky clearing at c. 1592 m, growing alongside Gleichenia truncata ferns and bamboo orchids; these plants have rather unremarkable red-speckled but otherwise plain green pitchers, though unusually the stem and underside of the midrib are almost black.[33]

The species is abundant on Mount Alab, the highest peak of the Crocker Range, where a number of colour variants have been documented, including one with wholly green lower pitchers save for red mottling on the interior and underside of the lid.[33] An atypical yellow form grows on Mount Lumarku in southwestern Sabah.[8] Other notable locations include the Kimanis–Keningau Road that runs through the Crocker Range[8] and the summit area of Mount Apo Dari (where it grows at 1500 m).[14]

On Mount Mulu in Sarawak, N. fusca (found below 1200 m) appears to occupy a discrete altitudinal zone from N. vogelii (1200–1500 m) and N. hurrelliana (above 1500 m), both of which are also epiphytes.[8] In the Hose Mountains of central Sarawak, N. fusca is typically found growing as an epiphyte in lower montane forest at 700–1200 m.[34] It is also an uncommon inhabitant of roadside embankments there.[34]

Conservation status edit

Charles Clarke noted that since substantial populations of N. fusca lie within the boundaries of national parks, they "are unlikely to become threatened in the foreseeable future".[6] Habitat destruction is considered to be the greatest threat to the species's survival in the wild.[35] Plant poaching is of far lesser concern, as this species is not particularly sought after in the carnivorous plant hobby and its epiphytic habit makes it largely inaccessible to collectors.[35]

Related species edit

 
 
Upper pitchers of N. hurrelliana (left) and N. vogelii (right)

Among the closest relatives of N. fusca are the Bornean species N. epiphytica,[13] N. hurrelliana, N. platychila, N. stenophylla, and N. vogelii.[8] More broadly, it belongs to the loosely defined "N. maxima complex", which also includes N. boschiana, N. chaniana, N. eymae, N. faizaliana, N. klossii, and N. maxima.[13] The enigmatic N. mollis, which some authors have suggested is conspecific with N. hurrelliana,[36] may also be closely allied.

The lower pitchers of N. hurrelliana are distinctive, but the upper ones bear a close resemblance to those of N. fusca. Of the Bornean pitcher plant flora, only these two species have such a narrowly triangular lid. The upper pitchers of N. hurrelliana differ in having a horizontal mouth that rises abruptly into a long neck at the back and in having a hirsute basal crest on the underside of the lid.[8]

Nepenthes hurrelliana is particularly similar to a form of N. fusca from the southern portion of the Crocker Range in Sabah. This form exhibits a wider peristome, longer neck, and a more triangular lid than most other examples of the species.[8] However, the peristome is still not as well developed as in N. hurrelliana and the plant lacks the dense indumentum of the latter. Furthermore, N. hurrelliana differs in the distribution of nectar glands on the lower surface of its lid.[8]

The first known collection of N. vogelii, made in 1961, was labelled as N. fusca.[8] In 1969, botanist Shigeo Kurata examined this specimen and noted that it did not fall within the known variation exhibited by N. fusca.[8] Nevertheless, the species remained undescribed until 2002.[37] Nepenthes vogelii differs in having much smaller pitchers and lacking appendages on the underside of the lid.[37] In addition, the lid of N. vogelii is broadly triangular as opposed to the narrowly triangular lid of N. fusca.[8][22] The colour of the pitchers—light cream with dark speckles—is also distinctive.[22]

Nepenthes faizaliana also bears a resemblance to N. fusca. In their description of the former, J. H. Adam and C. C. Wilcock distinguished these taxa on the basis of inflorescence structure, the size of the glandular region on the inner surface of upper pitchers, and the development and characteristics of the indumentum.[6][18] Nepenthes fusca also differs in having a very narrow pitcher lid, as opposed to the orbicular lid of N. faizaliana.[6]

Nepenthes platychila, another closely allied species, differs from N. fusca in having a much wider peristome and lid, and lacking appendages on its lower lid surface.[8][38] Nepenthes fusca is also thought to be closely related to Sulawesi's N. eymae, and N. maxima, which is widespread in Sulawesi, New Guinea, and the Maluku Islands.[39]

Natural hybrids edit

Due its widespread distribution throughout Borneo, N. fusca forms natural hybrids with a relatively large number of other species. However, like N. fusca itself, these are often hard to find due to the species's epiphytic growth habit.[8]

 
Lower pitchers of N. burbidgeae × N. fusca from Mesilau

N. burbidgeae × N. fusca edit

Nepenthes burbidgeae × N. fusca has been known since at least the early 1980s, when it was found during an expedition to Sabah.[26]

N. fusca × N. lowii edit

This hybrid was initially identified by Charles Clarke as a cross between N. chaniana (known as N. pilosa at the time[40]) and N. lowii.[6][41] However, in their 2008 book, Pitcher Plants of Borneo, Anthea Phillipps, Anthony Lamb and Ch'ien Lee pointed out that the plants exhibit influences of N. fusca, such as a triangular lid and an elongated neck.[8] The authors noted that both N. fusca and N. lowii are common on the summit area of Mount Alab where this plant is found, whereas N. chaniana is rare.[8] Another possible parent species, N. stenophylla, is apparently absent from the site.[8]

This cross was originally discovered by Rob Cantley and Charles Clarke on Bukit Batu Buli in Sarawak.[6] Clarke later found larger plants of this hybrid in the Crocker Range of Sabah, particularly near the summit of Mount Alab.[6] More recently a single plant has been recorded from Mount Trusmadi.[33] The pitchers of this cross have a slight constriction in the middle and range in colour from green to dark purple throughout.[6]

This hybrid differs from N. fusca in the presence of bristles on the underside of the lid. Conversely, it has a dense indumentum on the stem and at the margins of the lamina, compared to the virtually glabrous stem and leaves of N. lowii. It also differs from N. lowii in having a more developed peristome, which is circular in cross section. While lower pitchers of N. lowii have prominent teeth, those of N. fusca × N. lowii are indistinct. In addition, a glandular appendage is present on the underside of the lid,[6] a trait inherited from N. fusca.

Nepenthes fusca × N. lowii is difficult to confuse with its putative parent species, but is somewhat similar to N. chaniana × N. veitchii. The latter hybrid can be distinguished on the basis of its peristome, which is wider, more flared, and less cylindrical. In addition, this hybrid has a less ovate lid, which lacks the bristles characteristic of N. lowii, and a denser indumentum covering the stem and leaves.[6]

N. fusca × N. reinwardtiana edit

 
 
Lower pitchers of the putative natural hybrids N. fusca × N. reinwardtiana (left) and N. fusca × N. stenophylla (right)

A putative cross between N. fusca and N. reinwardtiana has been recorded.[12] Nepenthes naquiyuddinii is generally thought to be a heterotypic synonym of N. reinwardtiana,[16] but may also represent this hybrid, as both putative parent species grow in close proximity to it.[8]

N. fusca × N. stenophylla edit

This hybrid is known from the eastern slopes of Mount Trusmadi, where it grows in lower montane forest. Both N. fusca and N. stenophylla are common in this area.[8] It has also been recorded from the Mamut copper mine.[30] It more closely resembles N. stenophylla, but differs in the shape of the lid, which is more oval-shaped.[8]

N. fusca × N. tentaculata edit

Nepenthes fusca × N. tentaculata was discovered by Linus Gokusing near the summit of Mount Alab, where it grows in upper montane forest at elevations of around 1800 to 2000 m.[8] It is sympatric with both parent species, which are abundant in the area.[8]

Other hybrids edit

Natural crosses with N. platychila,[42] N. rajah,[6] and N. veitchii[6] have also been recorded.

