| Nepenthes edwardsiana | |
|---|---|
 | |
| Upper pitcher ofNepenthes edwardsiana fromMount Tambuyukon | |
Scientific classification | |
| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Angiosperms |
| Clade: | Eudicots |
| Order: | Caryophyllales |
| Family: | Nepenthaceae |
| Genus: | Nepenthes |
| Species: | N. edwardsiana |
| Binomial name | |
| Nepenthes edwardsiana | |
| Synonyms | |
Nepenthes edwardsiana/nɪˈpɛnθiːzɛdˌwɔːrdziˈɑːnə/, or thesplendid pitcher-plant,[4] is a carnivorous tropicalpitcher plantendemic toMount Kinabalu and neighbouringMount Tambuyukon inSabah,Malaysian Borneo. It is considered one of the most spectacular of allNepenthes, producing some of the largest pitchers and the most highly developedperistome ribs of any species in thegenus.[5]
Thetype specimen ofN. edwardsiana was collected onMount Kinabalu in 1858[4] byHugh Low andSpenser St. John.[6] Designated asLow s.n., the specimen is deposited at theRoyal Botanic Gardens, Kew.[7]
Nepenthes edwardsiana wasformally described[note a] in 1859 byJoseph Dalton Hooker.[2] Hooker named the species afterGeorge Edwardes, Governor of theCrown Colony of Labuan, at the request of his friendHugh Low.[4][2][note b] Hooker's original description and illustration were reproduced inSpenser St. John'sLife in the Forests of the Far East, published in 1862.[8] St. John wrote the following account ofN. edwardsiana on Mount Kinabalu:[8]
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 theNepenthes 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.
Alfred Russel Wallace made brief mention ofN. edwardsiana in his famous workThe 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".[9]
In subsequent years,N. edwardsiana was featured in a number of publications by eminent botanists such asFriedrich Anton Wilhelm Miquel (1870),[10]Joseph Dalton Hooker (1873),[11]Frederick William Burbidge (1882, 1897),[12][13]Odoardo Beccari (1886),[14]William E. Dixon (1888),[15]Ernst Wunschmann (1891),[16]Otto Stapf (1894),[17]Harry James Veitch (1897),[18]Jacob Gijsbert Boerlage (1900),[19]William Botting Hemsley (1905),[20] andElmer Drew Merrill (1921).[21]
However, most of these publications made only passing mention ofN. edwardsiana. The first work to include significant taxonomic revisions was that ofGünther Beck von Mannagetta und Lerchenau in 1895, "Die GattungNepenthes". Beck was the first to uniteN. edwardsiana andN. villosa, considering the former a variety or form of the latter.[3] He also published the nameNepenthes edgeworthii based on a specimen collected in Borneo byHeinrich Gustav Reichenbach. The specimen,Herb.Reichenbach s.n., is deposited at theUniversity of Viennaherbarium (WU).[7] Beck, like all subsequent authors, consideredN. edgeworthii to be conspecific withN. edwardsiana.[5][7]
Nepenthes edwardsiana was formally reinstated as a valid species inJohn Muirhead Macfarlane's1908 monograph, which included a revised description[note c] and illustration of the species.[22] Macfarlane also wrote aboutN. edwardsiana in theJournal of the Linnean Society in 1914[23] andThe Standard Cyclopedia of Horticulture in 1919.[24]
B. H. Danser treatedN. edwardsiana in synonymy withN. villosa in his seminal monograph, "The Nepenthaceae of the Netherlands Indies", published in 1928. The work included a revisedLatin diagnosis and botanical description ofN. villosa.[25] Eight years later,Hermann Harms once again elevatedN. edwardsiana to species status.[26] This treatment was supported byShigeo Kurata in 1976[27] and has not been challenged since.
