Chlorophyta: Difference between revisions

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{{About|a group of green algae|the genus with the common name 'spider plant'|Chlorophytum}}

~~{{Short description|Phylum of green algae}}~~

{{Automatic taxobox

| fossil_range = {{Long fossil range|1000|0|earliest=2040|ref={{Sfn|Tang|Pang|Yuan|Xiao|2020}}}}

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| subdivision_ref = {{Sfn|Guiry|2024}}

| subdivision = * [[Chloropicophyceae]]

* {{Extinct}}[[Chuariophyceae]]

* {{Extinct}}Chuariophyceae

* [[Mamiellophyceae]]

* [[Nephroselmidophyceae]]

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}}

'''Chlorophyta'''{{efn|{{Etymology|gre|''χλωρός'' chlōros|pale green||''φυτόν'' phyton|plant}}{{Sfn|Margulis|Chapman|2009|p=200}}}} is a [[division (botany)|division]] of [[green algae]] ~~informally called '''chlorophytes'''.~~{{Sfn|Rockwell|Martin|Li|Mathews|2017}}

'''Chlorophyta'''{{efn|{{Etymology|gre|''χλωρός'' chlōros|pale green||''φυτόν'' phyton|plant}}{{Sfn|Margulis|Chapman|2009|p=200}}}} or '''chlorophytes''' is a major [[division (botany)|division]] of [[green algae]],{{Sfn|Rockwell|Martin|Li|Mathews|2017}} and is [[sister taxon]] to the other major division [[Charophyta]] (a [[paraphyletic]] group of predominantly [[freshwater]] green algae, which form the [[monophyletic]] [[clade]] [[Streptophyta]] after including all [[land plant]]s) as well as the proposed [[basal (phylogenetics)|basal]] clade [[Prasinodermophyta]], together with whom they form the major [[primary algae]] clade [[Viridiplantae]] ([[Plantae]] ''[[sensu stricto]]'').

== Description ==

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[[Asexual reproduction]] is widely observed in chlorophytes. Among core chlorophytes, both unicellular groups can reproduce asexually through [[autospore]]s,{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=17-8}} wall-less zoospores,{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=17-11}} fragmentation, plain cell division, and exceptionally budding.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=17-9}} Multicellular thalli can reproduce asexually through motile zoospores,{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-8}} non-motile [[aplanospore]]s, autospores, filament fragmentation,{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=19-3}} differentiated resting cells,{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-19}} and even unmated gametes.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-29}} Colonial groups can reproduce asexually through the formation of autocolonies, where each cell divides to form a colony with the same number and arrangement of cells as the parent colony.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=19-14}}

Many chlorophytes exclusively conduct asexual reproduction, but some display [[sexual reproduction]], which may be [[isogamy|isogamous]] (i.e., [[gamete]]s of both sexes are identical), [[anisogamy|anisogamous]] (gametes are different) or [[oogamy|oogamous]] (gametes are sperm and egg cells), with an evolutionary tendency towards oogamy. Their gametes are usually specialized cells differentiated from [[vegetative cell]]s, although in unicellular [[Volvocales]] the vegetative cells can function simultaneously as gametes. Most chlorophytes have a [[~~Biological_life_cycle~~#~~Diplontic_life_cycle~~|diplontic life cycle]] (also known as zygotic), where the gametes fuse into a [[zygote]] which germinates, grows and eventually undergoes [[meiosis]] to produce [[haploid]] [[spore]]s (gametes), similarly to [[ochrophyte]]s and [[animal]]s. Some exceptions display a [[~~Biological_life_cycle~~#~~Haplodiplontic_life_cycle~~|haplodiplontic life cycle]], where there is an alternation of generations, similarly to land plants.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=16-13}} These generations can be isomorphic (i.e., of similar shape and size) or heteromorphic.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-14}} The formation of reproductive cells usually does not occur in specialized cells,{{Sfn|Lee|2018|p=318}} but some [[Ulvophyceae]] have specialized reproductive structures: gametangia, to produce gametes, and sporangia, to produce spores.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-14}}

Many chlorophytes exclusively conduct asexual reproduction, but some display [[sexual reproduction]], which may be [[isogamy|isogamous]] (i.e., [[gamete]]s of both sexes are identical), [[anisogamy|anisogamous]] (gametes are different) or [[oogamy|oogamous]] (gametes are sperm and egg cells), with an evolutionary tendency towards oogamy. Their gametes are usually specialized cells differentiated from [[vegetative cell]]s, although in unicellular [[Volvocales]] the vegetative cells can function simultaneously as gametes. Most chlorophytes have a [[Biological life cycle#Diplontic life cycle|diplontic life cycle]] (also known as zygotic), where the gametes fuse into a [[zygote]] which germinates, grows and eventually undergoes [[meiosis]] to produce [[haploid]] [[spore]]s (gametes), similarly to [[ochrophyte]]s and [[animal]]s. Some exceptions display a [[Biological life cycle#Haplodiplontic life cycle|haplodiplontic life cycle]], where there is an alternation of generations, similarly to land plants.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=16-13}} These generations can be isomorphic (i.e., of similar shape and size) or heteromorphic.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-14}} The formation of reproductive cells usually does not occur in specialized cells,{{Sfn|Lee|2018|p=318}} but some [[Ulvophyceae]] have specialized reproductive structures: gametangia, to produce gametes, and sporangia, to produce spores.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-14}}

