諾斯豬籠草
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諾斯豬籠草![]() | |
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產自馬來西亞砂拉越州石隆門縣的諾斯豬籠草的下位籠 | |
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: | 諾斯豬籠草 N. northiana
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二名法 | |
Nepenthes northiana | |
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異名 | |
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諾斯豬籠草(學名:Nepenthes northiana)是馬來西亞特有的熱帶食蟲植物[3],分布於海拔0至500米的地區。諾斯豬籠草得名於它的發現者——瑪麗安娜·諾斯。諾斯豬籠草是豬籠草屬最為著名的物種之一,在19世紀後半葉它被當作砂拉越特有植物的一個代表。[4]
植物學史
諾斯豬籠草最先由瑪麗安娜·諾斯帶入科學界,她也將馬來西亞砂拉越州石隆門縣的植株引入種植。哈里·維奇認定這是一個未描述的物種,並派查爾斯·柯蒂斯採集標本,並將諾斯豬籠草的種子寄回英國。1881年,約瑟夫·道爾頓·胡克用瑪麗安娜·諾斯的名字命名了諾斯豬籠草。[1]1876年,「M.North s.n.」號標本,即諾斯豬籠草的模式標本被採集於縣內的
瑪麗安娜·諾斯的自傳《快樂一生的回憶》(Recollections of a Happy Life)第一版的封面即是一幅鎏金的諾斯豬籠草作品。她對發現諾斯豬籠草這樣寫道:[注 1][6]
![](http://images.weserv.nl/?url=//upload.wikimedia.org/wikipedia/commons/thumb/a/a3/MN561_A_New_Pitcher_Plant_from_the_Limestone_Mountains_of_Sarawak%2C_Borneo.jpg/220px-MN561_A_New_Pitcher_Plant_from_the_Limestone_Mountains_of_Sarawak%2C_Borneo.jpg)
“ | 埃弗里特先生爬上附近的一座山,給我帶下來了一些具巨大捕蟲籠的標本,其為所以豬籠草中最大的。我給最大的一個捕蟲籠畫了一幅畫,這幅圖後來引起了維奇先生的好奇,使他派了一位訪者來尋找它的種子,並種植出了植株,之後約瑟夫·道爾頓·胡克爵士將其命名為諾斯豬籠草。這種豬籠草的捕蟲籠通常有一英尺長,上面覆蓋着深紅色的斑點。 | ” |
諾斯豬籠草的模式描述發表於《園丁紀事》中,並進一步闡述了諾斯豬籠草的發現:[注 2][7][8]
“ | 諾斯女士所繪的標本由婆羅洲公司的赫伯特·埃弗里特先生採集,在他『穿過沒路的森林時,儘管會遇見蛇和螞蟥,但他仍將其帶給了諾斯女士』。『只有這樣』,諾斯女士寫道,『只有身歷其境才能了解這是多麼艱難。其存在於沙撈越石灰岩山脈中海拔1000英尺以上的地區,附生於樹枝上。當我收到它們後,我將它們全部綁在陽台周圍,遺憾的是我僅將它們畫在了一張小小的紙上』。 | ” |
瑪麗安娜·諾斯所畫的諾斯豬籠草作品現陳列於英國國家植物的瑪麗安娜·諾斯畫廊(Marianne North Gallery)中。[9]
在發現諾斯豬籠草後的數十年間,諾斯豬籠草出現在各大植物學出版物上。弗雷德里克·威廉·伯比奇在1882年的一期《園丁紀事》中堅信諾斯豬籠草是血紅豬籠草(N. sanguinea)與維奇豬籠草(N. veitchii)的自然雜交種:[注 3][10]
“ | 你的諾斯豬籠草插圖非常棒。諾斯女士的圖畫,如果我沒記錯的話,其為底色為亮紅色-緋紅色,並具深色的斑點。這是個好東西,但我堅信其為血紅豬籠草與維奇豬籠草的自然雜交種。傾斜的籠口會讓人聯想到馬來王豬籠草,但其最高海拔僅為庭院,而就我所知,在京那峇魯山以北250英里,其海拔高度從未低於4500英尺,最高約至10000英尺。
諾斯豬籠草也許在早些時期就生長在草原上——無論如何也不會有人能將諾斯豬籠草、維奇豬籠草、血紅豬籠草給一一培植且不被它們捕蟲籠的相似度給混淆到。另外,乍一看諾斯豬籠草畫作會讓人聯想到馬來王豬籠草長甕外狀變型那傾斜、邊緣波浪狀的籠口,不過馬來王豬籠草的莖生捕蟲籠從未呈這樣。當瑪麗安娜·諾斯向我與威廉·伯比奇先生展示她的作品那一刻時;上述的推論立刻在我與伯比奇先生的腦海裡成形。當我從切爾西前往婆羅洲時,已得到了原生地的經緯度,但不幸的是我沒機會去往砂拉越;僅前往到了西北海岸,並找到了非常親善的海盜首領。 |
