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#REDIRECT [[Cryptomonad]]
{{Taxobox
| classification_status = [[incertae sedis]] within [[Eukaryota]]
| name = Cryptophytes
| image = Rhodomonas_salina_CCMP_322.jpg
| image_caption = ''Rhodomonas salina''
| domain = [[Eukaryota]]
| unranked_subregnum = [[Hacrobia]]
| unranked_phylum = [[Cryptista]]
| superclassis = [[Cryptomonada]] <br/><small>[[Thomas Cavalier-Smith|Cavalier-Smith]], 2004</small>
| class = '''Cryptophyceae'''
| subdivision_ranks = Orders
| subdivision =
* [[Tetragonidiales]]
* [[Pyrenomonadales]]
* [[Cryptomonadales]]
| synonyms =
* Cryptomonada <small>Senn, 1900</small>
* Cryptomonadinae <small>Pascher, 1913</small>
* Cryptomonadophyceae <small>Pascher ex Schoenichem, 1925</small>
}}


The '''cryptophyceae''' are a class of [[algae]],<ref name="pmid18397952">{{cite journal |author=Khan H, Archibald JM |title=Lateral transfer of introns in the cryptophyte plastid genome |journal=Nucleic Acids Res. |volume=36 |issue=9 |pages=3043–53 |date=May 2008 |pmid=18397952 |pmc=2396441 |doi=10.1093/nar/gkn095 |url=http://nar.oxfordjournals.org/cgi/pmidlookup?view=long&pmid=18397952}}</ref> most of which have [[chloroplast|plastids]]. They are common in freshwater, and also occur in marine and brackish habitats. Each cell is around [[1 E-5 m|10–50 μm]] in size and flattened in shape, with an anterior groove or pocket. At the edge of the pocket there are typically two slightly unequal [[flagellum|flagella]].
{{R taxon with possibilities}}


Some exhibit [[mixotrophy]].<ref name="urlCryptophyta - the cryptomonads">{{cite web|url=http://www.life.umd.edu/labs/delwiche/PSlife/lectures/Cryptophyta.html/wwwtax.cgi?mode=Info&id=412159&lvl=1|title=Cryptophyta - the cryptomonads|format=|website=|accessdate=2009-06-02|deadurl=yes|archiveurl=https://web.archive.org/web/20110610151135/http://www.life.umd.edu/labs/delwiche/PSlife/lectures/Cryptophyta.html/wwwtax.cgi?mode=Info&id=412159&lvl=1|archivedate=2011-06-10|df=}}</ref>
[[Category:Bikont classes]]

==Characteristics==
[[File:Cryptophyta_cell_scheme.svg|thumb|left|150px|Cell scheme: 1-[[vacuole|contractile vacuole]], 2-[[plastid]], 3-[[thylakoid]], 4-[[Eyespot apparatus|stigma]], 5-[[nucleomorph]], 6-[[starch]] granule, 7-[[70S ribosome]], 8-[[cell nucleus|nucleus]], 9-[[Eukaryotic ribosome (80S)|80S ribosome]], 10-[[flagella]], 11-invagination, 12-[[lipid]] globules, 13-[[ejectosome]]s, 14-[[mitochondrion]], 15-[[pyrenoid]], 16-[[Golgi apparatus]], 17-[[endoplasmic reticulum]], 18-[[endoplasmic reticulum|chloroplast-endoplasmic reticulum]].]]

Cryptophytes are distinguished by the presence of characteristic [[extrusome]]s called [[ejectosome]]s or ejectisomes, which consist of two connected spiral ribbons held under tension.<ref name="Graham">{{cite book |last=Graham |first=L. E. |last2=Graham |first2=J. M. |last3=Wilcox |first3=L. W. |year=2009 |title=Algae |edition=2nd |publisher=Benjamin Cummings (Pearson) |location=San Francisco, CA |isbn=9780321559654 }}</ref> If the cells are irritated either by mechanical, chemical or light stress, they discharge, propelling the cell in a zig-zag course away from the disturbance. Large ejectosomes, visible under the light microscope, are associated with the pocket; smaller ones occur underneath the [[periplast]], the cryptophyte-specific cell surrounding.<ref name="MorrallGreenwood">{{cite journal |last=Morrall |first=S. |last2=Greenwood |first2=A. D. |year=1980 |title=A comparison of the periodic sub-structures of the trichocysts of the Cryptophyceae and Prasinophyceae |journal=BioSystems |volume=12 |issue= 1–2|pages=71–83 |doi= 10.1016/0303-2647(80)90039-8|pmid=6155157 }}</ref><ref name="GrimStaehelin">{{cite journal |last=Grim |first=J. N. |last2=Staehelin |first2=L. A. |year=1984 |title=The ejectisomes of the flagellate ''Chilomonas paramecium'' - Visualization by freeze-fracture and isolation techniques |journal=[[Journal of Protozoology]] |volume=31 |issue=2 |pages=259–267 |doi=10.1111/j.1550-7408.1984.tb02957.x }}</ref>

