''Nepenthes fusca'' , or the dusky pitcher-plant,Phillipps, A. & A. Lamb 1996. ''

Pitcher-Plants of Borneo ''Pitcher-Plants of Borneo'' is a monograph by Anthea Phillipps and Anthony Lamb on the tropical pitcher plants of Borneo. It was first published in 1996 by Natural History Publications (Borneo), in association with the Royal Botanic Gardens, ...

''. Natural History Publications (Borneo), Kota Kinabalu. is a tropical

pitcher plant Pitcher plants are several different carnivorous plants which have modified leaves known as pitfall traps—a prey-trapping mechanism featuring a deep cavity filled with digestive liquid. The traps of what are considered to be "true" pitcher p ...

endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...

Borneo Borneo (; id, Kalimantan) is the third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java, west of Sulawesi, and ea ...

. It is found throughout a wide altitudinal range and is almost always

epiphytic An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...

in nature, primarily growing in

mossy forest A cloud forest, also called a water forest, primas forest, or tropical montane cloud forest (TMCF), is a generally tropical or subtropical, evergreen, montane, moist forest characterized by a persistent, frequent or seasonal low-level cloud c ...

. The specific epithet ''fusca'' is derived from the

Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...

word '' fuscus'', meaning "dark brown" or "dusky", and refers to the colour of the pitchers.

Botanical history

The first known collection of ''N. fusca'' was made by

Frederik Endert Frederik Hendrik Endert (1891 in Semarang, Java – 1953 in Bussum, Netherlands) was a Dutch botanist and plant collector.

on October 12, 1925, from Mount Kemul in

East Kalimantan East Kalimantan ( Indonesian: ) is a province of Indonesia. Its territory comprises the eastern portion of Borneo. It had a population of about 3.03 million at the 2010 census (within the current boundary), 3.42 million at the 2015 census, and 3 ...

, 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), on which Endert also made the only known collection of '' N. mollis''. 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 The Bogor Botanical Gardens ( id, Kebun Raya Bogor) is a botanical garden located in Bogor, Indonesia, 60 km south of central Jakarta. It is currently operated by Indonesian Institute of Sciences (Indonesian: ''Lembaga Ilmu Pengetahuan Ind ...

.Schlauer, J.
''Nepenthes fusca''
Carnivorous Plant Database. Endert wrote about this pitcher plant in a detailed 1927 account of the expedition, although he misidentified it as '' N. veitchii''. ''Nepenthes fusca'' was formally described in 1928 by

Dutch Dutch commonly refers to: * Something of, from, or related to the Netherlands * Dutch people () * Dutch language () Dutch may also refer to: Places * Dutch, West Virginia, a community in the United States * Pennsylvania Dutch Country People E ...

botanist B. H. Danser in his seminal monograph "

The Nepenthaceae of the Netherlands Indies "The Nepenthaceae of the Netherlands Indies" is a seminal monograph by B. H. Danser on the tropical pitcher plants of the Dutch East Indies and surrounding regions. It was originally published in the ''Bulletin du Jardin Botanique de Buitenzorg ...

". Danser based his description solely on ''Endert 3955'', citing no other specimens. He wrote of ''N. fusca'':

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.

Botanist

Jan Schlauer Jan, JaN or JAN may refer to: Acronyms * Jackson, Mississippi (Amtrak station), US, Amtrak station code JAN * Jackson-Evers International Airport, Mississippi, US, IATA code * Jabhat al-Nusra (JaN), a Syrian militant group * Japanese Article Numb ...

has noted differences between the type specimen of ''N. fusca'' and Sabah plants referred to this species, even interpreting plants illustrated in Kurata's ''

Nepenthes of Mount Kinabalu ''Nepenthes of Mount Kinabalu'' is a monograph by Shigeo Kurata on the tropical pitcher plants of Mount Kinabalu and the surrounding area of Kinabalu National Park in Sabah, Borneo. It was published in 1976 by Sabah National Parks Trustees as th ...

'' as representing '' N. stenophylla'' (as distinct from '' N. fallax'').

Matthew Jebb Matthew Hilary Peter Jebb (born 1958) is an Irish botanist and taxonomist specialising in the ant plant genera ''Squamellaria'', '' Myrmecodia'', '' Hydnophytum'', ''Myrmephytum'' and '' Anthorrhiza'', as well as the carnivorous plant genus '' ...

