Nepenthes edwardsiana 3834632 223638381 2008-07-05T00:46:49Z Lightbot 7178666 Units/dates/other {{Taxobox | name = ''Nepenthes edwardsiana'' | status = VU | status_system = iucn2.3 | image = Nepenthes edwardsiana entire ASR 052007 tambu.jpg | image_width = 200px | image_caption = Upper pitcher of ''Nepenthes edwardsiana'' from Mount Tambuyukon | regnum = [[Plant]]ae | divisio = [[Magnoliophyta]] | classis = [[Magnoliopsida]] | ordo = [[Caryophyllales]] | familia = [[Nepenthaceae]] | genus = ''[[Nepenthes]]'' | species = '''''N. edwardsiana''''' | binomial = ''Nepenthes edwardsiana'' | binomial_authority = [[Hugh Low|Low]] ''ex'' [[Joseph Dalton Hooker|Hook.f.]] (1859) | synonyms = *''Nepenthes edgeworthii''<br><small>[[Heinrich Gustav Reichenbach|Rchb.f.]] ''ex'' [[Günther von Mannagetta und Lërchenau Beck|Beck]] (1895)</small> }} '''''Nepenthes edwardsiana''''' ({{pronEng|nəˈpɛnθiːz ˈɛdwɔːrdziːˌɑːnə}}), or the '''Splendid Pitcher-Plant''',<ref name=P&L>Phillipps, A. & A. Lamb 1996. ''Pitcher-Plants of Borneo''. Natural History Publications (Borneo), Kota Kinabalu.</ref> is a tropical [[pitcher plant]] [[endemism|endemic]] to [[Mount Kinabalu]] and neighbouring [[Mount Tambuyukon]] in [[Sabah]], [[Malaysian Borneo]]. It is perhaps the most spectacular of all ''[[Nepenthes]]'', producing some of the largest pitchers and the most highly developed [[peristome]] ribs of any species in the [[genus]].<ref name=Clarke>Clarke, C.M. 1997. ''Nepenthes of Borneo''. Natural History Publications (Borneo), Kota Kinabalu.</ref> {{TOClimit|limit=2}} ==Botanical history== The [[type specimen]] of ''N. edwardsiana'' was collected on [[Mount Kinabalu]] in 1858<ref name=P&L /> by a climbing team consisting of [[Hugh Low]], [[Frederick William Burbidge]], and [[Spenser St. John]].<ref name=taxonomy>Turnbull, J. & A. Middleton 1984. {{PDFlink|[http://carnivorousplants.org/cpn/articles/CPNv13n3p61_67.pdf Tedious Trudge Through the Torturous Turmoils of Taxonomy.]|1.44&nbsp;[[Mebibyte|MiB]]<!-- application/pdf, 1520380 bytes -->}} ''[[Carnivorous Plant Newsletter]]'' '''13'''(3): 61–67.</ref> Designated as ''Low s.n.'', the specimen is deposited at the [[Royal Botanic Gardens, Kew]].<ref name=Schlauer>Schlauer, J. 2006. [http://www.omnisterra.com/bot/cp_home.cgi?name=Nepenthes+edwardsiana&submit=Submit+Query&search=all ''Nepenthes edwardsiana'']. Carnivorous Plant Database.</ref> ''N. edwardsiana'' was formally described{{Ref_label|A|I|none}} in 1859 by [[Joseph Dalton Hooker]].<ref name=Hooker>Hooker, J.D. 1859. On the origin and the development of the pitchers of ''Nepenthes'', with an account of some new Bornean plants of that genus. ''The Transactions of the Linnean Society of London'' '''22'''(4): 415–424.</ref> Hooker named the species after George Edwards, Governor of the [[Crown Colony of Labuan]], at the request of his friend [[Hugh Low]].<ref name=P&L />{{Ref_label|B|II|none}} Hooker's original description and illustration were reproduced in [[Spenser St. John]]'s ''Life in the Forests of the Far East'', published in 1862.<ref name=John>St. John, S. 1862. ''Life in the Forests of the Far East; or, Travels in northern Borneo''. 2 volumes. London: Smith, Elder & Co..</ref> St. John wrote the following account of ''N. edwardsiana'' on Mount Kinabalu:<ref name=John /> <blockquote>As we ascended, we left the brushwood and entered a tangled jungle, in which few of the trees were large. The spur of the mountain became very narrow, sometimes not much wider than the path, and was greatly encumbered at one part by the twining stems of the ''Nepenthes Edwardsiana''. This handsome plant was not, however, much diffused along the spur, but confined to a space about a quarter of a mile in length, and climbed upon the trees around, with its fine pitchers hanging from all the lower boughs. We measured one plant and it was twenty feet in length, quite smooth, and the leaves of a very acute shape at both ends. It is a long, cylindrical, finely-frilled pitcher, growing on every leaf; one we picked measured twenty-one inches and a half long, by two and a half in breadth. They swell out a little towards the base, which is bright pea green, the rest of the cylinder being of a brilliant brick-red colour. Its mouth is nearly circular, the border surrounding it being finely formed of thin plates about a sixth of an inch apart, and about the same in height, and both of a flesh colour; the handsome lid is of a circular shape. The dried specimen forwarded to Dr. Hooker only measured eighteen inches. The plant is epiphytal, growing on casuarinas (''species nova''). The pitchers of the young creepers precisely resemble those of the older ones, except in size.