Except for ''[[Chilomonas]]'', which has [[leucoplast]]s, cryptophytes have one or two chloroplasts. These contain [[chlorophyll]]s ''a'' and ''c'', together with [[phycobiliprotein]]s and other pigments, and vary in color (brown, red to blueish-green). Each is surrounded by four membranes, and there is a reduced [[cell nucleus]] called a [[nucleomorph]] between the middle two. This indicates that the plastid was derived from a [[eukaryote|eukaryotic]] symbiont, shown by genetic studies to have been a [[red alga]].<ref name="Douglas2002">{{cite journal |last=Douglas |first=S. |year=2002 |title=The highly reduced genome of an enslaved algal nucleus |journal=[[Nature (journal)|Nature]] |volume=410 |issue=6832 |pages=1091–1096 |doi=10.1038/35074092 |pmid=11323671|display-authors=etal}}</ref> However, the plastids are very different from red algal plastids: phycobiliproteins are present but only in the thylakoid lumen and are present only as phycoerythrin or [[phycocyanin]]. In the case of "Rhodomonas" the crystal structure has been determined to 1.63Å;<ref name="Wilk1999">{{cite journal |last=Wilk |first=K. |year=1999 |title=Evolution of a light-harvesting protein by addition of new subunits and rearrangement of conserved elements: Crystal structure of a cryptophyte phycoerythrin at 1.63Å resolution. |journal=PNAS |volume=96 |issue=16 |pages=8901–8906 |doi=10.1073/pnas.96.16.8901|pmid=10430868 |display-authors=etal|pmc=17705 }}</ref> and it has been shown that the alpha subunit bears no relation to any other known phycobiliprotein.

A few cryptophytes, such as ''[[Cryptomonas]]'', can form palmelloid stages, but readily escape the surrounding mucus to become free-living flagellates again. Some ''[[Cryptomonas]]'' species may also form immotile [[microbial cyst]]s–resting stages with rigid cell walls to survive unfavorable conditions. Cryptophyte flagella are inserted parallel to one another, and are covered by bipartite hairs called [[mastigonemes]], formed within the [[endoplasmic reticulum]] and transported to the cell surface. Small scales may also be present on the flagella and cell body. The [[mitochondrion|mitochondria]] have flat [[crista]]e, and [[mitosis]] is open; [[sexual reproduction]] has also been reported.

==Classification==
[[File:CSIRO ScienceImage 6743 SEM Cryptophyte.jpg|thumb|Cryptophytes under [[Scanning electron microscope|SEM]]]]
[[File:CSIRO ScienceImage 7234 microalgal cultures.jpg|thumb|Cryptophytes under [[light microscope]] ]]
{{further|Wikispecies:Cryptophyceae}}