A similartaxon fromMount Trusmadi was long considered to beN. edwardsiana.[28] It was described in 1987 asN. edwardsianasubsp.macrophylla byJohannes Marabini.[29] A decade later,Matthew Jebb andMartin Cheek recognised it as a separate species (N. macrophylla) intheir monograph on the genus.[30] This interpretation has been followed by subsequent authors.[4][5][31][32][33]
Nepenthes edwardsiana is a climbing plant. The stem can attain a length of 15 m and is up to 10 mm in diameter.Internodes are up to 35 cm long and circular in cross section.[5]
Leaves arecoriaceous andpetiolate. Thelamina is trulylanceolate in shape and may be up to 30 cm long by 7 cm wide. It has an acute-obtuse apex that is occasionallyacuminate. The base of the lamina is gradually or abruptly contracted at thepetiole. The petiole iscanaliculate and up to 15 cm long. It is generally winged and bears a partlyamplexicaul sheath that clasps the stem for two-thirds to three-quarters of its circumference.[27] One to three longitudinal veins are present on either side of themidrib.Pinnate veins are indistinct.Tendrils are generally between one and two times as long as the lamina[27] and grow to 35 cm in length.[5]
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,[5] although they are usually around 30 cm high.[34] Wings at the front of the pitcher cup are either greatly reduced or absent altogether. On the inner surface of the pitcher, theglandular region is present in the bulbous portion. The mouth is elongated into a neck and has an oblique insertion. Theperistome is cylindrical and up to 30 mm wide. It bears very highly developed teeth and ribs, the latter reaching 20 mm in diameter.[34] The pitcher lid oroperculum iscordate, up to 8 cm wide,[27] and lacks appendages. An unbranchedspur up to 20 mm long is inserted near the base of the lid.[5] Pitchers range in colour from light yellow to dark red.[34][35] 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.[36][37]
Nepenthes edwardsiana has aracemoseinflorescence. Thepeduncle may be up to 30 cm long, whereas theattenuaterachis reaches 20 cm in length.Pedicels are one-flowered,[27] up to 25 mm long, and do not possess abract.Sepals are round to elliptic in shape and up to 5 mm long.[5] A study of 100pollen samples taken from a herbarium specimen (Sands 3651, collected at an altitude of 2,600 m) found the mean pollen diameter to be 34.4μm (SE = 0.5;CV = 7.7%).[38]
Most parts of the plant bear anindumentum of very short hairs, although it is not conspicuous.[5]
Nepenthes edwardsiana varies relatively little across its range; consequently, noinfraspecific taxa have been described.[5]
Nepenthes edwardsiana isendemic to the highland slopes ofMount Kinabalu and the eastern side of neighbouringMount Tambuyukon inSabah,Borneo. On Mount Kinabalu, this species has been recorded from theMarai Paraiplateau, East Ridge,Upper Kolopis River,[27] and an area below the Kambarangoh Telekom station (below Pondok Lowii).[39]Nepenthes edwardsiana has an altitudinal distribution of 1,500–2,700 m above sea level.[4][5]
Anthea Phillipps andAnthony Lamb note that plants growing in theRacemobambosbamboo 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.[4] On theMarai Paraiplateau of Mount Kinabalu,N. edwardsiana grows amongst shrubs up to 5 m high. Pitchers on these plants rarely exceed 30 cm.[36] The species does not grow along the Kinabalu summit trail and so very few visitors to the park manage to see it.[34] A specimen is kept at the "Mountain Garden" near Kinabalu Park Headquarters.[40][41]
During thelast ice age, around 20,000 to 10,000 years ago, Mount Kinabalu had anice cap on its summit. As such, it appears thatN. edwardsiana is a relatively recent species in evolutionary terms.[42]
Contrary to the observations ofFrederick William Burbidge,[43]N. edwardsiana grows not only as anepiphyte, but also occurs terrestrially on moss-covered rocks. It typically grows amongst ridge-top vegetation in densemossy forest.[5] The natural habitat ofN. edwardsiana is constantly moist[44] as the slopes are often enveloped in clouds.[5] Despite generally occurring epiphytically,N. edwardsiana seems to grow mostly in areas withultramafic soils,[5] although it has also been recorded fromsandstonesubstrates.[4]
