Carex

Carex
Various species of sedges
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Monocots
Clade:	Commelinids
Order:	Poales
Family:	Cyperaceae
Genus:	Carex L.
Type species
Carex hirta L. [1]
Diversity
2000+ species
Global distribution of Carex (green)
Synonyms[2]
List Agistron Raf. Ammorrhiza Ehrh. Anithista Raf. Archaeocarex Börner Baeochortus Ehrh. Bitteria Börner Blysmocarex N.A.Ivanova Callistachys Heuff. Caricella Ehrh. Caricina St.-Lag. Caricinella St.-Lag. Chionanthula Börner Chordorrhiza Ehrh. Cobresia Pers. Coleachyron J.Gay ex Boiss. Cryptoglochin Heuff. Cymophyllus Mack. ex Britton & A.Br. Cyperoides Ség. Dapedostachys Börner Desmiograstis Börner Deweya Raf. Diemisa Raf. Diplocarex Hayata Dornera Heuff. ex Schur Drymeia Ehrh. Echinochlaenia Börner Edritria Raf. Elyna Schrad. Facolos Raf. Forexeta Raf. Froelichia Wulfen Genersichia Heuff. Heleonastes Ehrh. Hemicarex Benth. Heuffelia Opiz Holmia Börner Homalostachys Boeckeler Itheta Raf. Kobresia Willd. Kobria St.-Lag. Kolerma Raf. Kuekenthalia Börner Lamprochlaenia Börner Leptostachys Ehrh. Leptovignea Börner Leucoglochin Heuff. Limivasculum Börner Limonaetes Ehrh. Loncoperis Raf. Loxanisa Raf. Loxotrema Raf. Manochlaenia Börner Maukschia Heuff. Meltrema Raf. Neilreichia Kotula Neskiza Raf. Olamblis Raf. Olotrema Raf. Onkerma Raf. Osculisa Raf. Phaeolorum Ehrh. Phyllostachys Torr. Physiglochis Neck. Polyglochin Ehrh. Proteocarpus Börner Pseudocarex Miq. Psyllophora Ehrh. Ptacoseia Ehrh. Rhaptocalymma Borrer Rhynchopera Börner Schelhammeria Moench Schoenoxiphium Nees Temnemis Raf. Thysanocarex Börner Trasus Gray Ulva Adans. Uncinia Pers. Vesicarex Steyerm. Vignantha Schur Vignea P.Beauv. ex T.Lestib. Vignidula Börner

Carex is a vast genus of over 2,000 species^[2] of grass-like plants in the family Cyperaceae, commonly known as sedges (or seg, in older books). Other members of the family Cyperaceae are also called sedges, however those of genus Carex may be called true sedges, and it is the most species-rich genus in the family. The study of Carex is known as caricology.

Description

All species of Carex are perennial,^[3] although some species, such as C. bebbii and C. viridula can fruit in their first year of growth, and may not survive longer.^[4] They typically have rhizomes, stolons or short rootstocks, but some species grow in tufts (caespitose). The culm – the flower-bearing stalk – is unbranched and usually erect. It is usually distinctly triangular in section.^[3]

The leaves of Carex comprise a blade, which extends away from the stalk, and a sheath, which encloses part of the stalk. The blade is normally long and flat, but may be folded, inrolled, channelled or absent. The leaves have parallel veins and a distinct midrib. Where the blade meets the culm there is a structure called the ligule.^[3] The colour of foliage may be green, red or brown, and "ranges from fine and hair-like, sometimes with curled tips, to quite broad with a noticeable midrib and sometimes razor sharp edges".^[5]

In this Carex panicea, the upper spike contains male flowers, and the lower spike contains female flowers.