The pitchers of N. hurrelliana are roughly intermediate in appearance between those of N. fusca and N. veitchii. This has led to speculation regarding the lineage of this species, with a number of authors suggesting a possible hybridogenic origin.[22] However, N. hurrelliana is distinct from the natural hybrid N. fusca × N. veitchii and most authors now regard it as a valid species.[8][22][43]

Cultivation edit

 
In the wild, N. fusca is generally found is rather shady conditions (upper pitchers growing along a logging road to Mount Murud pictured) and this preference is reflected in cultivated plants, which show optimal growth under lower light levels than many other Nepenthes species.[44]

Little information has been published on the growing requirements of N. fusca. In 2004, professional horticulturist Robert Sacilotto wrote an article for the Carnivorous Plant Newsletter, summarising measured tolerances of several highland Nepenthes species based on experiments conducted between 1996 and 2001.[44]

Sacilotto found N. fusca to be tolerant of a wide range of conditions; with the exception of plants not treated with fungicides, no test groups showed a survival rate of less than 75%. Nepenthes fusca was found to tolerate temperatures in the range of 10 to 38 °C (50 to 100 °F). A nighttime drop in temperature below 21 °C (70 °F) was necessary for good growth; plants that were not exposed to such a drop grew poorly and produced fewer pitchers. The experiments suggested that N. fusca grows best when relative humidity is in the range of 65 to 90%.[44]

The species seemed to do best in growing media with a high proportion of organic matter (such as a mixture consisting of 10% peat moss chunks, 30% perlite, and 60% of any combination of Sphagnum moss and fir bark). Soil with a slightly acidic pH of 4.5 to 5.0 produced the best results. Optimal soil conductivity appeared to lie between 10 and 45 microsiemens.[44]

An illuminance of 6400–8600 lx (600–800 fc) proved to be optimal when plants were grown under sunlight, high pressure sodium, and metal halide lamps. However, specimens placed under an even combination of Gro-Lux and cool white fluorescent lamps at 5400–7500 lx (500–700 fc) exhibited the most vibrant colours (although growth rates remained the same). Plants moved from the former to the latter light set up showed a significant change in pigmentation; green leaf blades turned bronzy and speckles on the pitchers darkened markedly.[44]

Nepenthes fusca was found to respond well to a quarter-strength fertilizer that was applied to the pitchers. Ants were also an effective food source.[44]

Notes edit

a.^ During September and October 1925, Endert explored the hilly area around Mount Kemul, climbing the summit (1,847 m) several times, and the neighbouring valleys of the Long Mehiang, Long Kiau, and Long Petak.[45] He collected N. fusca on October 12 and N. mollis on October 17.[2]

b.^ The Latin description of N. fusca from Danser's monograph reads:[2]

Folia mediocria breviter petiolata, lamina lanceolata, nervis longitudinalibus utrinque c. 2, vagina caulis 1/2 amplectente; ascidia rosularum ignota; ascidia inferiora magnitudine mediocria, parte inferiore anguste ovata, os versus subcylindrica, parte superiore alis 2 fimbriatis; peristomio in collum elongato, applanato, 4-10 mm lato, costis c. 1/3-2/3 mm distantibus, dentibus c. tam longis quam latis; operculo anguste ovato, subcordato, facie inferiore appendice lateraliter applanata; ascidia superiora magnitudine mediocria, infundibuliformia, costis 2 prominentibus; peristomio in collum elongato, applanato, 3-8 mm lato, costis 1/3-1/4 mm distantibus, dentibus brevissimis; operculo anguste ovato, subcordato, facie inferiore prope basin appendice lateraliter applanata; inflorescentia racemis parvus, pedicillis inferioribus c. 8 mm longis, omnibus 1-floris v. partim 2-floris ; indumentum iuventute densissimum, denique passim densum, breve, e pilis patentibus crassis simplicibus v. basi ramosis compositum.

c.^ Some authors treat N. fallax in synonymy with N. stenophylla,[6][11] while others consider them to be two distinct species, with plants commonly referred to as N. stenophylla actually representing N. fallax.[46][47]

d.^ The subspecies was named after Zakri Abdul Hamid.[48] This taxon, along with N. curtisii (sensu J.H.Adam & Wilcock) itself, is considered a heterotypic synonym of N. stenophylla by some taxonomists.[15]

