Charales

Charales is an order of freshwater green algae in the division Charophyta,

Stoneworts
class Charophyceae, commonly known as stoneworts. Linnaeus established
the genus Chara in 1753.

Contents
Description
Ecology
Life history
Species
British Isles
Distribution
Ireland
References
Further reading
External links
Chara globularis

Scientific classification
Description
Kingdom: Plantae
The Charales grow in freshwater and brackish environments worldwide, and
have large, macroscopic thalli growing up to 120 cm long, they are branched, Division: Charophyta
multicellular, and use chlorophyll to photosynthesize. Their only diploid stage Class: Charophyceae
in the life cycle is the unicellular oospore.[2] They may be called
Order: Charales
stoneworts,[3] because the plants can become encrusted in lime (calcium
carbonate) after some time. The "stem" is actually a central stalk consisting of Families[1]
giant, multinucleated cells. They are unique in having a whorl of small
branchlets at each node in the stipe, this gives them a superficial resemblance Aclistocharaceae†
to the genus Equisetum. In these whorls it is possible to see the phenomenon Atopocharaceae†
of cytoplasmic streaming. In fact the streaming in Chara is the fastest
Characeae
recorded of any cell. Cytoplasmic streaming is caused by the microfilaments
found inside the cell, as proven by the use of cytochalasin B to stop streaming. Clavatoraceae†
Feistiellaceae
There are about 400 species worldwide, with 33 in Britain and Ireland
according to Groves and Bullock-Webster),[4][5] however Stewart and Church Porocharaceae
(1992) reduce this to 21.[6]

Characeae are the principal plant life of some of the volcanic crater lakes of Nicaragua, and can be found in excess of 20 meters
depth in some circumstances. Introduced tilapia (Oreochromis niloticus) consumed all the Characeae in Lake Apoyo.[7]

Ecology
Most Characeae are found in fresh water, usually in still, clear water where they attach to the substrate by rhizoids. They can be
pioneer colonizers or ephemerals.[8] They are usually found in low to medium nutrient-rich water and tend to disappear due to
eutrophication. Some stoneworts can survive in brackish or maritime habitats and occur in ephemeral saline lakes in Australia
that have twice the salinity of seawater.[9]

Life history
The antheridia and oogonia are protected by a layer of sterile cells when mature; the oogonium is oblong in shape and consists of
a single egg, while the spherical antheridium is packed with threadlike cells that produce spermatia. As a result, the Charales
have the most complex structure of all green algae, if indeed they should be so labelled.

The possible sister group of the land plants are also known as brittleworts or skunkweed. These curious labels arise from the
fragility of their lime-encrusted stems, and from the foul odor these produce when stepped on.

Many botanists propose that the stoneworts and their relatives be placed in a division, sub-kingdom, or even kingdom by
themselves, often named Charophyta.[10][11] Their classification by taxonomists is currently undergoing much cladistic scrutiny.
Further DNA and RNA analysis may prove the charophytes to be a crucial evolutionary link in the phylogenetic tree of life, the
critical developmental step from the algae toward the non-vascular and then vascular land plants.

Species

British Isles
Ref: Stewart & Church (1992).[6]

Chara baltica Bruz.

Chara canescens Desv. & Lois.
Chara connivens Salzm. ex A.Braun
Chara curta Nolta ex Kütz. (=C. aspera var. curta)
Chara denudata (A.Braun) R.D.Wood Submerse meadow of Chara vulgaris
Chara fragifera Durieu
Chara intermedia Braun (=C. papillosa Kütz. and C. contraria x
hispida)
Chara mucosa J.Groves & Bullock-Webster
Chara rudis (A.Braun) Leonh.
Chara tomentosa L.
Lamprothamnium papulosum (Wallr.) J.Groves
Nitella capillaris (Krocker) J.Groves & Bullock-Webster
Nitella gracilis (Smith) Agardh
Nitella hyalina (DC.)Agardh
Lime crust on Chara sp. in a spring
Nitella mucronata (A.Braun)Miquel
pond in Germany
Nitella spanioclema J.Groves & Bullock-Webster (Nitella flexilis var.
spanioclema (J.Groves & Bullock-Webster))
Nitella tenuissima (Desv.) Kütz.
Nitellopsis obtusa (Desv.) J.Groves
Tolypella intricata (Trent. ex Roth) Leonh.[12]
Tolypella nidifica (O.F.Müll.) Leonh. (=Tolypella nidifica var. nidifica)
Tolypella prolifera (Ziz. ex A.Braun) Leonh.

