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'{{Short description|Genus of extinct fishes}} {{Automatic taxobox | fossil_range = [[Late Devonian]] ([[Frasnian]] to [[Famennian]]), {{Fossil range|382|358}} | image = Dunkleosteus terrelli (fossil fish) (Cleveland Shale Member, Ohio Shale, Upper Devonian; Rocky River Valley, Cleveland, Ohio, USA) 21 (34001200911).jpg | image_caption = Partially reconstructed ''D. terrelli'' skull (specimen CMNH 5768), [[Cleveland Museum of Natural History]] | display_parents = 7 | taxon = Dunkleosteus | authority = [[Jean-Pierre Lehman|Lehman]], 1956 | type_species = †'''''Dunkleosteus terrelli''''' | type_species_authority = [[John Strong Newberry|Newberry]], 1873 | subdivision_ranks = [[Species]] | subdivision = {{collapsible list| ''D. terrelli'' <small>(Newberry, 1873 [originally ''[[Dinichthys]]''])</small><br /> ''D. ? belgicus'' <small>(Newberry) 1873</small><br /> ''D. denisoni'' <small>(Kulczycki) 1957</small><br /> ''D. marsaisi'' <small>Lehmann, 1956</small><br /> ''D. magnificus'' <small>(Hussakof & Bryant) 1919</small><br /> ''D. missouriensis'' <small>(Branson) 1914</small><br /> ''D. newberryi'' <small>(Clarke) 1885</small><br /> ''D. amblyodoratus'' <small>Carr & Hlavin, 2010</small><br /> ''D. raveri'' <small>Carr & Hlavin, 2010</small><br /> ''D. tuderensis'' <small>Lebedev et. al. 2023</small><br /> }} }} '''''Dunkleosteus''''' is an [[extinct]] [[genus]] of large [[arthrodira|arthrodire]] fish that existed during the [[Late Devonian]] period, about 382–358 [[million years ago]]. It consists of ten species, some of which are among the largest [[placoderm]]s to have ever lived: ''D. terrelli'', ''D. belgicus'', ''D. denisoni'', ''D. marsaisi'', ''D. magnificus'', ''D. missouriensis'', ''D. newberryi'', ''D. amblyodoratus'', ''D. raveri'', and ''D. tuderensis,'' and the largest and most well known species is ''D. terrelli''. Since body shape is not known, various methods of estimation put the living total length of the largest known specimen between {{convert|4.1|to|10|m|ft|abbr=on}} long and weigh around {{convert|1|-|4|MT|ST|abbr=on}}.<ref name=":0" /> ''Dunkleosteus'' could quickly open and close its jaw, like modern-day [[Aquatic feeding mechanisms#Suction feeding|suction feeders]], and had a bite force of {{convert|4414|-|6170|N|kg-f lb-f|0|abbr=on}} at the tip and {{convert|5363|-|7495|N|kg-f lb-f|0|abbr=on}} at the blade edge.<ref name=Anderson&Westneat2007/><ref name=Anderson&Westneat2009/> Numerous [[fossil]]s of the various species have been found in [[North America]], [[Poland]], [[Belgium]], and [[Morocco]]. ''Dunkleosteus'' was a [[pelagic fish]] inhabiting open waters, and an [[apex predator]] of its ecosystem. ==Discovery== ''Dunkleosteus'' fossils were first discovered in 1867 by Jay Terrell, a hotel owner and amateur paleontologist who collected fossils in the cliffs along [[Lake Erie]] near his home of [[Sheffield Lake, Ohio]] (due west of [[Cleveland]]). Terrell donated his fossils to [[John Strong Newberry]] and the Ohio Geological Survey, who in 1873 described all the material as belonging to a single new [[genus]] and [[species]]: ''[[Dinichthys|Dinichthys herzeri]]''. However, with later fossil discoveries by 1875 it became apparent multiple large fish species were present in the [[Ohio Shale]]. ''[[Dinichthys|Dinichthys herzeri]]'' came from the lowermost layer, the [[Huron Shale]], whereas most of the fossils were coming from the younger [[Cleveland Shale]] and represented a distinct species.<ref name=":1" /> Newberry named this more common species "''Dinichthys''" ''terrelli'', after Terrell. Most of Terrell's original collection does not survive, having been destroyed by a fire in [[Elyria, Ohio|Elyria]] in 1873.