The flowers of Carex are small and are combined into spikes, which are themselves combined into a larger inflorescence. The spike typically contains many flowers, but can hold as few as one in some species. Almost all Carex species are monoecious; each flower is either male (staminate) or female (pistillate).^[3] A few species are dioecious. Sedges exhibit diverse arrangements of male and female flowers. Often, the lower spikes are entirely pistillate and upper spikes staminate, with one or more spikes in between having pistillate flowers near the base and staminate flowers near the tip.^[6] In other species, all spikes are similar. In that case, they may have male flowers above and female flowers below (androgynous) or female flowers above and male flowers below (gynecandrous). In relatively few species, the arrangement of flowers is irregular.^{[citation needed]}

The defining structure of the genus Carex is the bottle-shaped bract surrounding each female flower. This structure is called the perigynium or utricle, a modified prophyll. It is typically extended into a "rostrum" or beak, which is often divided at the tip (bifid) into two teeth.^[6] The shape, venation, and vestiture (hairs) of the perigynium are important structures for distinguishing Carex species.^{[citation needed]}

The fruit of Carex is a dry, one-seeded indehiscent achene or nut^[3] which grows within the perigynium. Perigynium features aid in fruit dispersal.^{[citation needed]}

Ecology and distribution

Carex species are found across most of the world, albeit with few species in tropical lowlands, and relatively few in sub-Saharan Africa.^[4] Most (but not all) sedges are found in wetlands – such as marshes, calcareous fens, bogs and other peatlands, pond and stream banks, riparian zones, and even ditches.^[6] They are one of the dominant plant groups in arctic and alpine tundra, and in wetland habitats with a water depth of up to 50 cm (20 in).^[4]

Taxonomy and cytogenetics

Main article: List of Carex species

The genus Carex was established by Carl Linnaeus in his work Species Plantarum in 1753, and it is one of the largest genera of flowering plants.^[7] Estimates of the number of species vary from about 1100 to almost 2000.^[4] Carex displays the most dynamic chromosome evolution of all flowering plants. Chromosome numbers range from n = 6 to n = 66, and over 100 species are known to show variation in chromosome number within the species, with differences of up to 10 chromosomes between populations.^[8]

The genomes of Carex kokanica, Carex parvula and Carex littledalei have been sequenced.^[9][10]

Carex has been divided into subgenera in a number of ways. The most influential was Georg Kükenthal's classification using four subgenera – Carex, Vignea, Indocarex and Primocarex – based primarily on the arrangement of the male and female flowers.^[4] There has been considerable debate about the status of these four groups, with some species being transferred between groups and some authors, such as Kenneth Kent Mackenzie, eschewing the subgenera altogether and dividing the genus directly into sections.^[4] The genus is now divided into around four subgenera, some of which may not, however, be monophyletic:^[11]

• Carex subg. Carex – 1450 species, distributed globally^[12]
• Carex subg. Psyllophora (Degl.) Peterm. (equivalent to Kükenthal's "Primocarex") – 70 species^[11]
• Carex subg. Vignea (P. Beauv. ex T. Lestib.) Peterm. – 350 species, cosmopolitan^[12]
• Carex subg. Vigneastra (Tuckerman) Kükenthal (equivalent to Kükenthal's "Indocarex"^[11]) – 100 species, tropical and subtropical Asia^[12]

Fossil record

Several fossil fruits of two Carex species have been described from middle Miocene strata of the Fasterholt area near Silkeborg in Central Jutland, Denmark.^[13]

Uses

Ornamental

Carex species and cultivars are popular in horticulture, particularly in shady positions.^[14][15] Native species are used in wildland habitat restoration projects, natural landscaping, and in sustainable landscaping as drought-tolerant grass replacements for lawns and garden meadows.^[16] Some require damp or wet conditions, others are relatively drought-tolerant. Propagation is by seed or division in spring.^[17]

The cultivars Carex elata 'Aurea' (Bowles' golden sedge)^[18] and Carex oshimensis 'Evergold' ^[19] have received the Royal Horticultural Society's Award of Garden Merit.

