圓盾豬籠草

圓盾豬籠草
	圓盾豬籠草的捕蟲籠，人工栽培植株
保護狀況
	; 極危（IUCN 3.1）
科學分類
界：	植物界 Plantae
演化支：	維管束植物 Tracheophyta
演化支：	被子植物 Angiosperms
演化支：	真雙子葉植物 Eudicots
目：	石竹目 Caryophyllales
科：	豬籠草科 Nepenthaceae
屬：	豬籠草屬 Nepenthes
種：	圓盾豬籠草 N. clipeata
二名法
	Nepenthes clipeata; Danser (1928)
	圓盾豬籠草的分佈地

圓盾豬籠草（學名：Nepenthes clipeata），又稱盾葉毛豬籠草^[3]，是婆羅洲西加里曼丹黑山特有的熱帶食蟲植物。其生長於近乎垂直的花崗岩壁上^[3]，海拔約600米至800米。其種加詞「clipeata」來源於拉丁文「clipeus」，意為「圓盾形」，是指其葉片的形狀。

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圓盾豬籠草是豬籠草屬中極其瀕危的物種之一。1995年發現僅存在15棵野生植株。^[4]

植物學史

1894年1月30日至2月13日，約翰內斯·戈特弗里德·哈利爾先後5次登頂黑山，並首次採集到了圓盾豬籠草。^[5]他將該發現發表於B·H·丹瑟1928年的專著《荷屬東印度群島的豬籠草科植物》中：^{[注 1]}^[6]

“

在再次爬過一個長滿芒箕的陡坡之後，一個陡峭的岩壁出現在眼前。光滑的水磨石完全看不出任何結構上的變化，反覆整座山是由一塊巨大的岩石構成的。岩壁的底部架設了約45米高的陡峭藤梯；但其只位於岩壁的底部，而中部和頂部都是赤裸的岩石……藤梯的中部具一小塊腐殖土，僅能供一個人站立及休息片刻。也就是在這，藤梯的頂部生長着一個具有非常巨大捕蟲籠的豬籠草。捕蟲籠的基部膨大呈壺狀。因此，其一方面可以儲存大量的水，另一方面相對窄小的籠肩又可以防止落入籠內的昆蟲逃出。（翻譯自荷蘭語^[7]和德語^[2]^[8]，見於《婆羅洲的豬籠草》）

”

形態特徵

圓盾豬籠草最大的特點是其圓盾形的葉片，即其籠蔓是從距葉片末端一定距離的葉片背面穿出，而非從圓盾形葉尖伸出。其捕蟲籠體型較大，可高達30厘米。捕蟲籠基部為球形，上部為漏斗形。圓盾豬籠草的捕蟲籠只有一種形態，且莖沒有攀爬的能力，最多僅可長達2米。其花序較小，長度很少超過25厘米。植株的所有部分都披被着棕色的長毛。^[9]

B·H·丹瑟在他的專著中這樣寫道：^{[注 2]}^[2]

“

圓盾豬籠草是豬籠草屬中最特別且最引人注目的物種之一。特別是它接近圓形的厚實葉片，從距葉片末端一段距離伸出的不捲曲的粗短籠蔓，及無籠翼的捕蟲籠和拱形的籠蓋，都非常的特殊。類似的特殊葉片形態，僅出現於菲律賓特有的寶特豬籠草身上。目前還不知道圓盾豬籠草是以什麼方式生長的。以下是我覺得有可能的情況。圓盾豬籠草不會攀爬。短而健壯的莖、葉柄和籠蔓證明，該物種存在於開闊的地區。但我無法想像當葉片水平展開時，捕蟲籠是以什麼方式出現的。因此，我認為這種植物生長於黑山垂直的岩壁，且其葉片垂直，捕蟲籠位於葉片的後方。但是，圓盾豬籠草應該不可能只長在岩壁上，但目前尚未清楚，或許它還生長於其他地方。

”

尚未有關於圓盾豬籠草的變型或變種的描述。^[9]