<ref name=":1" /><ref>{{Cite journal |last=Claypole |first=E. W. |date=1893 |title=The three great fossil placoderms of Ohio |url=https://www.biodiversitylibrary.org/item/240810#page/107/mode/1up |journal=American Geologist |volume=12 |pages=89–99}}</ref> The largest collection of ''Dunkleosteus'' fossils in the world is housed at the [[Cleveland Museum of Natural History]],<ref>{{cite web |url=https://www.cmnh.org/dunk |title=Dunkleosteus terrelli: Fierce prehistoric predator |author=<!--Not stated--> |website=Cleveland Museum of Natural History |access-date=July 21, 2022 |archive-date=May 19, 2021 |archive-url=https://web.archive.org/web/20210519150010/https://www.cmnh.org/dunk |url-status=dead }}</ref> with smaller collections (in descending order of size) held at the [[American Museum of Natural History]],<ref>{{cite web |url=https://www.amnh.org/exhibitions/permanent/vertebrate-origins/dunkleosteus |title=Dunkleosteus |author=<!--Not stated--> |website=American Museum of Natural History |access-date=July 21, 2022}}</ref> [[Smithsonian National Museum of Natural History]],<ref name="Smithsonian">{{cite web |title=Collections Catalog of the Department of Paleobiology of the Smithsonian National Museum of Natural History |url=https://collections.nmnh.si.edu/search/paleo/ |access-date=12 August 2022 |publisher=Smithsonian Institute}}</ref> [[Yale Peabody Museum]],<ref>{{cite web |title=Collections Database of the Yale Peabody Museum |url=https://collections.peabody.yale.edu/search/Search/Results?sort=relevance&join=AND&lookfor0%5B%5D=dunkleosteus&type0%5B%5D=AllFields&lookfor0%5B%5D=&type0%5B%5D=AllFields&lookfor0%5B%5D=&type0%5B%5D=AllFields&bool0%5B%5D=AND&lookfor1%5B%5D=VP&type1%5B%5D=CatalogNumber&lookfor1%5B%5D=VPPU&type1%5B%5D=CatalogNumber&bool1%5B%5D=OR&filter%5B%5D=%7Ecollection%3A%22Vertebrate+Paleontology%22&limit=5&daterange%5B%5D=collecting_year_first&collecting_year_firstfrom=&collecting_year_firstto= |publisher=Yale Peabody Museum |access-date=12 August 2022}}</ref> the [[Natural History Museum in London]], and the [[Cincinnati Museum Center]]. Specimens of ''Dunkleosteus'' are on display in many museums throughout the world (see table below), most of which are casts of the same specimen: CMNH 5768, the largest well-preserved individual of ''D. terrelli''.<ref name="Ferron2017" /> The original CMNH 5768 is on display in the [[Cleveland Museum of Natural History]]. ==Taxonomy== ''Dunkleosteus'' was named in 1956 to honour [[David Dunkle]] (1911–1982), former curator of [[vertebrate paleontology]] at the [[Cleveland Museum of Natural History]]. The genus name ''Dunkleosteus'' combines [[David Dunkle]]'s surname with the Greek word {{wikt-lang|grc|ὀστέον}} ({{transliteration|grc|ostéon}} 'bone'), literally meaning 'Dunkle's-bone'. The [[type species]] ''D. terrelli'' was originally described in 1873 as a species of ''[[Dinichthys]]'', its specific epithet chosen in honor of Jay Terrell, the fossil's discoverer.<ref>"Dunkleosteus terrelli: Fierce prehistoric predator" page at ''Cleveland Museum of Natural History''. https://www.cmnh.org/dunk {{Webarchive|url=https://web.archive.org/web/20210519150010/https://www.cmnh.org/dunk |date=2021-05-19 }}</ref> Originally thought to be a member of the genus ''[[Dinichthys]]'', ''Dunkleosteus'' was later recognized as belonging to its own genus in 1956. It was thought to be closely related to ''Dinichthys'', and they were grouped together in the [[family (biology)|family]] [[Dinichthyidae]]. However, in the 2010 Carr & Hlavin [[phylogenetic]] study, ''Dunkleosteus'' and ''Dinichthys'' were found to belong to two separate [[clade]]s. Carr & Hlavin resurrected the family [[Dunkleosteidae]] and placed ''Dunkleosteus'', ''[[Eastmanosteus]]'', and a few other genera from Dinichthyidae within it.