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當發現婆羅洲並不存在血紅豬籠草後,學者們就意識到弗雷德里克·威廉·伯比奇的雜種假說是完全錯誤的。[11]1884年,愛德華·奧古斯特·馮里格爾在《Gartenflora》中發表了一篇關於諾斯豬籠草的短文。[12]1895年,君特·貝克·馮·曼那哥塔-勒馳奈在其1895年的專著《豬籠草屬》中描述了將一種豬籠草命名為偽豬籠草(N. spuria)。[2]該名稱為不合法名,現在被認為是諾斯豬籠草的同物異名。[5]雅各布·海斯伯特·博爾拉吉(Jacob Gijsbert Boerlage)在其1900年的著作《荷屬東印度群島植物指南》(Handleiding tot de kennis der flora van Nederlandsch Indië)中將諾斯豬籠草的學名「N. northiana」錯印為「N. nordtiana」。 [13]其為印刷錯誤。[5]
1908年,約翰·繆爾黑德·麥克法蘭在其專著《豬籠草科》中對豬籠草屬進行修訂時,也對諾斯豬籠草進行了分類處理,並提供了諾斯豬籠草的修訂描述。[9]
![](http://images.weserv.nl/?url=//upload.wikimedia.org/wikipedia/commons/thumb/c/cc/Nepenthes_northiana_pulchra.jpg/220px-Nepenthes_northiana_pulchra.jpg)
1909年,R·傑瑞-迪斯洛吉斯(R. Jarry-Desloges)描述了諾斯豬籠草的一個變種——美麗諾斯豬籠草(Nepenthes northiana var. pulchra)。[5][15]其顏色鮮艷,捕蟲籠為紫紅色,唇有更醒目的紅黃條紋。相比之下,原變種的捕蟲籠多為黃色,並帶有棕色或紅色的斑點。[14]但美麗諾斯豬籠草現已不被認為是有效的分類。[16]
下延豬籠草
1925年,約翰·繆爾黑德·麥克法蘭描述了下延豬籠草(N. decurrens)}}。[17]該描述基於由約翰·休伊特於1907年9月採集於美里峇南河的「Hewitt 100」號標本。[18]其與諾斯豬籠草的模式標本一樣,都存放於英國皇家植物園。[5][17]
B·H·丹瑟在其1928年的開創性著作《荷屬東印度群島的豬籠草科植物》中認為這兩個類群是不同的物種,但仍不確定。[注 4][注 5]他對其分類處理這樣寫道:[注 6][11]
“ | 我已在砂拉越博物館植物標本館中看到了該物種(下延豬籠草)的標本:兩片帶捕蟲籠的葉片,其由莖部扯下,連接方式不再可見。
其捕蟲籠與1881年2月《園丁紀事》第724頁至725頁的諾斯豬籠草繪畫作品很相似。該圖所示,其籠蓋具兩根籠蓋骨,整個捕蟲籠具籠翼,彎曲部亦如此,但這些差異並不顯著。根據其描述,諾斯豬籠草的莖較下延豬籠草細,葉片無柄,但這並不是重要的區別。其最重要的區別在花序。諾斯豬籠草的花序為鬆散的三角形總狀花序,花梗長2至3毫米,而下延豬籠草的花序更長,且花梗也更長(對這兩個物種花序的描述都不全面)。因此我無法對埃弗里特於1892年採集且存放於砂拉越植物標本館中的上述3個下位籠進行判斷。埃弗里特先生的這份標本可能與瑪麗安娜·諾斯採集到的諾斯豬籠草是相同的。因此上述的3個捕蟲籠也許就來源於某些諾斯豬籠草植株。其為卵形-橢圓形,高分別為23厘米、24厘米和26厘米,寬分別為10厘米、11厘米和10厘米,最寬處位於或略低於中部。唇與下延豬籠草很相似,寬分別為3厘米、4厘米和2.5厘米。籠口非常傾斜,約占捕蟲籠高的一半。籠蓋下表面中線具一根籠蓋骨,但和下延豬籠草一樣為褶皺狀,不顯著。 |
” |
之後的學者已不認為這些區別足以將其作為一個物種,所以下延豬籠草被視為是諾斯豬籠草的一個同物異名。[19][20][21][22]
形態特徵
![](http://images.weserv.nl/?url=//upload.wikimedia.org/wikipedia/commons/thumb/1/12/Nepenthes_northiana.jpg/280px-Nepenthes_northiana.jpg)
諾斯豬籠草為藤本植物。莖可長達10米,直徑寬至15毫米,呈圓柱形至三稜柱形[23]。節間距可長達25厘米。[21]
諾斯豬籠草的葉片紙質,薄[3],無柄至具小葉柄。葉片呈長圓形至倒卵形,可長達40厘米,寬至10厘米。葉尖急尖,漸尖至葉基。葉基半抱莖,下延成一對翼。中脈兩側各有4條縱脈。羽狀脈不明顯。葉片下表面約有20至60個蜜腺。[11]籠蔓可長達100厘米。[21]
諾斯豬籠草的下位籠通常為卵形,偶爾上部為圓柱形。[21]諾斯豬籠草是豬籠草屬中捕蟲籠較大的一種,其高達40厘米[7],寬至15厘米[21]。個別特別大的下位籠的容積可超過1品脫(946毫升)。[24]腹面的籠翼不寬於15毫米。[21]內表面蠟質區稀少。[25]籠口為卵形,傾斜。唇兩側擴展,可寬至25毫米,邊緣常為波浪狀。內緣具短小但明顯的唇齒。籠蓋為卵形至長圓形,無附屬物,具急尖。籠蓋基部的後側有一根不分叉籠蔓尾,可長達20毫米。上位籠的形態與下位籠類似,但呈漏斗狀。籠翼常保留,但也可能縮小為一對隆起。[21]
諾斯豬籠草的花序為總狀花序。總花梗可長達60厘米,花序軸可長達40厘米。通常雄性的花序較短。每根花梗帶兩朵花,可長達50毫米。
諾斯豬籠草的種子與其他豬籠草一樣為梭狀,兩端有兩個木質的附屬物,中部為巨型的胚。這樣的結構使得種子可以隨風播撒但又不至於飛得太遠而飛到不適宜生長的地方。[21]一項對來自同一副標本(J.H.Adam 2378,採集於海拔30米處)花粉大小的研究顯示,其120粒花粉的平均直徑為29.8微米(標準誤=0.4,變異係數=6.0%)。[26]