</blockquote> [[Alfred Russel Wallace]] made brief mention of ''N. edwardsiana'' in his famous work ''[[The Malay Archipelago]]'', first published in 1869: "Another, ''Nepenthes Edwardsiania'', has a narrow pitcher twenty inches long; while the plant itself grows to a length of twenty feet".<ref>Wallace, A.R. 1869. ''The Malay Archipelago, Volume I''. Macmillan, London.</ref> In subsequent years, ''N. edwardsiana'' was featured in a number of publications by eminent botanists such as [[Friedrich Anton Wilhelm Miquel]] (1870),<ref>Miquel, F.A.G. 1870. ''Nepenthes''. ''Illustrations de la flore l'Archipel Indien'' '''1''': 1–48.</ref> [[Joseph Dalton Hooker]] (1873),<ref>Hooker, J.D. 1873. Nepenthaceae. In: A. De Candolle. ''Prodromus systematis universalis regni vegetabilis'' '''17''': 90–105.</ref> [[Frederick William Burbidge]] (1882, 1897),<ref name=Burbidge>Burbidge, F.W. 1882. Notes on the new ''Nepenthes''. ''The Gardeners' Chronicle'' '''17'''(1): 56.</ref><ref>Burbidge, F.W. 1897. Note on ''Nepenthes''. ''Journal of the Royal Horticultural Society'' '''21'''(2): 256–262.</ref> [[Odoardo Beccari]] (1886),<ref>Beccari, O. 1886. Rivista delle specie del genere ''Nepenthes''. ''Malesia'' '''3''': 1–15.</ref> [[William E. Dixon]] (1888),<ref>Dixon, W.E. 1888. ''Nepenthes'' notes. ''The Gardeners' Chronicle'' '''3'''(1): 170.</ref> [[Ernst Wunschmann]] (1891),<ref>Wunschmann, E. 1891. Nepenthaceae. In: A. Engler & K. Prantl. ''Die natürlichen Pflanzenfamilien'' '''3'''(2): 253–260.</ref> [[Otto Stapf]] (1894),<ref>Stapf, O. 1894. On the flora of Mount Kinabalu, in North Borneo. ''The Transactions of the Linnean Society of London'' '''4''': 96–263.</ref> [[Harry James Veitch]] (1897),<ref>Veitch, H.J. 1897. ''Nepenthes''. ''Journal of the Royal Horticultural Society'' '''21'''(2): 226–255.</ref> [[Jacob Gijsbert Boerlage]] (1900),<ref>Boerlage, J.G. 1900. ''Nepenthes''. In: ''Handleiding tot de kennis der flora van Nederlandsch Indië'', Volume 3, Part 1. pp. 53–54.</ref> [[William Botting Hemsley]] (1905),<ref>Hemsley, W.B. 1905. The pitchers of ''Nepenthes''. ''The Gardeners' Chronicle'' '''27'''(1): 241–242 & 260.</ref> and [[Elmer Drew Merrill]] (1921).<ref>Merrill, E.D. 1921. A bibliographic enumeration of Bornean plants. ''Journal of the Straits branch of the Royal Asiatic Society'', special number. pp. 281–295.</ref> However, most of these publications made only passing mention of ''N. edwardsiana''. The first work to include significant taxonomic revisions was that of [[Günther von Mannagetta und Lërchenau Beck]] in 1895. Beck was the first to unite ''N. edwardsiana'' and ''N. villosa'', making the former a [[heterotypic synonym]] of the latter.<ref name=Beck>Beck, G.v.M.u.L. 1895. Die Gattung ''Nepenthes''. ''Wiener Illustrierte Gartenzeitung''.</ref> He also described ''Nepenthes edgeworthii'' based on a specimen collected in Borneo by [[Heinrich Gustav Reichenbach]]. The specimen, ''Herb.Reichenbach s.n.'', is deposited at the [[University of Vienna]] [[herbarium]] (WU).<ref name=Schlauer /> ''N. edgeworthii'' is now considered conspecific with ''N. edwardsiana''.<ref name=Clarke /><ref name=Schlauer /> {| align=right style="margin: 0 0 0 1em" | [[Image:Nepenthes edwardsiana Life in the forests of the Far East.png|thumb|center|182px|Illustration of ''N. edwardsiana'' upper pitchers from St. John's ''Life in the forests of the Far East'']] | [[Image:Nepenthes edwardsiana Macfarlane illustration.jpg|thumb|center|150px|Line drawing from Macfarlane's monograph on the Nepenthaceae]] |} ''N. edwardsiana'' was formally reinstated as a valid species in [[John Muirhead Macfarlane]]'s 1908 monograph, which included a revised description{{Ref_label|C|III|none}} and illustration of the species.<ref name=Macfarlane>Macfarlane, J.M. 1908. Nepenthaceae. In: A. Engler. ''Das Pflanzenreich IV'', III, Heft 36: 1–91.</ref> Macfarlane also wrote about ''N. edwardsiana'' in the ''Journal of the Linnean Society'' in 1914<ref>Macfarlane, J.M. 1914. Nepenthaceae. In: L.S. Gibbs. A contribution to the flora and the plant formations of Mount Kinabalu and the Highlands of British North Borneo. ''Journal of the Linnean Society'' '''42''': 125–127.</ref> and ''The Standard Cyclopedia of Horticulture'' in 1919.<ref>Macfarlane, J.M. 1919. ''Nepenthes''. In: L.H. Bailey. ''The Standard Cyclopedia of Horticulture'', Volume 4. pp. 2122–2130.