紙莎草

紙草（Cyperus papyrus，papyrus），又稱紙草、埃及莎草、埃及紙草、蒲草（bulrush）、埃及蒲草、印度叢草（Indian matting plant）等。屬於莎草科莎草屬。是北非洲原產的一種多年生草本植物。

生長型態

紙莎草是一種高而堅韌的挺水植物，約可長到 4 至 5 公尺高。它綠色、長稈狀的莖橫切面為三角形，從粗厚的地下莖中長出。每一條莖的頂端都會長出一簇濃密的、放射狀（像煙火一般）的葉。葉子細長、通常呈亮綠色，長度約 10 至 30 公分。當植物還年幼的時候，這簇葉子的生長型態就像雞毛撢一樣。花是綠褐色，生長在葉簇的末端。果實呈褐色。地下莖在剛長出時會被一層紅褐色的紙質層覆蓋，這層三角形的鱗狀層也覆蓋了莖的基部。

紙莎草生長在熱帶至亞熱帶的環境中，不論是潮濕的森林或乾燥的沙漠，只要全年均溫在 20 到 30℃且土壤 pH 值在 6.0-8.5 就可以生長。紙莎草的花在暮夏盛開，並且傾向於在全日照到半陰涼的環境開花。像許多典型的熱帶植物一樣，紙莎草對霜很敏感。

紙莎草的分佈遍及在沼澤、淺水湖和溪畔等非洲的潮濕部分，但在尼羅河三角洲已逐漸變得稀少。在較深的水中，紙莎草是構成植被的主要植物（被稱為 sudd）。現今它的野生分佈存在於剛果、馬達加斯加島、烏幹達、衣索匹亞和西西里島。

紙莎草羽毛狀的花是許多社會性鳥類築巢的地點。像許多莎草科植物一樣，它是風媒花，且果實由流水帶走。

今天，紙莎草被培育、改良為裝飾性的植物。其中 C. nanus（矮紙莎草）、C. profiler 兩個品種只會長到約 1 公尺高。

用途

古埃及人使用紙莎草的用途有很多。其中，最有名的應該是莎草紙的製作。在希臘文和英文中，「莎草紙」這個字的語源被許多人認為是來自於古埃及語，但這是謬傳。紙莎草現在被廣泛的運用於製作裝飾品，例如莎草紙畫。在古代，紙莎草在尼羅河三角洲有著相當廣泛的分佈，然而現在它在尼羅河三角洲的原生種群卻瀕臨絕滅。在泰奧弗拉斯托斯的《植物史》（Historia Plantarum，卷十）一書中寫到紙莎草在敘利亞也有分佈。而根據老普林尼的《博物志》一書，紙莎草是尼羅河及幼發拉底河流域的原生植物。

此外，古埃及人利用多種莎草科的植物的花來製作敬神的花圈。而嫩枝則被當作食物。它多纖維的地下莖則用來製作碗等多種器具，有時也當作燃料。而莖則用來編製葦舟（在埃及第四王朝的浮雕中，描繪了壯年男子採集莎草製作葦舟的情形），類似的船在蘇丹也能發現。

延伸閲讀

Boar, R. R., D. M. Harper and C. S. Adams. 1999. Biomass Allocation in Cyperus papyrus in a Tropical Wetland, Lake Naivasha, Kenya. 1999. Biotropica 3: 411.

Chapman, L.J., C.A. Chapman, R. Ogutu-Ohwayo, M. Chandler, L. Kaufman and A.E. Keiter. 1996. Refugia for endangered fishes from an introduced predator in Lake Nabugabo, Uganda. Conservation Biology 10: 554-561.

Chapman, L.J., C.A. Chapman, P.J. Schofield, J.P. Olowo, L. Kaufman, O. Seehausen and R. Ogutu-Ohwayo. 2003. Fish faunal resurgence in Lake Nabugabo, East Africa. Conservation Biology 17: 500-511.

Gaudet, John. 1975. Mineral concentrations in papyrus in various African swamps. Journal of Ecology 63: 483-491.

Gaudet, John. 1976. Nutrient relationships in the detritus of a tropical swamp.Archiv für Hydrobiologie 78: 213-239.

Gaudet, John. 1977. Natural drawdown on Lake Naivasha, Kenya and the formation of papyrus swamps. Aquatic Botany 3: 1-47.

Gaudet, John. 1977. Uptake and loss of mineral nutrients by papyrus in tropical swamps. Ecology 58: 415-422.

Gaudet, John. 1978. Effect of a tropical swamp on water quality. Verh. Internat. Ver. Limnol. 20: 2202-2206.