保護狀況

20世紀80年代起，植物採集者開始頻繁造訪黑山，對圓盾豬籠草野外種群造成了巨大壓力。當地嚮導也開始採集生長在他們村莊附近的圓盾豬籠草，尤其是那些生長在山腳下的。但大部分的植株都無法存活，從而導致需要採集更多植株來替代它們。此外，1997年至1998年因厄爾尼諾現象造成的乾旱和森林大火，又給圓盾豬籠草的原生地造成了更大的破壞。這些因素導致圓盾豬籠草在黑山的數量大幅度減少。^[10]1995年，圓盾豬籠草被認為在野外僅存在約15株^[4]，而到2001年，查爾斯·克拉克給出了更低的估計，約為2至6株。^[11]不過，2010年的一項研究顯示，在黑山海拔749至874米處共發現了260株圓盾豬籠草，其分散於黑山45個地點。^[12]該研究的作者指出，幾乎所有觀察到的植株都生長於人跡罕至的崖壁上。所觀察到的開花植株中，大部分都為雄性。^[12]

圓盾豬籠草已正式列入《瀕危野生動植物種國際貿易公約》附錄Ⅱ中，自然保護聯盟將其保護狀況從數據缺乏更改為極危。圓盾豬籠草被認為在野外長期生存的希望渺茫。國際食蟲植物協會已經設立了圓盾豬籠草倖存計劃（The Nepenthes clipeata Survival Project），以對其進行遷地保護。^[13]^[14]^[15]目前只有三到四個售賣圓盾豬籠草的商家具有合法種源。^[10]

2010年，珍稀豬籠草採集（Rare Nepenthes Collection）項目成立，以保護4種極其瀕危的豬籠草屬物種：馬兜鈴豬籠草（N. aristolochioides）、圓盾豬籠草、印度豬籠草（N. khasiana）和硬葉豬籠草（N. rigidifolia）。^[16]^[17]^[18]

伊爾萬·洛瓦迪（Irwan Lovadi）2010年2月倡議，拉福德小額基金（The Rufford Small Grants Foundation）出資。^[12]2011年10月，在伊爾萬·洛瓦迪和拉福德基金的支持下開始了對圓盾豬籠草的就地保護項目。^[19]

自然雜交種

已發現了以下關於圓盾豬籠草的自然雜交種：

N. albomarginata × N. clipeata^[9]
N. clipeata × N. rafflesiana^[6]
N. clipeata × N. reinwardtiana^[9]

註釋

[注 1]

《荷屬東印度群島的豬籠草科植物》中約翰內斯·戈特弗里德·哈利爾對其發現的敘述原文：^[6]

“

After once again climbing a steep slope with Gleichenia thickets, one stands suddenly beneath the high enclosing rock wall of the mountain ring. The smooth water-washed stone seamed with water channels shows no variation in structure, and it appears almost as if the whole mountain was composed of a single monstrous block of rock. On this wall has been erected the steep 45 metre high rattan ladder; it is secured only at the bottom, in the middle and in the solid earth at the top, the rest lying free against the stone... Just above the middle of the ladder a small thin patch of humus is found, just sufficient to allow one to stand and rest for a moment. Both here, and at the top of the ladder a Nepenthes plant with unusually large pitchers has established itself. In the basal part, the pitchers are expanded into a jug shape. They are thus able, on the one hand, to take up a large quantity of water, and on the other, to hinder the escape of insects which have fallen inside, by means of the relatively narrow neck.

”

[注 2]

B·H·丹瑟在其專著中論述圓盾豬籠草的原文：^[6]

“

N. clipeata is one of the most aberrant and striking species of its genus. Especially the almost orbicular leaves, the thick, short, never curved tendrils, which are inserted far from the apex, the peculiar-shaped pitcher without wings and the strongly vaulted lid are very remarkable. A leaf form as aberrant as this, only occurs in the Philippine species N. truncata. It is not known, in what manner N. clipeata grows. The following seems probable to me. The plant does not climb. The short and robust stems, petioles and tendrils prove, that the mentioned specimens are found in an open place. I can not imagine, in what manner the pitchers have been placed when the leaves were spread horizontally. Therefore I suggest, that the plant has grown against the perpendicular wall of the G. Kelam, and that the leaves stood vertically, the pitchers behind it. It is, however, improbable, that N. clipeata can grow only against perpendicular walls, but it is not clear, what may be the manner of growing in other habitats.

