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Hydroidolina

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Hydroidolina
Siphonophorae from Ernst Haeckel's 1904 Kunstformen der Natur
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Cnidaria
Class: Hydrozoa
Subclass: Hydroidolina
Marques & Collins, 2004
Orders
  • Anthoathecata
  • Leptothecata
  • Siphonophorae

See text.

Synonyms
  • Hydroida
  • Hydroidae
  • Hydroidolinae Marques & Collins, 2004
  • Hydroidomedusa
  • Hydroidomedusae
  • Leptolida Haeckel, 1879
  • Leptolina Haeckel, 1879
  • Leptolinae Haeckel, 1879
  • Hydroidida

Hydroidolina[1] is a subclass of Hydrozoa and makes up 90% of the class.[2] Controversy surrounds who the sister groups of Hydroidolina are, but research has shown that three orders remain consistent as direct relatives: Siphonophorae, Anthoathecata, and Leptothecata.[3]

Description and background

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The phylum Cnidaria contains two clades: Anthozoa and Medusozoa. There are around 3800 species within the clade Medusozoa and it consists of Cubozoans, Scyphozoans, and Hydrozoans.[4]

Hydroidolina are small predatory animals, ranging in 8-30 millimeters in size,[5] exhibiting radial symmetry and are diploblastic (developed from two embryonic layers: ectoderm and endoderm).[4]

The classification below is based on the World Register of Marine Species:[6]

Subclass Hydroidolina

Distribution

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Hydroidolina are commonly found in a variety of marine environments across the world such as deepwater caves or[7][8] brackish and fresh shallow waters,[9] and can exist as solitary or colonial.[citation needed] Benthic polyps can be found on a variety of hard substrates, including both natural and artificial surfaces. Many of them live on other organisms such as fish, tunicates, algae, and crustaceans. Furthermore, they prefer not to settle on sand or similarly textured surfaces unless fauna or flora is present.[10]

Because Hydroidomedusian polyps often settle on other organisms, they are also subject to partake in symbiotic relationships.[11][12] For example, the bivalve mantle cavity of a mollusk provides a sheltered environment, transporting food due to the current. In exchange, the hydroid protects against intruders.[13]

Diet

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Hydroidolina are carnivorous suspension feeders.[citation needed] Motile medusa use their cnidocytes and tentacles to capture prey.[citation needed]

Anatomy and morphology

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Cnidarians are united by the common characteristic of having a specialized cell called a cnidocyte, which contains an explosive organelle called a cnidocyst, or stinging cell.[1] In Hydrozoans, the cnidocysts are formed from interstitial stem cells in the ectoderm[14] and are used for prey capture and anti-predator defense.[15]

Cnidarians are known to occur in two body forms: the polyp form which is benthic and “stalk-like,” and the medusae form, which is commonly known as the “bell” form.[4]

Polyp forms are sessile as adults, with a single opening (the mouth/anus) to the digestive cavity facing up with tentacles surrounding it. Medusa forms are motile, with the mouth and tentacles hanging down from an umbrella-shaped bell.[4]

Though some outlier Hydrozoans go through a polyploid (polyp) and medusa stage, Hydroidolina, which comprises almost all hydrozoans, goes through an asexual polypoid stage where the polyp fixed to a substrate and a sexual hydroid stage varying from free-swimming medusa to a gonophore that remains attached to the polyp.[4][16]

An important characteristic of the Hydroidolina is the presence and formation of an exoskeleton.[17] The exoskeleton varies in chemical composition, structural rigidity, thickness, and coverage within the different regions of the colony and protects the coenosarc of the polypoid stage. It originates as epidermal secretions, with the exosarc being produced first by glandular epidermal cells. The exoskeleton can either be bilayered and contain both the exosarc (outer layer) and perisarc (inner layer) or corneous (just perisarc). The exoskeleton contains anchoring structures such as desmocytes and "perisarc extensions."[17]

Life cycle and reproduction

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The Hydroidolina follows a biphasic life cycle, which alternates in occurrence as planula larva, asexual colonial sessile polyps and free-swimming sexual medusa, not all of which may be present in the one life cycle of the Hydroidolina.[4]

Within its benthic phase, polyps of these hydroids attach to soft tissues on organisms, such as the mantle of a mollusk, and reproduce asexually by budding[18][19][20]

In the sexual medusa stage, gonophores, which are the reproductive organ that produces gametes, and will stay attached to the polyp as a reduced medusa stage but will sometimes, often rarely, form to become their own medusae.[21]

Taxonomy

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Alternate classifications

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Other hydrozoan classifications, which are beset by paraphyly however, are still often seen. They do not unite the Leptolinae in a monophyletic taxon and thus do not have any merit according to modern understanding of hydrozoan phylogeny. The alternate name Leptolinae (or Leptolina) was used in older sources for Hydroidolina.