References edit

  1. ^ a b Clarke, C.M. (2018). "Nepenthes fusca". IUCN Red List of Threatened Species. 2018: e.T40109A143967109. doi:10.2305/IUCN.UK.2018-1.RLTS.T40109A143967109.en. Retrieved 19 November 2021.
  2. ^ a b c d e f g h i Danser, B.H. 1928. 13. Nepenthes fusca Dans., nova spec.. In: The Nepenthaceae of the Netherlands Indies. Bulletin du Jardin Botanique de Buitenzorg, Série III, 9(3–4): 249–438.
  3. ^ a b Adam, J.H. & C.C. Wilcock 1998 ['1996']. Pitcher plants of Mt. Kinabalu in Sabah. The Sarawak Museum Journal 50(71): 145–171.
  4. ^ a b c (in Dutch) Endert, F.H. 1927. Botanisch en floristisch verslag. In: D.W. Buijs, H. Witkkamp, F.H. Endert, H.C. Siebers & D.F.K. Bosch. Midden-Oost-Borneo Expeditie 1925. G. Kolff & Co., Weltevreden.
  5. ^ a b c d e f g h i Kurata, S. 1976. Nepenthes of Mount Kinabalu. Sabah National Parks Publications No. 2, Sabah National Parks Trustees, Kota Kinabalu.
  6. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
  7. ^ a b Phillipps, A. & A. Lamb 1996. Pitcher-Plants of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
  8. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af Phillipps, A., A. Lamb & C.C. Lee 2008. Pitcher Plants of Borneo. Second Edition. Natural History Publications (Borneo), Kota Kinabalu.
  9. ^ a b c d e f g h Schlauer, J. N.d. Nepenthes fusca 2011-06-17 at the Wayback Machine. Carnivorous Plant Database.
  10. ^ Clarke, C.M. 2006. Introduction. In: Danser, B.H. The Nepenthaceae of the Netherlands Indies. Natural History Publications (Borneo), Kota Kinabalu. pp. 1–15.
  11. ^ a b c d e f Jebb, M.H.P. & M.R. Cheek 1997. A skeletal revision of Nepenthes (Nepenthaceae). Blumea 42(1): 1–106.
  12. ^ a b c McPherson, S.R. 2009. Pitcher Plants of the Old World. 2 volumes. Redfern Natural History Productions, Poole.
  13. ^ a b c d Robinson, A.S., J. Nerz & A. Wistuba 2011. Nepenthes epiphytica, a new pitcher plant from East Kalimantan. In: McPherson, S.R. New Nepenthes: Volume One. Redfern Natural History Productions, Poole. pp. 36–51.
  14. ^ a b Adam, J.H., C.C. Wilcock & M.D. Swaine 1992. The ecology and distribution of Bornean Nepenthes. 2011-07-22 at the Wayback Machine Journal of Tropical Forest Science 5(1): 13–25.
  15. ^ a b Schlauer, J. N.d. Nepenthes curtisii 2011-06-17 at the Wayback Machine. Carnivorous Plant Database.
  16. ^ a b Rice, B.A. 2006. Do you want to tell me about a species I missed? The Carnivorous Plant FAQ.
  17. ^ a b c Steiner, H. 2002. Borneo: Its Mountains and Lowlands with their Pitcher Plants. Toihaan Publishing Company, Kota Kinabalu.
  18. ^ a b Adam, J.H. & C.C. Wilcock 1991. A new species of Nepenthes (Nepenthaceae) from Sarawak. Blumea 36(1): 123–125.
  19. ^ Phillipps, A. & A. Lamb 1988. Pitcher-plants of East Malaysia and Brunei. Nature Malaysiana 13(4): 8–27.
  20. ^ a b Bauer, U., C.J. Clemente, T. Renner & W. Federle 2012. Form follows function: morphological diversification and alternative trapping strategies in carnivorous Nepenthes pitcher plants. Journal of Evolutionary Biology 25(1): 90–102. doi:10.1111/j.1420-9101.2011.02406.x
  21. ^ Adam, J.H. & C.C. Wilcock 1999. Palynological study of Bornean Nepenthes (Nepenthaceae). Pertanika Journal of Tropical Agricultural Science 22(1): 1–7.
  22. ^ a b c d e f Clarke, C.M. & C.C. Lee 2004. Pitcher Plants of Sarawak. Natural History Publications (Borneo), Kota Kinabalu.
  23. ^ a b c Clarke, C.M. 2001. A Guide to the Pitcher Plants of Sabah. Natural History Publications (Borneo), Kota Kinabalu.
  24. ^ Bourke, G. 2010. The climbing pitcher plants of the Kelabit highlands. 2013-04-09 at the Wayback Machine Captive Exotics Newsletter 1(1): 4–7.
  25. ^ Clarke, C.M. 2001. Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu.
  26. ^ a b Lowrie, A. 1983. Sabah Nepenthes Expeditions 1982 & 1983. Carnivorous Plant Newsletter 12(4): 88–95.
  27. ^ Triplitt, R. 1985. Nepenthes—Color Them Unique. Carnivorous Plant Newsletter 14(2): 40–42, 48–49.
  28. ^ Malouf, P. 1995. A visit to Kinabalu Park. Carnivorous Plant Newsletter 24(3): 64–69.
  29. ^ Yeo, J. 1996. A trip to Kinabalu Park. Bulletin of the Australian Carnivorous Plant Society, Inc. 15(4): 4–5.
  30. ^ a b c Thong, J. 2006. Travels around North Borneo – Part 1. 2011-07-07 at the Wayback Machine Victorian Carnivorous Plant Society Inc. 81: 12–17.
  31. ^ Marabini, J. 1984. A field trip to Gunong Trusmadi. Carnivorous Plant Newsletter 13(2): 38–40.
  32. ^ Damit, A. 2014. A trip to Mount Trus Madi – the Nepenthes wonderland. Carnivorous Plant Newsletter 43(1): 19–22.
  33. ^ a b c Fretwell, S. 2013. Back in Borneo to see giant Nepenthes. Part 3: Mt. Trusmadi and Mt. Alab. Victorian Carnivorous Plant Society Inc. 109: 6–15.
  34. ^ a b Lee, C.C. 2002. Nepenthes species of the Hose Mountains in Sarawak, Borneo. Proceedings of the 4th International Carnivorous Plant Conference, Hiroshima University, Tokyo: 25–30.
  35. ^ a b Amazing Species: Nepenthes fusca. IUCN Red List.
  36. ^ Salmon, B.[R.] 1999. Nepenthes mollis (Nepenthaceae)—Rediscovered? Carnivorous Plant Newsletter 28(1): 24–26.
  37. ^ a b Schuiteman, A. & E.F. de Vogel 2002. Nepenthes vogelii (Nepenthaceae): a new species from Sarawak. Blumea 47(3): 537–540.
  38. ^ Lee, C.C. 2002. Nepenthes species of the Hose Mountains in Sarawak, Borneo. [video] The 4th International Carnivorous Plant Conference, Tokyo, Japan. (video by Irmgard & Siegfried R. H. Hartmeyer)
  39. ^ D'Amato, P. 1993. Nepenthes eymai. Carnivorous Plant Newsletter 22(1–2): 21.
  40. ^ Clarke, C.M., C.C. Lee & S. McPherson 2006. Nepenthes chaniana (Nepenthaceae), a new species from north-western Borneo. Sabah Parks Journal 7: 53–66.
  41. ^ (in Czech) Macák, M. 2000. Portréty rostlin - Nepenthes lowii Hook. F.. Trifid 2000(3–4): 51–55. (page 2, page 3, page 4, page 5)
  42. ^ Lee, C.C. 2002. Nepenthes platychila (Nepenthaceae), a new species of pitcher plant from Sarawak, Borneo. The Gardens' Bulletin Singapore 54: 257–261.
  43. ^ Cheek, M., M. Jebb, C.C. Lee, A. Lamb & A. Phillipps. 2003. Nepenthes hurrelliana (Nepenthaceae), a new species of pitcher plant from Borneo. Sabah Parks Nature Journal 6: 117–124.
  44. ^ a b c d e f Sacilotto, R. 2004. Experiments with highland Nepenthes seedlings: a summary of measured tolerances. Carnivorous Plant Newsletter 33(1): 26–31.
  45. ^ van Steenis-Kruseman, M.J., et al. 2006. Cyclopaedia of Malesian Collectors: Frederik Hendrik Endert. Nationaal Herbarium Nederland.
  46. ^ Schlauer, J. N.d. Nepenthes fallax 2011-06-17 at the Wayback Machine. Carnivorous Plant Database.
  47. ^ Schlauer, J. 1996. N.stenophylla, once again. Carnivorous Plant Mailing List, May 31, 1996.
  48. ^ Wei-Shen, W. 2012. Up close and personal with Professor Emeritus Datuk Zakri Abdul Hamid. The Star, May 26, 2012.