</ref> [[B. H. Danser]] treated ''N. edwardsiana'' in synonymy with ''N. villosa'' in his seminal monograph "[[The Nepenthaceae of the Netherlands Indies]]", published in 1928. The work included a revised [[Latin]] diagnosis and botanical description of ''N. villosa''.<ref name=Danser>Danser, B.H. 1928. The Nepenthaceae of the Netherlands Indies. ''Bulletin de Jardin de Botanique, Buitenzorg'', Série III, '''9'''(3–4): 249–438.</ref> Eight years later, [[Hermann Harms]] once again elevated ''N. edwardsiana'' to species status.<ref name=Harms>Harms, H. 1936. Nepenthaceae. In: A. Engler & K. Prantl. ''Die natürlichen Pflanzenfamilien'', 2 aufl. band 17b.</ref> This treatment was supported by [[Shigeo Kurata]] in 1976<ref name=Kurata>Kurata, S. 1976. ''Nepenthes of Mount Kinabalu''. Sabah National Parks Publications No. 2, Sabah National Parks Trustees, Kota Kinabalu.</ref> and has not been challenged since. A similar [[taxon]] from [[Mount Trus Madi]] was long considered to be ''N. edwardsiana''.<ref>Marabini, J. 1984. {{PDFlink|[http://www.carnivorousplants.org/cpn/articles/CPNv13n2p38_40.pdf A Field Trip to Gunong Trusmadi.]|442&nbsp;[[Kibibyte|KiB]]<!-- application/pdf, 453522 bytes -->}} ''[[Carnivorous Plant Newsletter]]'' '''13'''(2): 38–40.</ref> It was described in 1987 as ''N. edwardsiana'' [[subspecies|subsp.]] ''macrophylla'' by [[Johannes Marabini]].<ref name=Marabini>Marabini, J. 1987. Eine neue Unterart von ''Nepenthes edwardsiana'' Hook.fil. sowie Anmerkungen zur Taxonomie der Gattung ''Nepenthes'' L.. ''Mitt. Bot. Staatssamml. München'' '''23''': 423–429.</ref> A decade later, [[Matthew Jebb]] and [[Martin Cheek]] recognised it as a separate species (''[[Nepenthes macrophylla|N. macrophylla]]'') in their monograph on the genus.<ref name=J&C>Jebb, M.H.P. & M.R. Cheek 1997. A Skeletal Revision of ''Nepenthes'' (Nepenthaceae). ''Blumea'' '''42'''(1): 1–106.</ref> This interpretation has been followed by subsequent authors.<ref name=P&L /><ref name=Clarke /> ==Description== ''N. edwardsiana'' is a climbing plant. The stem can attain a length of 15 m and is up to 10 mm in diameter. [[Internode]]s are up to 35 cm long and cylindrical in cross section.<ref name=Clarke /> [[Image:Nepenthes edwardsiana ASR 052007 tambu.jpg|thumb|left|Detail of the peristome on an upper pitcher]] Leaves are [[wikt:coriaceous|coriaceous]] and [[wikt:petiolate|petiolate]]. The [[lamina]] is truly [[wikt:lanceolate|lanceolate]] in shape and may be up to 30 cm long by 7 cm wide. It has an acute-obtuse apex that is occasionally [[wikt:acuminate|acuminate]]. The base of the lamina is gradually or abruptly contracted at the [[Petiole (botany)|petiole]]. The petiole is [[wikt:canaliculate|canaliculate]] and up to 15 cm long. It is generally winged and bears a partly [[wikt:amplexicaul|amplexicaul]] sheath that clasps the stem for two-thirds to three-quarters of its circumference.<ref name=Kurata /> One to three longitudinal veins are present on either side of the [[midrib]]. [[wikt:pinnate|Pinnate]] veins are indistinct. [[Tendril]]s are generally between one and two times as long as the lamina<ref name=Kurata /> and grow to 35 cm in length.<ref name=Clarke /> The pitcher base is bulbous to ovoid, with the pitcher cup becoming cylindrical in the upper two-thirds to three-quarters. The pitchers are some of the largest in the genus, sometimes exceeding 50 cm in height and 15 cm in width,<ref name=Clarke /> although they are usually around 30 cm high.<ref name=Sabah>Clarke, C.M. 2001. ''A Guide to the Pitcher Plants of Sabah''. Natural History Publications (Borneo), Kota Kinabalu.</ref> Wings at the front of the pitcher cup are either greatly reduced or absent altogether. On the inner surface of the pitcher, the [[gland]]ular region is present in the bulbous portion. The mouth is elongated into a neck and has an oblique insertion. The [[peristome]] is cylindrical and up to 30 mm wide. It bears very highly developed teeth and ribs, the latter reaching 20 mm in diameter.<ref name=Sabah /> The pitcher lid or [[operculum (botany)|operculum]] is [[wikt:cordate|cordate]], up to 8 cm wide,<ref name=Kurata /> and lacks appendages. An unbranched [[spur (biology)|spur]] up to 20 mm long is inserted near the base of the lid.<ref name=Clarke /> Pitchers range in colour from light yellow to dark red.<ref name=Sabah /><ref>D'Amato, P. 1998. ''The Savage Garden — Cultivating Carnivorous Plants''. Ten Speed Press, Berkley, California.