TheEl Niño climatic phenomenon of 1997 to 1998 had a catastrophic effect on theNepenthes species of Mount Kinabalu.[5] The dry period that followed severely depleted some natural populations.Forest fires broke out in 9 locations in Kinabalu Park, covering a total area of 25 square kilometres and generating large amounts ofsmog.Hugo Steiner recalls being unable to find any pitchers ofN. edwardsiana during a trip to Kinabalu in March 1999.[45] During the El Niño period, many plants were temporarily transferred to the park nursery. These were later replanted in the "Nepenthes Garden" inMesilau. 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, includingN. edwardsiana.[45]
Theconservation status ofN. edwardsiana is listed asVulnerable on the2006 IUCN Red List of Threatened Species.[1] The species has also been classified as Vulnerable by theWorld Conservation Monitoring Centre.[46] However,Charles Clarke notes that since all known populations ofN. edwardsiana lie within the boundaries ofKinabalu National Park and are inaccessible to collectors, they "are unlikely to become threatened in the foreseeable future".[5] Taking this into account, Clarke suggests a revised assessment ofConservation Dependent based on theIUCN criteria.[5]
Nepenthes edwardsiana is most closely related toN. macrophylla andN. villosa. There has been much taxonomic confusion surrounding the status of these threetaxa.[6]
Joseph Dalton Hooker, who described bothN. edwardsiana andN. villosa, noted the similarity between the two species as follows:[2]
This most remarkable plant [N. villosa] resembles that ofedwardsiana 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.
Günther Beck von Mannagetta und Lerchenau was the first to treatN. edwardsiana in synonymy withN. villosa when he published his monograph on the genus in 1895.[3]
In his 1908 monograph,John Muirhead Macfarlane treated the two taxa as distinct species, writing: "Examinatione microscopica probatur, illas species distinctas esse".[22] This was probably based on the scientific view at the time, which held that plants differing anatomically cannot be forms of the same species.[25]
B. H. Danser united the species "[w]ith some hesitation" underN. villosa in his 1928 monograph "The Nepenthaceae of the Netherlands Indies". He suggested thatN. villosa is a stunted form ofN. edwardsiana from higher altitudes, which flowers at a "juvenile stage of development".[25] Danser acknowledged that theindumentum ofN. villosa is more dense than that ofN. edwardsiana, but noted that it "is a difference only of degree".[25]
In 1936,Hermann Harms once again split the two species.[26] InNepenthes of Mount Kinabalu, published in 1976,Shigeo Kurata supported this interpretation based on field observations and reference to the type descriptions.[27]
Nepenthes macrophylla was originally described in 1987 as asubspecies ofN. edwardsiana byJohannes Marabini.[29] It was later elevated to species status byMatthew Jebb andMartin Cheek.[30] This interpretation was supported byCharles Clarke, who noted thatN. edwardsiana andN. villosa "have more in common" thanN. edwardsiana andN. macrophylla.[5]
Nepenthes edwardsiana andN. villosa differ in a number of morphological features. The peristome ofN. villosa is more intricate and its pitchers are not elongated above the hip, unlike those ofN. edwardsiana. InN. edwardsiana, the apex of the lamina is usually acute, compared to the typicallyemarginate apex found inN. villosa. As noted by Danser, theindumentum of these species also differs, withN. villosa being denselyhirsute throughout andN. edwardsiana having an inconspicuous covering of very short hairs. The two taxa can also be distinguished on the basis of their floral morphology; thepedicels ofN. villosa have afiliformbract, while those ofN. edwardsiana do not.[5]
Additionally,N. edwardsiana andN. villosa differ considerably in their altitudinal distributions. The latter species generally occurs at ultra-highland elevations (2,300–3,240 m),[33] whereasN. edwardsiana is found between 1,500 and 2,700 m.[5] Where their altitudinal distributions overlap, they are still identifiable as distinct species.
