centrarchidae

As their name suggests, sunfishes are most closely related to the pygmy sunfishes (Elassomatidae), which also occur in North America. Together, these two families are sister to the Chinese perches (Sinipercidae) of East Asia. These three families are among the few temperate Northern Hemisphere members of the order Centrarchiformes (primarily found in the Southern Hemisphere and the tropics), and share the suborder Centrarchoidei with a group of temperate Southern Hemisphere fishes that include the jutjaw, oldwife, and the Australian temperate perches.{{Cite journal |last=Lavoué |first=Sébastien |last2=Nakayama |first2=Kouji |last3=Jerry |first3=Dean R. |last4=Yamanoue |first4=Yusuke |last5=Yagishita |first5=Naoki |last6=Suzuki |first6=Nobuaki |last7=Nishida |first7=Mutsumi |last8=Miya |first8=Masaki |date=2014-10-01 |title=Mitogenomic phylogeny of the Percichthyidae and Centrarchiformes (Percomorphaceae): comparison with recent nuclear gene-based studies and simultaneous analysis |url=https://linkinghub.elsevier.com/retrieve/pii/S0378111914008166 |journal=Gene |volume=549 |issue=1 |pages=46–57 |doi=10.1016/j.gene.2014.07.033 |issn=0378-1119}}

The earliest fossils of Centrarchidae are from latest Eocene to early Oligocene deposits from Montana and South Dakota, belonging to several as yet undescribed species. Two extinct genera are also known: †Plioplarchus from the Early Oligocene to the Middle Miocene of the Dakotas & Oregon, and †Boreocentrarchus from the Late Oligocene or Early Miocene of southern Alaska (making it one of the northernmost known members of the family). Both Plioplarchus and Boreocentrarchus are classified in the subfamily Centrarchinae, because these species possess more than three anal fin spines.{{Cite book |url=https://onlinelibrary.wiley.com/doi/book/10.1002/9781444316032 |title=Centrarchid Fishes |date=2009-05-28 |publisher=Wiley |isbn=978-1-4051-3342-5 |editor-last=Cooke |editor-first=S. J. |language=en |doi=10.1002/9781444316032 |editor-last2=Philipp |editor-first2=D. P.}}

The family is classified as follows:{{Cite web |title=CAS - Eschmeyer's Catalog of Fishes: |url=https://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp |access-date=2025-04-09 |website=researcharchive.calacademy.org |language=en}}

• Family Centrarchidae
• Subfamily Centrarchinae
• Acantharchus Gill, 1864
• Ambloplites Rafinesque, 1820
• Archoplites Gill, 1861
• Centrarchus Cuvier, 1829
• Enneacanthus Gill, 1864
• Pomoxis Rafinesque, 1818
• †Boreocentrarchus Schlaikjer, 1937 (fossil; late Oligocene/early Miocene of Alaska)
• †Plioplarchus Cope, 1883 (fossil; Oligocene to Miocene of western North America)
• Subfamily Lepominae
• Lepomis Rafinesque, 1816
• Micropterus Lacépède, 1802

Recent genetic evidence suggests the following taxonomy of the centrarchid genera and species:{{cite journal |last1=Roe |first1=Kevin J. |last2=Harris |first2=Phillip M. |last3=Mayden |first3=Richard L. |year=2002 |title=Phylogenetic relationships of the genera of North American sunfishes and basses (Percoidei: Centrarchidae) as evidenced by the mitochondrial cytochrome b gene |url=http://specifyassets.nhm.ku.edu/Ichthyology/originals/sp67800263772931021157.att.pdf |url-status=dead |journal=Copeia |volume=2002 |issue=4 |pages=897–905 |doi=10.1643/0045-8511(2002)002[0897:protgo]2.0.co;2 |s2cid=86167841 |archive-url=https://web.archive.org/web/20170914124819/http://specifyassets.nhm.ku.edu/Ichthyology/originals/sp67800263772931021157.att.pdf |archive-date=2017-09-14}}{{cite journal |author=Near, T. J. |author2=D. I. Bolnick |author3=P. C. Wainwright |name-list-style=amp |date=2005 |title=Fossil calibrations and molecular divergence time estimates in centrarchid fishes (Teleostei: Centrarchidae) |journal=Evolution |volume=59 |issue=8 |pages=1768–1782 |doi=10.1111/j.0014-3820.2005.tb01825.x |pmid=16329246 |doi-access=free}}{{clade| style=font-size:85%;line-height:80%