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. 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.

and

Martin Cheek Martin Roy Cheek (born 1960) is a botanist and taxonomist at the Royal Botanic Gardens, Kew.subspecies of ''N. fusca'' have been described, neither of which is presently thought to represent the species: *''Nepenthes fusca'' subsp. ''apoensis'' J.H.Adam &

Wilcock Wilcock is a Norman surname. It may refer to the following: *C. C. Wilcock (born 1946), American taxonomist * Clifford Wilcock (1898–1962), British engineer, company director and politician * Dennis Wilcock, second singer for the band Iron Maide ...

''ex'' Jebb &

Cheek The cheeks ( la, buccae) constitute the area of the face below the eyes and between the nose and the left or right ear. "Buccal" means relating to the cheek. In humans, the region is innervated by the buccal nerve. The area between the insi ...

(1997)Jebb, M.H.P. & M.R. Cheek 1997. A skeletal revision of ''Nepenthes'' (Nepenthaceae). ''Blumea'' 42(1): 1–106. '' nom.nud.'' N. stenophylla''.html" ;"title="'' N. stenophylla''">'' N. stenophylla''*''Nepenthes fusca'' subsp. ''kostermansiana'' J.H.Adam &

''ex'' Jebb &

(1997) '' nom.nud.'' N. epiphytica''.html" ;"title="epenthes_epiphytica.html" ;"title="'' N. epiphytica''">epenthes_epiphytica.html"_;"title="''Nepenthes_epiphytica">N. epiphytica'' Both_were_originally_coined_by_J._H._Adam.html" ;"title="Nepenthes epiphytica">N. epiphytica''">epenthes_epiphytica.html" ;"title="''Nepenthes epiphytica">N. epiphytica'' Both were originally coined by J. H. Adam">Nepenthes epiphytica">N. epiphytica''">epenthes_epiphytica.html" ;"title="''Nepenthes epiphytica">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)". As these names were published without an adequate description, they are both considered ''

nomina nuda In taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published with an adequate descr ...

''. The former is likely based on ''Chai 35939'', a specimen collected from

Mount Apo Mount Apo, also known locally as Apo Sandawa, is a large solfataric, dormant stratovolcano on the island of Mindanao, Philippines. With an elevation of above sea level, it is the highest-mountain in the Philippine Archipelago, Mindanao and 2 ...

. Schlauer considers it synonymous with '' N. fallax'', a taxon that is in turn considered conspecific with '' N. stenophylla'' by most authors. ''Nepenthes fusca'' subsp. ''kostermansiana'' was named from the herbarium material ''Kostermans 21495'', which was collected by

André Joseph Guillaume Henri Kostermans Dr. André Joseph Guillaume Henri 'Dok' Kostermans (Purworejo, 1 July 1906 – Jakarta, 10 July 1994) was an Indonesian botanist of Dutch ancestry. He was born in Purworejo, Java, Dutch East Indies, and educated at Utrecht University, taking ...

on October 25, 1963, at 1000 m altitude along the Kelai River on Mount Nyapa (Njapa),

Berau Regency Berau Regency ( id, Kabupaten Berau) is one of the seven regencies in East Kalimantan province in Indonesia. The capital is the town of Tanjung Redeb. It has an area of 36,962.37 km2 and had a population of 179,079 at the 2010 census and 2 ...

.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. It is deposited at the

National Herbarium of the Netherlands National may refer to: Common uses * Nation or country ** Nationality – a ''national'' is a person who is subject to a nation, regardless of whether the person has full rights as a citizen Places in the United States * National, Maryland, c ...

Leiden Leiden (; in English and archaic Dutch also Leyden) is a city and municipality in the province of South Holland, Netherlands. The municipality of Leiden has a population of 119,713, but the city forms one densely connected agglomeration wi ...

. 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 A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...

''Nepenthes maxima''

'' Nepenthes maxima'', a species native to Sulawesi,

New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; id, Papua, or , historically ) is the world's second-largest island with an area of . Located in Oceania in the southwestern Pacific Ocean, the island is separated from Australia by the wide Torr ...

