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| subdivision_ranks = Species
| subdivision_ranks = Species
| subdivision_ref = <ref name="Algaebase">{{cite web | url = http://www.algaebase.org/browse/taxonomy/?id=8295 | publisher = AlgaeBase version 4.2 World-wide electronic publication, National University of Ireland, Galway | accessdate = 2007-09-23 | title = Genus: ''Caulerpa'' taxonomy browser |author1=Guiry, M.D. |author2=Guiry, G.M. | year = 2007 }}</ref>
| subdivision_ref = <ref name="Algaebase">{{cite web | url = http://www.algaebase.org/browse/taxonomy/?id=8295 | publisher = AlgaeBase version 4.2 World-wide electronic publication, National University of Ireland, Galway | accessdate = 2007-09-23 | title = Genus: ''Caulerpa'' taxonomy browser |author1=Guiry, M.D. |author2=Guiry, G.M. | year = 2007 }}</ref>
| subdivision = About 86
| subdivision = About 101
}}
}}


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Referring to its [[thalli]]'s crawling [[growth habit#Structure|habit]], the name means 'stem (that) creeps', from the Ancient Greek ''{{transl|grc|kaulos}}'' ({{lang|grc|[[wikt:καυλός|καυλός]]}}, ‘stalk’) and ''{{transl|grc|herpo}}'' ({{lang|grc|[[wikt:ἕρπω|ἕρπω]]}}, ‘to creep’).<ref>{{cite book|last1=Bold|first1=Harold Charles|last2=Wynne|first2=Michael James|title=Introduction to the algae|url=https://books.google.com/books?id=J4XwAAAAMAAJ|year=1985|publisher=Prentice-Hall|page=229|isbn=9780134777467}}</ref>
Referring to its [[thalli]]'s crawling [[growth habit#Structure|habit]], the name means 'stem (that) creeps', from the Ancient Greek ''{{transl|grc|kaulos}}'' ({{lang|grc|[[wikt:καυλός|καυλός]]}}, ‘stalk’) and ''{{transl|grc|herpo}}'' ({{lang|grc|[[wikt:ἕρπω|ἕρπω]]}}, ‘to creep’).<ref>{{cite book|last1=Bold|first1=Harold Charles|last2=Wynne|first2=Michael James|title=Introduction to the algae|url=https://books.google.com/books?id=J4XwAAAAMAAJ|year=1985|publisher=Prentice-Hall|page=229|isbn=9780134777467}}</ref>

== Taxonomy and Nomenclature ==
First described by Jean Vincent Lamouroux in 1809, ''Caulerpa'' is the only genus under the family Caulerpaceae, from the order Bryopsidales, class Ulvophyceae, and phylum Chlorophyta.<ref name=":0" />

Species discrimination via morphology-based identification is often hampered by the high degree of variation among traits. Thus, several species are often misidentified without the use if molecular analyses.<ref>{{Cite journal|last=Dumilag|first=Richard|last2=Aguinaldo|first2=Zae-Zae|date=2019|title=DNA Barcodes of Caulerpa Species (Caulerpaceae,
Chlorophyta) from the Northern Philippines|journal=Philippine Journal of Science|volume=148|issue=2|pages=343-353|via=researchgate.net}}</ref> As of 2019, there are 101 accepted species, with 40 varieties and 67 forms.<ref>{{Cite journal|last=Belton|first=Gareth S.|last2=Draisma|first2=Stefano G.A.|last3=Prud’homme van Reine|first3=Willem F.|last4=Huisman|first4=John M.|last5=Gurgel|first5=C. Frederico D.|date=2019-05-04|title=A taxonomic reassessment of Caulerpa (Chlorophyta, Caulerpaceae) in southern Australia, based on tuf A and rbc L sequence data|url=https://www.tandfonline.com/doi/full/10.1080/00318884.2018.1542851|journal=Phycologia|language=en|volume=58|issue=3|pages=234–253|doi=10.1080/00318884.2018.1542851|issn=0031-8884}}</ref><ref>{{Cite web|last=Guiry|first=MD|last2=Guiry|first2=GM|date=2019|title=Algaebase|url=http://www.algaebase.org|url-status=live|website=Algaebase}}</ref>