Further reading edit

  • Beaman, J.H. & C. Anderson 2004. The Plants of Mount Kinabalu: 5. Dicotyledon Families Magnoliaceae to Winteraceae. Natural History Publications (Borneo), Kota Kinabalu.
  • Benz, M.J., E.V. Gorb & S.N. Gorb 2012. Diversity of the slippery zone microstructure in pitchers of nine carnivorous Nepenthes taxa. Arthropod-Plant Interactions 6(1): 147–158. doi:10.1007/s11829-011-9171-2
  • Bonhomme, V., H. Pelloux-Prayer, E. Jousselin, Y. Forterre, J.-J. Labat & L. Gaume 2011. Slippery or sticky? Functional diversity in the trapping strategy of Nepenthes carnivorous plants. New Phytologist 191(2): 545–554. doi:10.1111/j.1469-8137.2011.03696.x
  • Bourke, G. 2011. The Nepenthes of Mulu National Park. Carniflora Australis 8(1): 20–31.
  • Buch, F., M. Rott, S. Rottloff, C. Paetz, I. Hilke, M. Raessler & A. Mithöfer 2012. Secreted pitfall-trap fluid of carnivorous Nepenthes plants is unsuitable for microbial growth. Annals of Botany 111(3): 375–383. doi:10.1093/aob/mcs287
  • Corner, E.J.H. 1996. Pitcher-plants (Nepenthes). In: K.M. Wong & A. Phillipps (eds.) Kinabalu: Summit of Borneo. A Revised and Expanded Edition. The Sabah Society, Kota Kinabalu. pp. 115–121. ISBN 9679994740.
  • Fretwell, S. 2013. Back in Borneo for giant Nepenthes. Part 1: Mesilau Nature Reserve, Ranau. Victorian Carnivorous Plant Society Inc. 107: 6–13.
  • Fretwell, S. 2013. Back in Borneo to see giant Nepenthes. Part 2: Mt Tambuyukon and Poring. Victorian Carnivorous Plant Society Inc. 108: 6–15.
  • Kurata, S. 1969. Mindoro/North Borneo Expedition. Part 3. The Journal of Insectivorous Plant Society No. 47.
  • Lee, C.C. 2000. Recent Nepenthes Discoveries. [video] The 3rd Conference of the International Carnivorous Plant Society, San Francisco, USA.
  • McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Borneo. Redfern Natural History Productions, Poole.
  • Meimberg, H., A. Wistuba, P. Dittrich & G. Heubl 2001. Molecular phylogeny of Nepenthaceae based on cladistic analysis of plastid trnK intron sequence data. Plant Biology 3(2): 164–175. doi:10.1055/s-2001-12897
  • (in German) Meimberg, H. 2002. 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
  • Meimberg, H., S. Thalhammer, A. Brachmann & G. Heubl 2006. Comparative analysis of a translocated copy of the trnK intron in carnivorous family Nepenthaceae. Molecular Phylogenetics and Evolution 39(2): 478–490. doi:10.1016/j.ympev.2005.11.023
  • (in Japanese) Oikawa, T. 1992. Nepenthes fusca Dans.. In: Muyū kusa – Nepenthes (無憂草 – Nepenthes). [The Grief Vanishing.] Parco Co., Japan. pp. 42–45.
  • Renner, T. & C.D. Specht 2011. A sticky situation: assessing adaptations for plant carnivory in the Caryophyllales by means of stochastic character mapping. International Journal of Plant Sciences 172(7): 889–901. doi:10.1086/660882
  • Thorogood, C. 2010. The Malaysian Nepenthes: Evolutionary and Taxonomic Perspectives. Nova Science Publishers, New York.