|label1=Centrarchidae

|1={{clade

|1=Acantharchus pomotis 70 px

|2={{clade

|label1=Centrarchinae

|1={{clade

|1={{clade

|label1=Centrarchus

|1=Centrarchus macropterus 70 px}}

|2={{clade

|label1=Enneacanthus

|1={{clade

|1={{clade

|1=Enneacanthus chaetodon

|2={{clade

|1=Enneacanthus gloriosus

|2=Enneacanthus obesus }} }} }}

|label2=Tribe Archoplitini

|2={{clade

|label1=Pomoxis (crappies)

|1={{clade

|1=Pomoxis annularis 70 px

|2=Pomoxis nigromaculatus 70 px}}

|2={{clade

|label1=Archoplites

|1={{clade

|1=Archoplites interruptus}}

|label2=Ambloplites (rock basses)

|2={{clade

|1={{clade

|1=Ambloplites ariommus

|2=Ambloplites rupestris 70 px}}

|2={{clade

|1=Ambloplites cavifrons

|2=Ambloplites constellatus }} }} }} }} }} }}

|label2=Lepominae

|2={{clade

|label1=Micropterus (black basses)

|1={{clade

|1={{clade

|1=Micropterus dolomieu 70 px

|2=Micropterus punctulatus}}

|2={{clade

|1=Micropterus cataractae

|2={{clade

|1=Micropterus coosae

|2={{clade

|1=Micropterus notius 70 px

|2={{clade

|1=Micropterus treculi

|2={{clade

|1=Micropterus salmoides 70 px

|2=Micropterus floridanus}} }} }} }} }} }}

|label2=Lepomis (true sunfish)

|2={{clade

|1={{clade

|1={{clade

|1=Lepomis humilis 70 px

|2=Lepomis macrochirus 70 px}}

|2={{clade

|1={{clade

|1=Lepomis gulosus 70 px}}

|2={{clade

|1=Lepomis symmetricus

|2=Lepomis cyanellus 70 px}} }} }}

|2={{clade

|1={{clade

|1={{clade

|1=Lepomis auritus 70 px

|2={{clade

|1=Lepomis marginatus

|2={{clade

|1=Lepomis peltastes

|2=Lepomis megalotis 70 px}} }} }}

|2={{clade

|1=Lepomis gibbosus 70 px

|2={{clade

|1=Lepomis microlophus 70 px

|2={{clade

|1=Lepomis punctatus

|2=Lepomis miniatus }} }} }} }} }} }} }} }} }} }}

The 5th edition of Fishes of the World recognises three subfamilies; the Lepominae, which contains the genera Acantharchus, Lepomis, and Micropterus; the Elassomatinae, which consists solely of the highly distinct genus Elassoma (treated by Fishbase as a separate family, the Elassomatidae);{{FishBase family|family=Elassomatidae|month=December|year=2019}} and the Centrarchinae, which contains all of the remaining genera.{{cite book |author1=J. S. Nelson |url=https://sites.google.com/site/fotw5th/ |title=Fishes of the World |author2=T. C. Grande |author3=M. V. H. Wilson |publisher=Wiley |year=2016 |isbn=978-1-118-34233-6 |edition=5th |pages=444–446 |access-date=2020-05-18 |archive-url=https://web.archive.org/web/20190408194051/https://sites.google.com/site/fotw5th/ |archive-date=2019-04-08 |url-status=dead}} The more recent Eschmeyer's Catalog of Fishes again treats Elassomatidae as a distinct family, with Centrarchidae composed of Lepominae and Centrarchinae.