諾斯豬籠草無明顯的毛被,整個植株幾乎無毛。[21]
諾斯豬籠草的莖和葉片為淺綠色,捕蟲籠為綠白色,並具大量紅色斑點。唇為白色至紅色,且具深色條紋。[21]
生態關係
![](http://images.weserv.nl/?url=//upload.wikimedia.org/wikipedia/commons/thumb/d/d7/Bau_N._northiana_habitat_2.jpg/280px-Bau_N._northiana_habitat_2.jpg)
諾斯豬籠草是砂拉越古晉省[4],特別是石隆門附近的山上特有的熱帶食蟲植物。其分布於海拔0至500米的石灰岩基質地區。[21]
諾斯豬籠草一般生長於近乎垂直的石灰岩壁上,並有水浸潤。[27]少見於一些小山丘的次級植被中。[21]其與其他石灰岩基質植物同域分布,如勞氏箭葉海芋(Alocasia longiloba var. lowii)。[28]
根據2000年的評估,諾斯豬籠草已被列入《2006年世界自然保護聯盟瀕危物種紅色名錄》中,保護狀況為易危。[27]這與查爾斯·克拉克在1997年的非正式評估一致。[21]但世界保護監測中心則將其列為瀕危。[29]
原生地的採石活動已破壞了幾個存在諾斯豬籠草的山丘。[21]另外,諾斯豬籠草的自然種群還遭受着過度採集的威脅。[24]諾斯豬籠草具體極高的商業價值,使其倍受植物收藏家青睞。[30]安西婭·飛利浦和安東尼·蘭姆在其1996年出版的專著《婆羅洲的豬籠草》中這樣寫道:「過度的採集使得諾斯豬籠草已經到了滅絕的邊緣。」[7]儘管如此,因採集者無法觸及大部分餘下的植株,所以諾斯豬籠草在短時間內似乎是安全的。[21]
相關物種
諾斯豬籠草與產於婆羅洲東加里曼丹石灰岩山上的馬普魯山豬籠草(N. mapuluensis)十分相似。[31]雖然它們之間的差別不大,但仍有一些穩定的差異可將其區別開來。與諾斯豬籠草相比,馬普魯岩山豬籠草攀援莖上的葉片更接近於線形,捕蟲籠的顏色更深,且上位籠更窄。[21][28]值得注意的是諾斯豬籠草只分布於砂拉越的石隆門縣,與馬普魯岩山豬籠草的分布地相隔數百公里。
諾斯豬籠草與大型平庸豬籠草(N. macrovulgaris)相比[23][32],它們具有形狀相同的葉片和葉柄,不同之處在於諾斯豬籠草的攀援莖為三稜柱形,而大型平庸豬籠草為圓柱形。此外,後者葉柄的基部不下延。[21][23]
胡瑞爾豬籠草(N. hurrelliana)與維奇豬籠草(N. veitchii)表面上也與諾斯豬籠草有些類似,但它們更小型,顏色也沒有這麼鮮艷。[4]
自然雜交種
白環豬籠草與諾斯豬籠草的自然雜交種
雪線豬籠草由於諾斯豬籠草分布局限,僅生長在砂拉越石隆門的少數地區,所以雪線豬籠草更為罕見。白環豬籠草(N. albomarginata)在雜交中很強勢,使得與諾斯豬籠草雜交的後代完全失去了寬大的唇。其捕蟲籠為窄漏斗形,籠身為奶油色至暗紫色,並具紅色或黑色斑點。[4][33]
小豬籠草與諾斯豬籠草的自然雜交種
石隆門豬籠草的形態表現為其親本的中間型。它繼承了小豬籠草(N. gracilis)喜攀爬的特徵,但其葉片和莖又相當的大型。此外,它還繼承了諾斯豬籠草捕蟲籠的形態特徵。特別是唇比小豬籠草要寬,並具有紅色的條紋。其捕蟲籠可高達15厘米,為暗綠色至紫紅色。[34]
石隆門豬籠草與親本一樣,是一種低地豬籠草,存在於海拔約100米的地區。它常陸生於特有諾斯豬籠草的石灰岩基質山丘附近的沼澤地區。
石隆門豬籠草十分罕見,僅發現過少量植株,都來源於同一原生地。
奇異豬籠草與諾斯豬籠草的自然雜交種
奇異豬籠草與諾斯豬籠草的自然雜交種(N. mirabilis × N. northiana)是一種較為罕見的自然雜交種,僅發現於2007年。[35][36]
種植方法
想要種好諾斯豬籠草相當困難,現在的諾斯豬籠草雖然並不是特別昂貴,卻在種植者間它以難種出名。[37][38]因此,其在17世紀引種後不久就消失於歐洲園藝界。[3]有一段時間內,人們猜測諾斯豬籠草必須紮根於石灰岩基質中才能健康生長,但事實並非如此。[38]諾斯豬籠草需要一個少光照的環境,陽光直射會使得葉片上出現褐色的壞死斑點,並使捕蟲籠生長減緩或停止生長。諾斯豬籠草對空氣相對濕度很敏感,需要種植於空氣濕度較高的環境中,在天氣較為溫暖的條件下,諾斯豬籠草擁有着相對較快的生長速度。[3][38]
彼得·達馬托在專著《薩維奇花園》一書中寫道,泥炭和泥炭蘚會阻礙諾斯豬籠草的生長。他認為使用兩份粗蛭石、一份珍珠岩、一份浮石和一份河沙組成的混合基質更有利於諾斯豬籠草的生長。[39]其他種植者也報道了一些明顯無必要的基質配方。[38]
注釋
- ^ 瑪麗安娜·諾斯在其自傳中描述發現諾斯豬籠草的原文:[6]
