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. & A. Lamb 1996. ''Pitcher-Plants of Borneo''. Natural History Publications (Borneo), Kota Kinabalu. is a tropical pitcher plant with a patchy distribution around Mount Kinabalu and neighbouring Mount Tambuyukon in Sabah, Borneo.Clarke, C.M. 1997. '' Nepenthes of Borneo''. Natural History Publications (Borneo), Kota Kinabalu.

Botanical history

''Nepenthes burbidgeae'' was discovered on Mount Kinabalu in 1858 by Hugh Low and Spenser St. John. St. John wrote the following account of finding the species near the Marai Parai plateau:

Crossing the Hobang, a steep climb led us to the western spur, along which our path lay; here, at about 4000 ft 200 m Mr. Low found a beautiful white and spotted pitcher-plant which he considered the prettiest of the twenty-two species of ''Nepenthes'' with which he was then acquainted; the pitchers are white and covered in a most beautiful manner with spots of an irregular form, of a rosy pink colour.

Frederick William Burbidge was one of the first to collect the plant in 1878, although he did not succeed in introducing it into cultivation. The type specimen of ''N. burbidgeae'', ''Burbidge s.n.'', was collected on the Marai Parai plateau of Mount Kinabalu and is deposited at the Royal Botanic Gardens, Kew.Schlauer, J. 2006
''Nepenthes burbidgeae''
Carnivorous Plant Database. A duplicate specimen is held at the New York Botanical Garden.Specimen Details: ''Nepenthes burbidgei'' Hook. f. ex Burb.
The New York Botanical Garden.

''Nepenthes burbidgeae'' appeared as an unnamed species in Burbidge's 1880 book ''The Gardens of the Sun''. Joseph Dalton Hooker named ''N. burbidgeae'' after Burbidge's wife, though the name only appeared in an unpublished manuscript. The specific epithet is attributed to Burbidge as he used it in a letter to '' The Gardeners' Chronicle'' in 1882. It reads:

Nepenthes Burbidgeae, Hook. f. ''MSS.'', is a lovely thing, as yet unintroduced : pitchers pure white, semi-translucent like egg-shell, porcelain-white, with crimson or blood-tinted blotches. Lid blotched and dotted with crimson-purple. It is a very distinct plant, with triangular stems, 50 feet long, and the margins of the leaves decurrent.

In 1894, Otto Stapf identified specimens belonging to ''N. burbidgeae'' as ''N. phyllamphora'', a taxon that is now considered synonymous with '' N. mirabilis''.

In two articles authored by Burbidge in 1894 and 1896, the name of this species was written as ''N. burbidgei''. This name is considered a ''sphalma typographicum'' (misprint) of ''N. burbidgeae'', although it appeared in a number of other works by authors such as Odoardo Beccari (1886), John Muirhead Macfarlane (1908), and Elmer Drew Merrill (1921). Herbarium material also bears this spelling of the name.

Seventy years after its discovery, ''N. burbidgeae'' remained a poorly known species. This is reflected in the writing of B. H. Danser in his seminal 1928 monograph, "The Nepenthaceae of the Netherlands Indies", where he suggests a close relative in '' N. pilosa'':Danser, B.H. 1928
7. ''Nepenthes Burbidgeae'' BURB.
In: The Nepenthaceae of the Netherlands Indies. ''Bulletin du Jardin Botanique de Buitenzorg'', Série III, 9(3–4): 249–438.

This species has only been found twice on Mt. Kinabalu and is very insufficiently known. I have not ventured to unite it with any other. ''N. pilosa'', though doubtless the most nearly related species, is certainly different.

In 1981, Australian botanist Allen Lowrie reported that the fluid in unopened pitchers of ''N. burbidgeae'' is effective in stopping external bleeding. Lowrie cited two examples of researchers in the field successfully using this fluid on cuts and wounds.

Description

''Nepenthes burbidgeae'' is a strong climber that quickly enters the vining stage. The stem reaches 15 m in length and is up to 18 mm in diameter. Internodes are cylindrical to triangular in cross section and up to 12 cm long.

The leaves of this species are coriaceous and petiolate. The lamina or leaf blade is oblong in shape and up to 40 cm long by 10 cm wide. It has an acute apex and its base is typically abruptly attenuate. The petiole is winged, up to 15 cm long, and clasps the stem. It is often decurrent into two narrow wings that extend down the stem. Three to four longitudinal veins are present on either side of the midrib. Pinnate veins are inconspicuous. Tendrils are up to 30 cm long.