</ref> The inner surface of the pitcher is usually white, contrasting sharply with the rich colouration of the outer surface. Most parts of the pitcher are very flexible, including the peristome ribs, with only the pitcher base, where the digestive zone is located, being rigid.<ref name=Anfraix>Anfraix, R. 2005. Discovery of ''Nepenthes edwardsiana'' at Marai Parai. ''Acta Botanica Gallica'' '''152'''(2): 205–214.</ref> ''N. edwardsiana'' has a [[raceme|racemose]] [[inflorescence]]. The [[peduncle (botany)|peduncle]] may be up to 30 cm long, where as the [[wikt:attenuate|attenuate]] [[rachis]] reaches 20 cm in length. [[Pedicel (botany)|Pedicels]] are one-flowered,<ref name=Kurata /> up to 25 mm long, and do not possess a [[bract]]. [[Sepal]]s are round to elliptic in shape and up to 5 mm long.<ref name=Clarke /> Most parts of the plant bear an [[indumentum]] of very short hairs, although it is not conspicuous.<ref name=Clarke /> ''N. edwardsiana'' varies relatively little across its range; consequently, no [[infraspecific taxon|infraspecific taxa]] have been described.<ref name=Clarke /> ==Ecology== ===Habitat and distribution=== {{Location map | Borneo Topography | label = | lat = 6.08333 | long = 116.55 | position = left | width = 300 | float = right | caption = Location of Mount Kinabalu }} [[Image:Nepenthes on Mount Kina Balu diagram Macfarlane.jpg|thumb|350px|Diagram from Macfarlane's 1908 monograph showing hot and cold air currents on Mount Kinabalu. The approximate altitudinal distribution of ''N. edwardsiana'' is indicated on the right.]] ''N. edwardsiana'' is [[endemism|endemic]] to the highland slopes of [[Mount Kinabalu]] and the eastern side of neighbouring [[Mount Tambuyukon]] in [[Sabah]], [[Borneo]]. On Mount Kinabalu, this species has been recorded from the [[Marai Parai]] [[plateau]], East Ridge, [[Kolopis River|Upper Kolopis River]], and an area below the [[Radio Sabah]] [[relay station]].<ref name=Kurata /> ''N. edwardsiana'' has an altitudinal distribution of 1500 to 2700 m above sea level.<ref name=P&L /><ref name=Clarke /> [[Anthea Phillipps]] and [[Anthony Lamb]] note that plants growing in the ''[[Racemobambos]]'' [[bamboo]] forest on Mount Tambuyukon produce some of the longest and finest pitchers, having a pronounced waist and ranging in colour from pink to reddish-orange.<ref name=P&L /> On the [[Marai Parai]] [[plateau]] of Mount Kinabalu, ''N. edwardsiana'' grows amongst shrubs up to 5 m high. Pitchers on these plants rarely exceed 30 cm.<ref name=Anfraix /> The species does not grow along the Kinabalu summit trail and so very few visitors to the park manage to see it.<ref name=Sabah /> A specimen is kept at the "Mountain Garden" near Kinabalu Park Headquarters.<ref>Malouf, P. 1995. {{PDFlink|[http://carnivorousplants.org/cpn/articles/CPNv24n3p64_69.pdf A visit to Kinabalu Park.]}} ''[[Carnivorous Plant Newsletter]]'' '''24'''(3): 64–69.</ref><ref>Malouf, P. 1995. {{PDFlink|[http://carnivorousplants.org/cpn/articles/CPNv24n4p104_108.pdf A visit to Kinabalu Park.]}} ''[[Carnivorous Plant Newsletter]]'' '''24'''(4): 104–108.</ref> Mount Kinabalu was only formed around 1 million years ago and during the [[last ice age]], around 20,000 to 10,000 years ago, it had an [[ice cap]] on its summit. As such, it appears that ''N. edwardsiana'' is a relatively recent species in evolutionary terms.<ref>Risner, J.K. 1987. {{PDFlink|[http://carnivorousplants.org/cpn/articles/CPNv16n4p115_118.pdf The Mystery of the ''Nepenthes'', or Just How Did They Get There?]}} ''[[Carnivorous Plant Newsletter]]'' '''16'''(4): 115–118.</ref> Contrary to the observations of [[Frederick William Burbidge]],<ref>Burbidge, F.W. 1880. ''The Gardens of the Sun''. Murray, London.</ref> ''N. edwardsiana'' grows not only as an [[epiphyte]], but also occurs terrestrially on moss-covered rocks. It typically grows amongst ridge-top vegetation in dense [[mossy forest]].<ref name=Clarke /> The natural habitat of ''N. edwardsiana'' is constantly moist<ref>Toyoda, Y. 1972. {{PDFlink|[http://carnivorousplants.org/cpn/articles/CPNv01n4p62_63.pdf ''Nepenthes'' and I - Mt. Kinabalu (Borneo, Malaysia) Trip.]|116&nbsp;[[Kibibyte|KiB]]<!-- application/pdf, 118897 bytes -->}} ''[[Carnivorous Plant Newsletter]]'' '''1'''(4): 62–63.</ref> as the slopes are often enveloped in clouds.<ref name=Clarke /> Despite generally occurring epiphytically, ''N. edwardsiana'' seems to grow mostly in areas with [[Ultramafic rock|ultramafic]] soils,<ref name=Clarke /> although it has also been recorded from [[sandstone]] [[substrate]]s.