Enchodus

{{Automatic taxobox

| fossil_range = Albian-Maastrichtian
~{{Fossil range|105|66|latest=55|earliest=121}} Possible Barremian & Paleogene records

| image = Enchodus Swimming Down.png

| image_caption = E. petrosus mounted skeleton cast, Rocky Mountain Dinosaur Resource Center

| taxon = Enchodus

| authority = Agassiz, 1835

| type_species = {{extinct}}Esox lewesiensis

| type_species_authority = Mantell, 1822

| subdivision_ranks = Species

| subdivision = ~26+, see text

| synonyms = *Isodus Heckel, 1849

Phasganodus Leidy, 1857
Ischyrocephalus von der Marck, 1858
Solenodon Kramberger, 1881 (preocc.)
?Tetheodus Cope, 1874
Holcodon Kramberger, 1885
Eurygnathus Davis, 1887 (preocc.)

| synonyms_ref =

}}

Enchodus (from {{langx|el|ἔγχος}} {{Transliteration|el|enchos}}, 'spear' and {{langx|el|ὀδούς}} {{Transliteration|el|odoús}} 'tooth'){{cite book |last1=Roberts |first1=George |title=An etymological and explanatory dictionary of the terms and language of geology |date=1839 |publisher=Longman, Orme, Brown, Green, & Longmans |location=London |page=55 |url=https://archive.org/details/anetymologicala00robegoog |access-date=30 December 2021 |language=English}} is an extinct genus of aulopiform ray-finned fish related to lancetfish and lizardfish. Species of Enchodus flourished during the Late Cretaceous, where they were a widespread component of marine ecosystems worldwide, and there is some evidence that they may have survived to the Paleocene or Eocene; however, this may just represent reworked Cretaceous material.{{cite journal |last1=Fielitz |first1=Christopher |last2=González-Rodríguez |first2=Katia A. |title=A New Species of Enchodus (aulopiformes: Enchodontidae) from the Cretaceous (albian to Cenomanian) of Zimapán, Hidalgo, México |journal=Journal of Vertebrate Paleontology |date=2010 |volume=30 |issue=5 |pages=1343–1351 |doi=10.1080/02724634.2010.501438 |jstor=40864352 |bibcode=2010JVPal..30.1343F |s2cid=84281080 }}{{cite journal |last1=Rana |first1=R. S. |last2=Kumar |first2=K. |last3=Singh |first3=H. |last4=Rose |first4=K. D. |title=Lower vertebrates from the Late Palaeocene–Earliest Eocene Akli Formation, Giral Lignite Mine, Barmer District, western India |journal=Current Science |date=2005 |volume=89 |issue=9 |pages=1606–1613 |jstor=24110948 }}{{cite journal |last1=Davis |first1=Matthew P. |last2=Fielitz |first2=Christopher |title=Estimating divergence times of lizardfishes and their allies (Euteleostei: Aulopiformes) and the timing of deep-sea adaptations |journal=Molecular Phylogenetics and Evolution |date=December 2010 |volume=57 |issue=3 |pages=1194–1208 |doi=10.1016/j.ympev.2010.09.003 |pmid=20854916 |bibcode=2010MolPE..57.1194D }}

Description

Enchodus species were small to medium in size, with E. zinensis reaching {{convert|172.2|cm}} long.{{cite journal |last1=Díaz-Cruz |first1=Jesús Alberto |last2=Alvarado-Ortega |first2=Jesús |last3=Ramírez-Sánchez |first3=Marcia M. |last4=Bernard |first4=Emma Louise |last5=Allington-Jones |first5=Lu |last6=Graham |first6=Mark |title=Phylogenetic morphometrics, geometric morphometrics and the Mexican fossils to understand evolutionary trends of enchodontid fishes |journal=Journal of South American Earth Sciences |date=November 2021 |volume=111 |pages=103492 |doi=10.1016/j.jsames.2021.103492 |bibcode=2021JSAES.11103492D }} One of the genus' most notable attributes are the large "fangs" at the front of the upper and lower jaws and on the palatine bones, leading to its misleading nickname among fossil hunters and paleoichthyologists, "the saber-toothed herring". These fangs, along with a long sleek body and large eyes, suggest Enchodus was a predatory species.{{cite web|author=Everhart, Mike|date=2013|url=http://www.oceansofkansas.com/Enchodus.html|title=Enchodus sp. - The Sabre-Toothed Fish of the Cretaceous|website=Oceans of Kansas|archive-url=https://web.archive.org/web/20221118171151/http://oceansofkansas.com/enchodus.html|archive-date=November 18, 2022|url-status=live}}

