Cyperus papyrus
Encyclopedia
Cyperus papyrus is a monocot belonging to the sedge family Cyperaceae
. It is a herbaceous
perennial
native to Africa
, and forms tall stands of reed-like swamp vegetation in shallow water.
Papyrus sedge (and its close relatives) has a very long history of use by humans, notably by the Ancient Egypt
ians—it is the source of papyrus paper, parts of it can be eaten, and the highly buoyant stems can be made into boats. It is now often cultivated as an ornamental plant
.
s. Each stem is topped by a dense cluster of thin, bright green, thread-like stems around 10 to 30 cm (3.9 to 11.8 in) in length, resembling a feather duster when the plant is young. Greenish-brown flower
clusters eventually appear at the ends of the rays, giving way to brown, nut-like fruit
s.
The younger parts of the rhizome are covered by red-brown, papery, triangular scales, which also cover the base of the culms. Botanically these represent reduced leaves
, so strictly it is not quite correct to call this plant fully "leafless".
used the plant for many purposes, most famously for making papyrus
paper. Its name in Greek
and in English
is widely believed to have come from Egyptian. Cyperus papyrus is now used mainly for decoration, as it is nearly extinct
in its native habitat in the Nile Delta, where in ancient times it was widely cultivated. Theophrastus
' "History of Plants" (Book iv. 10) states that it grew in Syria
; and, according to Pliny's
Natural History, it was also a native plant of the Niger River
and the Euphrates
.
Aside from papyrus, several other members of the genus Cyperus
may actually have been involved in the multiple uses Egyptians found for the plant. Its flowering heads were linked to make garlands for the gods in gratitude. The pith of young shoots was eaten both cooked and raw. Its woody root made bowls and other utensils and was burned for fuel. From the stems were made reed boat
s (seen in bas-reliefs of the Fourth Dynasty showing men cutting papyrus to build a boat; similar boats are still made in the southern Sudan
), sails, mats, cloth, cordage, and sandals. Theophrastus states that King Antigonus made the rigging of his fleet of papyrus, an old practice illustrated by the ship's cable, wherewith the doors were fastened when Odysseus slew the suitors in his hall (Odyssey
xxi. 390).
The "rush" or "reed" basket
in which the Biblical figure Moses
was abandoned may have been made from papyrus.
The adventurer Thor Heyerdahl
built two boats from papyrus, Ra and Ra II, in an attempt to demonstrate that ancient African or Mediterranean people could have reached America
. He succeeded in sailing Ra II from Morocco
to Barbados
.
and has escaped from cultivation in Louisiana
, California
and Hawaii
.
Papyrus sedge forms vast stands in swamps, shallow lakes, and along stream banks throughout the wetter parts of Africa, but it has become rare in the Nile Delta
. In deeper waters it is the chief constituent of the floating, tangled masses of vegetation known as sudd. It also occurs in Madagascar
, and some Mediterranean regions such as Sicily
and the Levant
.
The "feather-duster" flowering heads make ideal nesting sites for many social species of bird
s. As in most sedges, pollination
is by wind, not insect
s, and the mature fruits after release are distributed by water.
In recent years papyrus has been the subject of intense ecological studies centered around its prodigious growth rate and ability to recycle nutrients. Much of this research was begun at Makerere University
in Uganda
in the early 1970s in the swamps on the edge of Lake Victoria
and continued in Kenya
(University of Nairobi) on Lake Naivasha
. John Gaudet’s work in Africa, supported by a National Geographic Society
grant, appeared in various scientific journals over the period 1975-1991. In addition, other pioneer researchers of papyrus at Makerere in the 1970s were: Keith Thompson, T. R. Milburn, and Mike Jones. Thompson’s studies of papyrus swamp development throughout Africa (1976–1985) later formed the basis for management and conservation at national levels.
Extensive research on the productive physiology of papyrus were carried out by Jones from the 1980s onward. He started his work in Uganda and later continued his research on Lake Naivasha in Kenya where he was joined by a new generation of African researchers including Frank Muthuri. Jones's latest research (2002) found that papyrus is a C4
sedge which forms highly productive monotypic stands over large areas of wetland in Africa. Jones and others measured eddy covariance from a stand of Cyperus papyrus, which formed a fringing swamp on the north-west shore of Lake Naivasha, Kenya. They determined that fluxes of CO2 and H2O vapor between the papyrus swamp and the atmosphere were large but variable, depending on the hydrology of the wetland system and the condition of the vegetation. These measurements, combined with simulation modeling of annual fluxes of CO2, showed that papyrus swamps have the potential to sequester large amounts of the carbon (1.6 kg C m-2 y-1) when detritus accumulates under water in anaerobic conditions, but they are a net source of carbon release to the atmosphere (1.0 kg C m-2 y-1) when water levels fall to expose detritus and rhizomes to aerobic conditions. Evapotranspiration from papyrus swamps (E) was frequently lower than evaporation from open water surfaces (E o) and plant factors have a strong influence on the flux of water to the atmosphere.