Distribution
Ireland
County Antrim[13]

Chara aspera var. aspera

Chara globularis var. globularis
Chara vulgaris var. papillata Wallr. ex A. Braun
Chara globularis var. virgata (Kützing) R.D.Wood
Chara vulgaris var. vulgaris
Chara vulgaris var. contraria (A.Braun ex Kützing) J.A. Moore
Chara vulgaris var. longibracteata (Kützing) J. Groves & Bullock-Webster
Chara vulgaris var. papillata Wallr. ex A. Braun
Nitella flexilis var. flexilis
Nitella translucens (Pers.) C.A. Ag.
Tolypella nidifica var. glomerata (Desv.) R.D.Wood
County Down[13]

Chara aspera var. aspera

Chara aspera var. curta (Nolte ex Kützing) Braun ex Leonh.
Chara globularis var. globularis
Chara vulgaris var. papillata Wallr. ex A. Braun
Chara globularis var. virgata (Kützing) R.D.Wood
Chara globularis var. annulata (Lilleblad) J.A.Moore
Chara hispida L.
Chara hispida var. hispida
Chara hispida var. major (Hartm.) R.D. Wood
Chara hispida var. rudis A. Braun
Chara pedunculata Kützing
Chara vulgaris var. vulgaris
Chara vulgaris var. contraria (A.Braun ex Kützing) J.A. Moore
Chara vulgaris var. longibracteata (Kützing) J. Groves & Bullock-Webster
Chara vulgaris var. papillata Wallr. ex A. Braun
Nitella flexilis var. flexilis
Nitella translucens (Pers) C.A. Ag.
Tolypella nidifica var. glomerata (Desv.) R.D.Wood
County Londonderry[13]

Chara aspera Deth. ex Willd. var. aspera

Chara vulgaris var. papillata Wallr. ex A. Braun
Chara globularis var. globularis
Chara globularis var. virgata (Kützing) R.D.Wood
Chara hispida L.
Chara hispida var. hispida
Chara vulgaris var. vulgaris
Chara vulgaris var. contraria (A.Braun ex Kützing) J.A. Moore
Chara vulgaris var. papillata Wallr. ex A. Braun
Nitella flexilis var. flexilis
Nitella translucens (Pers) C.A. Ag.
Tolypella nidifica var. glomerata (Desv.) R.D.Wood
County Mayo.Recent records have been published from Clare Island.[14]