External links edit

 
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January 01, 1970
nepenthes, fusca, this, article, needs, updated, reason, given, reflect, current, taxonomy, splitting, dactylifera, zakriana, please, help, update, this, article, reflect, recent, events, newly, available, information, october, 2019, dusky, pitcher, plant, tro. This article needs to be updated The reason given is to reflect current taxonomy splitting off of N dactylifera and N zakriana Please help update this article to reflect recent events or newly available information October 2019 Nepenthes fusca n ɪ ˈ p ɛ n 8 iː z ˈ f ʌ s k e or the dusky pitcher plant 7 is a tropical pitcher plant endemic to Borneo It is found throughout a wide altitudinal range and is almost always epiphytic in nature primarily growing in mossy forest Nepenthes fusca An intermediate pitcher of N fuscafrom the Crocker Range Conservation status Least Concern IUCN 3 1 1 Scientific classification Kingdom Plantae Clade Tracheophytes Clade Angiosperms Clade Eudicots Order Caryophyllales Family Nepenthaceae Genus Nepenthes Species N fusca Binomial name Nepenthes fuscaDanser 1928 2 Synonyms Synonyms Nepenthes curtisii subsp zakrianaJ H Adam amp Wilcock 1996 3 Nepenthes veitchiiauct non Hook f Endert 1927 4 Nepenthes zakriana J H Adam amp Wilcock J H Adam amp Hafiza 2006 citation needed Heterochresonyms Nepenthes fuscaauct non Danser Sh Kurata 1976 5 N stenophylla Informal names Nepenthes sp AC Clarke 1997 6 The specific epithet fusca is derived from the Latin word fuscus meaning dark brown or dusky and refers to the colour of the pitchers 5 7 Contents 1 Botanical history 1 1 Subspecies 1 2 Nepenthes maxima 1 3 Nepenthes zakriana 1 4 Nepenthes sp A 2 Description 3 Ecology 3 1 Locations 3 2 Conservation status 4 Related species 5 Natural hybrids 5 1 N burbidgeae N fusca 5 2 N fusca N lowii 5 3 N fusca N reinwardtiana 5 4 N fusca N stenophylla 5 5 N fusca N tentaculata 5 6 Other hybrids 6 Cultivation 7 Notes 8 References 9 Further reading 10 External linksBotanical history editThe first known collection of N fusca was made by Frederik Endert on October 12 1925 from Mount Kemul in East Kalimantan at an elevation of 1500 m It was discovered during an expedition to central Borneo by the Forest Research Institute of Bogor then known as Buitenzorg 4 on which Endert also made the only known collection of N mollis 8 note a The N fusca specimen designated as Endert 3955 includes male floral material and is deposited at Herbarium Bogoriense BO the herbarium of the Bogor Botanical Gardens 2 9 Endert wrote about this pitcher plant in a detailed 1927 account of the expedition 4 although he misidentified it as N veitchii 2 9 Nepenthes fusca was formally described b in 1928 by Dutch botanist B H Danser in his seminal monograph The Nepenthaceae of the Netherlands Indies 2 Danser based his description solely on Endert 3955 citing no other specimens He wrote of N fusca 2 This new species is together with N Veitchii and N stenophylla very nearly related to N maxima but can not be confounded with any of these species According to Endert it grew in the forest on a narrow stony mountain ridge covered with humus and was not rare nbsp The type material of N fusca Endert 3955 bears pitchers that appear to be intermediate between typical lower and upper forms Botanist Jan Schlauer has noted differences between the type specimen of N fusca and Sabah plants referred to this species 6 even interpreting plants illustrated in Kurata s Nepenthes of Mount Kinabalu as representing N stenophylla as distinct from N fallax 9 10 c Matthew Jebb does not consider these differences significant enough to merit distinction at the species level He suggests that the type specimen consists of intermediate lower and upper pitchers as opposed to true forms of either making them appear atypical 6 Much of this taxonomic uncertainty stems from the fact that N fusca has not been recollected from the type locality and many similar plants have been lumped under this taxon 8 Matthew Jebb and Martin Cheek attempted to resolve this confusion in their 1997 monograph by interpreting N fusca as a widespread and variable species 11 Subspecies edit Two subspecies of N fusca have been described neither of which is presently thought to represent the species Nepenthes fusca subsp apoensis J H Adam amp Wilcock ex Jebb amp Cheek 1997 11 nom nud N stenophylla Nepenthes fusca subsp kostermansiana J H Adam amp Wilcock ex Jebb amp Cheek 1997 11 nom nud N epiphytica Both were originally coined by J H Adam and C C Wilcock and subsequently published in Jebb and Cheek s 1997 monograph A skeletal revision of Nepenthes Nepenthaceae 11 As these names were published without an adequate description they are both considered nomina nuda 9 The former is likely based on Chai 35939 a specimen collected from Mount Apo 9 Schlauer considers it synonymous with N fallax 9 a taxon that is in turn considered conspecific with N stenophylla by most authors 12 Nepenthes fusca subsp kostermansiana was named from the herbarium material Kostermans 21495 which was collected by Andre Joseph Guillaume Henri Kostermans on October 25 1963 at 1000 m altitude along the Kelai River on Mount Nyapa Njapa Berau Regency East Kalimantan 9 13 It is deposited at the National Herbarium of the Netherlands in Leiden 9 This taxon was initially thought to fall within the variability of N fusca but in 2011 it was described as a distinct species N epiphytica with Kostermans 21495 designated as its holotype 13 Nepenthes maxima edit nbsp An upper pitcher of N maxima from Sulawesi Nepenthes maxima a species native to Sulawesi New Guinea and the Maluku Islands was once thought to extend to Borneo as well Some authors even wrote that it was widespread on the island 5 14 This confusion stemmed from the likeness of N fusca and N maxima and from apparently mislabelled seeds collected by Charles Curtis Curtis was not meticulous in recording where he located individual plants although it was originally believed that he collected N curtisii now considered synonymous with N maxima 15 in Borneo botanist Charles Clarke points out that he also visited Sulawesi on the same trip and N maxima is common there 6 Matthew Jebb and Martin Cheek resolved this confusion in their 1997 monograph by referring a number of Bornean plants identified as N maxima to N fusca thereby excluding the former from the island 11 Nepenthes zakriana edit In 1996 J H Adam and C C Wilcock described Nepenthes curtisii subsp zakriana 3 d Ten years later Adam and Hafiza A Hamid elevated it to species status as Nepenthes zakriana n ɪ ˈ p ɛ n 8 iː z ˌ z ɑː k r i ˈ ae n e not ˌzaekriˈaene citation needed The authors described the taxon as a Sabah endemic growing at elevations of 1200 to 1500 m citation needed Adam and Hafiza wrote that N zakriana consistently differed from Nepenthes fusca by prominent raised midribs extended beyond apex forming an apical glandular appendages on lower lid surface of both upper and lower pitchers and basal half portion of the midrib developed in nail shaped glandular crest citation needed However a number of authors soon voiced their doubts as to whether N zakriana merited species status 16 and in Pitcher Plants of Borneo by Anthea Phillipps Anthony Lamb and Ch ien Lee these features were considered to fall within the natural variability of N fusca 8 Nepenthes sp A edit In his 1997 monograph Nepenthes of Borneo Charles Clarke lists the undescribed taxon Nepenthes sp A which has been recorded from Gunung Mulu National Park in Sarawak 6 It bears a close resemblance to N fusca and may be conspecific with it 6 although its colouration is unusual for the species 17 The pitchers of this plant match J H Adam and C C Wilcock s description 18 of N faizaliana 6 but the latter is now known to have a round lid as opposed to narrowly triangular in N fusca and Nepenthes sp A suggesting that these two taxa are not closely related 6 Clarke proposes that this taxon might fall under Jebb and Cheek s more inclusive concept of N fusca but retains it as an undescribed species because it remains poorly known 6 Nepenthes sp A was first illustrated in a 1988 article by Anthea Phillipps and Anthony Lamb where it was considered an undescribed species 19 Description edit nbsp A young plant from Mesilau Mount Kinabalu Nepenthes fusca is a climbing plant The stem may attain a length of 10 m 8 and is up to 8 mm in diameter Internodes are circular in cross section and up to 7 cm long 6 The leaves of this species are petiolate and coriaceous in texture The lamina or leaf blade is obovate oblong in shape and measures up to 15 cm in length by 6 cm in width Its apex is acute to obtuse and may even be slightly peltate The base of the lamina is gradually attenuate towards the petiole The petiole 4 cm long 5 is grooved lengthwise and bears a pair of narrow wings that form a semi amplexicaul sheath around