<ref name="Carr+2010">{{cite journal| author =Carr R. K., Hlavin V. J.| title =Two new species of Dunkleosteus Lehman, 1956, from the Ohio Shale Formation (USA, Famennian) and the Kettle Point Formation (Canada, Upper Devonian), and a cladistic analysis of the Eubrachythoraci (Placodermi, Arthrodira)| journal =Zoological Journal of the Linnean Society| volume =159| issue =1| pages =195–222| year =2010| doi =10.1111/j.1096-3642.2009.00578.x| doi-access =free}}</ref> Dinichthyidae, in turn, is left a [[Monotypic taxon|monospecific]] family.<ref name=Carr1995>{{cite journal|last=Carr|first=Robert K.|author2=William J. Hlavin|title=Dinichthyidae (Placodermi):A paleontological fiction?|journal=Geobios|date=September 2, 1995|volume=28|pages=85–87|doi=10.1016/S0016-6995(95)80092-1|bibcode=1995Geobi..28...85C }}</ref> [[File:Dunkleosteus terrelli (fossil fish) (Cleveland Shale Member, Ohio Shale, Upper Devonian; Rocky River Valley, Cleveland, Ohio, USA) 25 (34091142396).jpg|thumb|right|Front view of ''D. terrelli'' skull]] The [[cladogram]] below from the 2013 Zhu & Zhu study shows the placement of ''Dunkleosteus'' within [[Dunkleosteidae]] and ''[[Dinichthys]]'' within the separate clade [[Aspinothoracidi]]:<ref name=Zhu2013>{{Cite journal |author1=You-An Zhu |author2=Min Zhu |title=A redescription of Kiangyousteus yohii (Arthrodira: Eubrachythoraci) from the Middle Devonian of China, with remarks on the systematics of the Eubrachythoraci | journal=[[Zoological Journal of the Linnean Society]] |volume=169 |issue=4 |pages=798–819 |year=2013 |doi=10.1111/zoj12089 |doi-access=free }}</ref> {{clade| style=font-size:85%;line-height:85% |grouplabel1={{clade labels |width=10em; |label1='''Pachyosteomorphi''' |top1=65% |color1=green}} |label1='''[[Eubrachythoraci]]''' |1={{clade |1={{clade |label1='''[[Coccosteomorphi]]''' |1={{clade |1=''[[Coccosteus cuspidatus]]'' |2={{clade |1=''[[Harrytoombsia elegans]]'' |2={{clade |1=''[[Mcnamaraspis kaprios]]'' |label2=[[Incisoscutoidea]] |2={{clade |1=''[[Compagopiscis croucheri]]'' |2={{clade |1=''[[Incisoscutum ritchiei]]'' |2=''[[Incisoscutum sarahae]]'' }} |label3=[[Camuropiscidae]] |3={{clade |1=''[[Latocamurus coulthardi]]'' |2=''[[Camuropiscis laidlawi]]'' |3={{clade |1=''[[Rolfosteus canningensis]]'' |2=''[[Tubonasus lennardensis]]'' |3=''[[Fallacosteus turneri]]'' }} }} }} }} }} }} |label2='''[[Aspinothoracidi]]''' |2={{clade |1=''[[Dinichthys herzeri]]'' |barbegin1=green |2={{clade |1=''[[Hadrosteus rapax]]'' |bar1=green |2={{clade |1=''[[Gorgonichthys clarki]]'' |bar1=green |2={{clade |1=''[[Heintzichthys gouldii]]'' |bar1=green |label2=[[Selenosteidae]] |2={{clade |1=''[[Stenosteus angustopectus]]'' |bar1=green |2={{clade |1=''[[Gymnotrachelus hydei]]'' |bar1=green |2={{clade |1=''[[Rhinosteus parvulus]]'' |bar1=green |2=''[[Pachyosteus bulla]]'' |bar2=green }} }} }} }} }} }} }} }} |label2='''[[Dunkleosteoidea]]''' |2={{clade |1=''[[Westralichthys uwagedensis]]'' |bar1=green |2={{clade |1={{clade |1=''[[Protitanichthys rockportensis]]'' |bar1=green |label2=[[Panxiosteidae]] |2={{clade |1=''[[Panxiosteus ocullus]]'' |bar1=green |2=''[[Janiosteus timanicus]]'' |bar2=green |3=''[[Plourdosteus canadensis]]'' |bar3=green }} }} |label2='''[[Dunkleosteidae]]''' |2={{clade |1={{clade |1=''[[Eastmanosteus calliaspis]]'' |bar1=green |2=''[[Xiangshuiosteus wui]]'' |bar2=green }} |2={{clade |1=''[[Eastmanosteus pustulosus]]'' |bar1=green |2=''[[Kiangyousteus yohii]]'' |bar2=green |3=''[[Golshanichthys asiatica]]'' |bar3=green |4={{clade |1='''''Dunkleosteus amblyodoratus''''' |bar1=green |2='''''Dunkleosteus terrelli''''' |bar2=green |3='''''Dunkleosteus raveri''''' |barend3=green }} }} }} }} }} }} }} Alternatively, the subsequent 2016 Zhu ''et al.'' study using a larger [[morphology (biology)|morphological]] dataset recovered [[Panxiosteidae]] well outside of [[Dunkleosteoidea]], leaving the status of [[Dunkleosteidae]] as a clade grouping separate from Dunkleosteoidea in doubt, as shown in the cladogram below:<ref name=Zhu2016>{{Cite journal |last1=Zhu |first1=You-An |last2=Zhu |first2=Min |last3=Wang |first3=Jun-Qing |date=April 1, 2016 |title=Redescription of Yinostius major (Arthrodira: Heterostiidae) from the Lower Devonian of China, and the interrelationships of Brachythoraci |journal=[[Zoological Journal of the Linnean Society]] |volume=176 |issue=4 |pages=806–834 |doi=10.1111/zoj.12356|issn=0024-4082 |doi-access=free }}</ref> {{clade| style=font-size:85%;line-height:85% |label1='''[[Eubrachythoraci]]''' |1={{clade |label1='''[[Coccosteomorphi]]''' |1={{clade |label1=[[Coccosteoidea]] |1={{clade |label1=[[Coccosteidae]] |1={{clade |1={{clade |1=''[[Millerosteus minor]]'' |2=''[[Coccosteus cuspidatus]]'' }} |2={{clade |1=''[[Dickosteus threiplandi]]'' |2=''[[Watsonosteus fletti]]'' |3=''[[Protitanichthys rockportensis]]'' }} }} |label2='''[[Panxiosteidae]]''' |2={{clade |1=''[[Plourdosteus canadensis]]'' |2=''[[Panxiosteus ocullus]]'' |3=''[[Janiosteus timanicus]]'' }} }} |label2=[[Incisoscutoidea]] |2={{clade |1=''[[Harrytoombsia elegans]]'' |2={{clade |1=''[[Torosteus tuberculatus]]'' |2=''[[Torosteus pulchellus]]'' }} |3={{clade |1=''[[Mcnamaraspis kaprios]]'' |2={{clade |1=''[[Compagopiscis croucheri]]'' |2=''[[Trematosteus fontanellus]]'' |label3=[[Camuropiscidae]] |3={{clade |1={{clade |1=''[[Incisoscutum ritchiei]]'' |2=''[[Incisoscutum sarahae]]'' }} |2={{clade |1=''[[Rolfosteus canningensis]]'' |2={{clade |1=''[[Tubonasus lennardensis]]'' |2={{clade |1=''[[Fallacosteus turneri]]'' |2={{clade |1=''[[Camuropiscis laidlawi]]'' |2=''[[Latocamurus coulthardi]]'' }} }} }} }} }} }} }} }} }} |label2='''[[Pachyosteomorphi]]''' |2={{clade |1=''[[Rhachiosteus pterygiatus]]'' |2={{clade |label1='''[[Dunkleosteoidea]]''' |1={{clade |1=''[[Eastmanosteus calliaspis]]'' |2={{clade |1={{clade |1=''[[Eastmanosteus pustulosus]]'' |2=''[[Kiangyousteus yohii]]'' }} |2={{clade |1=''[[Golshanichthys asiatica]]'' |2={{clade |1=''[[Westralichthys uwagedensis]]'' |2={{clade |1={{clade |1='''''Dunkleosteus raveri''''' |2='''''Dunkleosteus terrelli''''' }} |2={{clade |1='''''Dunkleosteus amblyodoratus''''' |label2=[[Heterostiidae]] |2={{clade |1=''[[Heterosteus ingens]]'' |2=''[[Yinostius major]]'' }} }} }} }} }} }} }} |label2='''[[Aspinothoracidi]]''' |2={{clade |1=''[[Tapinosteus heintzi]]'' |2={{clade |1={{clade |1=''[[Bullerichthys fascidens]]'' |2={{clade |1=''[[Kendrickichthys cavernosus]]'' |2=''[[Bruntonichthys multidens]]'' }} }} |2={{clade |1=''[[Dinichthys herzeri]]'' |2={{clade |1=''[[Hadrosteus rapax]]'' |2={{clade |1=''[[Gorgonichthys clarki]]'' |label2=[[Selenosteidae]] |2={{clade |1=''[[Heintzichthys gouldii]]'' |2=''[[Pachyosteus bulla]]'' |3=''[[Gymnotrachelus hydei]]'' |4=''[[Stenosteus angustopectus]]'' |5=''[[Brachyosteus dietrichi]]'' |6={{clade |1=''[[Melanosteus occitanus]]'' |2=''[[Rhinosteus parvulus]]'' }} }} }} }} }} }} }} }} }} }} }} === Species === At least ten different species<ref name="Carr+2010"/><ref name=Denison>{{cite book|last=Denison|first=Robert|chapter=Placodermi|volume=2|title=Handbook of Paleoichthyology|year=1978|publisher=Gustav Fischer Verlag|location=Stuttgart New York|isbn=978-0-89574-027-4|pages=128}}</ref> of ''Dunkleosteus'' have been described so far. [[File:Dunkleosteus skull steveoc.jpg|thumb|Labelled skull diagram of ''D. terrelli'']] The [[type species]], ''D. terrelli'', is the largest, best-known species of the genus, measuring {{convert|4.1|-|10|m|ft|abbr=on}} in length. It has a rounded snout. ''D. terrelli''{{'s}} fossil remains are found in Upper Frasnian to Upper Famennian Late Devonian strata of the United States (Huron and [[Cleveland Shale]] of Ohio, the Conneaut of [[Pennsylvania]], [[Chattanooga Shale]] of Tennessee, [[Lost Burro Formation]], California, and possibly Ives [[breccia]] of Texas<ref name=Denison/>) and [[Europe]]. ''D. belgicus'' (?) is known from fragments described from the [[Famennian]] of [[Belgium]]. The median dorsal plate is characteristic of the genus, but, a plate that was described as a suborbital is anterolateral.