The first mention of cryptophytes appears to have been made by [[Christian Gottfried Ehrenberg]] in 1831,<ref>{{cite journal |last=Novarino |first=G. |date=2012 |title=Cryptomonad taxonomy in the 21st century: The first 200 years |journal=Phycological Reports: Current Advances in Algal Taxonomy and its Applications: Phylogenetic, Ecological and Applied Perspective |pp=19–52 |url= https://www.researchgate.net/publication/265520014 |access-date=2018-10-16}}</ref> while studying [[Infusoria]]. Later, botanists treated them as a separate [[algae]] group, class Cryptophyceae or division Cryptophyta, while zoologists treated them as the [[flagellate]] [[protozoa]] order Cryptomonadina. In some classifications, the cryptomonads were considered close relatives of the [[dinoflagellate]]s because of their (seemingly) similar pigmentation, being grouped as the [[Pyrrhophyta]]. There is considerable evidence that cryptophyte chloroplasts are closely related to those of the [[heterokont]]s and [[haptophyte]]s, and the three groups are sometimes united as the [[Chromista]]. However, the case that the organisms themselves are closely related is not very strong, and they may have acquired plastids independently. Currently they are discussed to be members of the clade [[Diaphoretickes]] and to form together with the [[Haptophyta]] the group [[Hacrobia]]. Parfrey et al. and Burki et al. placed Cryptophyceae as a sister clade to the [[Green algae|Green Algae]].<ref>{{Cite journal|title = Estimating the timing of early eukaryotic diversification with multigene molecular clocks|journal = Proceedings of the National Academy of Sciences|date = 2011-08-16|issn = 0027-8424|pmc = 3158185|pmid = 21810989|pages = 13624–13629|volume = 108|issue = 33|doi = 10.1073/pnas.1110633108|first = Laura Wegener|last = Parfrey|first2 = Daniel J. G.|last2 = Lahr|first3 = Andrew H.|last3 = Knoll|first4 = Laura A.|last4 = Katz|bibcode = 2011PNAS..10813624P}}</ref><ref>{{Cite journal|last=Burki|first=Fabien|last2=Kaplan|first2=Maia|last3=Tikhonenkov|first3=Denis V.|last4=Zlatogursky|first4=Vasily|last5=Minh|first5=Bui Quang|last6=Radaykina|first6=Liudmila V.|last7=Smirnov|first7=Alexey|last8=Mylnikov|first8=Alexander P.|last9=Keeling|first9=Patrick J. |date=2016-01-27 |title=Untangling the early diversification of eukaryotes: a phylogenomic study of the evolutionary origins of Centrohelida, Haptophyta and Cryptista |url=http://rspb.royalsocietypublishing.org/content/283/1823/20152802 |journal=Proc. R. Soc. B |language=en |volume=283 |issue=1823 |pages=20152802 |doi=10.1098/rspb.2015.2802 |issn=0962-8452 |pmid=26817772 |pmc=4795036}}</ref>

One suggested grouping is as follows: (1) ''[[Cryptomonas]]'', (2) ''[[Chroomonas]]/[[Komma caudata|Komma]]'' and ''[[Hemiselmis]]'', (3) ''[[Rhodomonas]]/[[Rhinomonas]]/[[Storeatula]]'', (4) ''[[Guillardia]]/[[Hanusia]]'', (5) ''[[Geminigera]]/[[Plagioselmis]]/[[Teleaulax]]'', (6) ''[[Proteomonas sulcata]]'', (7) ''[[Falcomonas daucoides]]''.<ref name="urlCryptomonads">{{cite web |url=http://tolweb.org/Cryptomonads/2396 |title=Cryptomonads |format= |website= |accessdate=2009-06-24}}</ref>