the stem Up to 3 longitudinal veins are present on either side of the midrib 5 although they are indistinct Pinnate veins are numerous Tendrils measure up to 5 cm in length 6 Rosette and lower pitchers are cylindrical throughout They usually grow to 20 cm in height by 4 cm in width although exceptional specimens up to 28 cm have been recorded 8 A pair of fringed wings 5 mm wide runs down the ventral surface of the pitcher and bears fringe elements measuring up to 10 mm and spaced 6 mm apart 5 The glandular region is restricted to the lower portion of the pitcher s inner surface The glands are small overarched and occur at a density of 600 to 650 per square centimetre 2 The waxy zone is reduced 20 The pitcher mouth is positioned horizontally at the front becoming elongated into a neck at the rear The peristome is flattened and expanded 12 mm wide but bears only indistinct teeth 0 3 mm long 5 The inner portion of the peristome accounts for around 51 of its total cross sectional surface length 20 The pitcher lid or operculum is very narrowly ovate in shape and has a distinctive basal crest on its lower surface An unbranched spur measuring up to 10 mm in length is inserted near the base of the lid 6 nbsp nbsp nbsp nbsp nbsp Lower pitchers of N fusca from the Crocker Range CR and the area around Mount Kinabalu MK From left to right CR MK MK CR and CR Upper pitchers differ markedly in shape being narrowly infundibular in the lower two thirds and becoming widely infundibular above They are similar in size to their lower counterparts typically measuring up to 18 cm with some larger forms reaching 26 cm 8 The tiny digestive glands are overarched and number 1500 to 2000 per square centimetre 2 The pitcher lid is very narrowly triangular with the margins and apex curved downwards 8 In aerial pitchers the wings are reduced to ribs 6 nbsp nbsp nbsp nbsp nbsp Upper pitchers of N fusca from the Crocker Range CR and the area around Mount Kinabalu MK From left to right MK MK CR MK and CR Nepenthes fusca produces a compact racemose inflorescence The peduncle is up to 6 cm long while the rachis is not known to exceed 10 cm Partial peduncles are one or two flowered up to 8 mm long and lack a bract Sepals are elliptic and up to 4 mm long 6 A study of 120 pollen samples taken from the type specimen Endert 3955 found the mean pollen diameter to be 34 8 mm SE 0 6 CV 9 1 21 Developing parts of the plant bear an indumentum of long brown hairs However most of these disappear during the normal course of development and mature parts only have a sparse covering of short brown hairs 6 Ecology edit nbsp Nepenthes fusca N reinwardtiana and N stenophylla growing in an exposed site along a logging road to Mount Murud in Sarawak Nepenthes fusca is endemic to Borneo where its range stretches from Central Kalimantan to northwestern Sabah 6 The species is confined to north western Borneo 1 The species has a wide altitudinal distribution and is typically found at elevations of 1200 to 2500 m above sea level 6 However N fusca has occasionally been reported from lowland hills down to 600 m 22 and in Sarawak it has been found at only 300 m in hill forest 8 Nepenthes fusca is most commonly found as an epiphyte in shady mossy forest on ridge tops where it may grow 10 to 15 m off the ground 8 This makes it particularly difficult to find and often the only evidence of its presence are dead pitchers that have fallen to the forest floor 22 23 24 In this respect it can be considered the ecological equivalent of N bongso from Sumatra 25 More rarely N fusca grows terrestrially in exposed sites near montane forest 6 or along logging roads 8 It is often sympatric with species such as N reinwardtiana N stenophylla and N tentaculata 26 and natural hybrids with all of these have been recorded 8 12 nbsp Nepenthes fusca from Mount Alab in the Crocker Range Locations edit The species has been recorded from numerous mountains across Borneo It can be found at several sites on Mount Kinabalu 27 28 including Kambarangoh the Marai Parai plateau the mountain s East Ridge and the Bambangan River at around 1500 m 5 It also grows on the side of the road connecting the park headquarters and the power station at an elevation of around 1550 m 29 despite being frequently cut back as part of road maintenance 23 This is one of the few places where N fusca can be easily seen by visitors 23 These roadside plants were greatly affected by the El Nino climatic phenomenon of 1997 to 1998 The resulting dry period severely depleted the population such that almost all the plants were destroyed 17 Nearby plants from more sheltered locations fared better and appeared to have recovered by the following year 17 A number of N fusca plants have also been transplanted to the Mesilau nature trail 30 Nepenthes fusca can be viewed along the road leading to the disused Mamut Copper Mine adjacent to Mount Kinabalu There it is sympatric with N macrovulgaris N stenophylla and the natural hybrid N fusca N stenophylla N burbidgeae grows a short distance away 30 The species also occurs on nearby Mount Tambuyukon 5 On Mount Trusmadi N fusca has been observed growing epiphytically on Eleocarpus trees at an elevation of almost 1800 m 31 A lone plant growing on a ridge top at 1962 m was found to have a small bush frog probably Philautus aurantium in one of its partly dried 15 cm long pitchers 32 Scattered plants have also been recorded from a rocky clearing at c 1592 m growing alongside Gleichenia truncata ferns and bamboo orchids these plants have rather unremarkable red speckled but otherwise plain green pitchers though unusually the stem and underside of the midrib are almost black 33 The species is abundant on Mount Alab the highest peak of the Crocker Range where a number of colour variants have been documented including one with wholly green lower pitchers save for red mottling on the interior and underside of the lid 33 An atypical yellow form grows on Mount Lumarku in southwestern Sabah 8 Other notable locations include the Kimanis Keningau Road that runs through the Crocker Range 8 and the summit area of Mount Apo Dari where it grows at 1500 m 14 On Mount Mulu in Sarawak N fusca found below 1200 m appears to occupy a discrete altitudinal zone from N vogelii 1200 1500 m and N hurrelliana above 1500 m both of which are also epiphytes 8 In the Hose Mountains of central Sarawak N fusca is typically found growing as an epiphyte in lower montane forest at 700 1200 m 34 It is also an uncommon inhabitant of roadside embankments there 34 Conservation status edit Charles Clarke noted that since substantial populations of N fusca lie within the boundaries of national parks they are unlikely to become threatened in the foreseeable future 6 Habitat destruction is considered to be the greatest threat to the species s survival in the wild 35 Plant poaching is of far lesser concern as this species is not particularly sought after in the carnivorous plant hobby and its epiphytic habit makes it largely inaccessible to collectors 35 Related species edit nbsp nbsp Upper pitchers of N hurrelliana left and N vogelii right Among the closest relatives of N fusca are the Bornean species N epiphytica 13 N hurrelliana N platychila N stenophylla and N vogelii 8 More broadly it belongs to the loosely defined N maxima complex which also includes N boschiana N chaniana N eymae N faizaliana N klossii and N maxima 13 The enigmatic N mollis which some authors have suggested is conspecific with N hurrelliana 36 may also be closely allied The lower pitchers of N hurrelliana are distinctive but the upper ones bear a close resemblance to those of N fusca Of the Bornean pitcher plant flora only these two species have such a narrowly triangular lid The upper pitchers of N hurrelliana differ in having a horizontal mouth that rises abruptly into a long neck at the back and in having a hirsute basal crest on the underside of the lid 8 Nepenthes hurrelliana is particularly similar to a form of N fusca from the southern portion of the Crocker Range in Sabah This form exhibits a wider peristome longer neck and a more triangular lid than most other examples of the species 8 However the peristome is still not as well developed as in N hurrelliana and the plant lacks the dense indumentum of the latter Furthermore N hurrelliana differs in the distribution of nectar glands on the lower surface of its lid 8 The first known collection of N vogelii made in 1961 was labelled as N fusca 8 In 1969 botanist Shigeo Kurata examined this specimen and noted that it did not fall within the known variation exhibited by N fusca 8 Nevertheless the species remained undescribed until 2002 37 Nepenthes vogelii differs in having much smaller pitchers and lacking appendages on the underside of the lid 37 In addition the lid of N vogelii is broadly triangular as opposed to the narrowly triangular lid of N fusca 8 22 The colour of the pitchers light