Research on the papyrus swamp habitat has in recent years attracted the attention of many more African biologists, such as A. O. Owino, K. M. Mavuti, S. M. Muchiri and S. Njuguna. Increasingly the value of papyrus to other species is being recognized. Papyrus swamps provide hypoxic and structural refugia for cichlids from the large predatory Nile Perch
and are an important habitat for several endangered bird species (Chapman et al. 1996; 2003; Maclean et al. 2003a; 2006).
The late 1990s also saw the rise in research on the papyrus swamps of Lake Naivasha in Kenya by teams from the English Universities of Leicester and East Anglia, notably led by David Harper. Harper's extensive recent studies on the swamps and lakes have led to a worldwide awareness of the problems facing papyrus swamps in Africa today.
Cyperaceae
Cyperaceae are a family of monocotyledonous graminoid flowering plants known as sedges, which superficially resemble grasses or rushes. The family is large, with some 5,500 species described in about 109 genera. These species are widely distributed, with the centers of diversity for the group...
. It is a herbaceous
Herbaceous
A herbaceous plant is a plant that has leaves and stems that die down at the end of the growing season to the soil level. They have no persistent woody stem above ground...
perennial
Perennial plant
A perennial plant or simply perennial is a plant that lives for more than two years. The term is often used to differentiate a plant from shorter lived annuals and biennials. The term is sometimes misused by commercial gardeners or horticulturalists to describe only herbaceous perennials...
native to Africa
Africa
Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...
, and forms tall stands of reed-like swamp vegetation in shallow water.
Papyrus sedge (and its close relatives) has a very long history of use by humans, notably by the Ancient Egypt
Ancient Egypt
Ancient Egypt was an ancient civilization of Northeastern Africa, concentrated along the lower reaches of the Nile River in what is now the modern country of Egypt. Egyptian civilization coalesced around 3150 BC with the political unification of Upper and Lower Egypt under the first pharaoh...
ians—it is the source of papyrus paper, parts of it can be eaten, and the highly buoyant stems can be made into boats. It is now often cultivated as an ornamental plant
Ornamental plant
Ornamental plants are plants that are grown for decorative purposes in gardens and landscape design projects, as house plants, for cut flowers and specimen display...
.
Description
This tall, robust, leafless aquatic plant can grow 4 to 5 m (13.1 to 16.4 ft) high. It forms a grass-like clump of triangular green stems that rise up from thick, woody rhizomeRhizome
In botany and dendrology, a rhizome is a characteristically horizontal stem of a plant that is usually found underground, often sending out roots and shoots from its nodes...
s. Each stem is topped by a dense cluster of thin, bright green, thread-like stems around 10 to 30 cm (3.9 to 11.8 in) in length, resembling a feather duster when the plant is young. Greenish-brown flower
Flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants . The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs...
clusters eventually appear at the ends of the rays, giving way to brown, nut-like fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...
s.
The younger parts of the rhizome are covered by red-brown, papery, triangular scales, which also cover the base of the culms. Botanically these represent reduced leaves
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....
, so strictly it is not quite correct to call this plant fully "leafless".
Papyrus in history
EgyptiansEgyptians
Egyptians are nation an ethnic group made up of Mediterranean North Africans, the indigenous people of Egypt.Egyptian identity is closely tied to geography. The population of Egypt is concentrated in the lower Nile Valley, the small strip of cultivable land stretching from the First Cataract to...
used the plant for many purposes, most famously for making papyrus
Papyrus
Papyrus is a thick paper-like material produced from the pith of the papyrus plant, Cyperus papyrus, a wetland sedge that was once abundant in the Nile Delta of Egypt....
paper. Its name in Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...
and in English
English language
English is a West Germanic language that arose in the Anglo-Saxon kingdoms of England and spread into what was to become south-east Scotland under the influence of the Anglian medieval kingdom of Northumbria...
is widely believed to have come from Egyptian. Cyperus papyrus is now used mainly for decoration, as it is nearly extinct
Extinction
In biology and ecology, extinction is the end of an organism or of a group of organisms , normally a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and recover may have been lost before this point...
in its native habitat in the Nile Delta, where in ancient times it was widely cultivated. Theophrastus
Theophrastus
Theophrastus , a Greek native of Eresos in Lesbos, was the successor to Aristotle in the Peripatetic school. He came to Athens at a young age, and initially studied in Plato's school. After Plato's death he attached himself to Aristotle. Aristotle bequeathed to Theophrastus his writings, and...