* '''Class Cryptophyceae '''<small>Fritsch 1937</small> [Cryptomonadea <small>Stein 1878 emend. Schoenichen 1925</small>]
** Order [[Tetragonidiales]] <small>Kristiansen 1992</small>
*** Family ?[[Tetragonidiaceae]] <small>Bourelly ex Silva1980</small>
**** Genus ''[[Bjornbergiella]]'' <small>Bicudo 1966</small>
**** Genus ''[[Tetragonidium]]'' <small>Pascher 1914</small>
** Order [[Pyrenomonadales]] <small>Novarino & Lucas 1993</small>
*** Family [[Pyrenomonadaceae]] <small>Novarino & Lucas 1993</small>
**** Genus ''[[Rhinomonas]]'' <small>Hill & Wetherbee 1988</small>
**** Genus ''[[Rhodomonas]]'' <small>Karsten 1898</small> [''[[Pyrenomonas]]'' <small>Santore 1984</small>]
**** Genus ''[[Storeatula]]'' <small>Hill 1991</small>
** Order [[Cryptomonadales]] <small>Pascher 1913</small>
*** Family ?[[Butschliellaceae]] <small>Skvortzov 1968</small>
**** Genus ''[[Butschliella]]'' <small>Skvortzov 1968</small>
**** Genus ''[[Skvortzoviella]]'' <small>Bourelly 1970</small>
*** Family ?[[Cryptochrysidaceae]] <small>Pascher 1931</small>
**** Genus ''[[Cryptochrysis]]'' <small>Pascher 1911</small>
*** Family ?[[Hilleaceae]] <small>Pascher 1967</small>
**** Genus ''[[Calkinsiella]]'' <small>Skvortzov 1969</small>
**** Genus ''[[Hillea]]'' <small>Schiller 1925</small>
*** Family ?[[Pleuromastigaceae]] <small>Bourrelly ex Silva 1980</small>
**** Genus ''[[Pleuromastix]]'' <small>Scherffel 1912 non Namyslowski 1913</small>]
**** Genus ''[[Xanthodiscus]]'' <small>Schewiakoff 1892</small>
*** Clade
**** Genus ''[[Proteomonas]]'' <small>Hill & Wetherbee 1986</small>
*** Family [[Chroomonadaceae]] <small>Clay, Cugrens & Lee 1999</small> [Hemiselmidaceae <small>Butcher 1967</small>; Senniaceae <small>Skuja 1948</small>; Planonephraceae <small>Christensen 1967</small>]
**** Genus ?''[[Smithimastix]]'' <small>Skvortzov 1969</small> [''[[Smithiella]]'' <small>Skvortzov 1968 nom. illeg.</small>]
**** Genus ''[[Falcomonas]]'' <small>Hill 1991</small>
**** Genus ''[[Chroomonas]]'' <small>Hansgirg 1885</small>
**** Genus ''[[Komma (alga)|Komma]]'' <small>Hill 1991</small>
**** Genus ''[[Planonephros]]'' <small>Christensen 1978</small>
**** Genus ''[[Nodeana]]'' <small>Skvortzov 1968</small>
**** Genus ''[[Protochrysis]]'' <small>Pascher 1911</small>
**** Genus ''[[Hemiselmis]]'' <small>Parke 1949</small>
*** Family [[Baffinellaceae]] <small>Daugbjerg & Norlin 2018</small><ref>{{Cite journal |last=Daugbjerg |first=Niels |last2=Norlin |first2=Andreas |last3=Lovejoy |first3=Connie |date=2018-07-25 |title=''Baffinella frigidus'' gen. et sp. nov. (Baffinellaceae fam. nov., Cryptophyceae) from Baffin Bay: Morphology, pigment profile, phylogeny, and growth rate response to three abiotic factors |url=https://www1.bio.ku.dk/staff/ndaugbjerg/pdf/Daugbjerg_et_al_2018.pdf |journal=J. Phycol. |language=en |volume=54 |issue=5 |pages=665–680 |doi=10.1111/jpy.12766 |issn=1529-8817 |pmid=30043990}}</ref>
**** Genus ''[[Baffinella]]'' <small>Norlin & Daugbjerg 2018</small>
*** Family [[Geminigeraceae]] <small>Clay, Cugrens & Lee 1999</small>
**** Genus ''[[Geminigera]]'' <small>Hill 1991</small>
**** Genus ''[[Plagioselmis]]'' <small>Butcher 1967 ex Novarino, Lucas & Morrall 1994</small>
**** Genus ''[[Teleaulax]]'' <small>Hill 1991</small>
**** Genus ''[[Urgorri]]'' <small>Laza-Martinez 2012</small>
*** clade ''Guillardia'' group
**** Genus ''[[Guillardia]]'' <small>Hill & Wetherbee 1990</small>
**** Genus ''[[Phia]]'' <small>Özdikmen 2009</small> [''[[Hanusia]]'' <small>Deane et al. 1998 non Cripps 1989</small>]
*** Family [[Cryptomonadaceae]] <small>Ehrenberg 1831</small> [Campylomonadaceae <small>Clay, Kugrens & Lee 1999</small>]
**** Genus ''[[Cryptomonas]]'' <small>Ehrenberg 1832</small> [''[[Campylomonas]]'' <small>Hill 1991</small>]
**** Genus ''[[Cryptella]]'' <small>Pascher 1929</small>
**** Genus ''[[Cryptochloris]]'' <small>Schiller 1925</small>
**** Genus ''[[Cyanomastix]]'' <small>Lackey 1936</small>
**** Genus ''[[Isoselmis]]'' <small>Butcher 1967</small>
**** Genus ''[[Kisselevia]]'' <small>Skvortzov 1969</small>
**** Genus ''[[Meyeriella]]'' <small>Skvortzov 1968</small>
**** Genus ''[[Olivamonas]]'' <small>Skvortzov 1969</small>
**** Genus ''[[Protocryptomonas]]'' <small>Skvortzov 1969 ex Bicudo 1989</small>