Other uses

A mix of dried specimens of several species of Carex (including Carex vesicaria) have a history of being used as thermal insulation in footwear (such as nutukas used by Sámi people^[20]). Sennegrass is one of the names for such mixes.^[20] During the first human expedition to the South Pole in 1911, such a mix were used in skaller, when camps had been set (after each stretch of travelling had been completed).^[21] Carsten Borchgrevink of the British Antarctic Expedition 1898-1900 reported "I found the Lapps method of never using socks in their Finn boots answered well. Socks are never used in Finnmarken in winter time, but 'senne grass' which they, of course, had a special method of arranging in the 'komager' (Finn boots) … if you get wet feet while wearing the grass in the 'komager' you will be warmer than ever, as the fresh grass will, by the moisture and the heat of your feet, in a way start to burn or produce its own heat by spontaneous combustion. The great thing seems to be to arrange the grass properly in the boots, and although we all tried to imitate the Finns in their skill at this work, none of us felt as warm on our feet as when they had helped us."^[22]

Species serve as a food source for numerous animals,^[23] and some are used as a livestock hay.^[24][25]

Use by Native Americans

The Blackfoot put carex in moccasins to protect the feet during winter.^[26] The Cherokee use an infusion of the leaf to "check bowels".^[27] The Ohlone use the roots of many species for basketry.^[28] The Goshute use the root as medicine.^[29] The Jemez consider the plant sacred and use it in the kiva.^[30] The Klamath people weave the leaves into mats, use the juice of the pith as a beverage, eat the fresh stems for food and use the tuberous base of the stem for food.^[31] The indigenous people of Mendocino County, California use the rootstocks to make baskets and rope.^[32] The indigenous people of Montana also weave the leaves into mats and use the young stems as food.^[33] The Navajo of Kayenta, Arizona grind the seeds into mush and eat them.^[34] The Oregon Paiute weave it to make spoons.^[35] The Pomo use the roots to make baskets,^[36][37] and use it to tend fishing traps.^[38] They also use it to make torches.^[38] The Coast Salish use the leaves to make baskets and twine.^[39] The Songhees eat the leaves to induce abortions.^[39] The Nlaka'pamux used the leaves as brushes for cleaning things and use the leaves as forage for their livestock.^[40] The Wailaki weave the roots and leaves into baskets and use the leaves to weave mats.^[41] The Yuki people use the large roots to make baskets.^[42]