The earliest-diverging unicellular chlorophytes (prasinophytes) produce walled resistant stages called [[microbial cyst|cysts]] or 'phycoma' stages before reproduction; in some groups the cysts are as large as 230 μm in diameter. To develop them, the flagellate cells form an inner wall by discharging mucilage vesicles to the outside, increase the level of lipids in the cytoplasm to enhance [[buoyancy]], and finally develop an outer wall. Inside the cysts, the nucleus and cytoplasm undergo [[cell division|division]] into numerous flagellate cells that are released by rupturing the wall. In some species these daughter cells have been confirmed to be gametes; otherwise, sexual reproduction is unknown in prasinophytes.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=16-17}}

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* [[Chlorodendrophyceae]] (60 species, 15 extinct):{{Sfn|Guiry|2024|p=5}} unicellular [[flagellate]]s (monadoids) surrounded by an outer cell covering or [[theca]] of organic extracellular scales composed of [[protein]]s and ketosugars. Some of these scales make up hair-like structures. Capable of [[asexual reproduction]] through cell division inside the theca. No [[sexual reproduction]] has been described. Each cell contains a single chloroplast and exhibits two [[flagella]]. Present in marine and freshwater habitats.{{Sfn|Hori|Norris|Chihara|1986}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=37}}{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=17-2}}

* [[Chlorophyceae]] (3,974 species):{{Sfn|Guiry|2024|p=5}} either unicellular monadoids (flagellated) or [[wikt:coccoid|coccoid]]s (without flagella) living solitary or in varied colonial forms (including [[coenocyte|coenobial]]), or [[multicellular]] filamentous (branch-like) [[thalli]] that may be ramified, or foliose (leaf-like) thalli. Cells are surrounded by a crystalline covering composed of [[glycoprotein]]s abundant in [[glycine]] and [[hydroxyproline]], as well as [[pectin]]s, [[arabinogalactan protein]]s, and [[extensin]]. They exhibit a [[Biological life cycle#Haplontic life cycle|haplontic life cycle]] with [[isogamy]], [[anisogamy]] or [[oogamy]]. They are capable of asexual reproduction through flagellated [[zoospore]]s, [[aplanospore]]s, or [[autospore]]s. Each cell contains a single chloroplast, a variable number of [[pyrenoid]]s (including lack thereof), and from one to hundreds of flagella without [[mastigoneme]]s. Present in marine, freshwater and terrestrial habitats.{{Sfn|Domozych|Ciancia|Fangel|Mikkelsen|2012}}{{Sfn|Leliaert|Smith|Moreau|Herron|2012}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=36–37}}{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=19-2–19-5}}

* [[Chlorophyceae]] (3,974 species):{{Sfn|Guiry|2024|p=5}} either unicellular monadoids (flagellated) or [[wikt:coccoid|coccoid]]s (without flagella) living solitary or in varied colonial forms (including [[coenocyte|coenobial]]), or [[multicellular]] filamentous (branch-like) [[thalli]] that may be ramified, or foliose (leaf-like) thalli. Cells are surrounded by a crystalline covering composed of [[glycoprotein]]s abundant in [[glycine]] and [[hydroxyproline]], as well as [[pectin]]s, [[arabinogalactan protein]]s, and [[extensin]]. They exhibit a [[~~Biological_life_cycle~~#~~Haplontic_life_cycle~~|haplontic life cycle]] with [[isogamy]], [[anisogamy]] or [[oogamy]]. They are capable of asexual reproduction through flagellated [[zoospore]]s, [[aplanospore]]s, or [[autospore]]s. Each cell contains a single chloroplast, a variable number of [[pyrenoid]]s (including lack thereof), and from one to hundreds of flagella without [[mastigoneme]]s. Present in marine, freshwater and terrestrial habitats.{{Sfn|Domozych|Ciancia|Fangel|Mikkelsen|2012}}{{Sfn|Leliaert|Smith|Moreau|Herron|2012}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=36–37}}{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=19-2–19-5}}

* [[Chloropicophyceae]] (8 species):{{Sfn|Guiry|2024|p=5}} unicellular solitary coccoids. Cells are surrounded by a multi-layered [[cell wall]]. No sexual or asexual reproduction has been described. Each cell contains a single chloroplast with [[astaxanthin]] and [[loroxanthin]], and lacks pyrenoids or flagella. They are exclusively marine.{{Sfn|Lopes dos Santos|Pollina|Gourvil|Corre|2017}}

* [[Chuariophyceae]] (3 extinct species): exclusively fossil group containing carbonaceous megafossils found in [[Ediacaran]] rocks, such as ''[[Tawuia]]''.{{Sfn|Guiry|2024|p=5}}{{Sfn|Srivastava|2002}}

* [[Mamiellophyceae]] (25 species):{{Sfn|Guiry|2024|p=5}} unicellular solitary monadoids. Cells are naked or covered by one or two layers of flat scales, mainly with spiderweb-like or reticulate ornamentation. Each cell contains one or rarely two chloroplasts, almost always with [[prasinoxanthin]]; two equal or unequal flagella, or just one flagellum, or lacking any flagella. If flagella are present, they can be either smooth or covered in scales in the same manner as the cells. Present in marine and freshwater habitats.{{Sfn|Marin|Melkonian|2010}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=37}}

* [[Nephroselmidophyceae]] (29 species):{{Sfn|Guiry|2024|p=5}} unicellular monadoids. Cells are covered by scales. They are capable of sexual reproduction through [[hologamy]] (fusion of entire cells), and of asexual reproduction through [[binary fission]]. Each cell contains a single chloroplast, a pyrenoid, and two flagella covered by scales. Present in marine and freshwater habitats.{{Sfn|Nakayama|Suda|Kawachi|Inouye|2007}}{{Sfn|Yamaguchi|Suda|Nakayama|Pienaar|2010}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=37}}

