圆盾猪笼草

圆盾猪笼草
	圆盾猪笼草的捕虫笼，人工栽培植株
保护状况
	; 極危（IUCN 3.1）[1]
科学分类
界：	植物界 Plantae
演化支：	Tracheophyta
演化支：	被子植物 Angiosperms
演化支：	真双子叶植物 Eudicots
目：	石竹目 Caryophyllales
科：	猪笼草科 Nepenthaceae
属：	猪笼草属 Nepenthes
种：	圆盾猪笼草 N. clipeata
二名法
	Nepenthes clipeata; Danser (1928)[2]
	圆盾猪笼草的分布地

圆盾猪笼草（学名：），又称盾叶猪笼草[3]，是婆罗洲西加里曼丹黑山特有的热带食虫植物。其生长于近乎垂直的花岗岩壁上[3]，海拔约600米至800米。其种加词“clipeata”来源于拉丁文“clipeus”，意为“圆盾形”，是指其叶片的形状。

圆盾猪笼草是猪笼草属中极其濒危的物种之一。1995年发现仅存在15棵野生植株。[4]

植物学史

1894年1月30日至2月13日，约翰内斯·戈特弗里德·哈利尔先后5次登顶黑山，并首次采集到了圆盾猪笼草。[5]他将该发现发表于B·H·丹瑟1928年的专著《荷属东印度群岛的猪笼草科植物》中：[注 1][6]

在再次爬过一个长满芒箕的陡坡之后，一个陡峭的岩壁出现在眼前。光滑的水磨石完全看不出任何结构上的变化，反复整座山是由一块巨大的岩石构成的。岩壁的底部架设了约45米高的陡峭藤梯；但其只位于岩壁的底部，而中部和顶部都是赤裸的岩石……藤梯的中部具一小块腐殖土，仅能供一个人站立及休息片刻。也就是在这，藤梯的顶部生长着一个具有非常巨大捕虫笼的猪笼草。捕虫笼的基部膨大呈壶状。因此，其一方面可以储存大量的水，另一方面相对窄小的笼肩又可以防止落入笼内的昆虫逃出。（翻译自荷兰语[7]和德语[2][8]，见于《婆罗洲的猪笼草》）

形态特征

圆盾猪笼草最大的特点是其圆盾形的叶片，即其笼蔓是从距叶片末端一定距离的叶片背面穿出，而非从圆盾形叶尖伸出。其捕虫笼体型较大，可高达30厘米。捕虫笼基部为球形，上部为漏斗形。圆盾猪笼草的捕虫笼只有一种形态，且茎没有攀爬的能力，最多仅可长达2米。其花序较小，长度很少超过25厘米。植株的所有部分都披被着棕色的长毛。[9]

B·H·丹瑟在他的专著中这样写道：[注 2][2]

圆盾猪笼草是猪笼草属中最特别且最引人注目的物种之一。特别是它接近圆形的厚实叶片，从距叶片末端一段距离伸出的不卷曲的粗短笼蔓，及无笼翼的捕虫笼和拱形的笼盖，都非常的特殊。类似的特殊叶片形态，仅出现于菲律宾特有的宝特猪笼草身上。目前还不知道圆盾猪笼草是以什么方式生长的。以下是我觉得有可能的情况。圆盾猪笼草不会攀爬。短而健壮的茎、叶柄和笼蔓证明，该物种存在于开阔的地区。但我无法想象当叶片水平展开时，捕虫笼是以什么方式出现的。因此，我认为这种植物生长于黑山垂直的岩壁，且其叶片垂直，捕虫笼位于叶片的后方。但是，圆盾猪笼草应该不可能只长在岩壁上，但目前尚未清楚，或许它还生长于其他地方。

尚未有关于圆盾猪笼草的变型或变种的描述。[9]

保护状况

20世纪80年代起，植物采集者开始频繁造访黑山，对圆盾猪笼草野外种群造成了巨大压力。当地向导也开始采集生长在他们村庄附近的圆盾猪笼草，尤其是那些生长在山脚下的。但大部分的植株都无法存活，从而导致需要采集更多植株来替代它们。此外，1997年至1998年因厄尔尼诺现象造成的干旱和森林大火，又给圆盾猪笼草的原生地造成了更大的破坏。这些因素导致圆盾猪笼草在黑山的数量大幅度减少。[10]1995年，圆盾猪笼草被认为在野外仅存在约15株[4]，而到2001年，查尔斯·克拉克给出了更低的估计，约为2至6株。[11]不过，2010年的一项研究显示，在黑山海拔749至874米处共发现了260株圆盾猪笼草，其分散于黑山45个地点。[12]该研究的作者指出，几乎所有观察到的植株都生长于人迹罕至的崖壁上。所观察到的开花植株中，大部分都为雄性。[12]

圆盾猪笼草已正式列入《瀕危野生動植物種國際貿易公約》附录Ⅱ中，自然保护联盟将其保护状况从数据缺乏更改为极危。圆盾猪笼草被认为在野外长期生存的希望渺茫。国际食虫植物协会已经设立了圆盾猪笼草幸存计划（The Nepenthes clipeata Survival Project），以对其进行迁地保护。[13][14][15]目前只有三到四个售卖圆盾猪笼草的商家具有合法种源。[10]

2010年，珍稀猪笼草采集（Rare Nepenthes Collection）项目成立，以保护4种极其濒危的猪笼草属物种：马兜铃猪笼草（N. aristolochioides）、圆盾猪笼草、印度猪笼草（N. khasiana）和硬叶猪笼草（N. rigidifolia）。[16][17][18]

伊尔万·洛瓦迪（Irwan Lovadi）2010年2月倡议，拉福德小额基金（The Rufford Small Grants Foundation）出资。[12]2011年10月，在伊尔万·洛瓦迪和拉福德基金的支持下开始了对圆盾猪笼草的就地保护项目。[19]

自然杂交种

已发现了以下关于圆盾猪笼草的自然杂交种：

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

注释

《荷属东印度群岛的猪笼草科植物》中约翰内斯·戈特弗里德·哈利尔对其发现的叙述原文：[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.