E. petrosus, with standard length around {{convert|76.7|cm}} and sometimes over {{convert|1|m}},{{cite journal |last1=Chida |first1=Mori |last2=Brinkman |first2=Donald B. |last3=Murray |first3=Alison M. |title=A large, new dercetid fish (Teleostei: Aulopiformes) from the Campanian Bearpaw Formation of Alberta, Canada |journal=Cretaceous Research |date=October 2023 |volume=150 |pages=105579 |doi=10.1016/j.cretres.2023.105579 |bibcode=2023CrRes.15005579C |s2cid=258803963 }} is known from common remains coming from the Niobrara Chalk, the Mooreville Chalk Formation, the Pierre Shale, and other geological formations deposited within the Western Interior Seaway and the Mississippi Embayment. Large individuals of this species had fangs measuring over {{convert|6|cm}} in length, giving its skull an appearance somewhat reminiscent of modern deep-sea fishes, such as anglerfish and viperfish. Other species, such as E. parvus, were considerably smaller, measuring only some centimetres (a few inches) long.{{cite book|author=Everhart, M.J.|year=2017|title=Oceans of Kansas, Second Edition: A Natural History of the Western Interior Sea (Life of the Past)|publisher=Indiana University Press|isbn=978-0253026323|pages=117}}

Despite being a formidable predator, remains of Enchodus are commonly found among the stomach contents of larger predators, including sharks, other bony fish, mosasaurs, plesiosaurs and seabirds such as Baptornis advenus.{{citation needed|date=May 2018}}

Distribution

Enchodus fossils have been found all over the world. In North America, Enchodus remains have been recovered from most US states with fossiliferous Late Cretaceous rocks, including Kansas, Nebraska, Colorado, Alabama, Mississippi, Georgia, Tennessee, Wyoming, Texas, California, North Carolina, and New Jersey. Fossils also have been found in the Aguja and El Doctor Formations of Mexico and the Ashville, Vermillion River and Dinosaur Park Formations, and Brown Bed Member of Canada. The taxon is also known from coeval strata in Mexico, South America (Tiupampan Santa Lucía Formation and Maastrichtian El Molino Formation of Bolivia, Paraíba, Pernambuco and Sergipe states of Brazil, as well as Argentina, Chile, and Peru), Africa (Egypt, Morocco, the Congo, Angola, Niger, and Equatorial Guinea), the Middle East (Saudi Arabia, Lebanon, Israel, Palestine and Jordan), Europe (England, France, the Netherlands, Belgium, Spain, Italy, Germany, Sweden, the Czech Republic, Slovenia, Greece, Ukraine{{Cite journal |last=Kovalchuk |first=Oleksandr |last2=Barkaszi |first2=Zoltán |last3=Anfimova |first3=Galina |date=2022-03-31 |title=Records of Enchodus (Teleostei, Aulopiformes) from the Cenomanian of Ukraine in the light of European distribution of enchodontid fishes |url=https://www.schweizerbart.de//papers/njgpa/detail/303/101174/Records_of_Enchodus_Teleostei_Aulopiformes_from_th?af=crossref |journal=Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen |language=en |pages=295–307 |doi=10.1127/njgpa/2022/1049|url-access=subscription }} and Russia), India, and Japan.{{Cite web |title=PBDB Taxon |url=https://paleobiodb.org/classic/basicTaxonInfo?taxon_no=txn:35564 |access-date=2024-12-26 |website=paleobiodb.org}}{{Cite book |last=Geology |first=British Museum (Natural History) Department of |url=https://www.google.com/books/edition/Catalogue_of_the_Fossil_Fishes_in_the_Br/ZtEKAQAAIAAJ? |title=Catalogue of the Fossil Fishes in the British Museum (Natural History): Actinopterygian Teleostomi of the suborders Isospondyli (in part), Ostariophysi, Apodes, Percesoces, Hemibranchii, Acanthopterygii, and Anacanthini |last2=Woodward |first2=Arthur Smith |date=1901 |publisher=order of the Trustees |language=en}} Potentially the latest Enchodus remains are known from the earliest Eocene of Barmer, India. However, it has also been suggested that all post-Cretaceous Enchodus records are just reworked material.