Family members are distinguished by having a laterally compressed body shape, 6 to 9 anal spines, and 2 dorsal fins (spinous first dorsal and rayed second dorsal) which are fused.{{Cite book |title=Peterson field guide to freshwater fishes of North America north of Mexico |date=2011 |publisher=Houghton Mifflin Harcourt |last2=Burr |first2=Brooks M. |last1=Page |first1=Lawrence M. |isbn=9780547242064 |edition=2nd |location=Boston |oclc=651912235}} The number of dorsal spines varies from 6 to 13. All species in Micropterus and Lepomis have 3 anal spines, which distinguishes them from the other genera in the family. The pseudobranch is small and concealed. Body size varies widely within the family with the black-banded sunfish at just {{convert|8|cm|in|abbr=on}} in length, while the largemouth bass is reported to reach almost {{convert|1|m|ft|abbr=on}} in extreme cases.{{cite book|title=Encyclopedia of Fishes|author1=Johnson, G.D.|author2=Gill, A.C.|publisher=Academic Press|year=1998|isbn=978-0-12-547665-2|editor=Paxton, J.R. |location=San Diego|pages=187|editor2=Eschmeyer, W.N.}}

Many of the species within Centrarchidae can be separated into two main groups based on the two most common genera (Micropterus and Lepomis). Species in the genera Lepomis are defined by a deep or more round body shape, smaller mouths, and obtaining food through suction feeding.{{Cite web |last1=Soes |first1=Menno |last2=Cooke |first2=Steven |last3=van Kleef |first3=H.H. |last4=Broeckx |first4=P.B. |last5=Veenvliet |first5=P. |date=March 21, 2010 |title=A risk analysis of sunfishes (Centrarchidae) and pygmy sunfishes (Elassomatidae) in the Netherlands |url=https://www.researchgate.net/publication/276255387 |publisher=Bureau of Waardenburg bv |id=Report nr 11-042 |pages=1–110 |via=ResearchGate}} Species in the genera Micropterus are defined by a more streamlined body shape, larger mouths, and consuming prey primarily by ram feeding methods.

File:Micropterus dolomieu2.jpg (Micropterus dolomieu)]]

Centrarchids prefer clear, warm, and slower-moving water, and are commonly found in habitats such as lakes, ponds, medium to low flow streams and rivers, and swamps.{{Cite book|title=Freshwater Fish Distribution|last=Berra|first=Tim|publisher=The University of Chicago Press|year=2007|isbn=978-0-226-04442-2|location=Chicago and London|pages=390–400}} They also prefer to live in and around aquatic vegetation so they can get adequate coverage from predators. While few species in the family diverge from the aforementioned habitat list, the Sacramento perch can survive in habitats with unusually high alkalinity, salinity, and temperatures. Centrarchids can be found in various locations within the water column and their exact preference is species specific. For instance, bluegill (Lepomis macrochirus) mainly inhabit the deeper littoral zones, while green sunfish (Lepomis cyanellus) prefer habitats near the shoreline and shallower areas.{{Cite journal|last1=Werner|first1=Earl E.|last2=Hall|first2=Donald J.|date=1977-07-01|title=Competition and Habitat Shift in Two Sunfishes (Centrarchidae)|journal=Ecology|language=en|volume=58|issue=4|pages=869–876|doi=10.2307/1936222|issn=1939-9170|jstor=1936222|bibcode=1977Ecol...58..869W }} Suction feeders within the family (i.e. Lepomis) generally feed off the bottom of their habitat, while ram feeders (i.e. Micropterus) generally feed in more open areas known as the limnetic zone.{{Cite journal|last1=Carroll|first1=Andrew M.|last2=Wainwright|first2=Peter C.|last3=Huskey|first3=Stephen H.|last4=Collar|first4=David C.|last5=Turingan|first5=Ralph G.|date=2004-10-15|title=Morphology predicts suction feeding performance in centrarchid fishes|journal=Journal of Experimental Biology|language=en|volume=207|issue=22|pages=3873–3881|doi=10.1242/jeb.01227|issn=0022-0949|pmid=15472018|doi-access=free}} Centrarchids diet consists of macro-invertebrates (e.g. insects, snails and crayfish) and other fish found in their habitat.