==References==
{{Reflist|40em}}

==External links==
* [http://tolweb.org/Cryptomonads/2396 Tree of Life: Cryptomonads]

{{Eukaryota classification}}
{{Cryptophyta and haptophyta}}
{{Life on Earth}}
{{Taxonbar|from=Q21281984|from2=Q18668642}}

[[Category:Cryptomonads| ]]
[[Category:Taxa named by Thomas Cavalier-Smith]]

Revision as of 00:15, 26 May 2019

Cryptophytes
Rhodomonas salina
Scientific classification
(incertae sedis within Eukaryota)
Domain:
(unranked):
(unranked):
Superclass:
Orders
Synonyms
  • Cryptomonada Senn, 1900
  • Cryptomonadinae Pascher, 1913
  • Cryptomonadophyceae Pascher ex Schoenichem, 1925

The cryptophyceae are a class of algae,[1] most of which have plastids. They are common in freshwater, and also occur in marine and brackish habitats. Each cell is around 10–50 μm in size and flattened in shape, with an anterior groove or pocket. At the edge of the pocket there are typically two slightly unequal flagella.

Some exhibit mixotrophy.[2]

Characteristics

Cell scheme: 1-contractile vacuole, 2-plastid, 3-thylakoid, 4-stigma, 5-nucleomorph, 6-starch granule, 7-70S ribosome, 8-nucleus, 9-80S ribosome, 10-flagella, 11-invagination, 12-lipid globules, 13-ejectosomes, 14-mitochondrion, 15-pyrenoid, 16-Golgi apparatus, 17-endoplasmic reticulum, 18-chloroplast-endoplasmic reticulum.

Cryptophytes are distinguished by the presence of characteristic extrusomes called ejectosomes or ejectisomes, which consist of two connected spiral ribbons held under tension.[3] If the cells are irritated either by mechanical, chemical or light stress, they discharge, propelling the cell in a zig-zag course away from the disturbance. Large ejectosomes, visible under the light microscope, are associated with the pocket; smaller ones occur underneath the periplast, the cryptophyte-specific cell surrounding.[4][5]

Except for Chilomonas, which has leucoplasts, cryptophytes have one or two chloroplasts. These contain chlorophylls a and c, together with phycobiliproteins and other pigments, and vary in color (brown, red to blueish-green). Each is surrounded by four membranes, and there is a reduced cell nucleus called a nucleomorph between the middle two. This indicates that the plastid was derived from a eukaryotic symbiont, shown by genetic studies to have been a red alga.[6] However, the plastids are very different from red algal plastids: phycobiliproteins are present but only in the thylakoid lumen and are present only as phycoerythrin or phycocyanin. In the case of "Rhodomonas" the crystal structure has been determined to 1.63Å;[7] and it has been shown that the alpha subunit bears no relation to any other known phycobiliprotein.

A few cryptophytes, such as Cryptomonas, can form palmelloid stages, but readily escape the surrounding mucus to become free-living flagellates again. Some Cryptomonas species may also form immotile microbial cysts–resting stages with rigid cell walls to survive unfavorable conditions. Cryptophyte flagella are inserted parallel to one another, and are covered by bipartite hairs called mastigonemes, formed within the endoplasmic reticulum and transported to the cell surface. Small scales may also be present on the flagella and cell body. The mitochondria have flat cristae, and mitosis is open; sexual reproduction has also been reported.

Classification

Cryptophytes under SEM
Cryptophytes under light microscope

The first mention of cryptophytes appears to have been made by Christian Gottfried Ehrenberg in 1831,[8] while studying Infusoria. Later, botanists treated them as a separate algae group, class Cryptophyceae or division Cryptophyta, while zoologists treated them as the flagellate protozoa order Cryptomonadina. In some classifications, the cryptomonads were considered close relatives of the dinoflagellates because of their (seemingly) similar pigmentation, being grouped as the Pyrrhophyta. There is considerable evidence that cryptophyte chloroplasts are closely related to those of the heterokonts and haptophytes, and the three groups are sometimes united as the Chromista. However, the case that the organisms themselves are closely related is not very strong, and they may have acquired plastids independently. Currently they are discussed to be members of the clade Diaphoretickes and to form together with the Haptophyta the group Hacrobia. Parfrey et al. and Burki et al. placed Cryptophyceae as a sister clade to the Green Algae.[9][10]

