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{{Short description|Genus of euglenoids}}
{{Italic title}}
{{Italic title}}
{{Automatic taxobox
{{Taxobox
| name = ''Trachelomonas''
| taxon = Trachelomonas
| image = Trachelomonas_sp.jpg
| image = Trachelomonas_sp.jpg
⚫
| authority = [[Christian Gottfried Ehrenberg|Ehrenberg]], 1833<ref name=EOL/>
| domain = [[Eukaryote|Eukaryota]]
| unranked_regnum = [[Excavata]]
⚫
| phylum = [[Euglenozoa]]
| classis = [[Euglenid|Euglenoidea]]
| ordo = [[Euglenales]]
| familia = [[Euglenaceae]]
| genus = '''''Trachelomonas'''''
⚫
| genus_authority = [[Christian Gottfried Ehrenberg|Ehrenberg]], 1833<ref name=EOL/>
}}
}}
⚫
'''''Trachelomonas''''' is a genus of swimming, free-living [[euglenoid]]s characterized by the presence of a shell-like covering called a [[Lorica (biology)|lorica]].<ref name=EOL>{{cite web|url=http://eol.org/pages/11712/overview |accessdate=28 August 2018 |title=Trachelomonas |website=Encyclopedia of Life}}</ref> Details of lorica structure determine the classification of distinct species in the genus.<ref name=EOL/> The lorica can exist in spherical, elliptical, cylindrical, and pyriform (pear-shaped) forms. The lorica surface can be smooth, punctuate or striate and range from hyaline, to yellow, or brown. These colors are due to the accumulation of [[ferric hydroxide]] and [[manganic oxide]] deposited with the mucilage and minerals that comprise the lorica. In ''Trachelomonas'', the presence of a lorica obscures cytoplasmic details of the underlying cell. In each ''Trachelomonas'' cell, there is a gap at the apex of the lorica from which the flagellum protrudes. Thickening around this gap results in a rim-like or collar-like appearance. During asexual reproduction, the nucleus divides yielding two daughter cells one of which exits through the opening in the lorica. This new cell then synthesizes its own new lorica.


⚫
'''''Trachelomonas''''' is a genus of swimming, free-living [[euglenoid]]s characterized by the presence of a shell-like covering called a [[Lorica (biology)|lorica]].<ref name=EOL>{{cite web|url=http://eol.org/pages/11712/overview |access-date=28 August 2018 |title=Trachelomonas |website=Encyclopedia of Life}}</ref> Details of lorica structure determine the classification of distinct species in the genus.<ref name=EOL/> The lorica can exist in spherical, elliptical, cylindrical, and pyriform (pear-shaped) forms. The lorica surface can be smooth, punctuate or striate and range from hyaline, to yellow, or brown. These colors are due to the accumulation of [[ferric hydroxide]] and [[manganic oxide]] deposited with the mucilage and minerals that comprise the lorica. In ''Trachelomonas'', the presence of a lorica obscures cytoplasmic details of the underlying cell. In each ''Trachelomonas'' cell, there is a gap at the apex of the lorica from which the [[flagellum]] protrudes. Thickening around this gap results in a rim-like or collar-like appearance. During [[asexual reproduction]], the [[cell nucleus|nucleus]] divides yielding two daughter cells one of which exits through the opening in the lorica. This new cell then synthesizes its own new lorica.
==History of Knowledge==
⚫
''Trachelomonas'' was first described by C. G. Ehrenberg in 1834.<ref name="Guiry">Guiry, M. D.; Guiry, G. M. (2012). “Trachelomonas Ehrenberg, 1834”. Retrieved March 5, 2019, from [http://www.algaebase.org/search/genus/detail/?genus_id=43657& session=abv4:AC1F11E20766f396C8RX21DDBA4A]</ref> Its separation from the genus ''Strombomonas'' occurred in 2008 with the discovery of five subclades within ''Trachelomonas'' through nuclear SSU and LSU rDNA analyses.<ref name="Ciugulea">Ciugulea, Ionel; Nudelman, María A.; Brosnan, Stacy; Triemer, Richard E. (2008). “Phylogeny of the euglenoid loricate genera Trachelomonas and Strombomonas (Euglenophyta) inferred from nuclear SSU and LSU rDNA”. Journal of Phycology. 44 (2): 406-418. [https://doi.org/10.1111/j.1529-8817.2008.00472.x doi: 10.1111/j.1529-8817.2008.00472.x]</ref>