* [[Nephroselmidophyceae]] (29 species):{{Sfn|Guiry|2024|p=5}} unicellular monadoids. Cells are covered by scales. They are capable of sexual reproduction through [[hologamy]] (fusion of entire cells), and of asexual reproduction through [[binary fission]]. Each cell contains a single ~~cloroplast~~, a pyrenoid, and two flagella covered by scales. Present in marine and freshwater habitats.{{Sfn|Nakayama|Suda|Kawachi|Inouye|2007}}{{Sfn|Yamaguchi|Suda|Nakayama|Pienaar|2010}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=37}}

* [[Pedinophyceae]] (24 species):{{Sfn|Guiry|2024|p=5}} unicellular asymmetrical monadoids that undergo a coccoid palmelloid phase covered by [[mucilage]]. Cells lack extracellular scales, but in rare cases are covered on the posterior side by a theca. Each cell contains a single chloroplast, a pyrenoid, and a single flagellum usually covered in mastigonemes. Present in marine, freshwater and terrestrial habitats.{{Sfn|Marin|2012}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=37}}{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=17-3}}

* [[Picocystophyceae]] (1 species):{{Sfn|Guiry|2024|p=5}} unicellular coccoids, ovoid and trilobed in shape. Cells are surrounded by a multi-layered cell wall of poly-[[arabinose]], [[mannose]], [[galactose]] and [[glucose]]. No sexual reproduction has been described. They are capable of asexual reproduction through autosporulation, resulting in two or rarely four daughter cells. Each cell contains a single bilobed chloroplast with diatoxanthin and monadoxanthin, without any pyrenoid or flagella. Present in [[saline lake]]s.{{Sfn|Lewin|Krienitz|Goericke|Takeda|2000}}{{Sfn|Lopes dos Santos|Pollina|Gourvil|Corre|2017}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=37}}

* [[Pyramimonadophyceae]] (166 species, 59 extinct):{{Sfn|Guiry|2024|p=5}} unicellular monadoids or coccoids. Cells are covered by two or more layers of organic scales. No sexual reproduction has been described, but some cells with only one flagellum have been interpreted as potential gametes. Asexual reproduction has only been observed in the coccoid forms, via zoospores. Each cell contains a single chloroplast, a pyrenoid, and between 4 and 16 flagella. The flagella are covered in at least two layers of organic scales: a bottom layer of pentagonal scales organized in 24 rows, and a top layer of limuloid scales distributed in 11 rows. They are exclusively marine.{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=37}}{{Sfn|Daugbjerg|Fassel|Moestrup|2020}}

* [[Trebouxiophyceae]] (926 species, 1 extinct):{{Sfn|Guiry|2024|p=5}} unicellular monadoids occasionally without flagella, or colonial, or ramified filamentous thalli, or living as the [[photobiont]]s of [[lichen]]. Cells are covered by a cell wall of [[cellulose]], [[algaenan]]s, and β-galactofuranane. No sexual reproduction has been described with the exception of some observations of gamete fusion and presence of [[meiosis|meiotic]] [[gene]]s. They are capable of asexual reproduction through autospores or zoospores. Each cell contains a single chloroplast, a pyrenoid, and one or two pairs of smooth flagella. They are present in marine, freshwater and terrestrial habitats.{{Sfn|Domozych|Ciancia|Fangel|Mikkelsen|2012}}{{Sfn|Fučíková|Pažoutová|Rindi|2015}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=36}}{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=17-4–17-11}}

* [[Ulvophyceae]] (2,695 species, 990 extinct):{{Sfn|Guiry|2024|p=5}} macroscopic thalli, either filamentous (which may be ramified) or foliose (composed of monostromatic or distromatic layers) or even compact tubular forms, generally [[multinucleate]]. Cells surrounded by a cell wall that may be [[Marine biogenic calcification|calcified]], composed of cellulose, β-manane, β-xilane, sulphated or piruvilated polysaccharides or sulphated ramnogalacturonanes, arabinogalactan proteins, and extensin. They exhibit a [[Biological life cycle#Haplodiplontic life cycle|haplodiplontic life cycle]] where the alternating generations can be isomorphic or heteromorphic. They reproduce asexually via zoospores that may be covered in scales. Each cell contains a single chloroplast, and one or two pairs of flagella without mastigonemes but covered in scales. They are present in marine, freshwater and terrestrial habitats.{{Sfn|Domozych|Ciancia|Fangel|Mikkelsen|2012}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=36}}{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-2–18-24}}

* [[Ulvophyceae]] (2,695 species, 990 extinct):{{Sfn|Guiry|2024|p=5}} macroscopic thalli, either filamentous (which may be ramified) or foliose (composed of monostromatic or distromatic layers) or even compact tubular forms, generally [[multinucleate]]. Cells surrounded by a cell wall that may be [[Marine biogenic calcification|calcified]], composed of cellulose, β-manane, β-xilane, sulphated or piruvilated polysaccharides or sulphated ramnogalacturonanes, arabinogalactan proteins, and extensin. They exhibit a [[~~Biological_life_cycle~~#~~Haplodiplontic_life_cycle~~|haplodiplontic life cycle]] where the alternating generations can be isomorphic or heteromorphic. They reproduce asexually via zoospores that may be covered in scales. Each cell contains a single chloroplast, and one or two pairs of flagella without mastigonemes but covered in scales. They are present in marine, freshwater and terrestrial habitats.{{Sfn|Domozych|Ciancia|Fangel|Mikkelsen|2012}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=36}}{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-2–18-24}}