”

參考文獻

[1]
Clarke, C., Cantley, R., Nerz, J., Rischer, H. & Witsuba, A. Nepenthes clipeata. The IUCN Red List of Threatened Species 2000. [18-01-2013].
[2]
Danser, B.H. 1928. 43. 9. Nepenthes clipeata DANS., spec. nova. （頁面存檔備份，存於互聯網檔案館）. In: The Nepenthaceae of the Netherlands Indies. Bulletin du Jardin Botanique de Buitenzorg, Série III, 9(3–4): 249–438.
[3]
夏洛特. 食蟲植物觀賞&栽培圖鑑. 台北: 商周出版. 2007: 54–55. ISBN 978-986-124-850-9.
[4]
Simpson, R.B. 1995. Nepenthes and conservatioN. Curtis's Botanical Magazine 12: 111-118.
[5]
van Steenis-Kruseman, M.J., et al. 2006. Cyclopaedia of Malesian Collectors: Johann Gottfried ('Hans') Hallier （頁面存檔備份，存於互聯網檔案館）. Nationaal Herbarium Nederland.
[6]
Phillipps, A., A. Lamb & C.C. Lee 2008. Pitcher Plants of Borneo. Second Edition. Natural History Publications (Borneo), Kota Kinabalu.
[7]
（荷蘭文） Hallier, H. 1895. Rapport over de botanische tochten in Borneo's Westerafdeeling gedurende de Borneo-expeditie 1893–1894. Natuurkundig Tijdschrift voor Nederlandsch Indië 9(3): 406–449.
[8]
（德文） Hallier, H. 1896. Die botanische Erforschung Mittelborneos. Naturwissenschaftliche Wochenschrift 11(9): 75–79, 85–89, 97–101, 109–114.
[9]
Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
[10]
Cantley, R., C.M. Clarke, J. Cokendolpher, B. Rice & A. Wistuba 2004. Nepenthes clipeata Survival Project （頁面存檔備份，存於互聯網檔案館）. International Carnivorous Plant Society.
[11]
Cheek, M.R. & M.H.P. Jebb 2001. Nepenthaceae. Flora Malesiana 15: 1–157.
[12]
Conservation of critically endangered Nepenthes clipeata on Mount Kelam （頁面存檔備份，存於互聯網檔案館）. The Rufford Small Grants Foundation.
[13]
Cantley, R., C.M. Clarke, J. Cokendolpher, B. Rice & A. Wistuba 2005. Nepenthes clipeata Survival Project. Carnivorous Plant Newsletter 34(4): 116–120.
[14]
Rice, B. 2005. Nepenthes clipeata conservatioN. Carnivorous Plant Newsletter 34(4): 115.
[15]
Rice, B. 2007. What is the Nepenthes clipeata Survival Program? （頁面存檔備份，存於互聯網檔案館） The Carnivorous Plant FAQ.
[16]
Ziemer, B. 2010. Exciting conservation news: the Rare Nepenthes Collection project! Carnivorous Plant Newsletter 39(3): 67.
[17]
Nepenthes clipeata （頁面存檔備份，存於互聯網檔案館）. Ark of Life.
[18]
Bourke, G. 2010. A new conservation initiative: the Rare Nepenthes Collection project.PDF Captive Exotics Newsletter 1(2): 5–6.
[19]
Promoting the conservation of critically endangered Nepenthes clipeata in West Kalimantan, Indonesia （頁面存檔備份，存於互聯網檔案館）. The Rufford Small Grants Foundation.