== Biology ==
== Biology ==
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The genus produces a number of secondary metabolites thought to be related to its toxicity and peppery taste. These include the red pigment alkaloid [[caulerpin]] and its derivative caulerchlorin and the amine mixture [[caulerpicin]].
The genus produces a number of secondary metabolites thought to be related to its toxicity and peppery taste. These include the red pigment alkaloid [[caulerpin]] and its derivative caulerchlorin and the amine mixture [[caulerpicin]].

== Morphology ==

===== External Characteristics =====
The genus ''Caulerpa'' exhibits several different growth forms. They possess a stolon with rhizoids which grow downward, anchoring the plant to the substrate. The stolons support upright fronds or leaves known as assimilators. The structure of the assimilator may be ligulate (leaf-like, e.g. ''Caulerpa prolifera''), or they may have a central axis known as a rachis. The rachis may have lateral branchlets known as ramuli which themselves come in different forms (terete, turbinate, clavate, peltate, falcate, vesiculate) and arrangements: distichous - ramuli are arranged evenly opposite each other (e.g. ''Caulerpa taxifolia''), irregular - vesiculate ramuli with no distinct arrangement (e.g. ''Caulerpa racemosa''), verticillate - whorled ramuli (e.g. ''Caulerpa cuppresoides'').<ref name=":0" />

==== Internal anatomy ====
''Caulerpa'' is [[Coenocyte|coenocytic]], meaning it has a multinucleate thallus organization. It is also siphonous, meaning unlike other algae, the thallus and the nuclei are not separated by cell walls. They are instead one long mass of protoplasm surrounded by a single cell wall.<ref>{{Cite journal|last=Coneva|first=Viktoriya|last2=Chitwood|first2=Daniel H.|date=2015-04-24|title=Plant architecture without multicellularity: quandaries over patterning and the soma-germline divide in siphonous algae|url=http://dx.doi.org/10.3389/fpls.2015.00287|journal=Frontiers in Plant Science|volume=6|doi=10.3389/fpls.2015.00287|issn=1664-462X}}</ref> The genus also possesses trabeculae, which are inward growing cell wall extensions that pass through the central lumen of siphons. It is hypothesized that these provide the thallus with structural support, facilitate diffusion to the inner cytoplasm, and can possibly help in determining cell shape - which may contribute to the diversity of growth forms found in the genus.<ref>{{Cite journal|last=Fagerberg|first=Wayne R.|last2=Hodges|first2=Ellen (Lavoie)|last3=Dawes|first3=Clinton J.|date=2010-04|title=THE DEVELOPMENT AND POTENTIAL ROLES OF CELL-WALL TRABECULAE INCAULERPA MEXICANA(CHLOROPHYTA)|url=http://dx.doi.org/10.1111/j.1529-8817.2009.00784.x|journal=Journal of Phycology|volume=46|issue=2|pages=309–315|doi=10.1111/j.1529-8817.2009.00784.x|issn=0022-3646}}</ref>