The obsolete name Hydroida was used for a paraphyletic grouping that is now considered synonymous with Hydroidolina but did not include the colonial jellies of the order Siphonophorae.

Ecological Impact

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The complexity of fauna environments in shallow and deep waters is only increased by benthic polyp colonization. These hydroid colonies affect many spatial and temporal settlement patterns of other benthic species due to providing a habitat for a wide variety of organisms, thus promoting species richness and abundance.[22][23]

These sessile invertebrates could prove to be useful as a measure of environmental changes within their own colonies as well as for changes within near marine environments pertaining to temporal and spatial changes to species distribution and composition, temperature, and food.[24][25]

References

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  1. ^ a b Kayal E, Bentlage B, Cartwright P, Yanagihara AA, Lindsay DJ, Hopcroft RR, Collins AG (November 2015). "Phylogenetic analysis of higher-level relationships within Hydroidolina (Cnidaria: Hydrozoa) using mitochondrial genome data and insight into their mitochondrial transcription". PeerJ. 3: e1403. doi:10.7717/peerj.1403. PMC 4655093. PMID 26618080.
  2. ^ Collins AG, Schuchert P, Marques AC, Jankowski T, Medina M, Schierwater B (February 2006). Collins T (ed.). "Medusozoan phylogeny and character evolution clarified by new large and small subunit rDNA data and an assessment of the utility of phylogenetic mixture models". Systematic Biology. 55 (1): 97–115. doi:10.1080/10635150500433615. PMID 16507527. S2CID 15523821.
  3. ^ Bentlage B, Collins AG (September 2021). "Tackling the phylogenetic conundrum of Hydroidolina (Cnidaria: Medusozoa: Hydrozoa) by assessing competing tree topologies with targeted high-throughput sequencing". PeerJ. 9: e12104. doi:10.7717/peerj.12104. PMC 8435201. PMID 34589302.
  4. ^ a b c d e f "Phylum Cnidaria – Biology 2e". opentextbc.ca. Retrieved 2022-04-26.
  5. ^ Nagale P, Apte D (2014). "Intertidal hydroids (Cnidaria: Hydrozoa: Hydroidolina) from the Gulf of Kutch, Gujarat, India". Marine Biodiversity Records. 7: e116. doi:10.1017/S1755267214001146. ISSN 1755-2672.
  6. ^ "WoRMS - World Register of Marine Species - Hydroidolina". marinespecies.org. Retrieved 2018-03-15.
  7. ^ Daly M, Brugler MR, Cartwright P, Collins AG, Dawson MN, Fautin DG, et al. (December 2007). "The phylum Cnidaria: A review of phylogenetic patterns and diversity 300 years after Linnaeus". Zootaxa. 1668 (1): 127–182. doi:10.11646/zootaxa.1668.1.11. hdl:1808/13641. ISSN 1175-5334.
  8. ^ Collins AG (April 2002). "Phylogeny of Medusozoa and the evolution of cnidarian life cycles". Journal of Evolutionary Biology. 15 (3): 418–432. doi:10.1046/j.1420-9101.2002.00403.x. ISSN 1010-061X. S2CID 11108911.
  9. ^ Graf DL. "Subclass Hydroidolina". WInvertebrates. University of Wisconsin - Stevens Point. Retrieved 2022-04-26.
  10. ^ Calder DR (January 1991). "Associations between hydroid species assemblages and substrate types in the mangal at Twin Cays, Belize". Canadian Journal of Zoology. 69 (8): 2067–2074. doi:10.1139/z91-288. ISSN 0008-4301.
  11. ^ Piraino S, Todaro C, Geraci S, Boero F (March 1994). "Ecology of the bivalve-inhabiting hydroid Eugymnanthea inquilina in the coastal sounds of Taranto (Ionian Sea, SE Italy)". Marine Biology. 118 (4): 695–703. doi:10.1007/bf00347518. ISSN 0025-3162. S2CID 84762351.
  12. ^ Lemer S, Giribet G (November 2014). "Occurrence of a bivalve-inhabiting marine hydrozoan (Hydrozoa: Hydroidolina: Leptothecata) in the amber pen-shell Pinna carnea Gmelin, 1791 (Bivalvia: Pteriomorphia: Pinnidae) from Bocas del Toro, Panama". Journal of Molluscan Studies. 80 (4): 464–468. doi:10.1093/mollus/eyu059. ISSN 1464-3766.