== Evolution ==

In February 2020, the fossilized remains of a green alga, named ''[[Proterocladus antiquus]]'' were discovered in the northern province of [[Liaoning]], [[China]]. At around a billion years old, it is believed to be one of the oldest examples of a multicellular chlorophyte. It is currently classified as a member of order [[Cladophorales|Siphonocladales]], class [[Ulvophyceae]].{{Sfn|Tang|Pang|Yuan|Xiao|2020}} In 2023, a study calculated the [[molecular clock|molecular age]] of green algae as [[~~Molecular_clock~~#Calibration|calibrated]] by this fossil. The study estimated the origin of Chlorophyta within the [[Mesoproterozoic]] era, at around 2.04–1.23 billion years ago.{{Sfn|Yang|Ma|Wang|Tian|2023}}

In February 2020, the fossilized remains of a green alga, named ''[[Proterocladus antiquus]]'' were discovered in the northern province of [[Liaoning]], [[China]]. At around a billion years old, it is believed to be one of the oldest examples of a multicellular chlorophyte. It is currently classified as a member of order [[Cladophorales|Siphonocladales]], class [[Ulvophyceae]].{{Sfn|Tang|Pang|Yuan|Xiao|2020}} In 2023, a study calculated the [[molecular clock|molecular age]] of green algae as [[Molecular clock#Calibration|calibrated]] by this fossil. The study estimated the origin of Chlorophyta within the [[Mesoproterozoic]] era, at around 2.04–1.23 billion years ago.{{Sfn|Yang|Ma|Wang|Tian|2023}}

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[[Category:Chlorophyta|Chlorophyta]]

~~[[Category:Plant divisions]]~~

[[Category:Taxa named by Ludwig Reichenbach]]

[[Category:Green algae phyla]]

