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Charophyta () is a group of
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does in ...
green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ( Embryophytes) have emerged deep in the Charophyte alg ...
, called charophytes (), sometimes treated as a
division Division or divider may refer to: Mathematics *Division (mathematics), the inverse of multiplication *Division algorithm, a method for computing the result of mathematical division Military * Division (military), a formation typically consisting ...
, yet also as a superdivision or an unranked clade. The terrestrial plants, the
Embryophyta The Embryophyta (), or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes () have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as sis ...
emerged within Charophyta, possibly from terrestrial unicellular charophytes, with the class
Zygnematophyceae Zygnematophyceae (or Conjugatophyceae) is a class of green algae in the paraphylum streptophyte algae, also referred to as Charophyta, consisting of more than 4000 described species. It contains five orders: the Spirogloeales, the Serritaeniales, ...
as a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
. The clade
Streptophyta Streptophyta (), informally the streptophytes (, from the Greek ''strepto'' 'twisted', for the morphology of the sperm of some members), is a clade of plants. The composition of the clade varies considerably between authors, but the definition ...
may be formed by placing Embryophyta within the Charophyta. The Embryophyta may already be included in the Charophyta, in which case it is a synonym of the Streptophyta. The sister group of the charophytes are the
Chlorophyta Chlorophyta or Prasinophyta is a taxon of green algae informally called chlorophytes. The name is used in two very different senses, so care is needed to determine the use by a particular author. In older classification systems, it refers to a ...
. In some charophyte groups, such as the
Zygnematophyceae Zygnematophyceae (or Conjugatophyceae) is a class of green algae in the paraphylum streptophyte algae, also referred to as Charophyta, consisting of more than 4000 described species. It contains five orders: the Spirogloeales, the Serritaeniales, ...
or conjugating green algae,
flagella A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
are absent and sexual reproduction does not involve free-swimming flagellate sperm. Flagellate sperm, however, are found in
stonewort Charales is an order of freshwater green algae in the division Charophyta, class Charophyceae, commonly known as stoneworts. Depending on the treatment of the genus ''Nitellopsis'', living (extant) species are placed into either one family (Ch ...
s (
Charales Charales is an order of freshwater green algae in the division Charophyta, class Charophyceae, commonly known as stoneworts. Depending on the treatment of the genus '' Nitellopsis'', living (extant) species are placed into either one family (C ...
) and
Coleochaetales Coleochaetaceae is a family of algae. It is the only family in the Coleochaetales, an order of parenchymous charophyte algae, that includes some of the closest multicellular relatives of land plants. They questionably include the fossil genus ...
, orders of parenchymatous charophytes that are the closest relatives of the land plants, where flagellate sperm are also present in all except the
conifers Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All ex ...
and
flowering plants Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. They include all forbs (flowering plants without a woody stem), grasses and grass-like plants, a vast majority of broa ...
. Fossil stoneworts of early
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, w ...
age that are similar to those of the present day have been described from the
Rhynie chert The Rhynie chert is a Lower Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness (a Lagerstätte). It is exposed near the village of Rhynie, Aberdeenshire, Scotland; a second unit, the Windyfield chert, is locate ...
of Scotland. Somewhat different charophytes have also been collected from the Late Devonian (Famennian)
Waterloo Farm lagerstätte The Waterloo Farm lagerstätte is a Famennian lagerstätte in South Africa that constitutes the only known record of a near-polar Devonian coastal ecosystem. History and discovery The Waterloo Farm Lagerstätte is an approximately 360 millio ...
of South Africa. These include two species each of ''
Octochara ''Octochara'' is a genus of fossil charophyte (aquatic green alga) from the Famennian (Late Devonian). It is one of two genera of charophyte described from the Waterloo Farm lagerstätte in southern Africa. It and ''Hexachara'', from the same loca ...
'' and ''
Hexachara ''Hexachara'' is a genus of fossil charophyte (aquatic green alga) that is likely to have formed meadows within sheltered oligohaline reaches of lakes. Etymology ''Hexachara'' is derived from Greek "hex", meaning six, a reference to the hexaradi ...
'', which are the oldest fossils of Charophyte axes bearing in situ
oogonia An oogonium (plural oogonia) is a small diploid cell which, upon maturation, forms a primordial follicle in a female fetus or the female (haploid or diploid) gametangium of certain thallophytes. In the mammalian fetus Oogonia are formed in larg ...
. The name comes from the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
'' Chara''.


Description

The Zygnematophyceae or, as they used to be called, Conjugatophyceae, generally possess two fairly elaborate chloroplasts in each cell, rather than many discoid ones. They reproduce asexually by the development of a septum between the two cell-halves or semi-cells (in unicellular forms, each daughter-cell develops the other semi-cell afresh) and sexually by conjugation, or the fusion of the entire cell-contents of the two conjugating cells. The saccoderm desmids and the placoderm or true desmids, unicellular or filamentous members of the Zygnematophyceae, are dominant in non-calcareous, acid waters of oligotrophic or primitive lakes (e.g. Wastwater), or in lochans, tarns and bogs, as in the West of Scotland, Eire, parts of Wales and of the Lake District. '' Klebsormidium'', the type of the
Klebsormidiophyceae Klebsormidiales is an order of charophyte algae. It is the only order in the class Klebsormidiophyceae, sister of the Phragmoplastophyta. , AlgaeBase accepted two families in the order: * Elakatotrichaceae Hindák *Klebsormidiaceae The Klebsorm ...
, is a simple filamentous form with circular, plate-like chloroplasts, reproducing by fragmentation, by dorsiventral, biciliate swarmers and, according to Wille, a twentieth-century algologist, by aplanospores. Sexual reproduction is simple and
isogamous Isogamy is a form of sexual reproduction that involves gametes of the same morphology (indistinguishable in shape and size), found in most unicellular eukaryotes. Because both gametes look alike, they generally cannot be classified as male or f ...
(the male and female gametes are outwardly indistinguishable). The various groups included in the Charophyta have diverse and idiosyncratic reproductive systems, sometimes with complex reproductive organs. The unique habit among the algae of protecting the overwintering
zygote A zygote (, ) is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism. In multicell ...
within the tissues of the parent
gametophyte A gametophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has on ...
is one of several characteristics of Coleochaetales that suggest that they are a sister group to the embryophytes. The Charales or stoneworts are freshwater algae with slender green or grey stems; the grey colour of many species results from the deposition of lime on the walls, masking the green colour of the chlorophyll. The main stems are slender and branch occasionally. Lateral
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