<ref name=P&L /> ===Threats and conservation status=== The [[El Niño]] climatic phenomenon of 1997 to 1998 had a catastrophic effect on the ''Nepenthes'' species of Mount Kinabalu.<ref name=Clarke /> The dry period that followed severely depleted some natural populations. [[Wildfire|Forest fires]] broke out in 9 locations in Kinabalu Park, covering a total area of 25 square kilometres and generating large amounts of [[smog]]. [[Hugo Steiner]] recalls being unable to find any pitchers of ''N. edwardsiana'' during a trip to Kinabalu in March 1999.<ref name=Steiner>Steiner, H. 2002.'' Borneo: Its Mountains and Lowlands with their Pitcher Plants''. Toihaan Publishing Company, Kota Kinabalu.</ref> During the El Niño period, many plants were temporarily transferred to the park nursery. These were later replanted in the "''Nepenthes'' Garden" in [[Mesilau]]. Since then, Ansow Gunsalam has established a nursery close to the Mesilau Lodge at the base of Kinabalu Park to protect the endangered species of that area, including ''N. edwardsiana''.<ref name=Steiner /> The [[conservation status]] of ''N. edwardsiana'' is listed as [[Vulnerable species|Vulnerable]] on the [[IUCN Red List|2006 IUCN Red List of Threatened Species]].<ref name=IUCN>Schnell, D., P. Catling, G. Folkerts, C. Frost, R. Gardner, ''et al.'' 2000. [http://www.iucnredlist.org/search/details.php/40107/all ''Nepenthes edwardsiana'']. ''2006 [[IUCN Red List|IUCN Red List of Threatened Species]]''. [[World Conservation Union|IUCN]] 2006. Retrieved on 11 May 2006. Listed as Vulnerable (VU B1+2e, D1 v2.3).</ref> The species has also been classified as Vulnerable by the [[World Conservation Monitoring Centre]].<ref>Simpson, R.B. 1995. ''Nepenthes'' and Conservation. ''Curtis's Botanical Magazine'' '''12''': 111–118.</ref> However, [[Charles Clarke (botanist)|Charles Clarke]] notes that since all known populations of ''N. edwardsiana'' lie within the boundaries of [[Kinabalu National Park]] and are inaccessible to collectors, "they are unlikely to become threatened in the forseeable<sup>''<nowiki>[</nowiki>[[sic]]]''</sup> future".<ref name=Clarke /> Taking this into account, Clarke suggests a revised assessment of [[Conservation Dependent]] based on the [[World Conservation Union|IUCN]] criteria.<ref name=Clarke /> ==Related species== ''N. edwardsiana'' is most closely related to ''[[Nepenthes macrophylla|N. macrophylla]]'' and ''[[Nepenthes villosa|N. villosa]]''. There has been much taxonomic confusion surrounding the status of these three [[taxon|taxa]].<ref name=taxonomy /> ===Changing taxonomic status=== [[Joseph Dalton Hooker]], who described both ''N. edwardsiana'' and ''N. villosa'', noted the similarity between the two species as follows:<ref name=Hooker /> <blockquote>This most remarkable plant [''N. villosa''] resembles that of ''edwardsiana'' in so many respects, especially in the size, form and disposition of the distant lamellae of the mouth, that I am inclined to suspect that it may be produced by young plants of that species, before it arrives at a stage when the pitchers have elongated necks.</blockquote> [[Günther von Mannagetta und Lërchenau Beck]] was the first to treat ''N. edwardsiana'' in synonymy with ''N. villosa'' when he published his monograph on the genus in 1895.<ref name=Beck /> In his 1908 monograph, [[John Muirhead Macfarlane]] treated the two taxa as distinct species, writing: "Examinatione microscopica probatur, illas species distinctas esse".<ref name=Macfarlane /> This was probably based on the scientific view at the time, which held that plants differing anatomically cannot be forms of the same species.<ref name=Danser /> [[B. H. Danser]] united the species "[w]ith some hesitation" in his 1928 monograph "[[The Nepenthaceae of the Netherlands Indies]]". He suggested that ''N. villosa'' is a stunted form of ''N. edwardsiana'' from higher altitudes, which flowers at a "juvenile stage of development".<ref name=Danser /> Danser acknowledged that the [[indumentum]] of ''N. villosa'' is more dense than that of ''N. edwardsiana'', but noted that it "is a difference only of degree".<ref name=Danser /> In 1936, [[Hermann Harms]] once again split the two species.<ref name=Harms /> In ''Nepenthes of Mount Kinabalu'', published in 1976, [[Shigeo Kurata]] supported this interpretation based on field observations and reference to the type descriptions.<ref name=Kurata /> ''[[Nepenthes macrophylla|N. macrophylla]]'' was originally described in 1987 as a [[subspecies]] of ''N. edwardsiana'' by [[Johannes Marabini]].