“ Mr E. [Everett] went up a mountain near and brought me down some grand trailing specimens of the largest of all pitcher-plants, which I festooned round the balcony by its yards of trailing stems. I painted a portrait of the largest, and my picture afterwards induced Mr Veitch to send a traveller to seek the seeds, from which he raised plants and Sir Joseph Hooker named the species Nepenthes northiana. These pitchers are often over a foot long, and richly covered with crimson blotches. ” - ^ 《園丁紀事》中對諾斯豬籠草進一步闡述的原文:[1][7]
“ "The specimen from which Miss North's drawing was made was procured by Mr. Herbert Everett of the Borneo Company who "traversed pathless forests amid snakes and leeches to find and bring it down to the artist." "Only those", writes Miss North, "who have been in such places can understand the difficulties of progress there. The specimens grew on the branches of a tree about 1000 feet above the sea on the limestone mountains of Sarawak. When I received them I tied them in festoons all round the verandah and grumbled at having only one small half-sheet of paper left to paint them on."" ” - ^ 弗雷德里克·威廉·伯比奇在1882年的一期《園丁紀事》中論述的原文:[10]
“ "Your figure of Nepenthes Northiana was very good. Miss North's drawing, however, has, if I recollect right, a ground-tint of bright reddish-crimson on which darker blotches are laid. It is a fine thing, and, as I firmly believe, a natural hybrid between N. sanguinea × N. Veitchii. The oblique mouth of the urns would suggest N. Rajah as one of the parents, but then his highness only holds court, so far as we know at present, on Kina Balu, 250 miles further north, and never at a less altitude than 4500 feet, rising to near 10,000 feet. In earlier times he may have been an inhabitant of the plains—at any rate no one can place the pitchers of N. Northiana, N. Veitchii, and N. sanguinea side by side without being struck by their affinity. Again, a glance at your engraving of N. Northiana reminds one of a long-urned form of N. Rajah in obliquity of mouth and its wavy-margined frill. The cauline pitchers of N. Rajah have never yet been figuted. I was with Mr. Harry Veitch when Miss North first showed him the picture of N. Northiana, and it was a revelation to us both. I had the latitude and longitude of its habitat in my portfolio when I left Chelsea for Borneo, but unfortunately never had the chance of seeing Sarawak ; my lot was the wild north-west coast, among the pirate chiefs, and very good genial fellows I found them !"