Rosette and lower pitchers are rounded-infundibular or conical in shape. Unlike the pitchers of many other ''Nepenthes'' species, those of ''N. burbidgeae'' have no obvious constriction in the middle. The lower pitchers are relatively large, being up to 25 cm high by 10 cm wide. A pair of fringed wings, measuring up to 10 mm in width, runs down the front of each pitcher. The glandular region, which bears minute overarched glands, covers the basal half of the pitcher's inner surface. The pitcher mouth is round and elongated into a short neck at the rear. The peristome is flattened and expanded, measuring up to 30 mm in width. Its inner margin is lined with a series of small but distinct teeth. The inner portion of the peristome accounts for around 49% 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 ovate and up to 8 cm wide. It bears a distinct keel as well as a characteristic hooked appendage on its lower surface. An unbranched spur (≤12 mm long) is inserted near the base of the lid.

Upper pitchers are similar to their terrestrial counterparts in most respects, even retaining the same colouration. However, they are smaller, reaching only 13 cm in height and 7 cm in width. They are infundibular in the basal third and globose above. In aerial pitchers, a pair of ribs is present in place of wings.

''Nepenthes burbidgeae'' has a racemose inflorescence. The peduncle is up to 25 cm long, while the rachis reaches 30 cm in length. Partial peduncles may be one- or two-flowered and are up to 15 mm long. Sepals are ovate and up to 5 mm long.

Most parts of the plant are covered in a sparse indumentum of short hairs. The margins of the lamina are lined with brown hairs up to 3 mm long.

''Nepenthes burbidgeae'' has a very restricted range and exhibits relatively little variability. As such, no infraspecific taxa have been described.

Ecology

Habitat and distribution

''Nepenthes burbidgeae'' is endemic to Kinabalu National Park, where it has a patchy distribution around Mount Kinabalu and neighbouring Mount Tambuyukon. Specifically, it has been recorded from the Marai Parai plateau, Mamut copper mine, and Pig Hill.Thong, J. 2006
Travels around North Borneo – Part 1.
''Victorian Carnivorous Plant Society Inc.'' 81: 12–17. On Pig Hill, it grows at 1900–1950 mAdam, 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. and is sympatric with '' N. rajah'', '' N. tentaculata'', and the natural hybrid '' N. Ã— alisaputrana''.Thong, J. 2006
Travels around North Borneo – Part 2.
''Victorian Carnivorous Plant Society Inc.'' 82: 6–12. The altitudinal range of this species is often quoted as 1200–1800 m above sea level,McPherson, S.R. 2009. ''Pitcher Plants of the Old World''. 2 volumes. Redfern Natural History Productions, Poole. but some sources give a lower limit of 1100 m and upper limit of 2250 mCheek, M.R. & M.H.P. Jebb 2001. Nepenthaceae. ''Flora Malesiana'' 15: 1–157. or even 2300 m.

Mount Kinabalu was only formed around 1 million years ago and, during the last ice age (approximately 20,000 to 10,000 years ago), it had an ice cap on its summit. As such, it appears that ''N. burbidgeae'' is a relatively recent species in evolutionary terms.

''Nepenthes burbidgeae'' is probably the rarest of the ''Nepenthes'' species native to Mount Kinabalu. Its typical habitat consists of mossy forest or montane forest, where it often grows in low scrub and exposed areas on the tops of steep ridges. The species is restricted to ultramafic soils.Jebb, M.H.P. & M.R. Cheek 1997. A skeletal revision of ''Nepenthes'' (Nepenthaceae). ''Blumea'' 42(1): 1–106. In more exposed areas, ''N. burbidgeae'' is often found climbing amongst bushes of ''Leptospermum javanicum''. At some localities it has also been recorded from bamboo forest.

''Nepenthes burbidgeae'' can often be found growing amongst populations of '' N. edwardsiana'', '' N. rajah'', and '' N. tentaculata'', and hybrids with all of these species have been recorded.

Threats and conservation status

The El Niño climatic phenomenon of 1997 to 1998 had a catastrophic effect on the ''Nepenthes'' species of Mount Kinabalu. The dry period that followed severely depleted some natural populations. Forest fires broke out in 9 locations in Kinabalu Park, covering a total area of 25 square kilometres and generating large amounts of smog. Hugo Steiner recalls being struck by the scarcity of ''N. burbidgeae'' pitchers observed on Mount Kinabalu during a trip in 1999.Steiner, H. 2002.'' Borneo: Its Mountains and Lowlands with their Pitcher Plants''. Toihaan Publishing Company, Kota Kinabalu. At the time of the El Niño, 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. burbidgeae''.