' "History of Plants" (Book iv. 10) states that it grew in Syria
Syria
Syria , officially the Syrian Arab Republic , is a country in Western Asia, bordering Lebanon and the Mediterranean Sea to the West, Turkey to the north, Iraq to the east, Jordan to the south, and Israel to the southwest....
; and, according to Pliny's
Pliny the Elder
Gaius Plinius Secundus , better known as Pliny the Elder, was a Roman author, naturalist, and natural philosopher, as well as naval and army commander of the early Roman Empire, and personal friend of the emperor Vespasian...
Natural History, it was also a native plant of the Niger River
Niger River
The Niger River is the principal river of western Africa, extending about . Its drainage basin is in area. Its source is in the Guinea Highlands in southeastern Guinea...
and the Euphrates
Euphrates
The Euphrates is the longest and one of the most historically important rivers of Western Asia. Together with the Tigris, it is one of the two defining rivers of Mesopotamia...
.
Aside from papyrus, several other members of the genus Cyperus
Cyperus
Cyperus is a large genus of about 600 species of sedges, distributed throughout all continents in both tropical and temperate regions. They are annual or perennial plants, mostly aquatic and growing in still or slow-moving water up to 0.5 m deep. The species vary greatly in size, with small species...
may actually have been involved in the multiple uses Egyptians found for the plant. Its flowering heads were linked to make garlands for the gods in gratitude. The pith of young shoots was eaten both cooked and raw. Its woody root made bowls and other utensils and was burned for fuel. From the stems were made reed boat
Reed boat
Reed boats and rafts, along with dugout canoes and other rafts, are among the oldest known types of boats. Often used as traditional fishing boats, they are still used in a few places around the world, though they have generally been replaced with planked boats. Reed boats can be distinguished from...
s (seen in bas-reliefs of the Fourth Dynasty showing men cutting papyrus to build a boat; similar boats are still made in the southern Sudan
Sudan
Sudan , officially the Republic of the Sudan , is a country in North Africa, sometimes considered part of the Middle East politically. It is bordered by Egypt to the north, the Red Sea to the northeast, Eritrea and Ethiopia to the east, South Sudan to the south, the Central African Republic to the...
), sails, mats, cloth, cordage, and sandals. Theophrastus states that King Antigonus made the rigging of his fleet of papyrus, an old practice illustrated by the ship's cable, wherewith the doors were fastened when Odysseus slew the suitors in his hall (Odyssey
Odyssey
The Odyssey is one of two major ancient Greek epic poems attributed to Homer. It is, in part, a sequel to the Iliad, the other work ascribed to Homer. The poem is fundamental to the modern Western canon, and is the second—the Iliad being the first—extant work of Western literature...
xxi. 390).
The "rush" or "reed" basket
Ark of bulrushes
The ark of bulrushes in which the infant Moses was laid is called in the Hebrew teiva, a word similar to the Egyptian teb, meaning "a chest". It is also the same word used for Noah's Ark....
in which the Biblical figure Moses
Moses
Moses was, according to the Hebrew Bible and Qur'an, a religious leader, lawgiver and prophet, to whom the authorship of the Torah is traditionally attributed...
was abandoned may have been made from papyrus.
The adventurer Thor Heyerdahl
Thor Heyerdahl
Thor Heyerdahl was a Norwegian ethnographer and adventurer with a background in zoology and geography. He became notable for his Kon-Tiki expedition, in which he sailed by raft from South America to the Tuamotu Islands...
built two boats from papyrus, Ra and Ra II, in an attempt to demonstrate that ancient African or Mediterranean people could have reached America
Americas
The Americas, or America , are lands in the Western hemisphere, also known as the New World. In English, the plural form the Americas is often used to refer to the landmasses of North America and South America with their associated islands and regions, while the singular form America is primarily...