@@ Line 1: / Line 1: @@
+{{Short description|Phylum of green algae}}
 {{About|a group of green algae|the genus with the common name 'spider plant'|Chlorophytum}}
-{{Short description|Phylum of green algae}}
 {{Automatic taxobox
 | fossil_range = {{Long fossil range|1000|0|earliest=2040|ref={{Sfn|Tang|Pang|Yuan|Xiao|2020}}}}
@@ Line 10: / Line 10: @@
 | subdivision_ref = {{Sfn|Guiry|2024}}
 | subdivision = * [[Chloropicophyceae]]
-* {{Extinct}}[[Chuariophyceae]]
+* {{Extinct}}Chuariophyceae
 * [[Mamiellophyceae]]
 * [[Nephroselmidophyceae]]
@@ Line 30: / Line 30: @@
 }}
-'''Chlorophyta'''{{efn|{{Etymology|gre|''χλωρός'' chlōros|pale green||''φυτόν'' phyton|plant}}{{Sfn|Margulis|Chapman|2009|p=200}}}} is a [[division (botany)|division]] of [[green algae]] informally called '''chlorophytes'''.{{Sfn|Rockwell|Martin|Li|Mathews|2017}}
+'''Chlorophyta'''{{efn|{{Etymology|gre|''χλωρός'' chlōros|pale green||''φυτόν'' phyton|plant}}{{Sfn|Margulis|Chapman|2009|p=200}}}} or  '''chlorophytes''' is a major [[division (botany)|division]] of [[green algae]],{{Sfn|Rockwell|Martin|Li|Mathews|2017}} and is [[sister taxon]] to the other major division [[Charophyta]] (a [[paraphyletic]] group of predominantly [[freshwater]] green algae, which form the [[monophyletic]] [[clade]] [[Streptophyta]] after including all [[land plant]]s) as well as the proposed [[basal (phylogenetics)|basal]] clade [[Prasinodermophyta]], together with whom they form the major [[primary algae]] clade [[Viridiplantae]] ([[Plantae]] ''[[sensu stricto]]'').
 == Description ==
@@ Line 53: / Line 53: @@
 [[Asexual reproduction]] is widely observed in chlorophytes. Among core chlorophytes, both unicellular groups can reproduce asexually through [[autospore]]s,{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=17-8}} wall-less zoospores,{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=17-11}} fragmentation, plain cell division, and exceptionally budding.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=17-9}} Multicellular thalli can reproduce asexually through motile zoospores,{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-8}} non-motile [[aplanospore]]s, autospores, filament fragmentation,{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=19-3}} differentiated resting cells,{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-19}} and even unmated gametes.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-29}} Colonial groups can reproduce asexually through the formation of autocolonies, where each cell divides to form a colony with the same number and arrangement of cells as the parent colony.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=19-14}}
-Many chlorophytes exclusively conduct asexual reproduction, but some display [[sexual reproduction]], which may be [[isogamy|isogamous]] (i.e., [[gamete]]s of both sexes are identical), [[anisogamy|anisogamous]] (gametes are different) or [[oogamy|oogamous]] (gametes are sperm and egg cells), with an evolutionary tendency towards oogamy. Their gametes are usually specialized cells differentiated from [[vegetative cell]]s, although in unicellular [[Volvocales]] the vegetative cells can function simultaneously as gametes. Most chlorophytes have a [[Biological_life_cycle#Diplontic_life_cycle|diplontic life cycle]] (also known as zygotic), where the gametes fuse into a [[zygote]] which germinates, grows and eventually undergoes [[meiosis]] to produce [[haploid]] [[spore]]s (gametes), similarly to [[ochrophyte]]s and [[animal]]s. Some exceptions display a [[Biological_life_cycle#Haplodiplontic_life_cycle|haplodiplontic life cycle]], where there is an alternation of generations, similarly to land plants.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=16-13}} These generations can be isomorphic (i.e., of similar shape and size) or heteromorphic.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-14}} The formation of reproductive cells usually does not occur in specialized cells,{{Sfn|Lee|2018|p=318}} but some [[Ulvophyceae]] have specialized reproductive structures: gametangia, to produce gametes, and sporangia, to produce spores.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-14}}
+Many chlorophytes exclusively conduct asexual reproduction, but some display [[sexual reproduction]], which may be [[isogamy|isogamous]] (i.e., [[gamete]]s of both sexes are identical), [[anisogamy|anisogamous]] (gametes are different) or [[oogamy|oogamous]] (gametes are sperm and egg cells), with an evolutionary tendency towards oogamy. Their gametes are usually specialized cells differentiated from [[vegetative cell]]s, although in unicellular [[Volvocales]] the vegetative cells can function simultaneously as gametes. Most chlorophytes have a [[Biological life cycle#Diplontic life cycle|diplontic life cycle]] (also known as zygotic), where the gametes fuse into a [[zygote]] which germinates, grows and eventually undergoes [[meiosis]] to produce [[haploid]] [[spore]]s (gametes), similarly to [[ochrophyte]]s and [[animal]]s. Some exceptions display a [[Biological life cycle#Haplodiplontic life cycle|haplodiplontic life cycle]], where there is an alternation of generations, similarly to land plants.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=16-13}} These generations can be isomorphic (i.e., of similar shape and size) or heteromorphic.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-14}} The formation of reproductive cells usually does not occur in specialized cells,{{Sfn|Lee|2018|p=318}} but some [[Ulvophyceae]] have specialized reproductive structures: gametangia, to produce gametes, and sporangia, to produce spores.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-14}}
 The earliest-diverging unicellular chlorophytes (prasinophytes) produce walled resistant stages called [[microbial cyst|cysts]] or 'phycoma' stages before reproduction; in some groups the cysts are as large as 230 μm in diameter. To develop them, the flagellate cells form an inner wall by discharging mucilage vesicles to the outside, increase the level of lipids in the cytoplasm to enhance [[buoyancy]], and finally develop an outer wall. Inside the cysts, the nucleus and cytoplasm undergo [[cell division|division]] into numerous flagellate cells that are released by rupturing the wall. In some species these daughter cells have been confirmed to be gametes; otherwise, sexual reproduction is unknown in prasinophytes.{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=16-17}}