B·H·丹瑟在其专著中论述圆盾猪笼草的原文：[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.

参考文献

Clarke, C., Cantley, R., Nerz, J., Rischer, H. & Witsuba, A. . The IUCN Red List of Threatened Species 2000. [18-01-2013].
Danser, B.H. 1928. 43. 9. Nepenthes clipeata DANS., spec. nova. （页面存档备份，存于）. In: The Nepenthaceae of the Netherlands Indies. Bulletin du Jardin Botanique de Buitenzorg, Série III, 9(3–4): 249–438.
夏洛特. . 台北: 商周出版. 2007: 54–55. ISBN 978-986-124-850-9.
Simpson, R.B. 1995. Nepenthes and conservatioN. Curtis's Botanical Magazine 12: 111-118.
van Steenis-Kruseman, M.J., et al. 2006. Cyclopaedia of Malesian Collectors: Johann Gottfried ('Hans') Hallier （页面存档备份，存于）. Nationaal Herbarium Nederland.
Phillipps, A., A. Lamb & C.C. Lee 2008. Pitcher Plants of Borneo. Second Edition. Natural History Publications (Borneo), Kota Kinabalu.
（荷兰文） 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.
（德文） Hallier, H. 1896. Die botanische Erforschung Mittelborneos. Naturwissenschaftliche Wochenschrift 11(9): 75–79, 85–89, 97–101, 109–114.
Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
Cantley, R., C.M. Clarke, J. Cokendolpher, B. Rice & A. Wistuba 2004. Nepenthes clipeata Survival Project （页面存档备份，存于）. International Carnivorous Plant Society.
Cheek, M.R. & M.H.P. Jebb 2001. Nepenthaceae. Flora Malesiana 15: 1–157.
Conservation of critically endangered Nepenthes clipeata on Mount Kelam （页面存档备份，存于）. The Rufford Small Grants Foundation.
Cantley, R., C.M. Clarke, J. Cokendolpher, B. Rice & A. Wistuba 2005. Nepenthes clipeata Survival Project. Carnivorous Plant Newsletter 34(4): 116–120.
Rice, B. 2005. Nepenthes clipeata conservatioN. Carnivorous Plant Newsletter 34(4): 115.
Rice, B. 2007. What is the Nepenthes clipeata Survival Program? （页面存档备份，存于） The Carnivorous Plant FAQ.
Ziemer, B. 2010. Exciting conservation news: the Rare Nepenthes Collection project! Carnivorous Plant Newsletter 39(3): 67.
Nepenthes clipeata （页面存档备份，存于）. Ark of Life.
Bourke, G. 2010. A new conservation initiative: the Rare Nepenthes Collection project.PDF Captive Exotics Newsletter 1(2): 5–6.
Promoting the conservation of critically endangered Nepenthes clipeata in West Kalimantan, Indonesia （页面存档备份，存于）. 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. . 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. （页面存档备份，存于） International Journal of Biodiversity and Conservation 3(13): 705–708.
Lee, C.C. 2000. Recent Nepenthes Discoveries（页面存档备份，存于）. [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

扩展阅读

圆盾猪笼草幸存计划
食虫植物照片搜寻引擎中圆盾猪笼草的照片（页面存档备份，存于）

维基共享资源上的相關多媒體資源：圆盾猪笼草維基物種上的相關：圆盾猪笼草

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[P&L2008-7] 《荷属东印度群岛的猪笼草科植物》中约翰内斯·戈特弗里德·哈利尔对其发现的叙述原文：[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] B·H·丹瑟在其专著中论述圆盾猪笼草的原文：[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] Clarke, C., Cantley, R., Nerz, J., Rischer, H. & Witsuba, A. . The IUCN Red List of Threatened Species 2000. [18-01-2013].

[Danser-2] Danser, B.H. 1928. 43. 9. Nepenthes clipeata DANS., spec. nova. （页面存档备份，存于）. 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] 夏洛特. . 台北: 商周出版. 2007: 54–55. ISBN 978-986-124-850-9.

[Simpson-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 （页面存档备份，存于）. Nationaal Herbarium Nederland.

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

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

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

[NcSP-12] Cantley, R., C.M. Clarke, J. Cokendolpher, B. Rice & A. Wistuba 2004. Nepenthes clipeata Survival Project （页面存档备份，存于）. International Carnivorous Plant Society.

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

[Rufford-14] Conservation of critically endangered Nepenthes clipeata on Mount Kelam （页面存档备份，存于）. The Rufford Small Grants Foundation.

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

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

[17] Rice, B. 2007. What is the Nepenthes clipeata Survival Program? （页面存档备份，存于） The Carnivorous Plant FAQ.

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

[19] Nepenthes clipeata （页面存档备份，存于）. Ark of Life.

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

[21] Promoting the conservation of critically endangered Nepenthes clipeata in West Kalimantan, Indonesia （页面存档备份，存于）. The Rufford Small Grants Foundation.