  13. ^ Rees WJ (April 1967). "A brief survey of the symbiotic associations of Cnidaria with Mollusca". Journal of Molluscan Studies. 37 (4): 213–231. doi:10.1093/oxfordjournals.mollus.a064991.
  14. ^ Gold DA, Lau CL, Fuong H, Kao G, Hartenstein V, Jacobs DK (March 2019). "Mechanisms of cnidocyte development in the moon jellyfish Aurelia". Evolution & Development. 21 (2): 72–81. doi:10.1111/ede.12278. PMC 7211293. PMID 30623570.
  15. ^ Sunagar K, Columbus-Shenkar YY, Fridrich A, Gutkovich N, Aharoni R, Moran Y (September 2018). "Cell type-specific expression profiling unravels the development and evolution of stinging cells in sea anemone". BMC Biology. 16 (1): 108. doi:10.1186/s12915-018-0578-4. PMC 6161364. PMID 30261880.
  16. ^ Hedgpeth JW (April 1954). "The Medusae of the British Isles: Anthomedusae, Leptomedusae, Limnomedusae, Trachymedusae, and Narcomedusae. E. T. Browne Monograph of the Marine Biological Association of the United Kingdom. Frederick Stratten Russell. Cambridge Univ. Press, New York, 1953. 530 pp. Illus. + 35 plates. $22.50". Science. 119 (3095): 562. doi:10.1126/science.119.3095.562. ISSN 0036-8075.
  17. ^ a b Mendoza-Becerril MD (2016). Padrões de diversificação de Bougainvilliidae no contexto evolutivo de Medusozoa (Cnidaria) (Ph.D. thesis). Universidade de Sao Paulo, Agencia USP de Gestao da Informacao Academica (AGUIA). doi:10.11606/t.41.2016.tde-04112015-142910.
  18. ^ Migotto AE, Caobelli JF, Kubota S (November 2004). "Redescription and life cycle of Eutima sapinhoa Narchi and Hebling, (Cnidaria: Hydrozoa, Leptothecata): a hydroid commensal with Tivela mactroides (Born) (Mollusca, Bivalvia, Veneridae)". Journal of Natural History. 38 (20): 2533–2545. doi:10.1080/00222930310001647316. ISSN 0022-2933. S2CID 85268652.
  19. ^ Piraino S, Todaro C, Geraci S, Boero F (March 1994). "Ecology of the bivalve-inhabiting hydroid Eugymnanthea inquilina in the coastal sounds of Taranto (Ionian Sea, SE Italy)". Marine Biology. 118 (4): 695–703. doi:10.1007/bf00347518. ISSN 0025-3162. S2CID 84762351.
  20. ^ Kubota S (December 2000). "Parallel, paedomorphic evolutionary processes of the bivalve-inhabiting hydrozoans (Leptomedusae, Eirenidae) deduced from the morphology, life cycle and biogeography, with special reference to taxonomic treatment of Eugymnanthea". Scientia Marina. 64 (S1): 241–247. doi:10.3989/scimar.2000.64s1241. hdl:2433/240921. ISSN 1886-8134.
  21. ^ Prudkovsky AA, Ekimova IA, Neretina TV (October 2019). "A case of nascent speciation: unique polymorphism of gonophores within hydrozoan Sarsia lovenii". Scientific Reports. 9 (1): 15567. Bibcode:2019NatSR...915567P. doi:10.1038/s41598-019-52026-7. PMC 6820802. PMID 31664107.
  22. ^ Blanco R, Shields MA, Jamieson AJ (December 2013). "Macrofouling of deep-sea instrumentation after three years at 3690m depth in the Charlie Gibbs fracture zone, mid-Atlantic ridge, with emphasis on hydroids (Cnidaria: Hydrozoa)". Deep Sea Research Part II: Topical Studies in Oceanography. 98: 370–373. Bibcode:2013DSRII..98..370B. doi:10.1016/j.dsr2.2013.01.019. ISSN 0967-0645.
  23. ^ Bradshaw C, Collins P, Brand AR (October 2003). "To what extent does upright sessile epifauna affect benthic biodiversity and community composition?". Marine Biology. 143 (4): 783–791. doi:10.1007/s00227-003-1115-7. ISSN 0025-3162. S2CID 86250065.
  24. ^ Topcu NE, Martell LF, Yilmaz IN, Isinibilir M (2018-06-18). "Benthic Hydrozoans as Potential Indicators of Water Masses and Anthropogenic Impact in the Sea of Marmara". Mediterranean Marine Science: 273. doi:10.12681/mms.15117. ISSN 1791-6763. S2CID 92144750.
  25. ^ Ronowicz M, Kukliński P, Mapstone GM (2015-03-20). "Trends in the diversity, distribution and life history strategy of Arctic Hydrozoa (Cnidaria)". PLOS ONE. 10 (3): e0120204. Bibcode:2015PLoSO..1020204R. doi:10.1371/journal.pone.0120204. PMC 4368823. PMID 25793294.
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