Taxonomy

Species of Enchodus are generally classified into two different clades, the North American and the Mediterranean. It has been proposed that this distinction is the result of several isolated events between the two populations over the Late Cretaceous. The earliest known species is E. zimapanensis from the late Albian or earliest Cenomanian of Mexico. Potentially earlier remains are known from the late Barremian/early Aptian of Brazil (Morro de Chaves Formation), but these specimens are too fragmentary to confidently assign to this genus.{{Cite web |last=Díaz-Cruz|first=Jesús Alberto|last2=Alvarado-Ortega|first2=Jesús|last3=Giles|first3=Sam|date=2020-06-21 |title=A long snout enchodontid fish (Aulopiformes: Enchodontidae) from the Early Cretaceous deposits at the El Chango quarry, Chiapas, southeastern Mexico: A multi-approach study |url=https://palaeo-electronica.org/content/current-in-press-articles/3063-a-long-snout-enchodontid-fish |access-date=2024-12-27 |website=Palaeontologia Electronica |language=en}}

= Species =

File:Enchodus gracilis.jpg

File:Enchodus petrosus 2.jpg

File:Enchodus faujasi.jpg

Enchodus was a diverse, long-lived genus with many species known throughout its temporal and geographic range. The following valid species are known:{{Cite journal |last=Silva |first=Hilda M. A. |last2=Gallo |first2=Valéria |date=2011 |title=Taxonomic review and phylogenetic analysis of Enchodontoidei (Teleostei: Aulopiformes) |url=https://www.scielo.br/j/aabc/a/vPvQZ4v493QvhCLL9wh9zDb/ |journal=Anais da Academia Brasileira de Ciências |language=en |volume=83 |pages=483–511 |doi=10.1590/S0001-37652011000200010 |issn=0001-3765}}[https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=35564 Enchodus] at Fossilworks.org