<ref name=Denison/> Lelièvre (1982) considers this taxon a [[nomen dubium]] and suggests the material may actually pertain to ''[[Ardennosteus]]''.<ref>{{Cite journal |last=Lelièvre |first=Hervé |date=1982 |title=Ardennosteus ubaghsi n.g., n. sp. Brachythoraci primitif (vertébré, placoderme) du Famennien d'Esneux (Belgique) |url=https://popups.uliege.be/0037-9395/index.php?id=3220 |journal=Annales de la Société géologique de Belgique |volume=105 |issue=1}}</ref> ''D. denisoni'' is known from a small median dorsal plate, typical in appearance for ''Dunkleosteus'', but much smaller than normal. It is comparable in skull structure to ''D. marsaisi''.<ref name=Denison/> [[File:Dunkleosteus marsaisi 45.JPG|thumb|''D. marsaisi'' skull]] ''D. marsaisi'' refers to the ''Dunkleosteus'' fossils from the Lower Famennian Late Devonian strata of the [[Atlas Mountains]] in [[Morocco]]. It differs in size, the known skulls averaging a length of {{convert|35|cm|ft}} and in form to ''D. terrelli''. In ''D. marsaisi'', the snout is narrower, and a postpineal fenestra may be present. Many researchers and authorities consider it a synonym of ''D. terrelli''.<ref>{{cite journal|last=Murray|first=A.M.|title=The Palaeozoic, Mesozoic and Early Cenozoic fishes of Africa|journal=Fish and Fisheries|year=2000|volume=1|issue=2|pages=111–145|doi=10.1046/j.1467-2979.2000.00015.x}}</ref> H. Schultze regards ''D. marsaisi'' as a member of ''[[Eastmanosteus]]''.<ref name=Denison/><ref name=Schultz>{{cite journal|last=Schultz|first=H|title=Large Upper Devonian arthrodires from Iran|journal=Fieldiana Geology|year=1973|volume=23|pages=53–78|doi=10.5962/bhl.title.5270|doi-access=free}}</ref> ''D. magnificus'' is a large placoderm from the Frasnian [[Rhinestreet Shale]] of New York. It was originally described as ''[[Dinichthys]] magnificus'' by Hussakof and Bryant in 1919, then as "'' Dinichthys mirabilis''" by Heintz in 1932. Dunkle and Lane moved it to ''Dunkleosteus'' in 1971.<ref name=Denison/> ''D. missouriensis'' is known from fragments from Frasnian [[Missouri]]. Dunkle and Lane regard them as being very similar to ''D. terrelli''.<ref name=Denison/> ''D. newberryi'' is known primarily from a {{convert|28|cm|in}} long infragnathal with a prominent anterior cusp, found in the Frasnian portion of the [[Genesee Group]] of New York, and originally described as ''Dinichthys newberryi''.<ref name=Denison/> ''D. amblyodoratus'' is known from some fragmentary remains from Late Devonian strata of [[Kettle Point Formation]], Ontario. The species name means 'blunt spear' and refers to the way the [[nuchal]] and [[paranuchal]] plates in the back of the head form the shape of a blunted spearhead.<ref name="Carr+2010"/> ''D. raveri'' is a small species, possibly 1 meter long, known from an uncrushed skull roof found in a carbonate concretion from near the bottom of the Huron Shale, of the Famennian [[Ohio Shale]] strata. Besides its small size, it had comparatively large eyes. Because ''D. raveri'' was found in the strata directly below the strata where the remains of ''D. terrelli'' are found, ''D. raveri'' may have given rise to ''D. terrelli''. The species name commemorates Clarence Raver of [[Wakeman, Ohio]], who discovered the concretion where the [[holotype]] was found.