See also

• List of Carex species

References

1. Ilkka Kukkonen; Heikki Toivonen (1988). "Taxonomy of wetland carices". Aquatic Botany. 30 (1–2): 5–22. Bibcode:1988AqBot..30....5K. doi:10.1016/0304-3770(88)90003-4.
2. "Carex L." Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 11 May 2024.
3. Robert H. Mohlenbrock; Paul Wayne Nelson (1999). "Introduction". Sedges: Carex. Volume 14 of The Illustrated flora of Illinois. Southern Illinois University Press. pp. 3–7. ISBN 978-0-8093-2074-5.
4. Peter W. Ball; A. A. Reznicek (2002). "Carex Linnaeus, Sp. Pl. 2: 972. 1753; Gen. Pl. ed. 5, 420. 1754". Magnoliophyta: Commelinidae (in part): Cyperaceae. Flora of North America North of Mexico. Vol. 23. Oxford University Press. pp. 254–258. ISBN 978-0-19-515207-4.
5. Amjad Almusaed (2010). Biophilic and Bioclimatic Architecture: Analytical Therapy for the Next Generation of Passive Sustainable Architecture. Springer. p. 52. ISBN 978-1-84996-534-7.
6. A. C. Jermy; D. A. Simpson; M. J. Y. Foley; M. S. Porter (2007). "General structure of Cyperaceae". Sedges of the British Isles. BSBI Handbook No. 1 (3rd ed.). Botanical Society of the British Isles. pp. 2–26. ISBN 978-0-901158-35-2.
7. David G. Frodin (2004). "History and concepts of big plant genera". Taxon. 53 (3): 753–776. doi:10.2307/4135449. JSTOR 4135449.
8. Andrew L. Hipp; Paul E. Rothrock; Eric H. Roalson (2009). "The evolution of chromosome arrangements in Carex (Cyperaceae)" (PDF). The Botanical Review. 75 (1): 96–109. Bibcode:2009BotRv..75...96H. doi:10.1007/s12229-008-9022-8. S2CID 4489708. Archived from the original (PDF) on 2015-02-13. Retrieved 2012-10-21.
9. Can, Muyou; Wei, Wei; Zi, Hailing; Bai, Magaweng; Liu, Yunfei; Gao, Dan; Tu, Dengqunpei; Bao, Yuhong; Wang, Li; Chen, Shaofeng; Zhao, Xing; Qu, Guangpeng (2020-06-11). "Genome sequence of Kobresia littledalei, the first chromosome-level genome in the family Cyperaceae". Scientific Data. 7 (1): 175. Bibcode:2020NatSD...7..175C. doi:10.1038/s41597-020-0518-3. ISSN 2052-4463. PMC 7289886. PMID 32528014.
10. Qu, Guangpeng; Bao, Yuhong; Liao, Yangci; Liu, Can; Zi, Hailing; Bai, Magaweng; Liu, Yunfei; Tu, Dengqunpei; Wang, Li; Chen, Shaofeng; Zhou, Gang; Can, Muyou (2022-03-23). "Draft genomes assembly and annotation of Carex parvula and Carex kokanica reveals stress-specific genes". Scientific Reports. 12 (1): 4970. Bibcode:2022NatSR..12.4970Q. doi:10.1038/s41598-022-08783-z. ISSN 2045-2322. PMC 8943043. PMID 35322069.
11. Julian R. Starr; Stephen A. Harris; David A. Simpson (2008). "Phylogeny of the unispicate taxa in Cyperaceae Tribe Cariceae II: the limits of Uncinia". In Robert F. C. Naczi; Bruce A. Ford (eds.). Sedges: Uses, Diversity, and Systematics of the Cyperaceae (PDF). Monographs in Systematic Botany. Vol. 180. Missouri Botanical Garden Press. ISBN 978-1-930723-72-6. Archived from the original (PDF proof) on 2017-11-10. Retrieved 2013-06-06.
12. Dai Lunkai; Liang Songyun; Zhang Shuren; Tang Yancheng; Tetsuo Koyama; Gordon C. Tucker. "33. Carex Linnaeus, Sp. Pl. 2: 972. 1753. 薹草属 tai cao shu". Acoraceae through Cyperaceae (PDF). Flora of China. Vol. 23. Harvard University Press. pp. 285–461.
13. Angiosperm Fruits and Seeds from the Middle Miocene of Jutland (Denmark) by Else Marie Friis, The Royal Danish Academy of Sciences and Letters 24:3, 1985
14. Judy Lowe (2012). "Carex". Tennessee & Kentucky Garden Guide: the Best Plants for a Tennessee or Kentucky Garden (2nd ed.). Cool Springs Press. p. 178. ISBN 978-1-59186-537-7.
15. Frances Tenenbaum, ed. (2003). "Carex". Taylor's Encyclopedia of Garden Plants. Houghton Mifflin Harcourt. pp. 74–75. ISBN 978-0-618-22644-3.
16. "Grasses and grasslike plants". Native Sons. Retrieved May 22, 2010.
17. RHS A-Z encyclopedia of garden plants. United Kingdom: Dorling Kindersley. 2008. p. 1136. ISBN 978-1-4053-3296-5.
18. "Carex elata 'Aurea' (Bowles' golden sedge)". RHS. Retrieved 12 April 2020.