==Habitat and Ecology==
==History of knowledge==
⚫
''Trachelomonas'' was first described by [[C. G. Ehrenberg]] in 1834.<ref name="Guiry">Guiry, M. D.; Guiry, G. M. (2012). “Trachelomonas Ehrenberg, 1834”. Retrieved March 5, 2019, from [http://www.algaebase.org/search/genus/detail/?genus_id=43657& session=abv4:AC1F11E20766f396C8RX21DDBA4A]</ref> The name comes from the Greek roots ''trachelos'', meaning "neck" (referring to the neck-like opening of the lorica), and ''monas'', meaning "unit", now used conventionally for unicellular flagellates.<ref name=CarolinaLucidKey>{{cite web|url=https://fmp.conncoll.edu/Silicasecchidisk/LucidKeys3.5/Keys_v3.5/Carolina35_Key/Media/Html/Trachelomonas_Main.html|title=Trachelomonas|first1= Hannah|last1= Shayler|first2=Peter A.|last2=Siver|work=Carolina Lucid Key to Freshwater Algae|publisher=Connecticut College | date=2004 |access-date=2024-11-28 }}</ref> Its separation from the genus ''[[Strombomonas]]'' occurred in 2008 with the discovery of five subclades within ''Trachelomonas'' through nuclear SSU and LSU rDNA analyses.<ref name="Ciugulea">Ciugulea, Ionel; Nudelman, María A.; Brosnan, Stacy; Triemer, Richard E. (2008). “Phylogeny of the euglenoid loricate genera Trachelomonas and Strombomonas (Euglenophyta) inferred from nuclear SSU and LSU rDNA”. Journal of Phycology. 44 (2): 406-418. [https://doi.org/10.1111/j.1529-8817.2008.00472.x doi: 10.1111/j.1529-8817.2008.00472.x]</ref>
⚫
''Trachelomonas'' is a common, cosmopolitan genus found in acidic to neutral [[fresh water]] (pH 4.5-7), often in habitats rich in iron and manganese, and pools rich in [[organic matter]] such as peat.<ref name="Guiry" /> These euglenoids have also been observed to prefer warm, eutrophic waters, increasing in abundance during [[harmful algal blooms]] of ''Planktothrix agardhii''.<ref>Grabowksa, M.; Wołowski, K. (2013). “Development of Trachelomonas species (Euglenophyta) during blooming of Planktothrix agardhii (Cyanoprokaryota)”. International Journal of Limnology. 50: 49-57. [https://doi.org/10.1051/limn/2013070 doi: 10.1051/limn/2013070]</ref> Most species are photosynthetic; therefore, contributing to global [[primary production]] and some species have been observed to be [[osmotrophy|osmotrophs]], having the ability to assimilate nutrients from its environment.<ref name="Guiry" />

==Habitat and ecology==
⚫
''Trachelomonas'' is a common, cosmopolitan genus found in acidic to neutral [[fresh water]] (pH 4.5-7), often in habitats rich in iron and manganese, and pools rich in [[organic matter]] such as peat.<ref name="Guiry" /> These euglenoids have also been observed to prefer warm, eutrophic waters, increasing in abundance during [[harmful algal blooms]] of ''Planktothrix agardhii''.<ref>Grabowksa, M.; Wołowski, K. (2013). “Development of Trachelomonas species (Euglenophyta) during blooming of Planktothrix agardhii (Cyanoprokaryota)”. International Journal of Limnology. 50: 49-57. [https://doi.org/10.1051/limn/2013070 doi: 10.1051/limn/2013070]</ref> Most species are photosynthetic; therefore, contributing to global [[primary production]] and some species have been observed to be [[osmotrophy|osmotrophs]], having the ability to assimilate nutrients from its environment.<ref name="Guiry" />