cream with dark speckles is also distinctive 22 Nepenthes faizaliana also bears a resemblance to N fusca In their description of the former J H Adam and C C Wilcock distinguished these taxa on the basis of inflorescence structure the size of the glandular region on the inner surface of upper pitchers and the development and characteristics of the indumentum 6 18 Nepenthes fusca also differs in having a very narrow pitcher lid as opposed to the orbicular lid of N faizaliana 6 Nepenthes platychila another closely allied species differs from N fusca in having a much wider peristome and lid and lacking appendages on its lower lid surface 8 38 Nepenthes fusca is also thought to be closely related to Sulawesi s N eymae and N maxima which is widespread in Sulawesi New Guinea and the Maluku Islands 39 Natural hybrids editDue its widespread distribution throughout Borneo N fusca forms natural hybrids with a relatively large number of other species However like N fusca itself these are often hard to find due to the species s epiphytic growth habit 8 nbsp Lower pitchers of N burbidgeae N fusca from Mesilau N burbidgeae N fusca edit Nepenthes burbidgeae N fusca has been known since at least the early 1980s when it was found during an expedition to Sabah 26 N fusca N lowii edit This hybrid was initially identified by Charles Clarke as a cross between N chaniana known as N pilosa at the time 40 and N lowii 6 41 However in their 2008 book Pitcher Plants of Borneo Anthea Phillipps Anthony Lamb and Ch ien Lee pointed out that the plants exhibit influences of N fusca such as a triangular lid and an elongated neck 8 The authors noted that both N fusca and N lowii are common on the summit area of Mount Alab where this plant is found whereas N chaniana is rare 8 Another possible parent species N stenophylla is apparently absent from the site 8 This cross was originally discovered by Rob Cantley and Charles Clarke on Bukit Batu Buli in Sarawak 6 Clarke later found larger plants of this hybrid in the Crocker Range of Sabah particularly near the summit of Mount Alab 6 More recently a single plant has been recorded from Mount Trusmadi 33 The pitchers of this cross have a slight constriction in the middle and range in colour from green to dark purple throughout 6 This hybrid differs from N fusca in the presence of bristles on the underside of the lid Conversely it has a dense indumentum on the stem and at the margins of the lamina compared to the virtually glabrous stem and leaves of N lowii It also differs from N lowii in having a more developed peristome which is circular in cross section While lower pitchers of N lowii have prominent teeth those of N fusca N lowii are indistinct In addition a glandular appendage is present on the underside of the lid 6 a trait inherited from N fusca Nepenthes fusca N lowii is difficult to confuse with its putative parent species but is somewhat similar to N chaniana N veitchii The latter hybrid can be distinguished on the basis of its peristome which is wider more flared and less cylindrical In addition this hybrid has a less ovate lid which lacks the bristles characteristic of N lowii and a denser indumentum covering the stem and leaves 6 N fusca N reinwardtiana edit nbsp nbsp Lower pitchers of the putative natural hybrids N fusca N reinwardtiana left and N fusca N stenophylla right A putative cross between N fusca and N reinwardtiana has been recorded 12 Nepenthes naquiyuddinii is generally thought to be a heterotypic synonym of N reinwardtiana 16 but may also represent this hybrid as both putative parent species grow in close proximity to it 8 N fusca N stenophylla edit This hybrid is known from the eastern slopes of Mount Trusmadi where it grows in lower montane forest Both N fusca and N stenophylla are common in this area 8 It has also been recorded from the Mamut copper mine 30 It more closely resembles N stenophylla but differs in the shape of the lid which is more oval shaped 8 N fusca N tentaculata edit Nepenthes fusca N tentaculata was discovered by Linus Gokusing near the summit of Mount Alab where it grows in upper montane forest at elevations of around 1800 to 2000 m 8 It is sympatric with both parent species which are abundant in the area 8 Other hybrids edit Natural crosses with N platychila 42 N rajah 6 and N veitchii 6 have also been recorded The pitchers of N hurrelliana are roughly intermediate in appearance between those of N fusca and N veitchii This has led to speculation regarding the lineage of this species with a number of authors suggesting a possible hybridogenic origin 22 However N hurrelliana is distinct from the natural hybrid N fusca N veitchii and most authors now regard it as a valid species 8 22 43 Cultivation edit nbsp In the wild N fusca is generally found is rather shady conditions upper pitchers growing along a logging road to Mount Murud pictured and this preference is reflected in cultivated plants which show optimal growth under lower light levels than many other Nepenthes species 44 Little information has been published on the growing requirements of N fusca In 2004 professional horticulturist Robert Sacilotto wrote an article for the Carnivorous Plant Newsletter summarising measured tolerances of several highland Nepenthes species based on experiments conducted between 1996 and 2001 44 Sacilotto found N fusca to be tolerant of a wide range of conditions with the exception of plants not treated with fungicides no test groups showed a survival rate of less than 75 Nepenthes fusca was found to tolerate temperatures in the range of 10 to 38 C 50 to 100 F A nighttime drop in temperature below 21 C 70 F was necessary for good growth plants that were not exposed to such a drop grew poorly and produced fewer pitchers The experiments suggested that N fusca grows best when relative humidity is in the range of 65 to 90 44 The species seemed to do best in growing media with a high proportion of organic matter such as a mixture consisting of 10 peat moss chunks 30 perlite and 60 of any combination of Sphagnum moss and fir bark Soil with a slightly acidic pH of 4 5 to 5 0 produced the best results Optimal soil conductivity appeared to lie between 10 and 45 microsiemens 44 An illuminance of 6400 8600 lx 600 800 fc proved to be optimal when plants were grown under sunlight high pressure sodium and metal halide lamps However specimens placed under an even combination of Gro Lux and cool white fluorescent lamps at 5400 7500 lx 500 700 fc exhibited the most vibrant colours although growth rates remained the same Plants moved from the former to the latter light set up showed a significant change in pigmentation green leaf blades turned bronzy and speckles on the pitchers darkened markedly 44 Nepenthes fusca was found to respond well to a quarter strength fertilizer that was applied to the pitchers Ants were also an effective food source 44 Notes edita During September and October 1925 Endert explored the hilly area around Mount Kemul climbing the summit 1 847 m several times and the neighbouring valleys of the Long Mehiang Long Kiau and Long Petak 45 He collected N fusca on October 12 and N mollis on October 17 2 b The Latin description of N fusca from Danser s monograph reads 2 Folia mediocria breviter petiolata lamina lanceolata nervis longitudinalibus utrinque c 2 vagina caulis 1 2 amplectente ascidia rosularum ignota ascidia inferiora magnitudine mediocria parte inferiore anguste ovata os versus subcylindrica parte superiore alis 2 fimbriatis peristomio in collum elongato applanato 4 10 mm lato costis c 1 3 2 3 mm distantibus dentibus c tam longis quam latis operculo anguste ovato subcordato facie inferiore appendice lateraliter applanata ascidia superiora magnitudine mediocria infundibuliformia costis 2 prominentibus peristomio in collum elongato applanato 3 8 mm lato costis 1 3 1 4 mm distantibus dentibus brevissimis operculo anguste ovato subcordato facie inferiore prope basin appendice lateraliter applanata inflorescentia racemis parvus pedicillis inferioribus c 8 mm longis omnibus 1 floris v partim 2 floris indumentum iuventute densissimum denique passim densum breve e pilis patentibus crassis simplicibus v basi ramosis compositum c Some authors treat N fallax in synonymy with N stenophylla 6 11 while others consider them to be two distinct species with plants commonly referred to as N stenophylla actually representing N fallax 46 47 d The subspecies was named after Zakri Abdul Hamid 48 This taxon along with N curtisii sensu J H Adam amp Wilcock itself is considered a heterotypic synonym of N stenophylla by some taxonomists 15 References edit a b Clarke C M 2018 Nepenthes fusca IUCN Red List of Threatened Species 2018 e T40109A143967109 doi 10 2305 IUCN UK 2018 1 RLTS T40109A143967109 en Retrieved 19 November 2021 a b c d e f g h i Danser B H 1928 13 Nepenthes fusca Dans nova spec In The Nepenthaceae of the Netherlands Indies Bulletin du