In freshwater systems, water temperature is determined by many abiotic factors, with air temperature being one of the most significant contributors.{{Cite journal|last1=Lyons|first1=J.|last2=Stewart|first2=J. S.|last3=Mitro|first3=M.|date=2010-11-01|title=Predicted effects of climate warming on the distribution of 50 stream fishes in Wisconsin, U.S.A.|journal=Journal of Fish Biology|language=en|volume=77|issue=8|pages=1867–1898|doi=10.1111/j.1095-8649.2010.02763.x|pmid=21078096|bibcode=2010JFBio..77.1867L |issn=1095-8649}} As in other ectotherms, many physiological processes and behaviors in Centrarchidae, such as feeding and reproduction, are heavily impacted by the temperature in their environment.{{Cite journal|last1=Shuter|first1=B. J.|last2=Finstad|first2=A. G.|last3=Helland|first3=I. P.|last4=Zweimüller|first4=I.|last5=Hölker|first5=F.|date=2012-10-01|title=The role of winter phenology in shaping the ecology of freshwater fish and their sensitivities to climate change|journal=Aquatic Sciences|language=en|volume=74|issue=4|pages=637–657|doi=10.1007/s00027-012-0274-3|bibcode=2012AqSci..74..637S |s2cid=18297554|issn=1015-1621}} All species in the family Centrarchidae are considered warmwater adapted species.{{Cite journal|last1=Carpenter|first1=Stephen R.|last2=Fisher|first2=Stuart G.|last3=Grimm|first3=Nancy B.|author-link3=Nancy Grimm|last4=Kitchell|first4=James F.|date=1992-11-01|title=Global Change and Freshwater Ecosystems|journal=Annual Review of Ecology and Systematics|volume=23|issue=1|pages=119–139|doi=10.1146/annurev.es.23.110192.001003|issn=0066-4162}} In general, warmwater adapted species are characterized as being larger at higher temperatures and lower latitudes.{{Cite journal|last=Rypel|first=Andrew L.|date=2014-01-01|title=The Cold-Water Connection: Bergmann's Rule in North American Freshwater Fishes.|journal=The American Naturalist|volume=183|issue=1|pages=147–156|doi=10.1086/674094|pmid=24334744|s2cid=22642325|issn=0003-0147}} The optimal temperature range of most species in the family is 28 °C(82 °F) to 32 °C(90 °F), although they can survive and reproduce in temperatures that are outside of this optimum range. Increases in temperature outside the optimal range for centrarchids can have negative effects, such as speeding up reproductive maturity or increasing mortality after the first reproductive event.{{Cite journal |last1=Dembski |first1=S. |last2=Masson |first2=G. |last3=Monnier |first3=D. |last4=Wagner |first4=P. |last5=Pihan|first5=J. C.|date=2006-08-01|title=Consequences of elevated temperatures on life-history traits of an introduced fish, pumpkinseed Lepomis gibbosus|journal=Journal of Fish Biology|language=en|volume=69|issue=2|pages=331–346 |doi=10.1111/j.1095-8649.2006.01087.x |bibcode=2006JFBio..69..331D |issn=1095-8649}} The lethal temperature range varies widely in the family, but some species have been seen to survive water temperatures as low as 1 °C(33 °F) or as high as 41 °C(106 °F).{{Cite journal|last1=Beitinger|first1=Thomas L.|last2=Bennett|first2=Wayne A.|last3=McCauley|first3=Robert W.|date=2000-07-01|title=Temperature Tolerances of North American Freshwater Fishes Exposed to Dynamic Changes in Temperature|journal=Environmental Biology of Fishes|language=en|volume=58|issue=3|pages=237–275|doi=10.1023/A:1007676325825|bibcode=2000EnvBF..58..237B |s2cid=35400804|issn=0378-1909}}