The conservation status of ''N. burbidgeae'' is listed as Endangered on the 2006 IUCN Red List of Threatened Species based on an assessment carried out in 2000. This does not agree with an informal assessment made by Charles Clarke in 1997, who classified the species as Vulnerable based on the IUCN criteria. However, Clarke noted that since all known populations of ''N. burbidgeae'' lie within the boundaries of Kinabalu National Park and are inaccessible to collectors, they "are unlikely to become threatened in the foreseeable future". Taking this into account, he suggested a revised assessment of Conservation Dependent.

Related species

''Nepenthes burbidgeae'' is easily distinguished from other species in the genus on the basis of its pitcher shape and colouration, as well as the hook-shaped appendage on the underside of the lid.Clarke, C.M. 2001. ''A Guide to the Pitcher Plants of Sabah''. Natural History Publications (Borneo), Kota Kinabalu. The only other Bornean ''Nepenthes'' species with a similarly developed appendage are '' N. chaniana'' and '' N. pilosa''.Clarke, C.M., C.C. Lee & S. McPherson 2006. ''Nepenthes chaniana'' (Nepenthaceae), a new species from north-western Borneo. ''Sabah Parks Journal'' 7: 53–66.

B. H. Danser suggested that ''N. burbidgeae'' is most closely related to ''N. pilosa''. The latter species is poorly known and was for a long time confused with ''N. chaniana''.

The glandular crest of ''N. chaniana'' is very similar to that of ''N. burbidgeae'', particularly in upper pitchers. However, it is difficult to confuse these species as the pitchers are otherwise markedly different in structure; the upper pitchers of ''N. burbidgeae'' are short and funnel-shaped, whereas those of ''N. chaniana'' are elongated and have a dense indumentum of white hair.

Natural hybrids

Natural hybrids involving ''N. burbidgeae'' appear to be relatively rare and only four have been recorded to date. Three of these (crosses with '' N. edwardsiana'', '' N. fusca'', and '' N. tentaculata'') have received little attention in the scientific literature, but ''N. burbidgeae'' × '' N. rajah'' has been described as '' N. Ã— alisaputrana'' and is famous for producing huge pitchers rivalling those of ''N. rajah'' in size.

''N. burbidgeae'' × ''N. rajah''

''Nepenthes × alisaputrana'' was described in 1992 by J. H. Adam and C. C. Wilcock and is named in honour of Datuk Lamri Ali, Director of Sabah Parks.Adam, J.H. & C.C. Wilcock 1992. A new natural hybrid of ''Nepenthes'' from Mt. Kinabalu (Sabah). ''Reinwardtia'' 11: 35–40. It is only known from a few remote localities within Kinabalu National Park, where it grows in stunted, open vegetation over serpentine soils at around 2000 m above sea level, often amongst populations of ''N. burbidgeae''.

This plant is notable for combining the best characters of both parent species, not least the size of its pitchers, which rival those of ''N. rajah'' in volume (≤35 cm high, ≤20 cm wide). The other hybrids involving ''N. rajah'' do not exhibit such impressive proportions. The pitchers of ''N. Ã— alisaputrana'' can be distinguished from those of ''N. burbidgeae'' by a broader peristome, larger lid and simply by their sheer size. The hybrid differs from its other parent, ''N. rajah'', by its lid structure, indumentum of short, brown hairs, narrower and more cylindrical peristome, and pitcher colour, which is usually yellow-green with red or brown flecking. For this reason, Anthea Phillipps and Anthony Lamb gave it the common name "Leopard Pitcher-Plant". The peristome is green to dark red and striped with purple bands. Leaves are often slightly peltate. The hybrid is a strong climber and frequently produces upper pitchers.

''Nepenthes × alisaputrana'' more closely resembles ''N. rajah'' than ''N. burbidgeae'', but it is difficult to confuse this plant with either. However, this mistake has previously been made on at least one occasion; a pitcher illustrated in Adrian Slack's ''Insect-Eating Plants and How to Grow Them'' as being ''N. rajah''Slack, A. 1986. ''Insect-Eating Plants and How to Grow Them''. Alphabooks, Dorset, UK. is in fact ''N. burbidgeae × N. rajah''.