== Distribution ==
== Distribution ==
''Caulerpa'' is mostly found in tropical regions, however its distribution may also extend up to temperate locations. Diversity is highest in the Caribbean and the Indo-Malay region, as well as in southern Australia, where a majority of ''Caulerpa'' species are endemic.<ref>{{Cite journal|last=Prud'Homme Van Reine|first=W. F.|last2=Verheij|first2=E.|last3=Coppejans|first3=E.|date=October 1996|title=Species and ecads of Caulerpa (Ulvophyceae, Chlorophyta) in Malesia (South-East Asia): Taxonomy, biogeography and biodiversity|url=http://dx.doi.org/10.1007/bf02272230|journal=Netherlands Journal of Aquatic Ecology|volume=30|issue=2-3|pages=83–98|doi=10.1007/bf02272230|issn=1380-8427}}</ref><ref>{{Cite journal|last=Zubia|first=Mayalen|last2=Draisma|first2=Stefano G. A.|last3=Morrissey|first3=Kathryn Lee|last4=Varela-Álvarez|first4=Elena|last5=De Clerck|first5=Olivier|date=2019-07-23|title=Concise review of the genus Caulerpa J.V. Lamouroux|url=http://dx.doi.org/10.1007/s10811-019-01868-9|journal=Journal of Applied Phycology|volume=32|issue=1|pages=23–39|doi=10.1007/s10811-019-01868-9|issn=0921-8971}}</ref>
''Caulerpa'' is mostly found in tropical regions, however its distribution may also extend up to temperate locations. Diversity is highest in the Caribbean and the Indo-Malay region, as well as in southern Australia, where a majority of ''Caulerpa'' species are endemic.<ref>{{Cite journal|last=Prud'Homme Van Reine|first=W. F.|last2=Verheij|first2=E.|last3=Coppejans|first3=E.|date=October 1996|title=Species and ecads of Caulerpa (Ulvophyceae, Chlorophyta) in Malesia (South-East Asia): Taxonomy, biogeography and biodiversity|url=http://dx.doi.org/10.1007/bf02272230|journal=Netherlands Journal of Aquatic Ecology|volume=30|issue=2-3|pages=83–98|doi=10.1007/bf02272230|issn=1380-8427}}</ref><ref name=":0">{{Cite journal|last=Zubia|first=Mayalen|last2=Draisma|first2=Stefano G. A.|last3=Morrissey|first3=Kathryn Lee|last4=Varela-Álvarez|first4=Elena|last5=De Clerck|first5=Olivier|date=2019-07-23|title=Concise review of the genus Caulerpa J.V. Lamouroux|url=http://dx.doi.org/10.1007/s10811-019-01868-9|journal=Journal of Applied Phycology|volume=32|issue=1|pages=23–39|doi=10.1007/s10811-019-01868-9|issn=0921-8971}}</ref>

== Ecology ==
The genus is typically found in shallow intertidal zones and can reach up to depths of 100 meters. They are known to be able to adapt and thrive in different environmental conditions, which contributes to their potential for becoming [[invasive species]]. Some of their traits include having a high tolerance for a wide range range of temperatures, their capacity for [[Asexual reproduction of plants|asexual reproduction]] via spore dispersal, their fast growth rate, as well as their capacity for nutrient intake through their rhizoids.<ref name=":0" /><ref>{{Cite web|last=Hoddle|first=Mark|date=2021|title=Caulerpa Taxifolia or Killer Alga|url=https://cisr.ucr.edu/invasive-species/caulerpa-taxifolia-or-killer-alga|url-status=live|access-date=14 January 2022|website=Center for Invasive Species Research, University of California Riverside}}</ref>


==Culinary use==
==Culinary use==

Revision as of 09:04, 14 January 2022

Caulerpa
Caulerpa prolifera
Scientific classification Edit this classification
(unranked): Viridiplantae
Division: Chlorophyta
Class: Ulvophyceae
Order: Bryopsidales
Family: Caulerpaceae
Genus: Caulerpa
J.V. Lamouroux, 1809
Type species
Caulerpa prolifera
(Forsskål) J.V. Lamouroux, 1809 [2]
Species[1]

About 101

Caulerpa is a genus of seaweeds in the family Caulerpaceae (among the green algae). They are unusual because they consist of only one cell with many nuclei, making them among the biggest single cells in the world. A species in the Mediterranean can have a stolon more than 3 metres (9.8 ft) long, with up to 200 fronds. This species can be invasive from time to time.