Jardin Botanique de Buitenzorg Serie III 9 3 4 249 438 a b Adam J H amp C C Wilcock 1998 1996 Pitcher plants of Mt Kinabalu in Sabah The Sarawak Museum Journal 50 71 145 171 a b c in Dutch Endert F H 1927 Botanisch en floristisch verslag In D W Buijs H Witkkamp F H Endert H C Siebers amp D F K Bosch Midden Oost Borneo Expeditie 1925 G Kolff amp Co Weltevreden a b c d e f g h i Kurata S 1976 Nepenthes of Mount Kinabalu Sabah National Parks Publications No 2 Sabah National Parks Trustees Kota Kinabalu a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad Clarke C M 1997 Nepenthes of Borneo Natural History Publications Borneo Kota Kinabalu a b Phillipps A amp A Lamb 1996 Pitcher Plants of Borneo Natural History Publications Borneo Kota Kinabalu a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af Phillipps A A Lamb amp C C Lee 2008 Pitcher Plants of Borneo Second Edition Natural History Publications Borneo Kota Kinabalu a b c d e f g h Schlauer J N d Nepenthes fusca Archived 2011 06 17 at the Wayback Machine Carnivorous Plant Database Clarke C M 2006 Introduction In Danser B H The Nepenthaceae of the Netherlands Indies Natural History Publications Borneo Kota Kinabalu pp 1 15 a b c d e f Jebb M H P amp M R Cheek 1997 A skeletal revision of Nepenthes Nepenthaceae Blumea 42 1 1 106 a b c McPherson S R 2009 Pitcher Plants of the Old World 2 volumes Redfern Natural History Productions Poole a b c d Robinson A S J Nerz amp A Wistuba 2011 Nepenthes epiphytica a new pitcher plant from East Kalimantan In McPherson S R New Nepenthes Volume One Redfern Natural History Productions Poole pp 36 51 a b Adam J H C C Wilcock amp M D Swaine 1992 The ecology and distribution of Bornean Nepenthes Archived 2011 07 22 at the Wayback Machine Journal of Tropical Forest Science 5 1 13 25 a b Schlauer J N d Nepenthes curtisii Archived 2011 06 17 at the Wayback Machine Carnivorous Plant Database a b Rice B A 2006 Do you want to tell me about a species I missed The Carnivorous Plant FAQ a b c Steiner H 2002 Borneo Its Mountains and Lowlands with their Pitcher Plants Toihaan Publishing Company Kota Kinabalu a b Adam J H amp C C Wilcock 1991 A new species of Nepenthes Nepenthaceae from Sarawak Blumea 36 1 123 125 Phillipps A amp A Lamb 1988 Pitcher plants of East Malaysia and Brunei Nature Malaysiana 13 4 8 27 a b Bauer U C J Clemente T Renner amp W Federle 2012 Form follows function morphological diversification and alternative trapping strategies in carnivorous Nepenthes pitcher plants Journal of Evolutionary Biology 25 1 90 102 doi 10 1111 j 1420 9101 2011 02406 x Adam J H amp C C Wilcock 1999 Palynological study of Bornean Nepenthes Nepenthaceae Pertanika Journal of Tropical Agricultural Science 22 1 1 7 a b c d e f Clarke C M amp C C Lee 2004 Pitcher Plants of Sarawak Natural History Publications Borneo Kota Kinabalu a b c Clarke C M 2001 A Guide to the Pitcher Plants of Sabah Natural History Publications Borneo Kota Kinabalu Bourke G 2010 The climbing pitcher plants of the Kelabit highlands Archived 2013 04 09 at the Wayback Machine Captive Exotics Newsletter 1 1 4 7 Clarke C M 2001 Nepenthes of Sumatra and Peninsular Malaysia Natural History Publications Borneo Kota Kinabalu a b Lowrie A 1983 Sabah Nepenthes Expeditions 1982 amp 1983 Carnivorous Plant Newsletter 12 4 88 95 Triplitt R 1985 Nepenthes Color Them Unique Carnivorous Plant Newsletter 14 2 40 42 48 49 Malouf P 1995 A visit to Kinabalu Park Carnivorous Plant Newsletter 24 3 64 69 Yeo J 1996 A trip to Kinabalu Park Bulletin of the Australian Carnivorous Plant Society Inc 15 4 4 5 a b c Thong J 2006 Travels around North Borneo Part 1 Archived 2011 07 07 at the Wayback Machine Victorian Carnivorous Plant Society Inc 81 12 17 Marabini J 1984 A field trip to Gunong Trusmadi Carnivorous Plant Newsletter 13 2 38 40 Damit A 2014 A trip to Mount Trus Madi the Nepenthes wonderland Carnivorous Plant Newsletter 43 1 19 22 a b c Fretwell S 2013 Back in Borneo to see giant Nepenthes Part 3 Mt Trusmadi and Mt Alab Victorian Carnivorous Plant Society Inc 109 6 15 a b Lee C C 2002 Nepenthes species of the Hose Mountains in Sarawak Borneo Proceedings of the 4th International Carnivorous Plant Conference Hiroshima University Tokyo 25 30 a b Amazing Species Nepenthes fusca IUCN Red List Salmon B R 1999 Nepenthes mollis Nepenthaceae Rediscovered Carnivorous Plant Newsletter 28 1 24 26 a b Schuiteman A amp E F de Vogel 2002 Nepenthes vogelii Nepenthaceae a new species from Sarawak Blumea 47 3 537 540 Lee C C 2002 Nepenthes species of the Hose Mountains in Sarawak Borneo video The 4th International Carnivorous Plant Conference Tokyo Japan video by Irmgard amp Siegfried R H Hartmeyer D Amato P 1993 Nepenthes eymai Carnivorous Plant Newsletter 22 1 2 21 Clarke C M C C Lee amp S McPherson 2006 Nepenthes chaniana Nepenthaceae a new species from north western Borneo Sabah Parks Journal 7 53 66 in Czech Macak M 2000 Portrety rostlin Nepenthes lowii Hook F Trifid 2000 3 4 51 55 page 2 page 3 page 4 page 5 Lee C C 2002 Nepenthes platychila Nepenthaceae a new species of pitcher plant from Sarawak Borneo The Gardens Bulletin Singapore 54 257 261 Cheek M M Jebb C C Lee A Lamb amp A Phillipps 2003 Nepenthes hurrelliana Nepenthaceae a new species of pitcher plant from Borneo Sabah Parks Nature Journal 6 117 124 a b c d e f Sacilotto R 2004 Experiments with highland Nepenthes seedlings a summary of measured tolerances Carnivorous Plant Newsletter 33 1 26 31 van Steenis Kruseman M J et al 2006 Cyclopaedia of Malesian Collectors Frederik Hendrik Endert Nationaal Herbarium Nederland Schlauer J N d Nepenthes fallax Archived 2011 06 17 at the Wayback Machine Carnivorous Plant Database Schlauer J 1996 N stenophylla once again Carnivorous Plant Mailing List May 31 1996 Wei Shen W 2012 Up close and personal with Professor Emeritus Datuk Zakri Abdul Hamid The Star May 26 2012 Further reading editBeaman J H amp C Anderson 2004 The Plants of Mount Kinabalu 5 Dicotyledon Families Magnoliaceae to Winteraceae Natural History Publications Borneo Kota Kinabalu Benz M J E V Gorb amp S N Gorb 2012 Diversity of the slippery zone microstructure in pitchers of nine carnivorous Nepenthes taxa Arthropod Plant Interactions 6 1 147 158 doi 10 1007 s11829 011 9171 2 Bonhomme V H Pelloux Prayer E Jousselin Y Forterre J J Labat amp L Gaume 2011 Slippery or sticky Functional diversity in the trapping strategy of Nepenthes carnivorous plants New Phytologist 191 2 545 554 doi 10 1111 j 1469 8137 2011 03696 x Bourke G 2011 The Nepenthes of Mulu National Park Carniflora Australis 8 1 20 31 Buch F M Rott S Rottloff C Paetz I Hilke M Raessler amp A Mithofer 2012 Secreted pitfall trap fluid of carnivorous Nepenthes plants is unsuitable for microbial growth Annals of Botany 111 3 375 383 doi 10 1093 aob mcs287 Corner E J H 1996 Pitcher plants Nepenthes In K M Wong amp A Phillipps eds Kinabalu Summit of Borneo A Revised and Expanded Edition The Sabah Society Kota Kinabalu pp 115 121 ISBN 9679994740 Fretwell S 2013 Back in Borneo for giant Nepenthes Part 1 Mesilau Nature Reserve Ranau Victorian Carnivorous Plant Society Inc 107 6 13 Fretwell S 2013 Back in Borneo to see giant Nepenthes Part 2 Mt Tambuyukon and Poring Victorian Carnivorous Plant Society Inc 108 6 15 Kurata S 1969 Mindoro North Borneo Expedition Part 3 The Journal of Insectivorous Plant Society No 47 Lee C C 2000 Recent Nepenthes Discoveries video The 3rd Conference of the International Carnivorous Plant Society San Francisco USA McPherson S R amp A Robinson 2012 Field Guide to the Pitcher Plants of Borneo Redfern Natural History Productions Poole Meimberg H A Wistuba P Dittrich amp G Heubl 2001 Molecular phylogeny of Nepenthaceae based on cladistic analysis of plastid trnK intron sequence data Plant Biology 3 2 164 175 doi 10 1055 s 2001 12897 in German Meimberg H 2002 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 amp 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 Meimberg H S Thalhammer A Brachmann amp G Heubl 2006 Comparative analysis of a translocated copy of the trnK intron in carnivorous family Nepenthaceae Molecular Phylogenetics and Evolution 39 2 478 490 doi 10 1016 j ympev 2005 11 023 in Japanese Oikawa T 1992 Nepenthes fusca Dans In Muyu kusa Nepenthes 無憂草 Nepenthes The Grief Vanishing Parco Co Japan pp 42 45 Renner T amp C D Specht 2011 A sticky situation assessing adaptations for plant carnivory in the Caryophyllales by means of stochastic character mapping International Journal of Plant Sciences 172 7 889 901 doi 10 1086 660882 Thorogood C 2010 The Malaysian Nepenthes Evolutionary and Taxonomic Perspectives Nova Science Publishers New York External links edit nbsp Wikimedia Commons has media related to Nepenthes fusca Nepenthes fusca at ARKive Retrieved from https en wikipedia org w index php title Nepenthes fusca amp oldid 1221075829, wikipedia, wiki, book, books, library,

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