<ref name=Marabini /> It was later elevated to species status by [[Matthew Jebb]] and [[Martin Cheek]].<ref name=J&C /> This interpretation was supported by [[Charles Clarke (botanist)|Charles Clarke]], who noted that ''N. edwardsiana'' and ''N. villosa'' "have more in common" than ''N. edwardsiana'' and ''N. macrophylla''.<ref name=Clarke /> ===Species differences=== [[Image:Nepenthes villosa.jpg|thumb|Upper pitcher of ''N. villosa'']] ''N. edwardsiana'' and ''N. villosa'' differ in a number of morphological features. The peristome of ''N. villosa'' is more intricate and its pitchers are not elongated above the hip, unlike those of ''N. edwardsiana''. In ''N. edwardsiana'', the apex of the lamina is usually acute, compared to the typically [[wikt:emarginate|emarginate]] apex found in ''N. villosa''. As noted by Danser, the [[indumentum]] of these species also differs, with ''N. villosa'' being densely [[wikt:hirsute|hirsute]] throughout and ''N. edwardsiana'' having an inconspicuous covering of very short hairs. The two taxa can also be distinguished on the basis of their floral morphology; the [[pedicel (botany)|pedicels]] of ''N. villosa'' have a [[wikt:filiform|filiform]] [[bract]], while those of ''N. edwardsiana'' do not.<ref name=Clarke /> Additionally, ''N. edwardsiana'' and ''N. villosa'' differ considerably in their altitudinal distributions. The latter species occurs at ultrahighland elevations (2340 to 3240 m),<ref name=P&L /> where as ''N. edwardsiana'' is found between 1500 and 2700 m.<ref name=Clarke /> Where their altitudinal distributions overlap, they are still identifiable as distinct species. ''N. edwardsiana'' differs from ''N. macrophylla'' in the structure of its peristome. Although highly developed, the peristome ribs and teeth of ''N. macrophylla'' are considerably shorter than those of either ''N. edwardsiana'' or ''N. villosa''. The pitcher mouth of ''N. macrophylla'' is distinctive in that it rises gradually towards the lid, while at the same time not forming a pronounced neck. In addition, the mouth of this species has a much more oblique insertion than its relatives. ''N. macrophylla'' is also distinguished by its broad, [[wikt:ovate|ovate]] lid. The lower pitchers of ''N. edwardsiana'' and ''N. macrophylla'' are quite similar in shape, although in the latter species the hip is always positioned in the upper portion of the pitcher cup. The upper pitchers of these species are more distinct, with those of ''N. macrophylla'' being more ovoid and less elongated.<ref name=Clarke /> As its name suggests, ''N. macrophylla'' has very large leaves and these may be twice as long as those of ''N. edwardsiana'' or ''N. villosa''.<ref name=P&L /> Where as ''N. edwardsiana'' and ''N. villosa'' are restricted to the Kinabalu area, ''N. macrophylla'' is only found near the summit of [[Mount Trus Madi]].<ref name=Clarke /> ==Natural hybrids== Natural hybrids involving ''N. edwardsiana'' appear to be relatively rare and only three have been recorded to date.<ref name=Clarke /> ''[[Nepenthes burbidgeae|N. burbidgeae]]'' × ''N. edwardsiana''<ref name=Clarke /> and ''N. edwardsiana'' × ''[[Nepenthes rajah|N. rajah]]''<ref name=Clarke /> have received little attention in the scientific literature, but ''N. edwardsiana'' × ''[[Nepenthes villosa|N. villosa]]'' has been known since the 19th century and was initially described as a separate species, ''N. harryana''.<ref name=Burbidge /> ===''N. edwardsiana'' × ''N. villosa''=== {{main|Nepenthes × harryana}} [[Image:Nepenthesxharryana.jpg|thumb|Lower pitcher of ''N. × harryana'']] ''N. × harryana'' is the natural hybrid between ''N. edwardsiana'' and ''[[Nepenthes villosa|N. villosa]]''. Its two parent species are very closely related and so ''N. × harryana'', which is roughly intermediate in form, may be difficult to distinguish from either of them.<ref name=Kurata /><ref name=Clarke /> It was originally described as a species in 1882 by [[Frederick William Burbidge]].<ref name=Burbidge /> [[John Muirhead Macfarlane]] was the first to realise its hybrid origin and described it as such in his monograph of 1908.<ref name=Macfarlane /> [[B. H. Danser]] wrote that ''N. × harryana'' might be a hybrid as Macfarlane suggested, or a form of ''N. villosa'' together with ''N. edwardsiana''.