. He succeeded in sailing Ra II from Morocco
Morocco
Morocco , officially the Kingdom of Morocco , is a country located in North Africa. It has a population of more than 32 million and an area of 710,850 km², and also primarily administers the disputed region of the Western Sahara...
to Barbados
Barbados
Barbados is an island country in the Lesser Antilles. It is in length and as much as in width, amounting to . It is situated in the western area of the North Atlantic and 100 kilometres east of the Windward Islands and the Caribbean Sea; therein, it is about east of the islands of Saint...
.
Ecology
Papyrus ranges from subtropical to tropical desert to wet forests, tolerating annual temperatures of 20 °C (68 °F) to 30 °C (86 °F) and a pH of 6.0 to 8.5. Papyrus flowers in late summer, and prefers full sun to partly shady conditions. Like most tropical plants, it is sensitive to frost. In the United States it has become invasive in FloridaFlorida
Florida is a state in the southeastern United States, located on the nation's Atlantic and Gulf coasts. It is bordered to the west by the Gulf of Mexico, to the north by Alabama and Georgia and to the east by the Atlantic Ocean. With a population of 18,801,310 as measured by the 2010 census, it...
and has escaped from cultivation in Louisiana
Louisiana
Louisiana is a state located in the southern region of the United States of America. Its capital is Baton Rouge and largest city is New Orleans. Louisiana is the only state in the U.S. with political subdivisions termed parishes, which are local governments equivalent to counties...
, California
California
California is a state located on the West Coast of the United States. It is by far the most populous U.S. state, and the third-largest by land area...
and Hawaii
Hawaii
Hawaii is the newest of the 50 U.S. states , and is the only U.S. state made up entirely of islands. It is the northernmost island group in Polynesia, occupying most of an archipelago in the central Pacific Ocean, southwest of the continental United States, southeast of Japan, and northeast of...
.
Papyrus sedge forms vast stands in swamps, shallow lakes, and along stream banks throughout the wetter parts of Africa, but it has become rare in the Nile Delta
Nile Delta
The Nile Delta is the delta formed in Northern Egypt where the Nile River spreads out and drains into the Mediterranean Sea. It is one of the world's largest river deltas—from Alexandria in the west to Port Said in the east, it covers some 240 km of Mediterranean coastline—and is a rich...
. In deeper waters it is the chief constituent of the floating, tangled masses of vegetation known as sudd. It also occurs in Madagascar
Madagascar
The Republic of Madagascar is an island country located in the Indian Ocean off the southeastern coast of Africa...
, and some Mediterranean regions such as Sicily
Sicily
Sicily is a region of Italy, and is the largest island in the Mediterranean Sea. Along with the surrounding minor islands, it constitutes an autonomous region of Italy, the Regione Autonoma Siciliana Sicily has a rich and unique culture, especially with regard to the arts, music, literature,...
and the Levant
Levant
The Levant or ) is the geographic region and culture zone of the "eastern Mediterranean littoral between Anatolia and Egypt" . The Levant includes most of modern Lebanon, Syria, Jordan, Israel, the Palestinian territories, and sometimes parts of Turkey and Iraq, and corresponds roughly to the...
.
The "feather-duster" flowering heads make ideal nesting sites for many social species of bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...
s. As in most sedges, pollination
Pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...
is by wind, not insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s, and the mature fruits after release are distributed by water.
In recent years papyrus has been the subject of intense ecological studies centered around its prodigious growth rate and ability to recycle nutrients. Much of this research was begun at Makerere University
Makerere University
Makerere University , Uganda's largest and second-oldest higher institution of learning, , was first established as a technical school in 1922. In 1963 it became the University of East Africa, offering courses leading to general degrees from the University of London...
in Uganda
Uganda
Uganda , officially the Republic of Uganda, is a landlocked country in East Africa. Uganda is also known as the "Pearl of Africa". It is bordered on the east by Kenya, on the north by South Sudan, on the west by the Democratic Republic of the Congo, on the southwest by Rwanda, and on the south by...
in the early 1970s in the swamps on the edge of Lake Victoria
Lake Victoria
Lake Victoria is one of the African Great Lakes. The lake was named for Queen Victoria of the United Kingdom, by John Hanning Speke, the first European to discover this lake....
and continued in Kenya
Kenya
Kenya , officially known as the Republic of Kenya, is a country in East Africa that lies on the equator, with the Indian Ocean to its south-east...