Centrarchids generally spawn in the spring, and juveniles emerge in the late spring to early summer.{{Cite news|url=http://z0ku333mvy924cayk1kta4r1-wpengine.netdna-ssl.com/wp-content/uploads/LakeLine/34-4/Full/34-4Full.pdf |archive-url=https://web.archive.org/web/20180403173502/http://z0ku333mvy924cayk1kta4r1-wpengine.netdna-ssl.com/wp-content/uploads/LakeLine/34-4/Full/34-4Full.pdf |archive-date=2018-04-03 |url-status=live|title=Fish in Winter – Changes in Latitudes, Changes in Attitudes|last=Miranda|first=L.E. (Steve)|date=Winter 2014|work=Lake Line (a publication of the North American Lake Management Society)|issue=4|volume=34|pages=28–31}} The transition from winter to spring conditions (i.e. melting of ice-cover, increase in day length, and increased food availability) is the main cue for centrarchids to begin preparing for reproduction. All species within Centrarchidae, except for those in the genus Micropterus, develop breeding coloration in both males and females (although less defined in females) during the breeding season. The process of courtship and reproduction is nearly identical for all species in the family, which is a major reason for the high levels of hybridization within Centrarchidae.{{Cite journal|last1=Jennings|first1=Martin J.|last2=Philipp|first2=David P.|last3=Montgomery|first3=W. L.|date=2002-12-01|title=Alternative Mating Tactics in Sunfishes (Centrarchidae): A Mechanism for Hybridization?|journal=Copeia|volume=2002|issue=4|pages=1102–1105|doi=10.1643/0045-8511(2002)002[1102:amtisc]2.0.co;2|s2cid=85691602 |issn=0045-8511}} With that said, there are some mechanisms in place to prevent hybridization, such as intricate morphology of the operculum in Lepomis, which assists in recognition of conspecific mates.

To initiate reproduction, males dig a deep circular depression in the substrate with their caudal fins to create a nest, which they will aggressively defend from intruding males. Males and females then undergo a courtship dancing ritual before the female deposits her eggs into the male's nest. Multiple females may deposit eggs in a single nest. Larger males usually attract more mates and also take better care of their offspring.{{Cite journal|last1=Danylchuk|first1=Andy J.|last2=Fox|first2=Michael G.|date=1996-10-01|title=Size- and age-related variation in the seasonal timing of nesting activity, nest characteristics, and female choice of parental male pumpkinseed sunfish (Lepomis gibbosus)|journal=Canadian Journal of Zoology|volume=74|issue=10|pages=1834–1840|doi=10.1139/z96-206|issn=0008-4301}} Male parental care includes nest building, nest guarding, guarding of eggs and fry, and nest fanning (aerating eggs).{{Cite journal|last=Blumer|first=Lawrence S.|date=1982-05-01|title=A bibliography and categorization of bony fishes exhibiting parental care|journal=Zoological Journal of the Linnean Society|language=en|volume=75|issue=1|pages=1–22|doi=10.1111/j.1096-3642.1982.tb01939.x|issn=0024-4082|hdl=2027.42/71841|hdl-access=free}}

Males unsuccessful at courtship may exhibit a cheater strategy where they sneak fertilizations of female's eggs by various behavioral methods.{{Cite journal|last1=Gross|first1=Mart R.|last2=Charnov|first2=Eric L.|date=1980-11-01|title=Alternative male life histories in bluegill sunfish|journal=Proceedings of the National Academy of Sciences|volume=77|issue=11|pages=6937–6940|doi=10.1073/pnas.77.11.6937|pmid=16592922|pmc=350407|bibcode=1980PNAS...77.6937G|doi-access=free}} This is commonly seen with smaller males in the genus Lepomis.

File:Centrarchidae_dis.png

The native range of Centrarchidae is confined within North America, covering most of the United States and stopping in southern Canada. The northern edge of the native range is heavily bound by temperature due to reduced foraging ability and growth in cold weather and subsequent starvation in winter months.{{Cite journal|last1=Alofs|first1=Karen M.|last2=Jackson|first2=Donald A. |last3=Lester |first3=Nigel P.|date=2014-02-01|title=Ontario freshwater fishes demonstrate differing range-boundary shifts in a warming climate |journal=Diversity and Distributions|language=en|volume=20|issue=2|pages=123–136 |doi=10.1111/ddi.12130 |bibcode= 2014DivDi..20..123A |s2cid=85651008 |issn=1472-4642|doi-access=free}} As a result, centrarchid distributions and range in any place they are found will be restricted by cold temperatures.

