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Notothenioidei

{{Short description|Suborder of fishes}}

{{See also|Antarctic fishes}}

{{Automatic taxobox

| fossil_range = {{fossil range|Eocene|present}}

| image = ANTARTIC FISH.png

| image_caption = Six Notothenioidei, illustrations based on "Antarctic Fish and Fisheries" (K.-H. Kock, 1992)

| taxon = Notothenioidei

| authority = Regan, 1913

| subdivision_ranks = Families

| subdivision = see text

}}

Notothenioidei is one of 19 suborders of the order Perciformes. The group is found mainly in Antarctic and Subantarctic waters, with some species ranging north to southern Australia and southern South America.{{Cite book|title=Antarctic Fish Biology: Evolution in a Unique Environment|last=Eastman|first=Joseph|publisher=Academic Press, Inc|year=1993|location=San Diego, California}} Notothenioids constitute approximately 90% of the fish biomass in the continental shelf waters surrounding Antarctica.{{Cite journal|last=Gon, O and Heemstra, PC|date=1992|title=Fishes of the Southern Ocean|journal=The Quarterly Review of Biology|volume=67|pages=220–221}}

Evolution and geographic distribution

The Southern Ocean has supported fish habitats for 400 million years; however, modern notothenioids likely appeared sometime after the Eocene epoch. This period marked the cooling of the Southern Ocean, resulting in the stable, frigid conditions that have persisted to the present day. Another key factor in the evolution of notothenioids is the preponderance of the Antarctic Circumpolar Current (ACC), a large, slow-moving current that extends to the seafloor and precludes most migration to and from the Antarctic region. The earliest known notothenioids are the fossils Proeleginops and Mesetaichthys from the Eocene La Meseta Formation of Seymour Island, the latter of which already shows close similarities with the extant Dissostichus.{{Cite journal |last1=Bieńkowska-Wasiluk |first1=M. |last2=Bonde |first2=N. |last3=Møller |first3=P. R. |last4=Gaździcki |first4=A. |date=2013 |title=Eocene relatives of cod icefishes (Perciformes: Notothenioidei) from Seymour Island, Antarctica |url=http://yadda.icm.edu.pl/baztech/element/bwmeta1.element.baztech-9cc2bb3b-c3ae-4f58-b99d-5845e309953e |journal=Geological Quarterly |language=EN |volume=57 |issue=4 |issn=1641-7291}}

These unique environmental conditions in concert with the key evolutionary innovation of Antifreeze glycoprotein promoted widespread radiation within the suborder, leading to the rapid development of new species.{{Cite journal|last=Clarke, A and Johnston, IA|date=1996|title=Evolution and adaptive radiation of Antarctic fishes|url=http://uahost.uantwerpen.be/funmorph/raoul/fylsyst/Clarke1996.pdf|journal=TREE|volume=11|issue=5|pages=212–218|doi=10.1016/0169-5347(96)10029-x|pmid=21237811|bibcode=1996TEcoE..11..212C }}{{Cite journal|last=Near, TJ, Dornburg, A, Kuhn, KA, Eastman, JT, Pennington, JT, Patarnello, T, Zane, L, Fernández, DA, and Jones, CD|date=2012|title=Ancient climate change, antifreeze, and the evolutionary diversification of antarctic fishes.|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=109|issue=9|pages=3434–9|doi=10.1073/pnas.1115169109|pmid=22331888|pmc=3295276|bibcode=2012PNAS..109.3434N|doi-access=free}} Their adaptive radiation is characterized by depth related diversification. Comparison studies between non-Antarctic and Antarctic species have revealed different ecological processes and genetic differences between the two groups of fish, such as the loss of hemoglobin (in the family Channichthyidae) and changes in buoyancy.

They are distributed mainly throughout the Southern Ocean around the coasts of New Zealand, southern South America, and Antarctica.{{Cite journal|last=Eastman, J and Grande, L|date=1989|title=Evolution of the Antarctic fish fauna with emphasis on the Recent notothenioids|journal= Geological Society, London, Special Publications|volume=47|issue=1|pages=241–252|citeseerx=10.1.1.897.9784|doi=10.1144/GSL.SP.1989.047.01.18|bibcode=1989GSLSP..47..241E|s2cid=84516955}} An estimated 79% of species reside within the Antarctic region. They primarily inhabit seawater temperatures between {{convert|-2|and|4|C|F}}; however, some of the non-Antarctic species inhabit waters that may be as warm as {{convert|10|°C|°F|abbr=on}} around New Zealand and South America.{{Cite web|url=https://sos.noaa.gov/datasets/surface-temperature/|title=Surface Temperature - NOAA's Science On a Sphere|date=2018|website=National Oceanic and Atmospheric Administration}} Seawater temperatures below the freezing point of freshwater (0 °C or 32 °F) are possible due to the greater salinity in the Southern Ocean waters.{{Cite news|url=https://www.science.org/doi/pdf/10.1126/science.1076252|title=The Salinity, Temperature, and δ18O of the Glacial Deep Ocean|last=Adkins, J|display-authors=etal|date=29 November 2002|work=Science Magazine|volume=298 |issue=5599 |pages=1769–1773 |doi=10.1126/science.1076252 |access-date=22 January 2019}} Notothenioids have an estimated depth range of about {{convert|0|–|1,500|m|ft|abbr=on}}.