Referring to its thalli's crawling habit, the name means 'stem (that) creeps', from the Ancient Greek kaulos (καυλός, ‘stalk’) and herpo (ἕρπω, ‘to creep’).[3]

Taxonomy and Nomenclature

First described by Jean Vincent Lamouroux in 1809, Caulerpa is the only genus under the family Caulerpaceae, from the order Bryopsidales, class Ulvophyceae, and phylum Chlorophyta.[4]

Species discrimination via morphology-based identification is often hampered by the high degree of variation among traits. Thus, several species are often misidentified without the use if molecular analyses.[5] As of 2019, there are 101 accepted species, with 40 varieties and 67 forms.[6][7]

Biology

Caulerpa supports its large size by having its cytoplasm circulate constantly, supported by a network of microtubules. This behavior was known in 1967.[8]

The cytoplasm does not leak out when the cell is cut. Regeneration is directional, with rhizoids at the bottom and fronds at the top.[9]

The genus produces a number of secondary metabolites thought to be related to its toxicity and peppery taste. These include the red pigment alkaloid caulerpin and its derivative caulerchlorin and the amine mixture caulerpicin.

Morphology

External Characteristics

The genus Caulerpa exhibits several different growth forms. They possess a stolon with rhizoids which grow downward, anchoring the plant to the substrate. The stolons support upright fronds or leaves known as assimilators. The structure of the assimilator may be ligulate (leaf-like, e.g. Caulerpa prolifera), or they may have a central axis known as a rachis. The rachis may have lateral branchlets known as ramuli which themselves come in different forms (terete, turbinate, clavate, peltate, falcate, vesiculate) and arrangements: distichous - ramuli are arranged evenly opposite each other (e.g. Caulerpa taxifolia), irregular - vesiculate ramuli with no distinct arrangement (e.g. Caulerpa racemosa), verticillate - whorled ramuli (e.g. Caulerpa cuppresoides).[4]

Internal anatomy

Caulerpa is coenocytic, meaning it has a multinucleate thallus organization. It is also siphonous, meaning unlike other algae, the thallus and the nuclei are not separated by cell walls. They are instead one long mass of protoplasm surrounded by a single cell wall.[10] The genus also possesses trabeculae, which are inward growing cell wall extensions that pass through the central lumen of siphons. It is hypothesized that these provide the thallus with structural support, facilitate diffusion to the inner cytoplasm, and can possibly help in determining cell shape - which may contribute to the diversity of growth forms found in the genus.[11]

Distribution

Caulerpa is mostly found in tropical regions, however its distribution may also extend up to temperate locations. Diversity is highest in the Caribbean and the Indo-Malay region, as well as in southern Australia, where a majority of Caulerpa species are endemic.[12][4]

Ecology

The genus is typically found in shallow intertidal zones and can reach up to depths of 100 meters. They are known to be able to adapt and thrive in different environmental conditions, which contributes to their potential for becoming invasive species. Some of their traits include having a high tolerance for a wide range range of temperatures, their capacity for asexual reproduction via spore dispersal, their fast growth rate, as well as their capacity for nutrient intake through their rhizoids.[4][13]

Culinary use

Edible fresh latô (Caulerpa lentillifera) sold at a fish market in the Philippines

Some species of Caulerpa are edible. The two most commonly eaten are Caulerpa lentillifera and Caulerpa racemosa, both called "sea grapes" in English. Both are traditionally harvested in the wild and sold in local markets in Southeast Asia, Oceania, and East Asia. They are eaten raw in salads and have a characteristic "sea" flavor and a crunchy texture.[14]

Only C. lentillifera is cultivated in aquaculture. Its cultivation began in the 1950s in Cebu, Philippines, after accidental introduction of C. lentillifera to fish ponds.[15] This was followed by Japan in 1986, where it was cultivated in tanks in the tropical waters of Okinawa.[16] Commercial cultivation has since spread to other countries, including Vietnam, Taiwan, and China (in Fujian and Hainan). Most are for domestic consumption, but they are also exported to Japan.[17]

Invasive behaviour

Another species, Caulerpa taxifolia, has become an invasive species in the Mediterranean Sea, Australia and southern California (where it has since been eradicated). In U.S. waters, the Mediterranean strain of Caulerpa taxifolia is listed as a federal noxious weed, under the Plant Protection Act. The Aquatic Nuisance Species Taskforce has also created a National Management Plan for the Genus Caulerpa. The state of California also prohibits possession of nine different species of Caulerpa.