, and the

Maluku Islands The Maluku Islands (; Indonesian: ''Kepulauan Maluku'') or the Moluccas () are an archipelago in the east of Indonesia. Tectonically they are located on the Halmahera Plate within the Molucca Sea Collision Zone. Geographically they are located ...

, was once thought to extend to Borneo as well. Some authors even wrote that it was widespread on the island.Adam, J.H., C.C. Wilcock & M.D. Swaine 1992
The ecology and distribution of Bornean ''Nepenthes''.
''Journal of Tropical Forest Science'' 5(1): 13–25. This confusion stemmed from the likeness of ''N. fusca'' and ''N. maxima'', and from apparently mislabelled seeds collected by

Charles Curtis Charles Curtis (January 25, 1860 – February 8, 1936) was an American attorney and Republican politician from Kansas who served as the 31st vice president of the United States from 1929 to 1933 under Herbert Hoover. He had served as the Sena ...

. 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'')Schlauer, J.
''Nepenthes curtisii''
Carnivorous Plant Database. in Borneo, botanist

Charles Clarke Charles Rodway Clarke (born 21 September 1950) is a British Labour Party politician, who was the Member of Parliament (MP) for Norwich South from 1997 until 2010, and served as Home Secretary from December 2004 until May 2006. Early life T ...

points out that he also visited Sulawesi on the same trip, and ''N. maxima'' is common there.

and

Martin Cheek Martin Roy Cheek (born 1960) is a botanist and taxonomist at the Royal Botanic Gardens, Kew.J. H. Adam and C. C. Wilcock described ''Nepenthes curtisii'' subsp. ''zakriana''. Ten years later, Adam and Hafiza A. Hamid elevated it to species status as ''Nepenthes zakriana'' (, not ). The authors described the

taxon In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...

as a

Sabah Sabah () is a state of Malaysia located in northern Borneo, in the region of East Malaysia. Sabah borders the Malaysian state of Sarawak to the southwest and the North Kalimantan province of Indonesia to the south. The Federal Territory o ...

growing at elevations of 1200 to 1500 m. 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". However, a number of authors soon voiced their doubts as to whether ''N. zakriana'' merited species statusRice, B.A. 2006
Do you want to tell me about a species I missed?
The Carnivorous Plant FAQ. and in ''

Pitcher Plants of Borneo ''Pitcher-Plants of Borneo'' is a monograph by Anthea Phillipps and Anthony Lamb on the tropical pitcher plants of Borneo. It was first published in 1996 by Natural History Publications (Borneo), in association with the Royal Botanic Gardens, Kew ...

'' by Anthea Phillipps, Anthony Lamb, and

Ch'ien Lee Ch'ien C. Lee ( Chinese: 李乾; pinyin: Lǐ Qián) is a photographer and botanist specialising in the carnivorous plant genus '' Nepenthes''. Lee has described several new ''Nepenthes'' species, including '' N. baramensis''Clarke, C., J.A. ...

, these features were considered to fall within the natural variability of ''N. fusca''.Phillipps, A., A. Lamb & C.C. Lee 2008. ''

''. Second Edition. Natural History Publications (Borneo), Kota Kinabalu.

''Nepenthes'' sp. A

In his 1997 monograph, ''

Nepenthes of Borneo ''Nepenthes of Borneo'' is a monograph by Charles Clarke on the tropical pitcher plants of Borneo.Clarke, C.M. 1997. ''Nepenthes of Borneo''. Natural History Publications (Borneo), Kota Kinabalu. It was first published in 1997 by Natural History ...

'',

lists the undescribed

"''Nepenthes'' sp. A", which has been recorded from

Gunung Mulu National Park The Gunung Mulu National Park is a national park in Miri Division, Sarawak, Malaysia. It is a UNESCO World Heritage Site that encompasses caves and karst formations in a mountainous equatorial rainforest setting. The park is famous for its ca ...

Sarawak Sarawak (; ) is a state of Malaysia. The largest among the 13 states, with an area almost equal to that of Peninsular Malaysia, Sarawak is located in northwest Borneo Island, and is bordered by the Malaysian state of Sabah to the northeast, ...