==Description==
==Description==
Trachelomonads are free-swimming, solitary, photosynthetic flagellates ranging in size from 5-100&nbsp;um, with an ovoid shape, sharing similar morphological characteristics with its sister group, ''Strombomonas''.<ref name="Guiry" /><ref>Brosnan, Stacy; Brown, Patrick J.; Farmer, Mark A.; Triemer, Richard E. (2005). “Morphological separation of the euglenoid genera Trachelomonas and Strombomonas (Euglenophyta) based on lorica development and posterior strip deduction”. Journal of Phycology. 41 (3): 590-605. [https://doi.org/10.1111/j.1529-8817.2005.00068.x doi: 10.1111/j.1529-8817.2005.00068.x]</ref> These cells are enclosed in a rigid, shell-like envelope, made up of minerals and [[polysaccharide]] mucilage, with a defined collar or truncate extension that surrounds an [[anterior]] apical pore where the flagellum emerges from, also known as a lorica.<ref name="Guiry" /><ref>Juráň, Josef (2016). “Trachelomonas bituricensis var. lotharingia M.L. Poucques 1952, a morphologically interesting, rare euglenoid new to the algal flora of the Czech Republic”. PhytoKeys. 61: 81-91. [https://doi.org/10.3897/phytokeys.61.7408 doi: 10.3897/phytokeys.61.7408]</ref> The lorica can be distinguished between different species by the orientation of spines or other ornamentations, such as pores, warts or ridges, and can range from being colourless to orange/brown or even black based on the nutrients in their surroundings.<ref name="Ehrenberg">[http://fmp.conncoll.edu/Silicasecchidisk/LucidKeys3.5/Keys_v3.5/Carolina35_Key/Media/Html/Trachelomonas_Main.html Trachelomonas Ehrenberg. (n.d.). Retrieved March 5, 2019, from]</ref><ref name="Guiry" />
Trachelomonads are free-swimming, solitary, photosynthetic flagellates ranging in size from 5-100&nbsp;um, with an ovoid shape, sharing similar morphological characteristics with its sister group, ''Strombomonas''.<ref name="Guiry" /><ref>Brosnan, Stacy; Brown, Patrick J.; Farmer, Mark A.; Triemer, Richard E. (2005). “Morphological separation of the euglenoid genera Trachelomonas and Strombomonas (Euglenophyta) based on lorica development and posterior strip deduction”. Journal of Phycology. 41 (3): 590-605. [https://doi.org/10.1111/j.1529-8817.2005.00068.x doi: 10.1111/j.1529-8817.2005.00068.x]</ref> These cells are enclosed in a rigid, shell-like envelope, made up of minerals and [[polysaccharide]] mucilage, with a defined collar or truncate extension that surrounds an [[anterior]] apical pore where the flagellum emerges from, also known as a lorica.<ref name="Guiry" /><ref>Juráň, Josef (2016). “Trachelomonas bituricensis var. lotharingia M.L. Poucques 1952, a morphologically interesting, rare euglenoid new to the algal flora of the Czech Republic”. PhytoKeys. 61: 81-91. [https://doi.org/10.3897/phytokeys.61.7408 doi: 10.3897/phytokeys.61.7408]</ref> The lorica can be distinguished between different species by the orientation of spines or other ornamentations, such as pores, warts or ridges, and can range from being colourless to orange/brown or even black based on the nutrients in their surroundings.<ref name="Ehrenberg">[http://fmp.conncoll.edu/Silicasecchidisk/LucidKeys3.5/Keys_v3.5/Carolina35_Key/Media/Html/Trachelomonas_Main.html Trachelomonas Ehrenberg. (n.d.). Retrieved March 5, 2019, from]</ref><ref name="Guiry" />