E. brevis {{small|Chalifa, 1989}} - Cenomanian of the West Bank, Palestine (Amminadav Formation), potentially Lebanon (Sannine Formation)
E. bursauxi {{small|(Arambourg, 1952)}} - Coniacian of Angola (Itombe Formation), Late Campanian of Egypt, Maastrichtian to potentially Danian of Morocco (Ouled Abdoun Basin)
E. dentex {{small|(Heckel, 1856)}} - Cenomanian of Slovenia (Komen Limestone)
E. dirus {{small|(Leidy, 1857)}} - Maastrichtian of the United States (Fox Hills Formation of North Dakota, Severn Formation of Maryland), potentially Gavdos, Greece{{Cite journal |last=Cavin |first=Lionel |last2=Alexopoulos |first2=Apostolos |last3=Piuz |first3=André |date=2012-12-01 |title=Late Cretaceous (Maastrichtian) ray-finned fishes from the island of Gavdos, southern Greece, with comments on the evolutionary history of the aulopiform teleost Enchodus |url=https://pubs.geoscienceworld.org/sgf/bsgf/article-abstract/183/6/561/314035/Late-Cretaceous-Maastrichtian-ray-finned-fishes?redirectedFrom=fulltext |journal=Bulletin de la Société Géologique de France |volume=183 |issue=6 |pages=561–572 |doi=10.2113/gssgfbull.183.6.561 |issn=0037-9409|url-access=subscription }}
E. elegans {{small|Dartevelle & Casier 1949}} - Coniacian of Angola (Itombe Formation), Maastrichtian of Brazil (Gramame Formation), Niger, Syria, and Jordan (Alhisa Phosphorite Formation); Maastrichtian to potentially Danian of Morocco (Ouled Abdoun Basin)
E. faujasi {{small|Agassiz, 1843}} - Coniacian of Angola (Itombe Formation), Campanian of Israel (Mishash Formation), Maastrichtian of France (Calcarintes du Jadet Formation), Maastrichtian/potentially Danian of the Netherlands (Maastricht Formation){{Cite journal |last=Friedman |first=M. |date=2012-01-01 |title=Ray-finned fishes (Osteichthyes, Actinopterygii) from the type Maastrichtian, the Netherlands and Belgium |url=https://repository.naturalis.nl/pub/428932 |journal=Scripta Geologica. Special Issue |language=en |volume=08 |pages=113–142 |issn=0922-4564}}
E. ferox {{small|Leidy, 1855}} - Santonian of Orenburg, Russia; Campanian to Maastrichtian (potentially Paleocene) of the United States (Marshalltown, Mount Laurel, Navesink, and Hornerstown Formations of New Jersey, Marshalltown Formation of Delaware, Severn Formation of Maryland, Arkansas, Fox Hills Formation of North Dakota); Maastrichtian of Argentina (Jagüel Formation) and India (Intertrappean Beds)
E. gladiolus {{small|(Cope, 1872)}} - Cenomanian to Maastrichtian of the United States (Greenhorn Limestone of Colorado, Kansas & Iowa, Graneros Shale & Carlile Shale of Nebraska, Mancos Shale of New Mexico, Carlile Shale of Kansas, Arkansas, and Merchantville, Navesink & Hornerstown Formations of New Jersey), Santonian to Campanian of Russia (Orenburg, Rybushka Formation), Maastrichtian of Argentina (Jagüel Formation), potentially Peru (Vivian Formation){{Cite journal |last=Gouiric Cavalli |first=Soledad |last2=Cione |first2=Alberto Luis |last3=Tineo |first3=David |last4=Pérez |first4=Leandro Martín |last5=Iribarne |first5=Martín |last6=Allcca Torres |first6=Miguel A. |last7=Poire |first7=Daniel Gustavo |date=2020 |title=The first Peruvian record of Enchodus (Osteichthyes, Aulopiformes, Enchodontidae) in the Upper Cretaceous Vivian Formation |url=https://ri.conicet.gov.ar/handle/11336/133320 |journal=Andean Geology |doi=10.5027/andgeoV48n2-3337 |issn=0718-7092|doi-access=free }}
E. gracilis {{small|(von der Marck, 1858)}} - Campanian of Germany (Ahlen Formation)
E. lewesiensis {{small|(Mantell, 1822)}} (type species) - Cenomanian to Coniacian of England (English Chalk, Seaford Formation), Cenomanian/Turonian of Germany (Hesseltal Formation){{Cite journal |last1=Amalfitano |first1=Jacopo |last2=Giusberti |first2=Luca |last3=Fornaciari |first3=Eliana |last4=Carnevale |first4=Giorgio |date=2020-04-03 |title=UPPER CENOMANIAN FISHES FROM THE BONARELLI LEVEL (OAE2) OF NORTHEASTERN ITALY |url=https://riviste.unimi.it/index.php/RIPS/article/view/13224 |journal=Rivista Italiana di Paleontologia e Stratigrafia |language=en |volume=126 |issue=2 |doi=10.13130/2039-4942/13224 |issn=2039-4942}} and the Czech Republic, potentially Maastrichtian of Germany (Gerhardsreit Formation)