Anatomy

Notothenioids display a morphology that is largely typical of other coastal perciform fishes. They are not distinguished by a single physical trait, but rather a distinctive set of morphological traits. These include the presence of three flat pectoral fin radials, nostrils located laterally on each side of the head, the lack of a swim bladder, and the presence of multiple lateral lines.

Because notothenioids lack a swim bladder, the majority of species are benthic or demersal in nature. However, a depth-related diversification has given rise to some species attaining increased buoyancy, using lipid deposits in tissues and reduced ossification of bony structures. This reduced ossification of the skeleton (observed in some notothenioids) changes the weight and creates neutral buoyancy in the water, where the fish neither sinks nor floats, and can thus adjust its depth with ease.

Physiology

Notothenioids have a variety of physiological and biochemical adaptations that either permit survival in, or are possible only because of, the generally cold, stable seawater temperatures of the Southern Ocean. These include highly unsaturated membrane lipids{{Cite journal|last=Logue, JA|display-authors=etal|date=2000|title=Lipid compositional correlates of temperature-adaptive interspecific differences in membrane physical structure.|journal=Journal of Experimental Biology|volume=203|issue=Pt 14|pages=2105–2115|doi=10.1242/jeb.203.14.2105|pmid=10862723}} and metabolic compensation in enzymatic activity.{{Cite journal|last=Kawall, HG|display-authors=etal|date=2002|title=Metabolic cold adaptation in Antarctic fishes: Evidence from enzymatic activities of the brain|journal=Marine Biology|volume=140|issue=2|pages=279–286|doi=10.1007/s002270100695|bibcode=2002MarBi.140..279H |s2cid=84943879}} Many notothenoids have lost the nearly universal heat shock response (HSR){{Cite journal|last=Bilyk, KT, Vargas-Chacoff, L, and Cheng CHC|date=2018|title=Evolution in chronic cold: varied loss of cellular response to heat in Antarctic notothenioid fish|journal=BMC Evolutionary Biology|volume=18|issue=1|pages=143|doi=10.1186/s12862-018-1254-6|pmid=30231868|pmc=6146603 |doi-access=free |bibcode=2018BMCEE..18..143B }} due to evolution at cold and stable temperatures.{{Cite journal|last=Beers, JM, and Jayasundara, N.|date=2015|title=Antarctic notothenioid fish: What are the future consequences of 'losses' and 'gains' acquired during long-term evolution at cold and stable temperatures?|journal=The Journal of Experimental Biology|volume=218|issue=Pt 12|pages=1834–45|doi=10.1242/jeb.116129|pmid=26085661|doi-access=free}}

Many notothenioid fishes are able to survive in the freezing, ice-laden waters of the Southern Ocean because of the presence of an antifreeze glycoprotein in blood and body fluids.{{Cite journal|last=Chen, L|display-authors=etal|date=1997|title=Evolution of antifreeze glycoprotein gene from a trypsinogen gene in Antarctic notothenioid fish|journal=PNAS|volume=94|issue=8|pages=3811–3816|doi=10.1073/pnas.94.8.3811|pmid=9108060|pmc=20523|bibcode=1997PNAS...94.3811C|doi-access=free}} Although many of the Antarctic species have antifreeze proteins in their body fluids, not all of them do. Some non-Antarctic species either produce no or very little antifreeze, and antifreeze concentrations in some species are very low in young, larval fish. They also possess aglomerular kidneys, an adaptation that aids the retention of these antifreeze proteins.{{Cite book |last1=Burton |first1=Derek |url=https://academic.oup.com/book/26668 |title=Essential Fish Biology |last2=Burton |first2=Margaret |date=2017-12-21 |publisher=Oxford University Press |isbn=978-0-19-878555-2 |volume=1 |language=en |doi=10.1093/oso/9780198785552.001.0001}}