Most species are phototrophic, having a characteristic green colour due to the discoid or flattened, shield-like [[chloroplast]], which usually bears sheathed, projecting or naked [[pyrenoid]]s.<ref name="Guiry" /><ref name="Ehrenberg" /> The few species that are osmotrophic, lack chloroplasts; therefore, they are colourless.<ref name="Guiry" /> Similar to other euglenoids, the cell has many [[paramylon]] bodies that are used for the storage of [[starch]]; these can be a distinguishing trait for species with similar lorica structures.<ref name="Ehrenberg" /> The structure and ornamentation of the lorica is very dependent on the growth conditions, especially the availability of nutrients. Therefore, the size, shape, collar form and the presence of spines and pores can vary, showing morphological plasticity within species.<ref name="Leedale">Leedale, Gordon F. (2007). “Envelope formation and structure in the Euglenoid genus Trachelomonas”. British Phycological Journal. 10 (1):17-41. [https://doi.org/10.1080/00071617500650031 doi: 10.1080/00071617500650031]</ref> This can make it difficult to describe species since morphological features can vary greatly. Trachelomonads also have an [[Eyespot apparatus|eyespot]], a feature of photosynthetic euglenoids, located outside the chloroplast with orange to red pigmentation.<ref name="Ehrenberg" /> These cells also have one long emergent flagellum that has previously been identified to emerge from the apical pore, and a shorter flagellum that is within the furrow and not used for motility. Under light microscopy, it is also possible to see condensed [[chromosome]]s.<ref name="Ehrenberg" />
Most species are phototrophic, having a characteristic green colour due to the discoid or flattened, shield-like [[chloroplast]], which usually bears sheathed, projecting or naked [[pyrenoid]]s.<ref name="Guiry" /><ref name="Ehrenberg" /> The few species that are osmotrophic, lack chloroplasts; therefore, they are colourless.<ref name="Guiry" /> Similar to other euglenoids, the cell has many [[paramylon]] bodies that are used for the storage of [[starch]]; these can be a distinguishing trait for species with similar lorica structures.<ref name="Ehrenberg" /> The structure and ornamentation of the lorica is very dependent on the growth conditions, especially the availability of nutrients. Therefore, the size, shape, collar form and the presence of spines and pores can vary, showing morphological plasticity within species.<ref name="Leedale">Leedale, Gordon F. (2007). “Envelope formation and structure in the Euglenoid genus Trachelomonas”. British Phycological Journal. 10 (1):17-41. [https://doi.org/10.1080/00071617500650031 doi: 10.1080/00071617500650031]</ref> This can make it difficult to describe species since morphological features can vary greatly. Trachelomonads also have an [[Eyespot apparatus|eyespot]], a feature of photosynthetic euglenoids, located outside the chloroplast with orange to red pigmentation.<ref name="Ehrenberg" /> These cells also have one long emergent flagellum that has previously been identified to emerge from the apical pore, and a shorter flagellum that is within the furrow and not used for motility. Under light microscopy, it is also possible to see condensed [[chromosome]]s.<ref name="Ehrenberg" />


==Life History==
==Life history==
Euglenoids have not been observed to undergo [[sexual reproduction]]; however, [[asexual reproduction]] does occur through [[mitosis]] followed by [[cytokinesis]].<ref>Esson, H. J.; Leander, B. S. (2006). “A model for the morphogenesis of strip reduction patterns in phototrophic euglenids: Evidence for heterochrony in pellicle evolution”. Evolution Development, 8 (4): 378-388. [https://doi.org/10.1111/j.1525-142x.2006.00110.x doi:10.1111/j.1525-142x.2006.00110.x]</ref> The formation of the lorica after asexual reproduction first occurs through the external skin and then a fibrillar layer is formed between the cell surface and the skin.<ref name="Leedale" /> Then manganese and ferric hydroxide compounds are precipitated on the inner fibrillar layer to produce a thick envelope and the original external skin is lost.<ref name="Leedale" /> However, differences in these processes exist among species.
Euglenoids have not been observed to undergo [[sexual reproduction]]; however, [[asexual reproduction]] does occur through [[mitosis]] followed by [[cytokinesis]].<ref>Esson, H. J.; Leander, B. S. (2006). “A model for the morphogenesis of strip reduction patterns in phototrophic euglenids: Evidence for heterochrony in pellicle evolution”. Evolution Development, 8 (4): 378-388. [https://doi.org/10.1111/j.1525-142x.2006.00110.x doi:10.1111/j.1525-142x.2006.00110.x]</ref> The formation of the lorica after asexual reproduction first occurs through the external skin and then a fibrillar layer is formed between the cell surface and the skin.<ref name="Leedale" /> Then manganese and ferric hydroxide compounds are precipitated on the inner fibrillar layer to produce a thick envelope and the original external skin is lost.<ref name="Leedale" /> However, differences in these processes exist among species.