. It bears a close resemblance to ''N. fusca'' and may be conspecific with it, although its colouration is unusual for the species. The pitchers of this plant match J. H. Adam and C. C. Wilcock's descriptionAdam, J.H. & C.C. Wilcock 1991. A new species of ''Nepenthes'' (Nepenthaceae) from Sarawak. ''Blumea'' 36(1): 123–125. of '' N. faizaliana'', 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. 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. "''Nepenthes'' sp. A" was first illustrated in a 1988 article by Anthea Phillipps and Anthony Lamb, where it was considered an undescribed species.

Description

''Nepenthes fusca'' is a climbing plant. The stem may attain a length of 10 m and is up to 8 mm in diameter. Internodes are circular in cross section and up to 7 cm long. The leaves of this species are petiolate and

coriaceous This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...

in texture. The

lamina Lamina may refer to: Science and technology * Planar lamina, a two-dimensional planar closed surface with mass and density, in mathematics * Laminar flow, (or streamline flow) occurs when a fluid flows in parallel layers, with no disruption betwee ...

or leaf blade is

obovate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular ...

-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 following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular ...

. The base of the lamina is gradually

attenuate In physics, attenuation (in some contexts, extinction) is the gradual loss of flux intensity through a medium. For instance, dark glasses attenuate sunlight, lead attenuates X-rays, and water and air attenuate both light and sound at vari ...

towards the petiole. The petiole (≤4 cm long) 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 This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...

, although they are indistinct.

Pinnate Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, and in ...

veins are numerous.

Tendril In botany, a tendril is a specialized stem, leaf or petiole with a threadlike shape used by climbing plants for support and attachment, as well as cellular invasion by parasitic plants such as '' Cuscuta''. There are many plants that have tend ...

s measure up to 5 cm in length. 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. 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. 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. The waxy zone is reduced. The pitcher mouth is positioned horizontally at the front, becoming elongated into a neck at the rear. The

peristome Peristome (from the Greek ''peri'', meaning 'around' or 'about', and ''stoma'', 'mouth') is an anatomical feature that surrounds an opening to an organ or structure. Some plants, fungi, and shelled gastropods have peristomes. In mosses In mosse ...

is flattened and expanded (≤12 mm wide), but bears only indistinct teeth (≤0.3 mm long). The inner portion of the peristome accounts for around 51% of its total cross-sectional surface length.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. The pitcher lid or operculum is very narrowly ovate in shape and has a distinctive basal crest on its lower surface. An unbranched

spur A spur is a metal tool designed to be worn in pairs on the heels of riding boots for the purpose of directing a horse or other animal to move forward or laterally while riding. It is usually used to refine the riding aids (commands) and to ba ...

measuring up to 10 mm in length is inserted near the base of the lid. 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. The tiny digestive glands are overarched and number 1500 to 2000 per square centimetre. The pitcher lid is very narrowly triangular with the margins and apex curved downwards. In aerial pitchers, the wings are reduced to ribs. ''Nepenthes fusca'' produces a compact

racemose A raceme ( or ) or racemoid is an unbranched, indeterminate type of inflorescence bearing flowers having short floral stalks along the shoots that bear the flowers. The oldest flowers grow close to the base and new flowers are produced as the s ...

inflorescence An inflorescence is a group or cluster of flowers arranged on a Plant stem, stem that is composed of a main branch or a complicated arrangement of branches. Morphology (biology), Morphologically, it is the modified part of the shoot of sperma ...

. The peduncle is up to 6 cm long, while the

rachis In biology, a rachis (from the grc, ῥάχις [], "backbone, spine") is a main axis or "shaft". In zoology and microbiology In vertebrates, ''rachis'' can refer to the series of articulated vertebrae, which encase the spinal cord. In this c ...

is not known to exceed 10 cm. Partial peduncles are one- or two-flowered, up to 8 mm long, and lack a bract.

Sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coine ...

s are elliptic and up to 4 mm long. A study of 120 pollen samples taken from the

type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes th ...

(''Endert 3955'') found the mean pollen diameter to be 34.8 μm ( SE = 0.6; CV = 9.1%). Developing parts of the plant bear an

indumentum In biology, an indumentum (Latin, literally: "garment") is a covering of trichomes (fine "hairs") on a plant Davis, Peter Hadland and Heywood, Vernon Hilton (1963) ''Principles of angiosperm taxonomy'' Van Nostrandpage, Princeton, New Jersey, pa ...