While the majority of animal species have up to 45% of hemoglobin (or other oxygen-binding and oxygen-transporting pigments) in their blood, the notothenioids of the family Channichthyidae do not express any globin proteins in their blood.{{Cite journal|last=Sidell, B and O'Brien, KM|date=2006|title=When bad things happen to good fish: the loss of hemoglobin and myoglobin expression in Antarctic icefishes|url=http://jeb.biologists.org/content/jexbio/209/10/1791.full.pdf|journal=The Journal of Experimental Biology|volume=209|issue=Pt 10|pages=1791–1802|doi=10.1242/jeb.02091|pmid=16651546|doi-access=free}} As a result, the oxygen-carrying capacity of their blood is reduced to less than 10% that of other fishes. This trait likely arose due to the high oxygen solubility of the Southern Ocean waters. At cold temperatures, the oxygen solubility of water is enhanced.{{Cite web|url=http://omp.gso.uri.edu/ompweb/doee/science/physical/choxy1.htm|title=Dissolved Oxygen|website=University of Rhode Island Office of Marine Programs|access-date=22 January 2019}} The loss of hemoglobin is partially compensated in these species by the presence of a large, slow-beating heart and enlarged blood vessels that transport a large volume of blood under low pressure to enhance cardiac output.{{Cite journal|last=Joyce, W|display-authors=etal|date=2019|title=Adrenergic and Adenosinergic Regulation of the Cardiovascular System in an Antarctic Icefish: Insight into Central and Peripheral Determinants of Cardiac Output|url=http://eprints.whiterose.ac.uk/140852/1/Adrenergic%20and%20Adenosinergic%20Regulation%20of%20the%20Cardiovascular%20System%20in%20an%20Antarctic%20Icefish.pdf|journal=Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology|volume=230|pages=28–38|doi=10.1016/j.cbpa.2018.12.012|pmid=30594528|s2cid=58589102 }} Despite these compensations, the loss of globin proteins still results in reduced physiological performance.

Systematics

=Naming=

Notothenioidei was first described as a separate grouping, as a "division" he named Nototheniiformes, by the British ichthyologist Charles Tate Regan in 1913.,{{cite journal | author = Charles T. Regan | year = 1913 | title = Antarctic fishes of the Scottish National Antarctic expedition | journal = Proceedings of the Royal Society of Edinburgh B | volume = 49 | issue = Part II (Part 2) | url = https://www.biodiversitylibrary.org/item/48647#page/9/mode/1up}} this subsequently has been considered as a suborder of the Percifomes.{{cite journal | author = Gosline, William A. | year = 1968 | title = The Suborders of Perciform Fishes | journal = Proceedings of the United States National Museum | volume = 124 | issue = 3647 | pages = 1–78 | doi = 10.5479/si.00963801.124-3647.1 | url = https://www.biodiversitylibrary.org/page/7510948#page/501/mode/1up}} The name is based on the genus Notothenia, a name coined by Sir John Richardson in 1841 and which means "coming from the south", a reference to the Antarctic distribution of the genus.{{cite web | url = https://etyfish.org/perciformes7/ | title = Order Perciformes: Suborder Notothenoididei: Families Bovichtidae, Pseaudaphritidae, Elegopinidae, Nototheniidae, Harpagiferidae, Artedidraconidae, Bathydraconidae, Channichthyidae and Percophidae | work = The ETYFish Project Fish Name Etymology Database | editor1= Christopher Scharpf | editor2 = Kenneth J. Lazara | name-list-style = amp |date = 12 April 2021 | access-date = 10 September 2021 | publisher = Christopher Scharpf and Kenneth J. Lazara}}

= Families =

This classification follows Eastman and Eakin, 2000{{cite journal | author = J. T. Eastman & R. R. Eakin | title = An updated species list for notothenioid fish (Percifomes; Notothenioidei), with comments on Antarctic species | journal = Arch. Fish. Mar. Res. | volume = 48 | issue = 1 | year = 2000 | pages = 11–20 | url = https://people.ohio.edu/eastman/reprints/058%20Eastman%20%26%20Eakin%202000.pdf | access-date = 2018-05-04 | archive-date = 2018-05-04 | archive-url = https://web.archive.org/web/20180504232017/https://people.ohio.edu/eastman/reprints/058%20Eastman%20%26%20Eakin%202000.pdf | url-status = dead }} and includes references to additional classified species.Last, P.R., A.V. Balushkin and J.B. Hutchins (2002): Halaphritis platycephala (Notothenioidei: Bovichtidae): a new genus and species of temperate icefish from southeastern Australia. Copeia 2002(2):433-440.Froese, Rainer, and Daniel Pauly, eds. (2013). Species of Channichthys in FishBase. February 2013 version. Most species are restricted to the vicinity of Antarctica.

References

Further reading

  • Macdonald, J. A. (2004). "Notothenioidei (Southern Cod-Icefishes)". In M. Hutchins, R. W. Garrison, V. Geist, P. V. Loiselle, N. Schlager, M. C. McDade, ...W. E. Duellman (Eds.), Grzimek's Animal Life Encyclopedia (2nd ed., Vol. 5, pp. 321–329). Detroit: Gale.

{{Taxonbar|from=Q571473}}

Category:Ray-finned fish suborders

Category:Taxa named by Charles Tate Regan