It is thought that Caulerpa species have such invasive properties in these regions due to their capability to thrive in temperate waters, along with their freedom from natural predators. Most Caulerpa species evolved in tropical waters, where herbivores have immunity to toxic compounds (mainly caulerpicin) within the alga. Temperate water herbivores have no natural immunity to these toxins, allowing Caulerpa to grow unchecked if introduced to temperate waters.

C. racemosa has recently been found in waters around Crete, where it is thought to have contributed to a significant reduction in fisheries. The alga has invaded the area from the warmer waters of the Red Sea.

C. cylindracea, which is native to Australia, has also become an invasive species in the Mediterranean.[18]

Use in aquariums

Caulerpa is common in the aquarium hobby as a nitrate absorber because of its rapid growth under relatively adverse conditions. It may also be used in refugiums for a long-term nitrite absorber. Many introductions of invasive Caulerpa to the wild are thought to have occurred via aquarium dumping although there is no proof that this is so. For this reason, some aquarium hobbyists have begun using Chaetomorpha or an algae scrubber instead.[19][20]

Species

Oval sea grapes, Caulerpa racemosa var. clavifera, at 5 metres' depth
Feather algae, Caulerpa sertularioides at 11 metres' depth on ridge

The species currently recognized are:[2]

References

  1. ^ Guiry, M.D.; Guiry, G.M. (2007). "Genus: Caulerpa taxonomy browser". AlgaeBase version 4.2 World-wide electronic publication, National University of Ireland, Galway. Retrieved 2007-09-23.
  2. ^ a b Caulerpa J.V. Lamouroux, 1809 World Register of Marine Species. Retrieved 2011-08-20
  3. ^ Bold, Harold Charles; Wynne, Michael James (1985). Introduction to the algae. Prentice-Hall. p. 229. ISBN 9780134777467.
  4. ^ a b c d Zubia, Mayalen; Draisma, Stefano G. A.; Morrissey, Kathryn Lee; Varela-Álvarez, Elena; De Clerck, Olivier (2019-07-23). "Concise review of the genus Caulerpa J.V. Lamouroux". Journal of Applied Phycology. 32 (1): 23–39. doi:10.1007/s10811-019-01868-9. ISSN 0921-8971.
  5. ^ Dumilag, Richard; Aguinaldo, Zae-Zae (2019). "DNA Barcodes of Caulerpa Species (Caulerpaceae, Chlorophyta) from the Northern Philippines". Philippine Journal of Science. 148 (2): 343–353 – via researchgate.net. {{cite journal}}: line feed character in |title= at position 48 (help)
  6. ^ Belton, Gareth S.; Draisma, Stefano G.A.; Prud’homme van Reine, Willem F.; Huisman, John M.; Gurgel, C. Frederico D. (2019-05-04). "A taxonomic reassessment of Caulerpa (Chlorophyta, Caulerpaceae) in southern Australia, based on tuf A and rbc L sequence data". Phycologia. 58 (3): 234–253. doi:10.1080/00318884.2018.1542851. ISSN 0031-8884.
  7. ^ Guiry, MD; Guiry, GM (2019). "Algaebase". Algaebase.{{cite web}}: CS1 maint: url-status (link)
  8. ^ Sabnis, DD; Jacobs, WP (December 1967). "Cytoplasmic streaming and microtubules in the coenocytic marine alga, Caulerpa prolifera". Journal of Cell Science. 2 (4): 465–72. PMID 6080540.