<ref name="Carr+2010"/> ''D. tuderensis'' is known from an infragnathal found in the lower-middle [[Famennian]]-aged [[Bilovo Formation]] of the [[Tver Region]] in northwest Russia. The specific name refers to the Maliy Tuder River as the holotyoe was found on its bank.<ref name=":3">{{Cite journal |last=Lebedev |first=Oleg A. |last2=Engelman |first2=Russell K. |last3=Skutschas |first3=Pavel P. |last4=Johanson |first4=Zerina |last5=Smith |first5=Moya M. |last6=Kolchanov |first6=Veniamin V. |last7=Trinajstic |first7=Kate |last8=Linkevich |first8=Valeriy V. |date=May 2023 |title=Structure, Growth and Histology of Gnathal Elements in Dunkleosteus (Arthrodira, Placodermi), with a Description of a New Species from the Famennian (Upper Devonian) of the Tver Region (North-Western Russia) |url=https://www.mdpi.com/1424-2818/15/5/648 |journal=Diversity |language=en |volume=15 |issue=5 |pages=648 |doi=10.3390/d15050648 |issn=1424-2818}}</ref> ==Description== ===Size and anatomy=== [[File:Dunkleosteus terrelli size.png|left|thumb|Comparison of multiple size estimates for ''D. terrelli'']] Mainly the armored frontal sections of specimens have been [[fossilized]], and consequently, the appearance of the other portions of the fish is mostly unknown.<ref>{{cite AV media |url=https://www.youtube.com/watch?v=lEEhm4rzxEg |title=Prehistoric Monsters Revealed |date=2008 |last=Dash |first=Sean |publisher=Workaholic Productions / History Channel |location=United States |access-date=December 18, 2015}}{{cbignore}}{{Dead YouTube link|date=February 2022}}</ref> In fact, only about 5% of ''Dunkleosteus'' specimens have more than a quarter of their skeleton preserved.<ref>Carr, R, & G.L. Jackson. 2008. [https://www.researchgate.net/publication/213769842_THE_VERTEBRATE_FAUNA_OF_THE_CLEVELAND_MEMBER_FAMENNIAN_OF_THE_OHIO_SHALE The Vertebrates fauna of the Cleveland member (Famennian) of the Ohio Shale. Society of Vertebrates Paleontology]. 1–17.</ref> Because of this, many reconstructions of the hindquarters are often based on fossils of smaller [[arthrodire]]s, such as ''[[Coccosteus]]'', which have preserved hind sections,<ref name=":0" /> leading to widely varying size estimates.<ref name=":0" /> ''Dunkleosteus'' is one of the largest known placoderms, with its maximum size being variably estimated as anywhere from {{convert|4.1|–|10|m|ft}} by different researchers.<ref name="Anderson&Westneat2007" /><ref name="Anderson&Westneat2009" /><ref name="Ferron2017" /><ref>{{cite journal|last=Williams|first=Nigel|year=2007|title=Force feeding|journal=Current Biology |volume=17 |pages=R3 |doi=10.1016/j.cub.2006.11.057|s2cid=36585467 }}</ref><ref name=":0">{{Cite journal |last=Engelman |first=Russell K. |date=2023|title=A Devonian Fish Tale: A New Method of Body Length Estimation Suggests Much Smaller Sizes for Dunkleosteus terrelli (Placodermi: Arthrodira) |journal=Diversity |language=en |volume=15 |issue=3 |pages=318 |doi=10.3390/d15030318 |issn=1424-2818 |doi-access=free }}</ref> However, most cited length estimates are speculative and lack quantitative or statistical backing.<ref name="Ferron2017" /><ref name=":0" /> Most studies that estimate the length of ''Dunkleosteus'' do not provide information as to how these estimates were calculated, the measurements used to scale them, or which specimens were examined. Most are implied to be based on either CMNH 5768 (the largest complete armor of ''Dunkleosteus'') or CMNH 5936 (the largest known jaw fragment). {{multiple image |align=right |perrow=1 |total_width=240 |image1=Dunkleosteus terrelli 2017 reconstruction.png |caption1=[[Paleoart|Life reconstruction]] of ''D. terrelli'', as presented by Ferrón ''et al.'' 2017<ref name="Ferron2017"/> |image2=Dunkleosteus terrelli 2023 reconstruction.png |caption2=Life reconstruction of ''D. terrelli'', as presented by Engelman 2023<ref name=":0"/> }} ===Revaluation of physiology=== Most of the studies with well-defined methods produce lengths of {{convert|5|m|ft}} or less for ''Dunkleosteus terrelli'',<ref name=":0" /> with the exception of Ferrón ''et al.'' (2017), which produces larger estimates of {{convert|6.88|–|8.79|m|ft}} based on upper jaw perimeter of modern sharks.