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.

Ecology

''Nepenthes fusca'' is

, where its range stretches from

Central Kalimantan Central Kalimantan ( id, Kalimantan Tengah) is a province of Indonesia. It is one of five provinces in Kalimantan, the Indonesian part of Borneo. Its provincial capital is Palangka Raya and in 2010 its population was over 2.2 million, while the 20 ...

to northwestern

. The species is confined to north-western Borneo. The species has a wide altitudinal distribution and is typically found at elevations of 1200 to 2500 m above sea level. However, ''N. fusca'' has occasionally been reported from lowland hills down to 600 mClarke, C.M. & C.C. Lee 2004. '' Pitcher Plants of Sarawak''. Natural History Publications (Borneo), Kota Kinabalu. and in Sarawak it has been found at only 300 m in hill forest. ''Nepenthes fusca'' is most commonly found as an epiphyte in shady

on ridge tops, where it may grow 10 to 15 m off the ground. 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.Clarke, C.M. 2001. ''

A Guide to the Pitcher Plants of Sabah This list of ''Nepenthes'' literature is a listing of major published works dealing with the tropical pitcher plants of the genus ''Nepenthes''. It includes specialised standalone publications and Biological classification, taxonomic monographs ...

''. Natural History Publications (Borneo), Kota Kinabalu. In this respect, it can be considered the "ecological equivalent" of '' N. bongso'' from Sumatra. More rarely, ''N. fusca'' grows terrestrially in exposed sites near

montane forest Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures fall as elevation increases, causing the ecosystem to stratify. This stratification is a crucial ...

or along logging roads. It is often sympatric with species such as '' N. reinwardtiana'', '' N. stenophylla'', and '' N. tentaculata'',Lowrie, A. 1983
Sabah ''Nepenthes'' Expeditions 1982 & 1983.
''

Carnivorous Plant Newsletter The ''Carnivorous Plant Newsletter'' is the official publication of the International Carnivorous Plant Society (ICPS), the largest such organization in the world. It is headquartered in Walnut Creek, California. History and editorship The newsle ...

'' 12(4): 88–95. and

natural hybrid In biology, a hybrid is the offspring resulting from combining the qualities of two organisms of different breeds, varieties, species or genera through sexual reproduction. Hybrids are not always intermediates between their parents (such as in ...

s with all of these have been recorded. Nepenthesfusca

Locations

The species has been recorded from numerous mountains across Borneo. It can be found at several sites on

Mount Kinabalu Mount Kinabalu ( ms, Gunung Kinabalu, Dusun: ''Gayo Ngaran or Nulu Nabalu'') is the highest mountain in Borneo and Malaysia. With an elevation of , it is third-highest peak of an island on Earth, and 20th most prominent mountain in the worl ...

, including Kambarangoh, the

Marai Parai Marai Parai or Marei Parei is a plateau on the northwestern side of Mount Kinabalu, in Malaysia. The mountain can be climbed from this side, although few parties attempt this route. The first recorded ascent of Mount Kinabalu via Marai Parai was do ...

plateau In geology and physical geography, a plateau (; ; ), also called a high plain or a tableland, is an area of a highland consisting of flat terrain that is raised sharply above the surrounding area on at least one side. Often one or more sides ...

, the mountain's East Ridge, and the Bambangan River at around 1500 m. It also grows on the side of the road connecting the park headquarters and the power station, at an elevation of around 1550 m, despite being frequently cut back as part of road maintenance. This is one of the few places where ''N. fusca'' can be easily seen by visitors. These roadside plants were greatly affected by the

El Niño El Niño (; ; ) is the warm phase of the El Niño–Southern Oscillation (ENSO) and is associated with a band of warm ocean water that develops in the central and east-central equatorial Pacific (approximately between the International Date ...