In 2002, phytochemical screening and analytical chromatography were used to study the presence of phenolic compounds and leucoanthocyanins in ''N. Ã— alisaputrana'' and its putative parent species. The research was based on leaf material from nine dry herbarium specimens. Eight spots containing phenolic acids, flavonols, flavones, leucoanthocyanins and 'unknown flavonoid' 1 and 3 were identified from chromatographic profiles. The distributions of these in the hybrid ''N. Ã— alisaputrana'' and its putative parental species ''N. burbidgeae'' and ''N. rajah'' are shown in the adjacent table. A specimen of ''N. Ã— alisaputrana'' grown from tissue culture (''in vitro'') was also tested.

Luteolin, cyanidin and 'Unknown Flavonoid 3' were undetected in ''N. burbidgeae'', while concentrations of 'Unknown Flavonoid 1' were found to be weak. Chromatographic patterns of the ''N. Ã— alisaputrana'' samples studied showed complementation of its putative parental species.

Myricetin was found to be absent from all studied taxa. This agrees with the findings of previous authors and suggests that the absence of a widely distributed compound like myricetin among the ''Nepenthes'' examined might provide additional diagnostic information for these taxa.

Cultivation

Little information has been published on the growing requirements of ''N. burbidgeae''. In ''Insect-Eating Plants and How to Grow Them'', Adrian Slack wrote that cuttings of ''N. burbidgeae'' were more difficult to root than those of other ''Nepenthes'' species.Marthaler, O. 1996. An addition to Adrian Slack's comments on ''Nepenthes burbidgeae'' cuttings. '' Bulletin of the Australian Carnivorous Plant Society, Inc.'' 15(1): 8–9.

In 2004, professional horticulturist Robert Sacilotto published a summary of measured tolerances of highland ''Nepenthes'' species, based on experiments conducted between 1996 and 2001.Sacilotto, R. 2004
Experiments with highland ''Nepenthes'' seedlings: A Summary of Measured Tolerances
'' Carnivorous Plant Newsletter'' 33(1): 26–31. ''Nepenthes burbidgeae'' was found to be tolerant of a fairly wide range of conditions, particularly in terms of temperature and soil composition; it grew in every substrate used in the experiment. However, plants showed stunted growth when grown in a mixture consisting of 50% silica gel, 20% '' Sphagnum'' moss, 20% fir bark, and 10% peat moss chunks. The highest growth rates were exhibited by specimens in 50% leached perlite, 30% long fiber ''Sphagnum'' moss, 10% peat moss chunks, and 10% fir bark, as well as media without fir bark and with a higher percentage of ''Sphagnum''. ''Nepenthes burbidgeae'' was found to tolerate temperatures in the range of 9 to 41 Â°C (48° to 105 Â°F). A nighttime drop in temperature below 18 Â°C (65 Â°F) was necessary for good growth; plants that were not exposed to such a drop grew around 50% slower and produced fewer pitchers. Optimal growth rates were observed with daytime temperatures of 20 to 29 Â°C (68° to 85 Â°F) and nighttime temperatures of 12 to 16 Â°C (54° to 60 Â°F). Soil with a pH of 4.8 to 5.5 produced the best results; values below 3.5 corresponded with slower growth. Optimal soil conductivity was between 10 and 24 microsiemens, and prolonged exposure of one week or more to levels of more than 60 microsiemens resulted in foliar burn. The experiments suggested that ''N. burbidgeae'' grows best when relative humidity is in the range of 68 to 95%. However, constant exposure to high humidity in excess of 90% resulted in disease outbreaks and increased plant death rates. Seedlings of less than one year proved to be particularly vulnerable to this. Optimal light levels varied depending on the light source used: 8100–11000 lx (750–1000 fc) in sunlight, 7000–9700 lx (650–900 fc) under high pressure sodium lamps, 6500–9100 lx (600–850 fc) under metal halide lamps, and 5400–7300 lx (500–680 fc) under fluorescent lamps. ''Nepenthes burbidgeae'' could be grown in lower light conditions, but such plants exhibited etiolated growth and reduced colouration. The species was found to respond well to a fertilizer that was applied to the pitchers on a monthly basis, but a foliar feed using the same solution produced no visible change in growth rate.

Notes

:a.The Latin description of ''N. burbidgeae'' from Danser's monograph reads:

''Folia mediocria'' petiolata, lamina elliptica, nervis longitudinalibus utrinque 3-4, vagina in alas 2 decurrente: ''ascidia rosularum'' et ''inferiora'' ignota ; ''ascidia superiora'' infundibuliformia, parte inferiore costis 2 prominentibus, os versus alis 2 fimbriatis ; peristomio operculum versus in collum ; 1-2 cm altum elevato, cylindrico, crebre costato, operculo late cordato, facie inferiore prope basin carina valida ; ''inflorescentia'' ignota ; ''indumentum'' in omnibus partibus iuventute pubescens, statu adulto parcum v. deciduum, in margine foliorum persistens.