Nepenthes edwardsiana differs fromN. macrophylla in the structure of its peristome. Although highly developed, the peristome ribs and teeth ofN. macrophylla are considerably shorter than those of eitherN. edwardsiana orN. villosa. The pitcher mouth ofN. 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.Nepenthes macrophylla is also distinguished by its broad,ovate lid. The lower pitchers ofN. edwardsiana andN. 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 ofN. macrophylla being more ovoid and less elongated.[5] As its name suggests,N. macrophylla has very large leaves and these may be twice as long as those ofN. edwardsiana orN. villosa.[4]
WhereasN. edwardsiana andN. villosa are restricted to the Kinabalu area,N. macrophylla is only found near the summit ofMount Trusmadi.[5]
BotanistsMatthew Jebb andMartin Cheek suggest thatN. edwardsiana is related toN. mira, a speciesendemic toPalawan in thePhilippines.[47][48]
Natural hybrids involvingN. edwardsiana appear to be relatively rare and only three have been recorded to date.[5]N. burbidgeae ×N. edwardsiana[5] andN. edwardsiana ×N. rajah[5] have received little attention in the scientific literature, butN. edwardsiana ×N. villosa has been known since the 19th century and was initially described as a separate species,N. harryana.[12]
Nepenthes × harryana is the natural hybrid betweenN. edwardsiana andN. villosa. Its two parent species are very closely related and soN. × harryana, which is roughly intermediate in form, may be difficult to distinguish from either of them.[5][27]
It was originally described as a species in 1882 byFrederick William Burbidge.[12]John Muirhead Macfarlane was the first to realise its hybrid origin and described it as such in his monograph of 1908.[22]B. H. Danser wrote thatN. × harryana might be a hybrid as Macfarlane suggested, or a form ofN. villosa together withN. edwardsiana.[25]
Nepenthes × harryana can be distinguished fromN. villosa on the basis of its pitcher morphology. The pitchers of the hybrid are more cylindrical than those ofN. villosa, whereas theindumentum is more dense than that ofN. edwardsiana. The hip of the pitcher cup, which is found just below the peristome inN. villosa and in the lower quarter ofN. edwardsiana pitchers, is located around the middle ofN. × harryana pitchers. However,N. villosa plants fromMount Tambuyukon are easier to confuse with this hybrid, as they produce pitchers that may be elongated slightly above the hip.[5]
Nepenthes × harryana is known from a ridge above theUpper Kolopis River and from two locations along the Kinabalu summit trail. SinceN edwardsiana does not grow along the summit trail, it cannot be confused with this hybrid there.[5] Burbidge wrote thatN. edwardsiana,N. × harryana, andN. villosa "are quite distinct in zone of the mountain".[12]
Nepenthes 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.[49]
In 2004, professional horticulturist Robert Sacilotto wrote a summary of measured tolerances of highlandNepenthes species, based on experiments conducted between 1996 and 2001.[50] 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 45microsiemens, and soilpH 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 liveSphagnum 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 84 °F), and cool nights, with temperatures of 13 to 16 °C (55 to 61 °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 underHigh Pressure Sodium lamps. Optimal light intensity seemed to be in the region of 7500-9100lx (700-850fc). Soil with apH of 4.8 to 5.4 and conductivity of less than 24 microsiemens produced the best results. Driedfruit flylarvae of the speciesDrosophila melanogaster were fed to the plants once their pitchers reached around 3 mm in height. As the pitchers increased in size, they were fed withants (Acanthomyops sp.).[50]
Ascidia magna, ore lamellis latis disciformibus annularibus remotis instructo.
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.)
Hab.—Kina Balu, north side; alt. 6,000–8,000 feet (Low).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.
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 ad2/3 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.
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).