Gaudet, John. 1978. Seasonal changes in nutrients in a tropical swamp. Journal of Ecology 67: 953-981.

Gaudet, John. 1980. Papyrus and the ecology of Lake Naivasha. National Geographic Society Research Reports. 12: 267-272.

Gaudet, J. and J. Melack. 1981. Major ion chemistry in a tropical African lake basin. Freshwater Biology 11: 309-333.

Gaudet, J. and C. Howard-Williams. 1985. “The structure and functioning of African swamps.” In (ed. Denny) The Ecology and Management of African Wetland Vegetation. Dr.w.Junk, Pub., Dordrecht (pp.154–175).

Gaudet, John. 1991. Structure and function of African floodplains. Journal of the East African Natural Historical Society. 82 (199): 1-32.

Harper, D.M., K.M. Mavuti and S. M. Muchiri. 1990: Ecology and management of Lake Naivasha, Kenya, in relation to climatic change, alien species introductions and agricultural development. Environmental Conservation 17: 328–336.

Harper, D. 1992. The ecological relationships of aquatic plants at Lake Naivasha, Kenya. Hydrobiologia. 232: 65-71.

Howard-Williams, C. and K. Thompson. 1985. The conservation and management of African wetlands. In (ed. Denny) The Ecology and Management of African Wetland Vegetation. Dr.w.Junk, Pub., Dordrecht (pp.203–230).

Jones, M.B. and T. R. Milburn. 1978. Photosynthesis in Papyrus (Cyperus papyrus L.), Photosynthetica. 12: 197 - 199.

Jones, M. B. and F. M. Muthuri. 1997. Standing biomass and carbon distribution in a papyrus (Cyperus Papyrus L) swamp on Lake Naivasha, Kenya. Journal of Tropical Ecology. 13: 347–356.

Jones M.B. and S. W. Humphries. 2002. Impacts of the C4 sedge Cyperus papyrus L. on carbon and water fluxes in an African wetland. Hydrobiologia, Volume 488, pp.107–113.

Maclean, I.M.D. 2004. An ecological and socio-economic analysis of biodiversity conservation in East African wetlands. Unpublished PhD thesis, University of East Anglia, Norwich.

Maclean, I.M.D., M. Hassall, M. R. Boar and I. Lake. 2006. Effects of disturbance and habitat loss on papyrus-dwelling passerines. Biological Conservation., 131: 349-358.

Maclean, I.M.D., M. Hassall, R. Boar, R. and O. Nasirwa. 2003a. Effects of habitat degradation on avian guilds in East African papyrus Cyperus papyrus L. swamps. Bird Conservation International, 13: 283-297.

Maclean, I.M.D., R. Tinch, M. Hassall and R.R. Boar, R.R. 2003b. Social and economic use of wetland resources: a case study from Lake Bunyonyi, Uganda. Environmental Change and Management Working Paper No. 2003-09, Centre for Social and Economic Research into the Global Environment, University of East Anglia, Norwich.

Maclean, I.M.D., R. Tinch, M. Hassall and R.R. Boar. 2003c. Towards optimal use of tropical wetlands: an economic evaluation of goods derived from papyrus swamps in southwest Uganda. Environmental Change and Management Working Paper No. 2003-10, Centre for Social and Economic Research into the Global Environment, University of East Anglia, Norwich.

Messenger Dally. 1908 How papyrus defeated South Sydney and assisted in making Eastern Suburbs great
Muthuri, F. M., M. B. Jones, and S.K. Imbamba. 1989. Primary productivity of papyrus (Cyperus papyrus) in a tropical swamp - Lake Naivasha, Kenya, Biomass, 18: 1 - 14.

Muthuri, F. M. and M. B. Jones. 1997. Nutrient distribution in a papyrus swamp: Lake Naivasha, Kenya. Aquatic Botany, 56: 35–50.

Owino, A. O. and P. G. Ryan. 2006. Habitat associations of papyrus specialist birds at three papyrus swamps in western Kenya. African Journal of Ecology 44: 438-443.

Thompson, K. 1976. Swamp development in the head waters of the White Nile. In (ed.J. Rzoska) ‘‘The Nile. Biology of an Ancient River.’’Monographiae Biologicae, 29. Dr.W. Junk b.v., The Hague.

Thompson, K., P.R. Shewry & H.W. Woolhouse. 1979. Papyrus swamp development in the Upemba Basin, Zaire: Studies of population structure in Cyperus papyrus stands. Botanical Journal of the Linn. Soc. 78: 299-316.