@@ Line 173: / Line 173: @@
 * [[Chlorodendrophyceae]] (60 species, 15 extinct):{{Sfn|Guiry|2024|p=5}} unicellular [[flagellate]]s (monadoids) surrounded by an outer cell covering or [[theca]] of organic extracellular scales composed of [[protein]]s and ketosugars. Some of these scales make up hair-like structures. Capable of [[asexual reproduction]] through cell division inside the theca. No [[sexual reproduction]] has been described. Each cell contains a single chloroplast and exhibits two [[flagella]]. Present in marine and freshwater habitats.{{Sfn|Hori|Norris|Chihara|1986}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=37}}{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=17-2}}
+* [[Chlorophyceae]] (3,974 species):{{Sfn|Guiry|2024|p=5}} either unicellular monadoids (flagellated) or [[wikt:coccoid|coccoid]]s (without flagella) living solitary or in varied colonial forms (including [[coenocyte|coenobial]]), or [[multicellular]] filamentous (branch-like) [[thalli]] that may be ramified, or foliose (leaf-like) thalli. Cells are surrounded by a crystalline covering composed of [[glycoprotein]]s abundant in [[glycine]] and [[hydroxyproline]], as well as [[pectin]]s, [[arabinogalactan protein]]s, and [[extensin]]. They exhibit a [[Biological life cycle#Haplontic life cycle|haplontic life cycle]] with [[isogamy]], [[anisogamy]] or [[oogamy]]. They are capable of asexual reproduction through flagellated [[zoospore]]s, [[aplanospore]]s, or [[autospore]]s. Each cell contains a single chloroplast, a variable number of [[pyrenoid]]s (including lack thereof), and from one to hundreds of flagella without [[mastigoneme]]s. Present in marine, freshwater and terrestrial habitats.{{Sfn|Domozych|Ciancia|Fangel|Mikkelsen|2012}}{{Sfn|Leliaert|Smith|Moreau|Herron|2012}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=36–37}}{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=19-2–19-5}}
-* [[Chlorophyceae]] (3,974 species):{{Sfn|Guiry|2024|p=5}} either unicellular monadoids (flagellated) or [[wikt:coccoid|coccoid]]s (without flagella) living solitary or in varied colonial forms (including [[coenocyte|coenobial]]), or [[multicellular]] filamentous (branch-like) [[thalli]] that may be ramified, or foliose (leaf-like) thalli. Cells are surrounded by a crystalline covering composed of [[glycoprotein]]s abundant in [[glycine]] and [[hydroxyproline]], as well as [[pectin]]s, [[arabinogalactan protein]]s, and [[extensin]]. They exhibit a [[Biological_life_cycle#Haplontic_life_cycle|haplontic life cycle]] with [[isogamy]], [[anisogamy]] or [[oogamy]]. They are capable of asexual reproduction through flagellated [[zoospore]]s, [[aplanospore]]s, or [[autospore]]s. Each cell contains a single chloroplast, a variable number of [[pyrenoid]]s (including lack thereof), and from one to hundreds of flagella without [[mastigoneme]]s. Present in marine, freshwater and terrestrial habitats.{{Sfn|Domozych|Ciancia|Fangel|Mikkelsen|2012}}{{Sfn|Leliaert|Smith|Moreau|Herron|2012}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=36–37}}{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=19-2–19-5}}
 * [[Chloropicophyceae]] (8 species):{{Sfn|Guiry|2024|p=5}} unicellular solitary coccoids. Cells are surrounded by a multi-layered [[cell wall]]. No sexual or asexual reproduction has been described. Each cell contains a single chloroplast with [[astaxanthin]] and [[loroxanthin]], and lacks pyrenoids or flagella. They are exclusively marine.{{Sfn|Lopes dos Santos|Pollina|Gourvil|Corre|2017}}
 * [[Chuariophyceae]] (3 extinct species): exclusively fossil group containing carbonaceous megafossils found in [[Ediacaran]] rocks, such as ''[[Tawuia]]''.{{Sfn|Guiry|2024|p=5}}{{Sfn|Srivastava|2002}}
 * [[Mamiellophyceae]] (25 species):{{Sfn|Guiry|2024|p=5}} unicellular solitary monadoids. Cells are naked or covered by one or two layers of flat scales, mainly with spiderweb-like or reticulate ornamentation. Each cell contains one or rarely two chloroplasts, almost always with [[prasinoxanthin]]; two equal or unequal flagella, or just one flagellum, or lacking any flagella. If flagella are present, they can be either smooth or covered in scales in the same manner as the cells. Present in marine and freshwater habitats.{{Sfn|Marin|Melkonian|2010}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=37}}
+* [[Nephroselmidophyceae]] (29 species):{{Sfn|Guiry|2024|p=5}} unicellular monadoids. Cells are covered by scales. They are capable of sexual reproduction through [[hologamy]] (fusion of entire cells), and of asexual reproduction through [[binary fission]]. Each cell contains a single chloroplast, a pyrenoid, and two flagella covered by scales. Present in marine and freshwater habitats.{{Sfn|Nakayama|Suda|Kawachi|Inouye|2007}}{{Sfn|Yamaguchi|Suda|Nakayama|Pienaar|2010}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=37}}
-* [[Nephroselmidophyceae]] (29 species):{{Sfn|Guiry|2024|p=5}} unicellular monadoids. Cells are covered by scales. They are capable of sexual reproduction through [[hologamy]] (fusion of entire cells), and of asexual reproduction through [[binary fission]]. Each cell contains a single cloroplast, a pyrenoid, and two flagella covered by scales. Present in marine and freshwater habitats.{{Sfn|Nakayama|Suda|Kawachi|Inouye|2007}}{{Sfn|Yamaguchi|Suda|Nakayama|Pienaar|2010}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=37}}
 * [[Pedinophyceae]] (24 species):{{Sfn|Guiry|2024|p=5}} unicellular asymmetrical monadoids that undergo a coccoid palmelloid phase covered by [[mucilage]]. Cells lack extracellular scales, but in rare cases are covered on the posterior side by a theca. Each cell contains a single chloroplast, a pyrenoid, and a single flagellum usually covered in mastigonemes. Present in marine, freshwater and terrestrial habitats.{{Sfn|Marin|2012}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=37}}{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=17-3}}