<ref name=Danser /> ''N. × harryana'' can be distinguished from ''N. villosa'' on the basis of its pitcher morphology. The pitchers of the hybrid are more cylindrical than those of ''N. villosa'', where as the [[indumentum]] is more dense than that of ''N. edwardsiana''. The hip of the pitcher cup, which is found just below the peristome in ''N. villosa'' and in the lower quarter of ''N. edwardsiana'' pitchers, is located around the middle of ''N. × harryana'' pitchers. However, ''N. villosa'' plants from [[Mount Tambuyukon]] are easier to confuse with this hybrid, as they produce pitchers that may be elongated slightly above the hip.<ref name=Clarke /> ''N. × harryana'' is known from a ridge above the [[Kolopis River|Upper Kolopis River]] and from two locations along the Kinabalu summit trail. Since ''N edwardsiana'' does not grow along the summit trail, it cannot be confused with this hybrid there.<ref name=Clarke /> Burbidge wrote that ''N. edwardsiana'', ''N. × harryana'', and ''N. villosa'' "are quite distinct in zone of the mountain".<ref name=Burbidge /> ==Cultivation== ''N. edwardsiana'' is very rare in cultivation and little information has been published on its growing requirements. Generally speaking, it is an alpine plant that requires highland conditions to grow well.<ref>Lowrie, A. 1983. {{PDFlink|[http://carnivorousplants.org/cpn/articles/CPNv12n4p88_95.pdf Sabah ''Nepenthes'' Expeditions 1982 & 1983.]}} ''[[Carnivorous Plant Newsletter]]'' '''12'''(4): 88–95.</ref> In 2004, professional horticulturist Robert Sacilotto wrote a summary of measured tolerances of highland ''Nepenthes'' species, based on experiments conducted between 1996 and 2001.<ref name=test>Sacilotto, R. 2004. [http://carnivorousplants.org/cpn/samples/v33n1p26_31.html Experiments with highland ''Nepenthes'' seedlings: A Summary of Measured Tolerances]. ''[[Carnivorous Plant Newsletter]]'' '''33'''(1): 26–31.</ref> Out of all of the studied species, ''N. edwardsiana'' proved to be the most challenging. [[Cotyledon]]-stage seedlings showed a 100% mortality rate when exposed to the following conditions: [[relative humidity]] constantly over 90%, water droplets present on the leaves, soil conductivity over 45 [[Siemens (unit)|microsiemens]], and soil [[pH]] above 6. However, several plants grew well in a substrate consisting of 50% [[perlite]], 30% ''[[Sphagnum]]'' moss, 10% [[peat moss]] chunks, and 10% [[fir]] bark. A top dressing of live ''Sphagnum'' was found to provide a good anchoring point for developing roots. Humidity levels of 65 to 85% appeared to be optimal, although more mature plants over 1 year old were able to tolerate exposure to relative humidity in the range of 90 to 99% for up to three days. The highest growth rate was exhibited by plants that experienced warm days, with temperatures of 21 to 29°C (70 to 85°F), and cool nights, with temperatures of 13 to 16°C (55 to 60°F). The seedlings grew very slowly during the first 8 months, but their growth rate increased significantly after they reached approximately 2 cm in diameter. The plants were grown under [[Sodium vapor lamp|High Pressure Sodium]] lamps. Optimal light intensity seemed to be in the region of 7500-9100 [[Lux|lx]] (700-850 [[Foot-candle|fc]]). Soil with a [[pH]] of 4.8 to 5.4 and conductivity of less than 24 microsiemens produced the best results. Dried [[fruit fly]] [[larva]]e of the species ''[[Drosophila melanogaster]]'' were fed to the plants once their pitchers reached around 3 mm in height. As the pitchers increased in size, they were fed with [[ant]]s (''[[Acanthomyops]]'' sp.).<ref name=test /> ==Notes== <div class="references-small"> '''<small>I</small>.''' {{Note_label|A|I|none}}Hooker's original description of ''N. edwardsiana'' from 1859 reads:<ref name=Hooker /> <blockquote> &nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;''Ascidia magna, ore lamellis latis disciformibus annularibus remotis instructo.''<br> &nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;Nepenthes Edwardsiana, Low. MSS. — Foliis (6" longis) crasse coriaceis longe petiolatis ellipticis, ascidiis magnis crasse pedunculatis cylindraceis basi ventricosis 8—18" longis, ore lamellis annularibus distantibus rigidis magnis cristato, collo elongato erecto operculo cordato-rotundato, racemo simplici, rachi pedicellisque ferrugineo-tomentosis.—(''Tab.'' LXX.)<br> &nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;''Hab.''—Kina Balu, north side; alt. 6,000–8,000 feet (''Low'').