E. libycus (Quaas, 1902) - Cenomanian to Maastrichtian of Brazil (Cotinguiba Formation, Gramame Formation), Campanian of Egypt, Maastrichtian to potentially Danian of Morocco (Ouled Abdoun Basin){{Cite web |title=Enchodus libycus |url=http://mundofosil.net/museo_virtual/fichas/chordata/enchodus_libycus.htm |url-status=dead |archive-url=https://web.archive.org/web/20141114004617/http://mundofosil.net/museo_virtual/fichas/chordata/enchodus_libycus.htm |archive-date=2014-11-14 |access-date=2015-05-13 |work=Mundo Fosil}}
E. longidens {{small|(Pictet, 1850)}} - Santonian of Lebanon (Sahel Alma), potentially Paleocene/early Eocene of India (Akli Formation)
E. longipectoralis {{small|(Schaeffer, 1947)}} - Cenomanian to Coniacian of Brazil (Cotinguiba Formation)
E. lycodon Kner, 1867 - Cenomanian of Slovenia (Komen Limestone)
E. macropterus (von der Marck, 1863) - Campanian of Germany (Baumberge Formation)
E. major {{small|Davis, 1887}} - Santonian of Lebanon (Sahel Alma)
E. marchesettii {{small|(Kramberger, 1895)}} - Cenomanian of Lebanon (Sannine Formation)
E. mecoanalis {{small|Forey et al., 2003}} - Cenomanian of Lebanon (Sannine Formation)
E. oliveirai {{small|Maury, 1930}} - Cenomanian to Maastrichtian of Brazil (Cotinguiba & Gramame Formations)
E. petrosus {{small|Cope, 1874}} - Cenomanian to late Campanian/early Maastrichtian of the United States (Tokio Formation of Arkansas, Carlile Shale of Kansas, Niobrara Formation of South Dakota, Mooreville & Demopolis Chalk of Alabama, Blufftown Formation of Georgia, Tar Heel Formation of North Carolina, Donoho Creek Formation of South Carolina, Navesink Formation of New Jersey), Turonian of Canada (Northwest Territories), Santonian to Campanian of Russia (Orenburg & Rybushka Formation)
E. shumardi {{small|Leidy, 1856}} - Cenomanian to Santonian of the United States (Greenhorn Limestone of Iowa, Kansas & Colorado, Carlile & Graneros Shale of Nebraska & Kansas, Niobrara Formation of Kansas & South Dakota) and Canada (Ashville Formation of Saskatchewan, Kaskapau Formation of Alberta)
E. subaequilateralis {{small|Cope, 1885}} - Maastrichtian of Brazil (Gramame Formation)
E. tineidae {{small|Holloway et al., 2017}} - Campanian of Egypt (Duwi Formation){{Cite journal |author1=Waymon L. Holloway |author2=Kerin M. Claeson |author3=Hesham M. Sallam |author4=Sanaa El-Sayed |author5=Mahmoud Kora |author6=Joseph J.W. Sertich |author7=Patrick M. O’Connor |year=2017 |title=A new species of the neopterygian fish Enchodus from the Duwi Formation, Campanian, Late Cretaceous, Western Desert, central Egypt |journal=Acta Palaeontologica Polonica |volume=62 |issue=3 |pages=603–611 |doi=10.4202/app.00331.2016 |doi-access=free}}
E. venator {{small|Arambourg, 1954}} - Cenomanian of Morocco (Jbel Tselfat), Italy (Scaglia Variegata Alpina Formation), and Germany (Hesseltal Formation)
E. zinensis Chalifa, 1996 - Campanian/Maastrichtian of Egypt
E. zimapanensis {{small|Fielitz & González-Rodríguez, 2010}} - Late Albian/Cenomanian of Mexico (El Doctor Formation){{Cite journal |last1=Fielitz |first1=Christopher |last2=GonzáLez-RodríGuez |first2=Katia A. |date=2010 |title=A new species of Enchodus (Aulopiformes: Enchodontidae) from the Cretaceous (Albian to Cenomanian) of Zimapán, Hidalgo, México |url=https://www.tandfonline.com/doi/full/10.1080/02724634.2010.501438 |journal=Journal of Vertebrate Paleontology |language=en |volume=30 |issue=5 |pages=1343–1351 |bibcode=2010JVPal..30.1343F |doi=10.1080/02724634.2010.501438 |issn=0272-4634|url-access=subscription }}

Many other dubious species based on insufficient remains have been described throughout its range. Even most of the valid Enchodus species are based on only isolated teeth and bones. The genus Parenchodus, considered to be the sister genus of Enchodus, has been synonymized with this genus based on some studies. However, more recent studies have found it to be a valid genus distinct from Enchodus.

= Phylogeny =

{{cladogram|align=center|style=font-size:100%; line-height:100%;width:500px;|title=Enchodus|caption=Phylogeny of the genus with some species|cladogram=

{{clade

|1={{clade

|label1=Enchodus

|1={{clade

|1=E. marchesettii

|2={{clade

|1={{clade

|1=Parenchodus

|2=E. brevis

|3=E. lewesiensis

}}

|2={{clade