Adam, J.H., C.C. Wilcock & M.D. Swaine 1992. The ecology and distribution of Bornean Nepenthes.PDF Journal of Tropical Forest Science 5(1): 13–25.
Adam, J.H. & C.C. Wilcock 1999. Palynological study of Bornean Nepenthes (Nepenthaceae).PDF Pertanika Journal of Tropical Agricultural Science 22(1): 1–7.
Bauer, U., C.J. Clemente, T. Renner & W. Federle 2012. Form follows function: morphological diversification and alternative trapping strategies in carnivorous Nepenthes pitcher plants. Journal of Evolutionary Biology 25(1): 90–102. doi:10.1111/j.1420-9101.2011.02406.x
Clarke, C.M., R. Cantley, J. Nerz, H. Rischer & A. Witsuba. Nepenthes clipeata. The IUCN Red List of Threatened Species 2000. [6 May 2006]. Listed as Critically Endangered (CR A1a, B1+2e, D v2.3).
Damayanti, F., M. Mansur & I. Roostika 2011. Diversity of Nepenthes spp. in West Kalimantan. （頁面存檔備份，存於互聯網檔案館） International Journal of Biodiversity and Conservation 3(13): 705–708.
Lee, C.C. 2000. Recent Nepenthes Discoveries（頁面存檔備份，存於互聯網檔案館）. [video] The 3rd Conference of the International Carnivorous Plant Society, San Francisco, USA.
Lee, C.C. 2011. A preliminary conservation assessment of Nepenthes clipeata (Nepenthaceae).PDF Ark of Life.
McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Borneo. Redfern Natural History Productions, Poole.
Meimberg, H., A. Wistuba, P. Dittrich & G. Heubl 2001. Molecular phylogeny of Nepenthaceae based on cladistic analysis of plastid trnK intron sequence data. Plant Biology 3(2): 164–175. doi:10.1055/s-2001-12897
（德文） Meimberg, H. 2002. Molekular-systematische Untersuchungen an den Familien Nepenthaceae und Ancistrocladaceae sowie verwandter Taxa aus der Unterklasse Caryophyllidae s. l..PDF Ph.D. thesis, Ludwig Maximilian University of Munich, Munich.
Meimberg, H. & G. Heubl 2006. Introduction of a nuclear marker for phylogenetic analysis of Nepenthaceae. Plant Biology 8(6): 831–840. doi:10.1055/s-2006-924676
Meimberg, H., S. Thalhammer, A. Brachmann & G. Heubl 2006. Comparative analysis of a translocated copy of the trnK intron in carnivorous family Nepenthaceae. Molecular Phylogenetics and Evolution 39(2): 478–490. doi:10.1016/j.ympev.2005.11.023

擴展閱讀

圓盾豬籠草倖存計劃
食蟲植物照片搜尋引擎中圓盾豬籠草的照片（頁面存檔備份，存於互聯網檔案館）

維基共享資源上的相關多媒體資源：圓盾豬籠草維基物種上的相關資訊：圓盾豬籠草

[P&L2008-7] [注 1]
《荷屬東印度群島的豬籠草科植物》中約翰內斯·戈特弗里德·哈利爾對其發現的敘述原文：^[6]

“ After once again climbing a steep slope with Gleichenia thickets, one stands suddenly beneath the high enclosing rock wall of the mountain ring. The smooth water-washed stone seamed with water channels shows no variation in structure, and it appears almost as if the whole mountain was composed of a single monstrous block of rock. On this wall has been erected the steep 45 metre high rattan ladder; it is secured only at the bottom, in the middle and in the solid earth at the top, the rest lying free against the stone... Just above the middle of the ladder a small thin patch of humus is found, just sufficient to allow one to stand and rest for a moment. Both here, and at the top of the ladder a Nepenthes plant with unusually large pitchers has established itself. In the basal part, the pitchers are expanded into a jug shape. They are thus able, on the one hand, to take up a large quantity of water, and on the other, to hinder the escape of insects which have fallen inside, by means of the relatively narrow neck. ”

[Danser-11] [注 2]
B·H·丹瑟在其專著中論述圓盾豬籠草的原文：^[6]

“ N. clipeata is one of the most aberrant and striking species of its genus. Especially the almost orbicular leaves, the thick, short, never curved tendrils, which are inserted far from the apex, the peculiar-shaped pitcher without wings and the strongly vaulted lid are very remarkable. A leaf form as aberrant as this, only occurs in the Philippine species N. truncata. It is not known, in what manner N. clipeata grows. The following seems probable to me. The plant does not climb. The short and robust stems, petioles and tendrils prove, that the mentioned specimens are found in an open place. I can not imagine, in what manner the pitchers have been placed when the leaves were spread horizontally. Therefore I suggest, that the plant has grown against the perpendicular wall of the G. Kelam, and that the leaves stood vertically, the pitchers behind it. It is, however, improbable, that N. clipeata can grow only against perpendicular walls, but it is not clear, what may be the manner of growing in other habitats. ”

[1] [1]
Clarke, C., Cantley, R., Nerz, J., Rischer, H. & Witsuba, A. Nepenthes clipeata. The IUCN Red List of Threatened Species 2000. [18-01-2013].