” - ^ B·H·丹瑟專著中對諾斯豬籠草的原拉丁文描述:[11]
“ Folia mediocria sessilia, lamina elliptica v. obovata, nervis longitudinalibus utrinque 4, basi lata semiamplexicauli in alas 2 decurrente ; ascidia rosularum ignota ; ascidia inferiora subovata, alis 2 fimbriatis ; peristomio applanato v. expanso, 10-50 mm lato, costis crebris, dentibus brevibus ; operculo ovato-oblongo, facie inferiore non appendiculato ; ascidia superiora infundibuliformia, alis 2 angustis fimbriatis, peristomio angustiore quam in ascidiis inferioribus, costis crebris, operculo angustiore quam in ascidiis inferioribus, facie inferiore inappendiculata ; inflorescentia racemus longus pedicellis 2-4 mm longis 2- v. 1-floris ; indumentum parcum (v. 0 ?). ” - ^ B·H·丹瑟專著中對下延豬籠草的原拉丁文描述:[11]
“ Folia mediocria petiolata, lanceolata, nervis longitudinalibus utrinque 5-6, vagina in alas 2 basi peltatas decurrente ; ascidia rosularum et inferiora ignota ; ascidia superiora magna, tubulosa v. infundibuliformia ; parte inferiore costis 2 prominentibus, os versus alis 2 fimbriatis ; peristomio expanso, 25-60 mm lato, costis c. 1 mm distantibus, dentibus vix longioribus quam latis ; operculo ovato, facie inferiore plana v. prope basin obtuse carinata ; inflorescentia racemus longus pedicellis longis fere omnibus 2-floris ; indumentum in caulibus foliisque fere 0, in ascidiis adpressum parcum in inflorescentiis tenue densum ferrugineum. ” - ^ B·H·丹瑟解釋其分類處理的原文:[11]
“ "I have seen type material of this species [N. decurrens] in the Herbarium of the Sarawak Museum: 2 pitcher-bearing leaves, torn from the stem in such a way, that the manner, in which they are inserted on it, is no longer visible. The pitchers show a great resemblance with those of the drawing of N. Northiana in The Gardeners' Chronicle, 1881, 2, between p. 724 and 725. This drawing shows 2 keels on the lid and wings over the whole pitchers, even over the curved part, but these are insignificant differences. According to the descriptions, the stems of N. Northiana are less thick than those of N. decurrens, and the leaves are sessile, but this too is not so important a difference as it seems. The most important difference is in the inflorescences. N. Northiana has a loose-flowered triangular raceme, with 2 to 3 mm long pedicels, N. decurrens has a long and coarse raceme, with long pedicels (the description of both inflorescences is very imperfect). Therefore it is impossible for me to determine the 3 above mentioned inferior pitchers, I found in the Sarawak Herbarium, and collected by Everett in 1892. This Mr. Everett may be the same which collected N. Northiana for Marianne North and therefore it seems possible that the 3 pitchers mentioned are the basal ones of the latter species. They are ovate-ellipsoidal, resp. 23, 24 and 26 cm high, 10, 11 and 10 cm wide, widest about, or somewhat below the middle ; the peristomes are as in N. decurrens, resp. 3, 4 and 2 1/2 cm broad, the mouth is very oblique, occupying about half the height of the pitcher, the lid has one median keel, but is crumpled, and the form, though not well visible, seems to be that of N. decurrens."
”
參考文獻
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