<ref name="Ferron2017" /> However, arthrodires have proportionally larger mouths than modern sharks, making the lengths estimated by Ferrón ''et al.'' (2017) unreliable.<ref name=":2">{{cite journal |last=Engelman |first=Russell |date=2023-04-10 |title=Giant, swimming mouths: oral dimensions of extant sharks do not accurately predict body size in ''Dunkleosteus terrelli'' (Placodermi: Arthrodira) |url=https://peerj.com/articles/15131 |journal=PeerJ |language=en |volume=11 |pages=e15131 |doi=10.7717/peerj.15131 |pmid=37065696 |pmc=10100833 }}</ref> Upper jaw perimeter overestimates the size of complete arthrodires like ''[[Coccosteus]]'' and the estimates of Ferrón ''et al.'' (2017) result in ''Dunkleosteus'' having an extremely small head and hyper-elongate trunk relative to the known dimensions of the fossils.<ref name=":2" /> If the reconstruction presented in Ferrón ''et al.'' (2017) is scaled to the known dimensions of CMNH 5768, it produces a length of {{convert|3.77|m|ft}}.<ref name=":2" /> Carr (2010) estimated a {{convert|4.6|m|ft}} long adult individual of ''Dunkleosteus terrelli'' to have weighed {{convert|665|kg|lb}}, assuming a shark-like body plan and a similar length-weight relationship.<ref name="Carr 20102">{{cite journal |last1=Carr |first1=Robert K. |date=2010 |title=Paleoecology of Dunkleosteus terrelli (Placodermi: Arthrodira). |url=https://www.researchgate.net/publication/235924093 |journal=Kirtlandia |volume=57}}</ref> Engelman (2023), using an [[ellipsoid]] [[Volume|volumetric]] method, estimated weights of {{convert|950|–|1200|kg|lb}} for typical ({{convert|3.41|m|ft}} long) adult ''Dunkleosteus'', and weights of {{convert|1490|–|1760|kg|lb}} for the largest ({{convert|4.1|m|ft|1}} in this study) individual.<ref name=":0" /> The higher weights by Engelman (2023) are mostly a result of the fact that arthrodires tend to have relatively deeper and wider bodies compared to sharks.<ref name=":0" /> An exceptionally preserved specimen of ''D. terrelli'' preserves a [[Fish fin|pectoral fin]] outline with [[ceratotrichia]], implying that the fin morphology of placoderms was much more variable than previously thought, and was heavily influenced by locomotory requirements. This knowledge, coupled with the knowledge that fish morphology is more heavily influenced by feeding niche than phylogeny, allowed a 2017 study to infer the [[Fish fin|caudal fin]] shape of ''D. terrelli'', reconstructing this fin with a strong ventral lobe, a high [[aspect ratio]], narrow [[caudal peduncle]], in contrast to previous reconstructions based on the [[Fish locomotion|anguilliform]] caudal fin of coccosteomorph placoderms.<ref name="Ferron2017">{{Cite journal|last1=Ferrón|first1=Humberto G.|last2=Martínez-Pérez|first2=Carlos|last3=Botella|first3=Héctor|date=2017|title=Ecomorphological inferences in early vertebrates: reconstructing Dunkleosteus terrelli (Arthrodira, Placodermi) caudal fin from palaeoecological data|journal=PeerJ|language=en|volume=5|pages=e4081|doi=10.7717/peerj.4081|pmid=29230354|pmc=5723140|issn=2167-8359}}</ref> ====Length estimations of ''D. terrelli''==== {| class="wikitable sortable" style="margin-left: auto; margin-right: auto; border: none;" |+ Length estimates of ''Dunkleosteus terrelli'' (modified from Table 7 of Engelman 2023<ref name=":0" />) ! Study (author) ! Year ! Length ! Method ! Reference |- | Newberry || 1875 || {{convert|4.5|–|5.5|m|ft}} || '''Extrapolated