One suggested grouping is as follows: (1) Cryptomonas, (2) Chroomonas/Komma and Hemiselmis, (3) Rhodomonas/Rhinomonas/Storeatula, (4) Guillardia/Hanusia, (5) Geminigera/Plagioselmis/Teleaulax, (6) Proteomonas sulcata, (7) Falcomonas daucoides.[11]

References

  1. ^ Khan H, Archibald JM (May 2008). "Lateral transfer of introns in the cryptophyte plastid genome". Nucleic Acids Res. 36 (9): 3043–53. doi:10.1093/nar/gkn095. PMC 2396441. PMID 18397952.
  2. ^ "Cryptophyta - the cryptomonads". Archived from the original on 2011-06-10. Retrieved 2009-06-02. {{cite web}}: Unknown parameter |deadurl= ignored (|url-status= suggested) (help)
  3. ^ Graham, L. E.; Graham, J. M.; Wilcox, L. W. (2009). Algae (2nd ed.). San Francisco, CA: Benjamin Cummings (Pearson). ISBN 9780321559654.
  4. ^ Morrall, S.; Greenwood, A. D. (1980). "A comparison of the periodic sub-structures of the trichocysts of the Cryptophyceae and Prasinophyceae". BioSystems. 12 (1–2): 71–83. doi:10.1016/0303-2647(80)90039-8. PMID 6155157.
  5. ^ Grim, J. N.; Staehelin, L. A. (1984). "The ejectisomes of the flagellate Chilomonas paramecium - Visualization by freeze-fracture and isolation techniques". Journal of Protozoology. 31 (2): 259–267. doi:10.1111/j.1550-7408.1984.tb02957.x.
  6. ^ Douglas, S.; et al. (2002). "The highly reduced genome of an enslaved algal nucleus". Nature. 410 (6832): 1091–1096. doi:10.1038/35074092. PMID 11323671.
  7. ^ Wilk, K.; et al. (1999). "Evolution of a light-harvesting protein by addition of new subunits and rearrangement of conserved elements: Crystal structure of a cryptophyte phycoerythrin at 1.63Å resolution". PNAS. 96 (16): 8901–8906. doi:10.1073/pnas.96.16.8901. PMC 17705. PMID 10430868.
  8. ^ Novarino, G. (2012). "Cryptomonad taxonomy in the 21st century: The first 200 years". Phycological Reports: Current Advances in Algal Taxonomy and its Applications: Phylogenetic, Ecological and Applied Perspective: 19–52. Retrieved 2018-10-16.
  9. ^ Parfrey, Laura Wegener; Lahr, Daniel J. G.; Knoll, Andrew H.; Katz, Laura A. (2011-08-16). "Estimating the timing of early eukaryotic diversification with multigene molecular clocks". Proceedings of the National Academy of Sciences. 108 (33): 13624–13629. Bibcode:2011PNAS..10813624P. doi:10.1073/pnas.1110633108. ISSN 0027-8424. PMC 3158185. PMID 21810989.
  10. ^ Burki, Fabien; Kaplan, Maia; Tikhonenkov, Denis V.; Zlatogursky, Vasily; Minh, Bui Quang; Radaykina, Liudmila V.; Smirnov, Alexey; Mylnikov, Alexander P.; Keeling, Patrick J. (2016-01-27). "Untangling the early diversification of eukaryotes: a phylogenomic study of the evolutionary origins of Centrohelida, Haptophyta and Cryptista". Proc. R. Soc. B. 283 (1823): 20152802. doi:10.1098/rspb.2015.2802. ISSN 0962-8452. PMC 4795036. PMID 26817772.
  11. ^ "Cryptomonads". Retrieved 2009-06-24.
  12. ^ Daugbjerg, Niels; Norlin, Andreas; Lovejoy, Connie (2018-07-25). "Baffinella frigidus gen. et sp. nov. (Baffinellaceae fam. nov., Cryptophyceae) from Baffin Bay: Morphology, pigment profile, phylogeny, and growth rate response to three abiotic factors" (PDF). J. Phycol. 54 (5): 665–680. doi:10.1111/jpy.12766. ISSN 1529-8817. PMID 30043990.