 * [[Picocystophyceae]] (1 species):{{Sfn|Guiry|2024|p=5}} unicellular coccoids, ovoid and trilobed in shape. Cells are surrounded by a multi-layered cell wall of poly-[[arabinose]], [[mannose]], [[galactose]] and [[glucose]]. No sexual reproduction has been described. They are capable of asexual reproduction through autosporulation, resulting in two or rarely four daughter cells. Each cell contains a single bilobed chloroplast with diatoxanthin and monadoxanthin, without any pyrenoid or flagella. Present in [[saline lake]]s.{{Sfn|Lewin|Krienitz|Goericke|Takeda|2000}}{{Sfn|Lopes dos Santos|Pollina|Gourvil|Corre|2017}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=37}}
 * [[Pyramimonadophyceae]] (166 species, 59 extinct):{{Sfn|Guiry|2024|p=5}} unicellular monadoids or coccoids. Cells are covered by two or more layers of organic scales. No sexual reproduction has been described, but some cells with only one flagellum have been interpreted as potential gametes. Asexual reproduction has only been observed in the coccoid forms, via zoospores. Each cell contains a single chloroplast, a pyrenoid, and between 4 and 16 flagella. The flagella are covered in at least two layers of organic scales: a bottom layer of pentagonal scales organized in 24 rows, and a top layer of limuloid scales distributed in 11 rows. They are exclusively marine.{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=37}}{{Sfn|Daugbjerg|Fassel|Moestrup|2020}}
 * [[Trebouxiophyceae]] (926 species, 1 extinct):{{Sfn|Guiry|2024|p=5}} unicellular monadoids occasionally without flagella, or colonial, or ramified filamentous thalli, or living as the [[photobiont]]s of [[lichen]]. Cells are covered by a cell wall of [[cellulose]], [[algaenan]]s, and β-galactofuranane. No sexual reproduction has been described with the exception of some observations of gamete fusion and presence of [[meiosis|meiotic]] [[gene]]s. They are capable of asexual reproduction through autospores or zoospores. Each cell contains a single chloroplast, a pyrenoid, and one or two pairs of smooth flagella. They are present in marine, freshwater and terrestrial habitats.{{Sfn|Domozych|Ciancia|Fangel|Mikkelsen|2012}}{{Sfn|Fučíková|Pažoutová|Rindi|2015}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=36}}{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=17-4–17-11}}
+* [[Ulvophyceae]] (2,695 species, 990 extinct):{{Sfn|Guiry|2024|p=5}} macroscopic thalli, either filamentous (which may be ramified) or foliose (composed of monostromatic or distromatic layers) or even compact tubular forms, generally [[multinucleate]]. Cells surrounded by a cell wall that may be [[Marine biogenic calcification|calcified]], composed of cellulose, β-manane, β-xilane, sulphated or piruvilated polysaccharides or sulphated ramnogalacturonanes, arabinogalactan proteins, and extensin. They exhibit a [[Biological life cycle#Haplodiplontic life cycle|haplodiplontic life cycle]] where the alternating generations can be isomorphic or heteromorphic. They reproduce asexually via zoospores that may be covered in scales. Each cell contains a single chloroplast, and one or two pairs of flagella without mastigonemes but covered in scales. They are present in marine, freshwater and terrestrial habitats.{{Sfn|Domozych|Ciancia|Fangel|Mikkelsen|2012}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=36}}{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-2–18-24}}
-* [[Ulvophyceae]] (2,695 species, 990 extinct):{{Sfn|Guiry|2024|p=5}} macroscopic thalli, either filamentous (which may be ramified) or foliose (composed of monostromatic or distromatic layers) or even compact tubular forms, generally [[multinucleate]]. Cells surrounded by a cell wall that may be [[Marine biogenic calcification|calcified]], composed of cellulose, β-manane, β-xilane, sulphated or piruvilated polysaccharides or sulphated ramnogalacturonanes, arabinogalactan proteins, and extensin. They exhibit a [[Biological_life_cycle#Haplodiplontic_life_cycle|haplodiplontic life cycle]] where the alternating generations can be isomorphic or heteromorphic. They reproduce asexually via zoospores that may be covered in scales. Each cell contains a single chloroplast, and one or two pairs of flagella without mastigonemes but covered in scales. They are present in marine, freshwater and terrestrial habitats.{{Sfn|Domozych|Ciancia|Fangel|Mikkelsen|2012}}{{Sfn|Adl|Bass|Lane|Lukeš|2019|p=36}}{{Sfn|Graham|Graham|Wilcox|Cook|2022|p=18-2–18-24}}
 == Evolution ==
-In February 2020, the fossilized remains of a green alga, named ''[[Proterocladus antiquus]]'' were discovered in the northern province of [[Liaoning]], [[China]]. At around a billion years old, it is believed to be one of the oldest examples of a multicellular chlorophyte. It is currently classified as a member of order [[Cladophorales|Siphonocladales]], class [[Ulvophyceae]].{{Sfn|Tang|Pang|Yuan|Xiao|2020}} In 2023, a study calculated the [[molecular clock|molecular age]] of green algae as [[Molecular_clock#Calibration|calibrated]] by this fossil. The study estimated the origin of Chlorophyta within the [[Mesoproterozoic]] era, at around 2.04–1.23 billion years ago.{{Sfn|Yang|Ma|Wang|Tian|2023}}
+In February 2020, the fossilized remains of a green alga, named ''[[Proterocladus antiquus]]'' were discovered in the northern province of [[Liaoning]], [[China]]. At around a billion years old, it is believed to be one of the oldest examples of a multicellular chlorophyte. It is currently classified as a member of order [[Cladophorales|Siphonocladales]], class [[Ulvophyceae]].{{Sfn|Tang|Pang|Yuan|Xiao|2020}} In 2023, a study calculated the [[molecular clock|molecular age]] of green algae as [[Molecular clock#Calibration|calibrated]] by this fossil. The study estimated the origin of Chlorophyta within the [[Mesoproterozoic]] era, at around 2.04–1.23 billion years ago.{{Sfn|Yang|Ma|Wang|Tian|2023}}
 == Usage ==
@@ Line 248: / Line 238: @@
   |display-authors=10
   |date=2019
-  |title=Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes |url=https://doi.org/10.1111%2FJEU.12691 |journal=Journal of Eukaryotic Microbiology