climatic phenomenon of 1997 to 1998. The resulting dry period severely depleted the population, such that "almost all the plants were destroyed".Steiner, H. 2002.'' Borneo: Its Mountains and Lowlands with their Pitcher Plants''. Toihaan Publishing Company, Kota Kinabalu. Nearby plants from more sheltered locations fared better and appeared to have recovered by the following year. A number of ''N. fusca'' plants have also been transplanted to the Mesilau nature trail.Thong, J. 2006
Travels around North Borneo – Part 1.
'' Victorian Carnivorous Plant Society Inc.'' 81: 12–17. ''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

''N. fusca'' × ''N. stenophylla''; '' N. burbidgeae'' grows a short distance away. The species also occurs on nearby Mount Tambuyukon. On

Mount Trusmadi Mount Trusmadi or Trus Madi ( ms, Gunung Trusmadi) is a mountain located at the Interior Division of Sabah, Malaysia. It is considered as the second highest mountain in both Sabah and Malaysia at , after Mount Kinabalu with Trusmadi offering a ...

, ''N. fusca'' has been observed growing epiphytically on '' Eleocarpus'' trees at an elevation of almost 1800 m.Marabini, J. 1984
A field trip to Gunong Trusmadi.
''

'' 13(2): 38–40. 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. Scattered plants have also been recorded from a rocky clearing at 1592 m, growing alongside '' Gleichenia truncata''

fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...

s 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.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. The species is abundant on

Mount Alab Mount Alab ( ms, Gunung Alab) is a mountain located at the West Coast Division of Sabah, Malaysia. The mountain is located approximately from Sabah's capital of Kota Kinabalu, where it can be highly visible from the city on a clear sky and eas ...

, the highest peak of the

Crocker Range Crocker Range ( ms, Banjaran Crocker) is a mountain range in West Coast Division of Sabah, Malaysia that separates the west and east coast of Sabah. At an average height of , it is the highest mountain range in the state with the range is named ...

, 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. An atypical yellow form grows on Mount Lumarku in southwestern Sabah. Other notable locations include the

Kimanis Kimanis (Chinese: 金馬利) is a town and also a parliamentary constituency in Papar District, West Coast Division of Sabah, Malaysia. It is located approximately 45 kilometres south of the city of Kota Kinabalu, halfway between Papar and B ...

–

Keningau Keningau ( ms, Pekan Keningau) is the capital of the Keningau District in the Interior Division of Sabah, Malaysia. It is the fifth-largest town in Sabah, as well one of the oldest. Keningau is between Tambunan and Tenom. The town had an es ...

Road that runs through the Crocker Range and the summit area of Mount Apo Dari (where it grows at 1500 m). On

Mount Mulu Mount Mulu ( ms, Gunung Mulu) is a sandstone and shale mountain. At 2376 m, it is the second highest mountain in the state of Sarawak, after Mount Murud. It is located within the boundaries of Gunung Mulu National Park, which is named after ...

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. In the Hose Mountains of central Sarawak, ''N. fusca'' is typically found growing as an epiphyte in lower

at 700–1200 m.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. It is also an uncommon inhabitant of roadside embankments there.

Conservation status

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".

Habitat destruction Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...

is considered to be the greatest threat to the species's survival in the wild.Amazing Species: ''Nepenthes fusca''
IUCN Red List. 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.

Related species

Among the closest relatives of ''N. fusca'' are the Bornean species '' N. epiphytica'', '' N. hurrelliana'', '' N. platychila'', '' N. stenophylla'', and '' N. vogelii''. 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''. The enigmatic '' N. mollis'', which some authors have suggested is conspecific with ''N. hurrelliana'',Salmon, B. .1999
''Nepenthes mollis'' (Nepenthaceae)—Rediscovered?
''Carnivorous Plant Newsletter'' 28(1): 24–26. 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

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 Hirsutism is excessive body hair on parts of the body where hair is normally absent or minimal. The word is from early 17th century: from Latin ''hirsutus'' meaning "hairy". It usually refers to a "male" pattern of hair growth in a female that ...

basal crest on the underside of the lid. ''Nepenthes hurrelliana'' is particularly similar to a form of ''N. fusca'' from the southern portion of the

. This form exhibits a wider peristome, longer neck, and a more triangular lid than most other examples of the species. 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. The first known collection of '' N. vogelii'', made in 1961, was labelled as ''N. fusca''. In 1969, botanist