The ability to adapt to cold temperatures at the edge of the sunfish range varies widely within the family. Largemouth bass (Micropterus salmoides) have no cold acclimation ability as seen through the strict maintenance of the northern boundary of the species range. Other species like smallmouth bass (Micropterus dolomieu) and green sunfish (Lepomis cyanellus) have exhibited signs of minor cold-water adaptation and have even experienced slight range expansions into colder habitats.{{Cite journal|last1=Tschantz|first1=Deidra R.|last2=Crockett|first2=Elizabeth L.|last3=Niewiarowski|first3=Peter H.|last4=Londraville|first4=Richard L.|date=2002-11-01|title=Cold Acclimation Strategy Is Highly Variable among the Sunfishes (Centrarchidae)|journal=Physiological and Biochemical Zoology|volume=75|issue=6|pages=544–556|doi=10.1086/344492|pmid=12601611|s2cid=36858254|issn=1522-2152}}

If air temperatures continue to rise in the next 50 to 100 years as predicted,IPCC, 2014: Climate Change 2014: Synthesis Report. Contribution of Working Groups I, II and III to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change [Core Writing Team, R.K. Pachauri and L.A. Meyer (eds.)]. IPCC, Geneva, Switzerland, 151 pp. https://www.ipcc.ch/report/ar5/ warmwater species like centrarchids will likely experience range expansions northward and see an overall increase in occupiable habitat.{{Cite journal|last1=COMTE|first1=LISE|last2=BUISSON|first2=LAËTITIA|last3=DAUFRESNE|first3=MARTIN|last4=GRENOUILLET|first4=GAËL|date=2013-04-01|title=Climate-induced changes in the distribution of freshwater fish: observed and predicted trends|journal=Freshwater Biology|language=en|volume=58|issue=4|pages=625–639|doi=10.1111/fwb.12081|bibcode=2013FrBio..58..625C |issn=1365-2427|url=https://hal.archives-ouvertes.fr/hal-00992708/file/Comte_11129.pdf |archive-url=https://web.archive.org/web/20190501141112/https://hal.archives-ouvertes.fr/hal-00992708/file/Comte_11129.pdf |archive-date=2019-05-01 |url-status=live|doi-access=free}} This range expansion can have grave consequences for other freshwater fishes however, as many centrarchids are dominant top predators which can severely alter the community structure of non-native ecosystems and drive the extinction of other native predators.{{Cite book|title=Centrarchid Fishes|last1=Near|first1=T. J.|last2=Koppelman|first2=J. B.|publisher=Wiley-Blackwell|pages=1–38|language=en|doi=10.1002/9781444316032.ch1|chapter = Species Diversity, Phylogeny and Phylogeography of Centrarchidae|year = 2009|isbn = 9781444316032}}

While centrarchids are native to only North America, they can be found worldwide due to introductions on multiple continents, including Europe, South America, Africa, and Asia. At least 18 species of Centrarchidae are North American exports. Its multi-continental spread is mostly due to the high popularity of the family (especially from the genera Micropterus) as freshwater game fish that are frequently stocked for recreational fishing all around Europe.

Across the globe, invasive and introduced centrarchids pose a great threat to native species in the areas they invade.{{Cite journal|last=Sterud|first=Erik|title=Pumpkinseed Lepomis gibbosus (Centrarchidae) and associated parasites introduced to Norway|journal=Aquatic Invasions|volume=1|issue=4|pages=278–280|doi=10.3391/ai.2006.1.4.10|year=2006|doi-access=free}} There are multiple confirmed instances of largemouth bass (Micropterus salmoides) severely altering and reducing native fish populations in Italy, South Africa, Japan, and Madagascar and even causing the local extinction of any species of the family Cyprinodontidae within the waterbodies they have invaded in Mexico.

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• {{FishBase_family|family=Centrarchidae|year=2004|month=October}}

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Category:Extant Miocene first appearances

Category:Taxa named by Pieter Bleeker

Category:Centrarchiformes families