19. "Carex oshimensis 'Evergold'". RHS. Retrieved 12 April 2020.
20. "Bruk av land og vann i Finnmark i historisk perspektiv" [The use of land and water in Finnmark in historical perspective]. Norges Offentlige Utredninger (in Norwegian). 1994 (21). Norwegian Ministry of Justice and Public Security. 1994. Retrieved 29 April 2020.
21. Ole Mathismoen (December 14, 2011). "Blir ikke varm i rått reinskinn". Aftenposten (in Norwegian). p. 17. ... skalder med senegress fra Kautokeino til bruk når de hadde slått leir.
22. Borchgrevink, Carston (1980) [1901]. First on the Antarctic continent. Being an account of the British Antarctic Expedition, 1898 – 1900. London: Hurst and Co.
23. Ronald W. Crites; E. Joe Middlebrooks; Sherwood C. Reed (2005). Natural Wastewater Treatment Systems. CRC Press. p. 263. ISBN 978-1-4200-2644-3.
24. Heinjo Lahring (2003). Water and Wetland Plants of the Prairie Provinces. University of Regina Press. p. 114. ISBN 978-0-88977-162-8.
25. Joel Greenberg (2010). Of Prairie, Woods, and Water: Two Centuries of Chicago Nature Writing. ReadHowYouWant.com. p. 206. ISBN 978-1-4596-0615-9.
26. Johnston, Alex, 1987, Plants and the Blackfoot, Lethbridge, Alberta. Lethbridge Historical Society, page 22
27. Hamel, Paul B. and Mary U. Chiltoskey, 1975, Cherokee Plants and Their Uses -- A 400 Year History, Sylva, N.C. Herald Publishing Co., page 54
28. Bocek, Barbara R., 1984, Ethnobotany of Costanoan Indians, California, Based on Collections by John P. Harrington, Economic Botany 38(2):240-255, page 255
29. Chamberlin, Ralph V., 1911, The Ethno-Botany of the Gosiute Indians of Utah, Memoirs of the American Anthropological Association 2(5):331-405., page 365
30. Cook, Sarah Louise, 1930, The Ethnobotany of Jemez Indians., University of New Mexico, M.A. Thesis, page 21
31. Coville, Frederick V., 1897, Notes On The Plants Used By The Klamath Indians Of Oregon., Contributions from the U.S. National Herbarium 5(2):87-110, page 92
32. Chestnut, V. K., 1902, Plants Used by the Indians of Mendocino County, California, Contributions from the U.S. National Herbarium 7:295-408., page 314
33. Blankinship, J. W., 1905, Native Economic Plants of Montana, Bozeman. Montana Agricultural College Experimental Station, Bulletin 56, page 9
34. Wyman, Leland C. and Stuart K. Harris, 1951, The Ethnobotany of the Kayenta Navaho, Albuquerque. The University of New Mexico Press, page 16
35. Mahar, James Michael., 1953, Ethnobotany of the Oregon Paiutes of the Warm Springs Indian Reservation, Reed College, B.A. Thesis, page 52
36. Merriam, C. Hart, 1966, Ethnographic Notes on California Indian Tribes, University of California Archaeological Research Facility, Berkeley, page 296
37. Gifford, E. W., 1967, Ethnographic Notes on the Southwestern Pomo, Anthropological Records 25:10-15, page 11
38. Gifford, E. W., 1967, Ethnographic Notes on the Southwestern Pomo, Anthropological Records 25:10-15, page 12
39. Turner, Nancy Chapman and Marcus A. M. Bell, 1971, The Ethnobotany of the Coast Salish Indians of Vancouver Island, I and II, Economic Botany 25(1):63-104, 335-339, page 73
40. Turner, Nancy J., Laurence C. Thompson and M. Terry Thompson et al., 1990, Thompson Ethnobotany: Knowledge and Usage of Plants by the Thompson Indians of British Columbia, Victoria. Royal British Columbia Museum, page 114
41. Chestnut, V. K., 1902, Plants Used by the Indians of Mendocino County, California, Contributions from the U.S. National Herbarium 7:295-408., page 315
42. Curtin, L. S. M., 1957, Some Plants Used by the Yuki Indians ... II. Food Plants, The Masterkey 31:85-94, page 93

External links

• Jones, T. M. (2010) Interactive Visual Identification to Carices of North America @ LSU Herbarium
• eMonocot Cyperaceae Archived 2013-07-15 at the Wayback Machine, a portal to updated classification, images, species descriptions, and vetted specimen data for the entire sedge family, with a strong focus on Carex.