Chara virgata Kützing

Nitella flexilis (Linnaeus) C.Agardh
Nitella translucens (Persoon) C.Agardh
References
1. Algaebase (http://www.algaebase.org/browse/taxonomy/?id=4526)
2. Charales - an overview | ScienceDirect Topics (https://www.sciencedirect.com/topics/agricultural-and-biological-s
ciences/charales)
3. Kapraun DF (April 2007). "Nuclear DNA content estimates in green algal lineages: chlorophyta and streptophyta"
(http://aob.oxfordjournals.org/cgi/pmidlookup?view=long&pmid=17272304). Ann. Bot. 99 (4): 677–701.
doi:10.1093/aob/mcl294 (https://doi.org/10.1093%2Faob%2Fmcl294). PMC 2802934 (https://www.ncbi.nlm.nih.g
ov/pmc/articles/PMC2802934). PMID 17272304 (https://www.ncbi.nlm.nih.gov/pubmed/17272304).
4. Groves, J. and Bullock-Webster, G.R. 1920. The British Charophyta. Vol.1, Nitelleae. London, The Ray Society
5. Groves, J. and Bullock-Webster, G.R. 1924. The British Charophyta. Vol.2, Characeae.. London, The Ray
Society
6. Stewart, N.F. and Church, J.M. 1992. Red Data Books of Britain and Ireland. The Joint Nature Conservation
Committee, Peterborough. ISBN 1-873701-24-1
7. McCrary JK, Murphy BR, Stauffer, Jr. JR, Hendrix SS (2007): Tilapia (Teleostei: Cichlidae) status in Nicaraguan
natural waters; Env. Biol. Fishes 78:107-114 "Archived copy" (https://web.archive.org/web/20111003200007/htt
p://www.gaianicaragua.org/EBF2007.pdf) (PDF). Archived from the original (http://www.gaianicaragua.org/EBF20
07.pdf) (PDF) on 3 October 2011. Retrieved 31 May 2011.
8. John, D.M., Whitton, B.A. and Brook, A.J. 2002. The Freshwater Algal Flora of the British Isles. Cambridge
University Press, London. ISBN 0-521-77051-3
9. Burne, R.V., Bauld, J. and de Decker, P. 1980. Saline lake charophytes and their geological significance. Journal
of Sedimentary Petrology, 50, 281-293. https://doi.org/10.1306/212F79D2-2B24-11D7-8648000102C1865D
10. Kenneth G. Karol, Richard M. McCourt, Matthew T. Cimino, Charles F. Delwiche (December 2001). "The Closest
Living Relatives of Land Plants" (http://science.sciencemag.org/content/294/5550/2351?sid=d9f712d8-5e6d-440e
-b157-64ede6b28cc2). Science Magazine.
11. "CHARA (MUSKGRASS; STONEWORT) Chara spp" (https://nature.mdc.mo.gov/discover-nature/field-guide/char
a-muskgrass-stonewort). Missouri Department of Conservation.
12. "Tolypella intricata (Trentepohl ex Roth) Leonhardi" (http://www.algaebase.org/search/species/detail/?species_id
=35602). AlgaeBase.
13. Morton, O. 1992 in Hackney, P. (Ed.) Stewart & Corry's Flora of the North-east of Ireland. Institute of Irish
Studies and The Queen's University of Belfast ISBN 0-85389-446-9 (HB)
14. Guiry, M.D., John, D.M., Rindi, F. and McCarthy, T.K 2007. New Survey of Clare Island, Volume 6: The
Freshwater and Terrestrial Algae. Royal Irish Academy. ISBN 978-1-904890-31-7

Further reading
Bryant, J. The stoneworts (Chlorophyta, Charales). In Guiry, M.D., John, D.M., Rindi, F. and McCarthy, T.K. 2007.
New Survey of Clare Island. Royal Irish Academy. ISBN 978-1-904890-31-7.
Lloyd, James. 2007. "Cytoskeletal Structures Responsible for Cytoplasmic Streaming in Chara." St. Vincent-St.
Mary High School in Accordance with Dr. Donald Ott of The University of Akron. (Science Inquiry)
Schaible, R. and Schubert, H. 2008. The occurrence of sexual Chara canesces populations (Charophyceae) is
not related to ecophysiological potentials with respect to salinity and irradiance. Eur. J. Phycol. 43: 309 - 316.
Desai, Udaysingh and Karande C.T. 2008. "Biodiversity of Charophytes from Kolhapur District, Maharashtra".
Shivaji University, Kolhapur.

External links
Images of Charales (https://web.archive.org/web/20110904203722/http://www.azote.se/index.asp?q=Charophyc
eae&lang=eng&ref=W)
University California (https://web.archive.org/web/20070701164144/http://plants.ifas.ufl.edu/charpic.html)
 Chara curta in Ireland (https://web.archive.org/web/20070930181245/http://www.habitas.org.uk/priority/species.a
sp?item=39846)

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