References

Further reading

* Beaman, J.H. & C. Anderson 2004. ''The Plants of Mount Kinabalu: 5. Dicotyledon Families Magnoliaceae to Winteraceae''. Natural History Publications (Borneo), Kota Kinabalu.
* Beck, G. 1895. Die Gattung ''Nepenthes''. ''Wiener Illustrirte Garten-Zeitung'' 20(3–6): 96–107, 141–150, 182–192, 217–229.
* Bonhomme, V., H. Pelloux-Prayer, E. Jousselin, Y. Forterre, J.-J. Labat & L. Gaume 2011. Slippery or sticky? Functional diversity in the trapping strategy of ''Nepenthes'' carnivorous plants. ''New Phytologist'' 191(2): 545–554.
* Corner, E.J.H. 1996. Pitcher-plants (''Nepenthes''). In: K.M. Wong & A. Phillipps (eds.
''Kinabalu: Summit of Borneo. A Revised and Expanded Edition.''
The Sabah Society, Kota Kinabalu. pp. 115–121. .
* Dixon, W.E. 1889
''Nepenthes''.
''The Gardeners' Chronicle'', series 3, 6(144): 354.
* Fretwell, S. 2013. Back in Borneo for giant ''Nepenthes''. Part 1: Mesilau Nature Reserve, Ranau. ''Victorian Carnivorous Plant Society Inc.'' 107: 6–13.
* Fretwell, S. 2013. Back in Borneo to see giant ''Nepenthes''. Part 2: Mt Tambuyukon and Poring. ''Victorian Carnivorous Plant Society Inc.'' 108: 6–15.
* Harms, H. 1936. Nepenthaceae. In: A. Engler & K. Prantl. ''Die natürlichen Pflanzenfamilien'', 2 aufl. band 17b.
* Mansur, M. 2001
Koleksi ''Nepenthes'' di Herbarium Bogoriense: prospeknya sebagai tanaman hias.
In: ''Prosiding Seminar Hari Cinta Puspa dan Satwa Nasional''. Lembaga Ilmu Pengetahuan Indonesia, Bogor. pp. 244–253.
* McPherson, S.R. & A. Robinson 2012. ''Field Guide to the Pitcher Plants of Borneo''. Redfern Natural History Productions, Poole.
* Meimberg, H., A. Wistuba, P. Dittrich & G. Heubl 2001. Molecular phylogeny of Nepenthaceae based on cladistic analysis of plastid trnK intron sequence data. ''Plant Biology'' 3(2): 164–175.
* Meimberg, H. 2002.â
Molekular-systematische Untersuchungen an den Familien Nepenthaceae und Ancistrocladaceae sowie verwandter Taxa aus der Unterklasse Caryophyllidae s. l..
Ph.D. thesis, Ludwig Maximilian University of Munich, Munich.
* Meimberg, H. & G. Heubl 2006. Introduction of a nuclear marker for phylogenetic analysis of Nepenthaceae. ''Plant Biology'' 8(6): 831–840.
* Meimberg, H., S. Thalhammer, A. Brachmann & G. Heubl 2006. Comparative analysis of a translocated copy of the ''trnK'' intron in carnivorous family Nepenthaceae. ''Molecular Phylogenetics and Evolution'' 39(2): 478–490.
* Moran, J.A., C. Clarke, M. Greenwood & L. Chin 2012. Tuning of color contrast signals to visual sensitivity maxima of tree shrews by three Bornean highland ''Nepenthes'' species. ''Plant Signaling & Behavior'' 7(10): 1267–1270.
* Thorogood, C. 2010. '' The Malaysian Nepenthes: Evolutionary and Taxonomic Perspectives''. Nova Science Publishers, New York.
* Yeo, J. 1996. A trip to Kinabalu Park. '' Bulletin of the Australian Carnivorous Plant Society, Inc.'' 15(4): 4–5.

External links

Photographs of ''N. burbidgeae''
at the Carnivorous Plant Photofinder

{{Taxonbar, from=Q4912319

Carnivorous plants of Asia
burbidgeae
Endemic flora of Borneo
Flora of Sabah
Endangered plants
Plants described in 1882