Shigeo Kurata is a Japanese botanist and '' Nepenthes'' taxonomist whose work in the 1960s and 1970s contributed much to the current popularity of these plants.Clarke, C.M. & C.C. Lee 2004. ''Pitcher Plants of Sarawak''. Natural History Publications (Borneo), ...

examined this specimen and noted that it did not fall within the known variation exhibited by ''N. fusca''. Nevertheless, the species remained undescribed until 2002.Schuiteman, A. & E.F. de Vogel 2002. ''Nepenthes vogelii'' (Nepenthaceae): a new species from Sarawak. ''Blumea'' 47(3): 537–540. ''Nepenthes vogelii'' differs in having much smaller pitchers and lacking appendages on the underside of the lid. In addition, the lid of ''N. vogelii'' is broadly triangular as opposed to the narrowly triangular lid of ''N. fusca''. The colour of the pitchers—light cream with dark speckles—is also distinctive. '' 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 In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...

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

. ''Nepenthes fusca'' also differs in having a very narrow pitcher lid, as opposed to the orbicular lid of ''N. faizaliana''. '' 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. ''Nepenthes fusca'' is also thought to be closely related to Sulawesi's '' N. eymae'', and '' N. maxima'', which is widespread in Sulawesi,

, and the

Natural hybrids

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

growth habit. Nepenthes burbidgeae x Nepenthes fusca

''N. burbidgeae'' × ''N. fusca''

Nepenthes burbidgeae ''Nepenthes burbidgeae'' , also known as the painted pitcher plantKurata, S. 1976. '' Nepenthes of Mount Kinabalu''. Sabah National Parks Publications No. 2, Sabah National Parks Trustees, Kota Kinabalu. or Burbidge's Pitcher-Plant,Phillipps, A. ...

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

''N. fusca'' × ''N. lowii''

This hybrid was initially identified by

as a cross between '' N. chaniana'' (known as '' N. pilosa'' at the timeClarke, C.M., C.C. Lee & S. McPherson 2006. ''Nepenthes chaniana'' (Nepenthaceae), a new species from north-western Borneo. ''Sabah Parks Journal'' 7: 53–66.) and '' N. lowii''. However, in their 2008 book, ''

'', Anthea Phillipps, Anthony Lamb and

pointed out that the plants exhibit influences of ''N. fusca'', such as a triangular lid and an elongated neck. The authors noted that both ''N. fusca'' and ''N. lowii'' are common on the summit area of

where this plant is found, whereas ''N. chaniana'' is rare. Another possible parent species, '' N. stenophylla'', is apparently absent from the site. This cross was originally discovered by Rob Cantley and

on Bukit Batu Buli in

. Clarke later found larger plants of this hybrid in the

, particularly near the summit of Mount Alab. More recently a single plant has been recorded from

. The pitchers of this cross have a slight constriction in the middle and range in colour from green to dark purple throughout. This hybrid differs from ''N. fusca'' in the presence of bristles on the underside of the lid. Conversely, it has a dense

on the stem and at the margins of the lamina, compared to the virtually

glabrous Glabrousness (from the Latin '' glaber'' meaning "bald", "hairless", "shaved", "smooth") is the technical term for a lack of hair, down, setae, trichomes or other such covering. A glabrous surface may be a natural characteristic of all or part of ...

stem and leaves of ''N. lowii''. It also differs from ''N. lowii'' in having a more developed

, 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, 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.

''N. fusca'' × ''N. reinwardtiana''

A putative cross between ''N. fusca'' and '' N. reinwardtiana'' has been recorded.McPherson, S.R. 2009. ''

Pitcher Plants of the Old World ''Pitcher Plants of the Old World'' is a two-volume monograph by Stewart McPherson on the pitcher plants of the genera '' Nepenthes'' and ''Cephalotus''. It was published in May 2009 by Redfern Natural History Productions and covers all species ...

''. 2 volumes. Redfern Natural History Productions, Poole. '' Nepenthes naquiyuddinii'' is generally thought to be a

heterotypic synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linn ...

of ''N. reinwardtiana'', but may also represent this hybrid, as both putative parent species grow in close proximity to it.