(University of Nairobi) on Lake Naivasha
Lake Naivasha
Lake Naivasha is a freshwater lake in Kenya, lying north west of Nairobi, outside the town of Naivasha. It is part of the Great Rift Valley. The name derives from the local Maasai name Nai'posha, meaning "rough water" because of the sudden storms which can arise...
. John Gaudet’s work in Africa, supported by a National Geographic Society
National Geographic Society
The National Geographic Society , headquartered in Washington, D.C. in the United States, is one of the largest non-profit scientific and educational institutions in the world. Its interests include geography, archaeology and natural science, the promotion of environmental and historical...
grant, appeared in various scientific journals over the period 1975-1991. In addition, other pioneer researchers of papyrus at Makerere in the 1970s were: Keith Thompson, T. R. Milburn, and Mike Jones. Thompson’s studies of papyrus swamp development throughout Africa (1976–1985) later formed the basis for management and conservation at national levels.
Extensive research on the productive physiology of papyrus were carried out by Jones from the 1980s onward. He started his work in Uganda and later continued his research on Lake Naivasha in Kenya where he was joined by a new generation of African researchers including Frank Muthuri. Jones's latest research (2002) found that papyrus is a C4
C4 carbon fixation
C4 carbon fixation is one of three biochemical mechanisms, along with and CAM photosynthesis, used in carbon fixation. It is named for the 4-carbon molecule present in the first product of carbon fixation in these plants, in contrast to the 3-carbon molecule products in plants. fixation is an...
sedge which forms highly productive monotypic stands over large areas of wetland in Africa. Jones and others measured eddy covariance from a stand of Cyperus papyrus, which formed a fringing swamp on the north-west shore of Lake Naivasha, Kenya. They determined that fluxes of CO2 and H2O vapor between the papyrus swamp and the atmosphere were large but variable, depending on the hydrology of the wetland system and the condition of the vegetation. These measurements, combined with simulation modeling of annual fluxes of CO2, showed that papyrus swamps have the potential to sequester large amounts of the carbon (1.6 kg C m-2 y-1) when detritus accumulates under water in anaerobic conditions, but they are a net source of carbon release to the atmosphere (1.0 kg C m-2 y-1) when water levels fall to expose detritus and rhizomes to aerobic conditions. Evapotranspiration from papyrus swamps (E) was frequently lower than evaporation from open water surfaces (E o) and plant factors have a strong influence on the flux of water to the atmosphere.
Research on the papyrus swamp habitat has in recent years attracted the attention of many more African biologists, such as A. O. Owino, K. M. Mavuti, S. M. Muchiri and S. Njuguna. Increasingly the value of papyrus to other species is being recognized. Papyrus swamps provide hypoxic and structural refugia for cichlids from the large predatory Nile Perch
Nile perch
The Nile perch is a species of freshwaterfish in family Latidae of order Perciformes. It is widespread throughout muchof the Afrotropic ecozone, being native to the Congo, Nile, Senegal, Niger, and Lake Chad, Volta, Lake Turkana and other river basins. It also occurs in the brackish waters of...
and are an important habitat for several endangered bird species (Chapman et al. 1996; 2003; Maclean et al. 2003a; 2006).
The late 1990s also saw the rise in research on the papyrus swamps of Lake Naivasha in Kenya by teams from the English Universities of Leicester and East Anglia, notably led by David Harper. Harper's extensive recent studies on the swamps and lakes have led to a worldwide awareness of the problems facing papyrus swamps in Africa today.
Further reading
- 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 fur Hydrobiologie 78: 213-239.
- Gaudet, John. 1977. Natural drawdown on Lake Naivasha, Kenya and the formation of papyrus swamps. Aquatic BotanyAquatic BotanyAquatic Botany is a peer-reviewed scientific journal dedicated to research on structure, function, dynamics, and classification of plant-dominated aquatic communities and ecosystems, as well as molecular, biochemical, and...
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. HydrobiologiaHydrobiologiaHydrobiologia: The International Journal of Aquatic Sciences is a scientific journal specialising in hydrobiology, including limnology and oceanography, systematics of aquatic organisms and aquatic ecology.-History:...
. 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. HydrobiologiaHydrobiologiaHydrobiologia: The International Journal of Aquatic Sciences is a scientific journal specialising in hydrobiology, including limnology and oceanography, systematics of aquatic organisms and aquatic ecology.-History:...
, 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.