+  |title=Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes |journal=Journal of Eukaryotic Microbiology
   |volume=66 |issue=1 |pages=4–119 |doi=10.1111/JEU.12691 |pmc=6492006 |pmid=30257078}}
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@@ Line 256: / Line 246: @@
   |date=2018
   |title=Mixotrophy in Chlorophytes and Haptophytes—Effect of Irradiance, Macronutrient, Micronutrient and Vitamin Limitation |journal=Frontiers in Microbiology
-  |volume=9 |page=1704 |doi=10.3389/fmicb.2018.01704 |pmc=6080504 |pmid=30108563 |doi-access=free}}
+  |volume=9 |article-number=1704 |doi=10.3389/fmicb.2018.01704 |pmc=6080504 |pmid=30108563 |doi-access=free}}
 * {{Cite journal
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@@ Line 274: / Line 264: @@
 * {{Cite book
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-  |title=Introduction to the algae : structure and reproduction
+  |title=Introduction to the algae: structure and reproduction
   |last2=Wynne |first2=Michael James |publisher=Prentice-Hall
   |date=1985 |isbn=978-0-1347-7746-7 |edition=2nd |location=Englewood Cliffs, N.J.}}
@@ Line 306: / Line 296: @@
   |date=2012
   |title=The Cell Walls of Green Algae: A Journey through Evolution and Diversity |journal=Frontiers in Plant Science
-  |volume=3 |pages=82 |doi=10.3389/fpls.2012.00082 |pmc=3355577 |pmid=22639667 |doi-access=free}}
+  |volume=3 |page=82 |doi=10.3389/fpls.2012.00082 |pmc=3355577 |pmid=22639667 |bibcode=2012FrPS....3...82D
+ |doi-access=free}}
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@@ Line 318: / Line 309: @@
   |title=The unexpected extremophile: Tolerance to fluctuating salinity in the green alga ''Picochlorum'' |journal=Algal Research
   |volume=16 |pages=465–472 |doi=10.1016/j.algal.2016.04.003|bibcode=2016AlgRe..16..465F
+ |osti=1619391
   }}
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@@ Line 365: / Line 357: @@
   |date=1971
   |title=Parasitic green algae |journal=Annual Review of Phytopathology
-  |volume=9 |pages=45–64 |doi=10.1146/annurev.py.09.090171.000401}}
+  |volume=9 |issue=1
+ |pages=45–64 |doi=10.1146/annurev.py.09.090171.000401
+ |bibcode=1971AnRvP...9...45J
+ }}
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@@ Line 410: / Line 405: @@
   |date=2004
   |title=Green algae and the origin of land plants |journal=American Journal of Botany
-  |volume=91 |issue=10 |pages=1535–1556 |doi=10.3732/ajb.91.10.1535 |pmid=21652308 |doi-access=free}}
+  |volume=91 |issue=10 |pages=1535–1556 |doi=10.3732/ajb.91.10.1535 |pmid=21652308 |bibcode=2004AmJB...91.1535L
+ |doi-access=free}}
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@@ Line 465: / Line 461: @@
   |date=2017
   |title=Chloropicophyceae, a new class of picophytoplanktonic prasinophytes |journal=Sci Rep
-  |volume=7 |issue=1 |pages=14019 |bibcode=2017NatSR...714019L |doi=10.1038/s41598-017-12412-5 |pmc=5656628 |pmid=29070840}}
+  |volume=7 |issue=1 |page=14019 |bibcode=2017NatSR...714019L |doi=10.1038/s41598-017-12412-5 |pmc=5656628 |pmid=29070840}}
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@@ Line 563: / Line 559: @@
   |date=2002
   |title=Carbonaceous Megafossils from the Dholpura Shale, Uppermost Vindhyan Supergroup, Rajasthan: An Age Implication |url=http://palaeontologicalsociety.in/vol47/v9.pdf |journal=Journal of the Palaeontological Society of India
-  |volume=47 |pages=97–105|doi=10.1177/0971102320020109
+  |volume=47 |issue=1
+ |pages=97–105|doi=10.1177/0971102320020109
+ |bibcode=2002JPalS..47...97S
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@@ Line 573: / Line 571: @@
   |title=A one-billion-year-old multicellular chlorophyte |journal=Nature Ecology & Evolution
   |volume=4 |issue=4
-  |pages=543–549 |doi=10.1038/s41559-020-1122-9 |pmc=8668152|bibcode=2020NatEE...4..543T
+  |pages=543–549 |doi=10.1038/s41559-020-1122-9 |pmid=32094536
+ |pmc=8668152|bibcode=2020NatEE...4..543T
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@@ Line 587: / Line 586: @@
   |date=October 2014
   |title=Green algae and the origins of multicellularity in the plant kingdom |journal=Cold Spring Harbor Perspectives in Biology
-  |volume=6 |issue=11 |pages=a016170 |doi=10.1101/cshperspect.a016170 |pmc=4413236 |pmid=25324214}}
+  |volume=6 |issue=11 |article-number=a016170 |doi=10.1101/cshperspect.a016170 |pmc=4413236 |pmid=25324214}}
 * {{Cite book
   |last1=van den Hoek |first1=Christiaan |url=https://books.google.com/books?id=xuUoiFesSHMC
@@ Line 609: / Line 608: @@
   |date=2023
   |title=Algal ocelloids and plant ocelli |journal=Plants
-  |volume=12 |pages=61 |doi=10.3390/plants12010061 |doi-access=free|pmc=9824129 }}
+  |volume=12 |issue=1
+ |page=61 |doi=10.3390/plants12010061 |pmid=36616190
+ |doi-access=free|pmc=9824129 }}
 * {{Cite journal
   |last1=Yang |first1=Zhiping
@@ Line 623: / Line 624: @@
   |date=September 2023
   |title=Phylotranscriptomics unveil a Paleoproterozoic-Mesoproterozoic origin and deep relationships of the Viridiplantae |journal=Nature Communications
-  |volume=14 |issue=1 |pages=5542 |doi=10.1038/s41467-023-41137-5 |pmc=10495350 |pmid=37696791 |doi-access=free|bibcode=2023NatCo..14.5542Y
+  |volume=14 |issue=1 |page=5542 |doi=10.1038/s41467-023-41137-5 |pmc=10495350 |pmid=37696791 |doi-access=free|bibcode=2023NatCo..14.5542Y
   }}
 {{Refend}}
@@ Line 632: / Line 633: @@
 {{Refbegin}}
 * {{Cite book |title=Seaweeds of the British Isles |vauthors=Burrows EM |date=1991 |publisher=Natural History Museum |isbn=978-0-5650-0981-6 |volume=2 (Chlorophyta) |location=London}}
-* {{Cite book |title=Green Algae. Structure, Reproduction and Evolution in Selected Genera |vauthors=Pickett-Heaps JD |date=1975 |publisher=Sinauer Assoc. |location=Stamford, CT |pages=606}}
+* {{Cite book |title=Green Algae. Structure, Reproduction and Evolution in Selected Genera |vauthors=Pickett-Heaps JD |date=1975 |publisher=Sinauer Assoc. |location=Stamford, CT |page=606}}
 {{Refend}}
@@ Line 641: / Line 642: @@
 [[Category:Chlorophyta|Chlorophyta]]
-[[Category:Plant divisions]]
 [[Category:Taxa named by Ludwig Reichenbach]]
 [[Category:Green algae phyla]]

Chlorophyta: Difference between revisions

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