==List of Species==
==Species==
A list of species in ''Trachelomonas'' (incomplete):
* ''T. acanthopohora'' Stokes
* ''T. americana'' Lemmermann
* ''T. acanthopohora'' <small>Stokes</small>
* ''T. argentina'' Frenguelli
* ''T. americana'' <small>Lemmermann</small>
* ''T. bernarddii'' Woloszynka
* ''T. argentina'' <small>Frenguelli</small>
* ''T. bituricensis'' Wurtz
* ''T. bernarddii'' <small>Woloszynka</small>
* ''T. bituricensis'' <small>Wurtz</small>
* [[Trachelomonas cervicula|''T. cervicula'']] Stokes
* [[Trachelomonas cervicula|''T. cervicula'']] <small>Stokes</small>
* ''T. foliata'' Skvortzov
* ''T. grandis'' K.P. Singh
* ''T. foliata'' <small>Skvortzov</small>
* ''T. volvocina'' (Ehrenberg) Ehrenberg
* ''T. grandis'' <small>K.P. Singh</small>
* ''T. hispida'' <small>(Perty) F.Stein</small>
* ''T. robusta'' <small>Svirenko</small>
* ''T. volvocina'' <small>(Ehrenberg) Ehrenberg</small>
* ''T. volvocinopsis'' <small>Svirenko</small>


== References ==
== References ==
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{{Taxonbar|from=Q146181}}
{{Taxonbar|from=Q146181}}
{{Authority control}}


[[Category:Euglenozoa genera]]
[[Category:Euglenozoa genera]]
[[Category:Taxa named by Christian Gottfried Ehrenberg]]
⚫
[[Category:Euglenozoa]]

Latest revision as of 04:34, 29 November 2024

Trachelomonas
Scientific classification Edit this classification
Domain: Eukaryota
Phylum: Euglenozoa
Class: Euglenida
Clade: Euglenophyceae
Order: Euglenales
Family: Euglenaceae
Genus: Trachelomonas
Ehrenberg, 1833[1]

Trachelomonas is a genus of swimming, free-living euglenoids characterized by the presence of a shell-like covering called a lorica.[1] Details of lorica structure determine the classification of distinct species in the genus.[1] The lorica can exist in spherical, elliptical, cylindrical, and pyriform (pear-shaped) forms. The lorica surface can be smooth, punctuate or striate and range from hyaline, to yellow, or brown. These colors are due to the accumulation of ferric hydroxide and manganic oxide deposited with the mucilage and minerals that comprise the lorica. In Trachelomonas, the presence of a lorica obscures cytoplasmic details of the underlying cell. In each Trachelomonas cell, there is a gap at the apex of the lorica from which the flagellum protrudes. Thickening around this gap results in a rim-like or collar-like appearance. During asexual reproduction, the nucleus divides yielding two daughter cells one of which exits through the opening in the lorica. This new cell then synthesizes its own new lorica.

History of knowledge

[edit]

Trachelomonas was first described by C. G. Ehrenberg in 1834.[2] The name comes from the Greek roots trachelos, meaning "neck" (referring to the neck-like opening of the lorica), and monas, meaning "unit", now used conventionally for unicellular flagellates.[3] Its separation from the genus Strombomonas occurred in 2008 with the discovery of five subclades within Trachelomonas through nuclear SSU and LSU rDNA analyses.[4]

Habitat and ecology

[edit]

Trachelomonas is a common, cosmopolitan genus found in acidic to neutral fresh water (pH 4.5-7), often in habitats rich in iron and manganese, and pools rich in organic matter such as peat.[2] These euglenoids have also been observed to prefer warm, eutrophic waters, increasing in abundance during harmful algal blooms of Planktothrix agardhii.[5] Most species are photosynthetic; therefore, contributing to global primary production and some species have been observed to be osmotrophs, having the ability to assimilate nutrients from its environment.[2]

Description

[edit]

Trachelomonads are free-swimming, solitary, photosynthetic flagellates ranging in size from 5-100 um, with an ovoid shape, sharing similar morphological characteristics with its sister group, Strombomonas.[2][6] These cells are enclosed in a rigid, shell-like envelope, made up of minerals and polysaccharide mucilage, with a defined collar or truncate extension that surrounds an anterior apical pore where the flagellum emerges from, also known as a lorica.[2][7] The lorica can be distinguished between different species by the orientation of spines or other ornamentations, such as pores, warts or ridges, and can range from being colourless to orange/brown or even black based on the nutrients in their surroundings.[8][2]