[Danser-2] [2]
Danser, B.H. 1928. 43. 9. Nepenthes clipeata DANS., spec. nova. （頁面存檔備份，存於互聯網檔案館）. In: The Nepenthaceae of the Netherlands Indies. Bulletin du Jardin Botanique de Buitenzorg, Série III, 9(3–4): 249–438.

[夏洛特2007-3] [3]
夏洛特. 食蟲植物觀賞&栽培圖鑑. 台北: 商周出版. 2007: 54–55. ISBN 978-986-124-850-9.

[Simpson-4] [4]
Simpson, R.B. 1995. Nepenthes and conservatioN. Curtis's Botanical Magazine 12: 111-118.

[5] [5]
van Steenis-Kruseman, M.J., et al. 2006. Cyclopaedia of Malesian Collectors: Johann Gottfried ('Hans') Hallier （頁面存檔備份，存於互聯網檔案館）. Nationaal Herbarium Nederland.

[P&L2008-6] [6]
Phillipps, A., A. Lamb & C.C. Lee 2008. Pitcher Plants of Borneo. Second Edition. Natural History Publications (Borneo), Kota Kinabalu.

[8] [7]
（荷蘭文） Hallier, H. 1895. Rapport over de botanische tochten in Borneo's Westerafdeeling gedurende de Borneo-expeditie 1893–1894. Natuurkundig Tijdschrift voor Nederlandsch Indië 9(3): 406–449.

[Hallier1896-9] [8]
（德文） Hallier, H. 1896. Die botanische Erforschung Mittelborneos. Naturwissenschaftliche Wochenschrift 11(9): 75–79, 85–89, 97–101, 109–114.

[Clarke-10] [9]
Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.

[NcSP-12] [10]
Cantley, R., C.M. Clarke, J. Cokendolpher, B. Rice & A. Wistuba 2004. Nepenthes clipeata Survival Project （頁面存檔備份，存於互聯網檔案館）. International Carnivorous Plant Society.

[C&J-13] [11]
Cheek, M.R. & M.H.P. Jebb 2001. Nepenthaceae. Flora Malesiana 15: 1–157.

[Rufford-14] [12]
Conservation of critically endangered Nepenthes clipeata on Mount Kelam （頁面存檔備份，存於互聯網檔案館）. The Rufford Small Grants Foundation.

[15] [13]
Cantley, R., C.M. Clarke, J. Cokendolpher, B. Rice & A. Wistuba 2005. Nepenthes clipeata Survival Project. Carnivorous Plant Newsletter 34(4): 116–120.

[16] [14]
Rice, B. 2005. Nepenthes clipeata conservatioN. Carnivorous Plant Newsletter 34(4): 115.

[17] [15]
Rice, B. 2007. What is the Nepenthes clipeata Survival Program? （頁面存檔備份，存於互聯網檔案館） The Carnivorous Plant FAQ.

[18] [16]
Ziemer, B. 2010. Exciting conservation news: the Rare Nepenthes Collection project! Carnivorous Plant Newsletter 39(3): 67.

[19] [17]
Nepenthes clipeata （頁面存檔備份，存於互聯網檔案館）. Ark of Life.

[20] [18]
Bourke, G. 2010. A new conservation initiative: the Rare Nepenthes Collection project.PDF Captive Exotics Newsletter 1(2): 5–6.

[21] [19]
Promoting the conservation of critically endangered Nepenthes clipeata in West Kalimantan, Indonesia （頁面存檔備份，存於互聯網檔案館）. The Rufford Small Grants Foundation.

[3]

[1]

[2]

[4]

[5]

[注 1]

[6]

[7]

[8]

[9]

[注 2]

[10]

[11]

[12]

[13]

[14]

[15]

[16]

[17]

[18]

[19]