  9. ^ Jacobs, William P. (December 1994). "Caulerpa". Scientific American. 271 (6): 100. Bibcode:1994SciAm.271f.100J. doi:10.1038/scientificamerican1294-100. This tropical alga is the world's largest single-celled organism. Yet it di›erentiates into a complex structure of leaves, stems and roots.
  10. ^ Coneva, Viktoriya; Chitwood, Daniel H. (2015-04-24). "Plant architecture without multicellularity: quandaries over patterning and the soma-germline divide in siphonous algae". Frontiers in Plant Science. 6. doi:10.3389/fpls.2015.00287. ISSN 1664-462X.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  11. ^ Fagerberg, Wayne R.; Hodges, Ellen (Lavoie); Dawes, Clinton J. (2010-04). "THE DEVELOPMENT AND POTENTIAL ROLES OF CELL-WALL TRABECULAE INCAULERPA MEXICANA(CHLOROPHYTA)". Journal of Phycology. 46 (2): 309–315. doi:10.1111/j.1529-8817.2009.00784.x. ISSN 0022-3646. {{cite journal}}: Check date values in: |date= (help)
  12. ^ Prud'Homme Van Reine, W. F.; Verheij, E.; Coppejans, E. (October 1996). "Species and ecads of Caulerpa (Ulvophyceae, Chlorophyta) in Malesia (South-East Asia): Taxonomy, biogeography and biodiversity". Netherlands Journal of Aquatic Ecology. 30 (2–3): 83–98. doi:10.1007/bf02272230. ISSN 1380-8427.
  13. ^ Hoddle, Mark (2021). "Caulerpa Taxifolia or Killer Alga". Center for Invasive Species Research, University of California Riverside. Retrieved 14 January 2022.{{cite web}}: CS1 maint: url-status (link)
  14. ^ Paul, Nicholas A.; Neveux, Nicolas; Magnusson, Marie; de Nys, Rocky (21 December 2013). "Comparative production and nutritional value of "sea grapes" — the tropical green seaweeds Caulerpa lentillifera and C. racemosa". Journal of Applied Phycology. doi:10.1007/s10811-013-0227-9. S2CID 15745994.
  15. ^ Trono, Gavino C., Jr. (December 1988). Manual on Seaweed Culture. ASEAN/UNDP/FAO Regional Small-Scale Coastal Fisheries Development Project.{{cite book}}: CS1 maint: multiple names: authors list (link)
  16. ^ Trono, G.C., Jr. "Caulerpa lentillifera (PROSEA)". Pl@ntUse. PROSEA (Plant Resources of South East Asia). Retrieved 26 October 2020.{{cite web}}: CS1 maint: multiple names: authors list (link)
  17. ^ Chen, Xiaolin; Sun, Yuhao; Liu, Hong; Liu, Song; Qin, Yukun; Li, Pengcheng (2019). "Advances in cultivation, wastewater treatment application, bioactive components of Caulerpa lentillifera and their biotechnological applications". PeerJ. 7: e6118. doi:10.7717/peerj.6118. PMC 6329336. PMID 30643691.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  18. ^ Montefalcone, Monica; Morri, Carla; Parravicini, Valeriano; Bianchi, Carlo Nike (26 May 2015). "A tale of two invaders: divergent spreading kinetics of the alien green algae Caulerpa taxifolia and Caulerpa cylindracea". Biological Invasions. 17 (9): 2717–2728. doi:10.1007/s10530-015-0908-1. S2CID 1973648.
  19. ^ Nutrient Cycling In The Great Barrier Reef Aquarium. Proceedings of the 6th International Coral Reef Symposium, Australia, 1988, Vol. 2
  20. ^ Reef Invertebrates, 2003, page 46

Further reading

External links

Wikispecies has information related to Caulerpa.
Retrieved from "https://en.wikipedia.org/w/index.php?title=Caulerpa&oldid=1065589988"