Most species are phototrophic, having a characteristic green colour due to the discoid or flattened, shield-like chloroplast, which usually bears sheathed, projecting or naked pyrenoids.[2][8] The few species that are osmotrophic, lack chloroplasts; therefore, they are colourless.[2] Similar to other euglenoids, the cell has many paramylon bodies that are used for the storage of starch; these can be a distinguishing trait for species with similar lorica structures.[8] The structure and ornamentation of the lorica is very dependent on the growth conditions, especially the availability of nutrients. Therefore, the size, shape, collar form and the presence of spines and pores can vary, showing morphological plasticity within species.[9] This can make it difficult to describe species since morphological features can vary greatly. Trachelomonads also have an eyespot, a feature of photosynthetic euglenoids, located outside the chloroplast with orange to red pigmentation.[8] These cells also have one long emergent flagellum that has previously been identified to emerge from the apical pore, and a shorter flagellum that is within the furrow and not used for motility. Under light microscopy, it is also possible to see condensed chromosomes.[8]

Life history

[edit]

Euglenoids have not been observed to undergo sexual reproduction; however, asexual reproduction does occur through mitosis followed by cytokinesis.[10] The formation of the lorica after asexual reproduction first occurs through the external skin and then a fibrillar layer is formed between the cell surface and the skin.[9] Then manganese and ferric hydroxide compounds are precipitated on the inner fibrillar layer to produce a thick envelope and the original external skin is lost.[9] However, differences in these processes exist among species.

Species

[edit]

A list of species in Trachelomonas (incomplete):

  • T. acanthopohora Stokes
  • T. americana Lemmermann
  • T. argentina Frenguelli
  • T. bernarddii Woloszynka
  • T. bituricensis Wurtz
  • T. cervicula Stokes
  • T. foliata Skvortzov
  • T. grandis K.P. Singh
  • T. hispida (Perty) F.Stein
  • T. robusta Svirenko
  • T. volvocina (Ehrenberg) Ehrenberg
  • T. volvocinopsis Svirenko

References

[edit]
  1. ^ a b c "Trachelomonas". Encyclopedia of Life. Retrieved 28 August 2018.
  2. ^ a b c d e f g h Guiry, M. D.; Guiry, G. M. (2012). “Trachelomonas Ehrenberg, 1834”. Retrieved March 5, 2019, from session=abv4:AC1F11E20766f396C8RX21DDBA4A
  3. ^ Shayler, Hannah; Siver, Peter A. (2004). "Trachelomonas". Carolina Lucid Key to Freshwater Algae. Connecticut College. Retrieved 2024-11-28.
  4. ^ Ciugulea, Ionel; Nudelman, María A.; Brosnan, Stacy; Triemer, Richard E. (2008). “Phylogeny of the euglenoid loricate genera Trachelomonas and Strombomonas (Euglenophyta) inferred from nuclear SSU and LSU rDNA”. Journal of Phycology. 44 (2): 406-418. doi: 10.1111/j.1529-8817.2008.00472.x
  5. ^ Grabowksa, M.; Wołowski, K. (2013). “Development of Trachelomonas species (Euglenophyta) during blooming of Planktothrix agardhii (Cyanoprokaryota)”. International Journal of Limnology. 50: 49-57. doi: 10.1051/limn/2013070
  6. ^ Brosnan, Stacy; Brown, Patrick J.; Farmer, Mark A.; Triemer, Richard E. (2005). “Morphological separation of the euglenoid genera Trachelomonas and Strombomonas (Euglenophyta) based on lorica development and posterior strip deduction”. Journal of Phycology. 41 (3): 590-605. doi: 10.1111/j.1529-8817.2005.00068.x
  7. ^ Juráň, Josef (2016). “Trachelomonas bituricensis var. lotharingia M.L. Poucques 1952, a morphologically interesting, rare euglenoid new to the algal flora of the Czech Republic”. PhytoKeys. 61: 81-91. doi: 10.3897/phytokeys.61.7408
  8. ^ a b c d e Trachelomonas Ehrenberg. (n.d.). Retrieved March 5, 2019, from
  9. ^ a b c Leedale, Gordon F. (2007). “Envelope formation and structure in the Euglenoid genus Trachelomonas”. British Phycological Journal. 10 (1):17-41. doi: 10.1080/00071617500650031
  10. ^ Esson, H. J.; Leander, B. S. (2006). “A model for the morphogenesis of strip reduction patterns in phototrophic euglenids: Evidence for heterochrony in pellicle evolution”. Evolution Development, 8 (4): 378-388. doi:10.1111/j.1525-142x.2006.00110.x
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