''N. fusca'' × ''N. stenophylla''

This hybrid is known from the eastern slopes of

, where it grows in lower

. Both ''N. fusca'' and '' N. stenophylla'' are common in this area. It has also been recorded from the Mamut copper mine. It more closely resembles ''N. stenophylla'', but differs in the shape of the lid, which is more oval-shaped.

''N. fusca'' × ''N. tentaculata''

''Nepenthes fusca'' × '' N. tentaculata'' was discovered by Linus Gokusing near the summit of

, where it grows in upper

at elevations of around 1800 to 2000 m. It is sympatric with both parent species, which are abundant in the area.

Other hybrids

Natural crosses with '' N. platychila'', '' N. rajah'', and '' N. veitchii'' 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. However, ''N. hurrelliana'' is distinct from the natural hybrid ''N. fusca'' × ''N. veitchii'' and most authors now regard it as a valid species.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.

Cultivation

Little information has been published on the growing requirements of ''N. fusca''. In 2004, professional horticulturist Robert Sacilotto wrote an article for the ''

'', summarising measured tolerances of several highland ''Nepenthes'' species based on experiments conducted between 1996 and 2001.Sacilotto, R. 2004
Experiments with highland ''Nepenthes'' seedlings: a summary of measured tolerances
''

'' 33(1): 26–31. 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 . A nighttime drop in temperature below 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 Humidity is the concentration of water vapor present in the air. Water vapor, the gaseous state of water, is generally invisible to the human eye. Humidity indicates the likelihood for precipitation, dew, or fog to be present. Humidity dep ...

is in the range of 65 to 90%. 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 ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, peat moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store wa ...

chunks, 30%

perlite Perlite is an amorphous volcanic glass that has a relatively high water content, typically formed by the hydration of obsidian. It occurs naturally and has the unusual property of greatly expanding when heated sufficiently. It is an industrial ...

, and 60% of any combination of ''

Sphagnum ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, peat moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store wa ...

'' 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. An

illuminance In photometry (optics), photometry, illuminance is the total luminous flux incident on a surface, per unit area. It is a measure of how much the incident light illuminates the surface, wavelength-weighted by the luminosity function to correlate w ...

of 6400–8600 lx (600–800 fc) proved to be optimal when plants were grown under sunlight,

high pressure sodium A sodium-vapor lamp is a gas-discharge lamp that uses sodium in an excited state to produce light at a characteristic wavelength near 589 nm. Two varieties of such lamps exist: low pressure and high pressure. Low-pressure sodium lamps are ...

, and

metal halide lamp A metal (from Greek μέταλλον ''métallon'', "mine, quarry, metal") is a material that, when freshly prepared, polished, or fractured, shows a lustrous appearance, and conducts electricity and heat relatively well. Metals are typicall ...

s. However, specimens placed under an even combination of Gro-Lux and cool white

fluorescent lamp A fluorescent lamp, or fluorescent tube, is a low-pressure mercury-vapor gas-discharge lamp that uses fluorescence to produce visible light. An electric current in the gas excites mercury vapor, which produces short-wave ultraviolet ligh ...

s 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. ''Nepenthes fusca'' was found to respond well to a quarter-strength

fertilizer A fertilizer (American English) or fertiliser (British English; see spelling differences) is any material of natural or synthetic origin that is applied to soil or to plant tissues to supply plant nutrients. Fertilizers may be distinct from ...

that was applied to the pitchers.

Ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...

s were also an effective food source.

Notes

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. He collected ''N. fusca'' on October 12 and '' N. mollis'' on October 17. b.The

description of ''N. fusca'' from Danser's monograph reads:

''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'', while others consider them to be two distinct species, with plants commonly referred to as ''N. stenophylla'' actually representing ''N. fallax''. d.The subspecies was named after Zakri Abdul Hamid.Wei-Shen, W. 2012
Up close and personal with Professor Emeritus Datuk Zakri Abdul Hamid
''The Star'', May 26, 2012. This taxon, along with ''N. curtisii'' (''sensu'' J.H.Adam & Wilcock) itself, is considered a

of '' N. stenophylla'' by some taxonomists.

References

External links

''Nepenthes fusca''
at

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{{Taxonbar, from=Q1106815 Carnivorous plants of Asia fusca Endemic flora of Borneo Epiphytes Plants described in 1928 Taxa named by Benedictus Hubertus Danser