<br> &nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;The leaves, ascidia, and pitchers sent by Mr. Low are all old, and nearly glabrous; but the young parts,—rachis, peduncles of the panicle, and the calyx—are covered with ferruginous tomentum. One of the pitchers sent is eighteen inches long from the base to the apex of the erect operculum; it is two and a half inches in diameter below the mouth, one and a half at the narrowest part (about one-third distant from the base) and the swollen part above the base is about two inches in diameter. The beautiful annular discs which surround the mouth are three-quarters of an inch in diameter.</blockquote> '''<small>II</small>.''' {{Note_label|B|II|none}}In ''Nepenthes of Mount Kinabalu'', [[Shigeo Kurata]] gives a different derivation for the name: "The specific epithet ''edwardsiana'' refers to S. Edwards—a botanical draughtsman, who made many drawings for the Botanical Magazine."<ref name=Kurata /> '''<small>III</small>.''' {{Note_label|C|III|none}}Macfarlane's revised description of ''N. edwardsiana'' from 1908 reads:<ref name=Macfarlane /> <blockquote> &nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;26. N. Edwardsiana Hook f. in Trans. Linn. Soc. XXII. (1859) 420 t. LXX; Spencer St. John, Life in For. Far East I. (1862) 335 t. 336; Burbidge, Gard. of Sun (1880) 100, 108, 280, 284, 344, et in Gard. Chron. ser. 2. XVII. (1882) 56; Macfarlane in Ann. of Bot. VII. (1893) 433; Stapf in Trans. Linn. Soc. Bot. IV. (1894) 69; G. Beck in Wien. Ill. Gart. Zeitg. (1895) 183; Burbidge in Journ. Roy. Hort. Soc. XXI. (1897) 258. — ''N. Edgeworthii'' Reichb. f. herb. ex G. Beck l. c. 183. — Planta saepe epiphytica, Casuarineas et arbores alias ascendens. Caulis 3—9 m longus X 8—12 mm crassus, cylindricus v. ± trigonus, juventute ferrugineopuberulus demum glaber. Folia 20—30 cm longa X 6—10 cm lata, coriacea, petiolata; petiolus 6—10 cm longus, validus, alatus, basis ad <sup>2</sup>/<sub>3</sub> amplexicaulis, alae non v. leviter decurrentes pilis longis fuscis sparsis obsitae; lamina elliptica v. oblonga inferne in petiolum angustata, ad apicem rotundata v. in cirrhum attenuata, supra glabra subtus glabra et punctata, marginibus et costa ± hirsuta, nervi longitudinales 4—5 obscuri, par intimum 25—35 mm, par secundum 29—40 mm, par tertium 30—44 mm, par quartum 31—43 mm a costa remotum, nervi transversi radiantes v. ascendentes, 2—3 mm inter se separati; cirrhus 30—60 cm longus X 3—6 mm crassus, cylindricus, in basim ascidii recurvatam abrupte ampliatus, juventute dense ferrugineo-pubescens demum glaber; ascidia 20—50 cm longa, X 4—7 cm lata, monomorphia, ± puberula v. glabra, dimidio inferiore leviter ventricosa et coeruleo-viridia, dimidio superiore cylindrica, rubra, alae ventrales 0, v. a basi sursum gradatim ampliatae et ± ciliatae, os subcirculare; peristomium magnum 1,5—2 cm latum, obliquum, postice in collum 3—5 cm altum elongatum, caro tinctum, superficie in lamellas transversas 6—8 mm profundas et 4—8 mm inter se separatas elevata, margo exterior recurvatus, interior in dentes magnos deflexus; operculum 6—10 cm longum X 5—9 cm latum, cordato-ovatum v. cordato-orbiculare, extus glabrum intus glandulosum, glandulae multae, parvae, dense aggregatae; ascidium intus per dimidium superius v. profundius glaucopurpureum opacum et deducens, inferne nitidum glandulosum et detinens, glandulae superne parvae discretae profunde immersae, inferne magnae approximatae et subexsertae. Inflorescentia 30—40 cm longa, rufo-tomentella; pedunculus 15—25 cm longus validus; racemus ± densiflorus; pedicelli 1,5—2 cm longi uniflori. Flores 7—12 mm lati, ♂ minores quam ♀. Sepala elliptica v. obovata, extus ferrugineo-pubescentia, intus glandulis paucis parvis mediis adspersa. Columna staminea 3 mm longa, inferne ferrugineo-pubescens superne glabra; antherae 8—12, uniseriatae v. antheris duabus superioribus transversis. Capsula 20—22 mm longa X 5 mm lata, fusiformis, breviter pedunculata, ferrugineo-puberula v. brunneo-nitida, valvae stigmatibus obtusis triangulis terminatae. Semina tenuia, 8—9 mm longa. — Fig. 16.<br> &nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;S. W. Malayische Provinz; Borneo: Auf der nördlichen Seite des Berges Kina Balu bis 2000 m (Low!); "on the sunny southern spur" up to 2700 m (Burbidge!); auf dem Kiau-Rücken, 1500 m (Burbidge!); "spur of Kina Balu" (Spencer St. John).</blockquote> </div> ==References== {{reflist|2}} ==External links== {{commons|Nepenthes edwardsiana}} * [http://www.arkive.org/species/GES/plants_and_algae/Nepenthes_edwardsiana/ ''Nepenthes edwardsiana''] at [[ARKive]] * [http://www.ars-grin.gov/cgi-bin/npgs/html/taxon.pl?431459 ''Nepenthes edwardsiana''] at [[Germplasm Resources Information Network|NPGS/GRIN]] {{Nepenthes}} [[Category:Carnivorous plants of Asia]] [[Category:Nepenthes]]