from ''Coccosteus cuspidatus'', measurements and specimen used unclear''' || <ref name=":1">{{cite book |last1=Newberry |first1=John S. |url=https://www.biodiversitylibrary.org/item/116994#page/15/mode/1up |title=Report of the Geological Survey of Ohio. Volume II. Geology and Paleontology |date=1875 |publisher=Nevins and Myers, State Printers |volume=2 |location=Columbus |page=24 |contribution=Descriptions of fossil fishes}}</ref> |- | Newberry || 1889 || {{convert|4.5|m|ft}} || Unstated (implied extrapolation from ''Coccosteus'') || <ref>{{cite journal |last1=Newberry |first1=John S. |date=1889 |title=Paleozoic fishes of North America |url=https://www.biodiversitylibrary.org/item/51025#page/5/mode/1up |journal=Monographs of the U.S. Geological Survey |volume=16 |page=24}}</ref> |- | Dean || 1895 || {{convert|3|m|ft}} || Methods, measurements, and specimens unstated || <ref>{{cite book |last1=Dean |first1=Bashford |title=Fishes, Living and Fossil: An Outline of Their Forms and Probable Relationships |date=1895 |publisher=Macmillan and Company |location=London |page=130}}</ref> |- | Hussakof || 1905 || {{convert|1.67|m|ft}} (AMNH FF 195) <br> {{convert|3.79|m|ft}} (extrapolated to CMNH 5768 by Engelman 2023<ref name=":0"/> assuming similar head-trunk proportions) || '''Entering angle of body''' || <ref>{{cite journal |last1=Hussakof |first1=Louis |title=Notes on the Devonian "placoderm" ''Dinichthys intermedius'' Newb. |journal=Bulletin of the American Museum of Natural History |date=1905 |volume=21 |pages=27–36}}</ref> |- | Anonymous|| 1923 || {{convert|7.6|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Anonymous |title=Cleveland shale fishes |journal=Bulletin of the Cleveland Museum of Natural History |date=1923 |volume=9 |pages=36}}</ref> |- | Hyde || 1926 || {{convert|4.5|–|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Hyde |first1=Jesse E. |title=Collecting fossil fishes from the Cleveland Shale |journal=Natural History |date=1926 |volume=26 |pages=497–504}}</ref> |- | Romer || 1966 || {{convert|9|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Romer |first1=Alfred S. |title=Vertebrate Paleontology |date=1966 |publisher=University of Chicago Press |location=Chicago |page=49 |edition=3rd}}</ref> |- | Colbert || 1969 || {{convert|9|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Colbert |first1=Edwin H. |title=Evolution of the Vertebrates: A History of Backboned Animals Through Time |date=1969 |publisher=John Wiley and Sons |location=Hoboken |page=36 |edition=2nd}}</ref> |- | Denison || 1978 || {{convert|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref name=Denison /> |- | Williams || 1992 || {{convert|5|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Williams |first1=Michael E. |title=Jaws: The Early Years |journal=Explorer |date=1992 |volume=34 |pages=4–8}}</ref> |- | Janvier || 2003 || {{convert|6|–|7|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Janvier |first1=P. |title=Early Vertebrates |date=2003 |publisher=Clarendon Press |location=Oxford |page=12}}</ref> |- | Young || 2003 || {{convert|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Young |first1=Gavin C. |title=Did placoderm fish have teeth? |journal=Journal of Vertebrate Paleontology |date=24 December 2003 |volume=23 |issue=4 |pages=987–990 |doi=10.1671/31|bibcode=2003JVPal..23..987Y |s2cid=85572061 }}</ref> |- | Anderson and Westneat || 2007 || {{convert|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref name="Anderson&Westneat2007" /> |- | Anderson and Westneat || 2009 || {{convert|10|m|ft}} || Methods, measurements, and specimens used not stated || <ref name="Anderson&Westneat2009" /> |- | Carr || 2010 || {{convert|4.5|–|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref name="Carr 2010">{{cite journal |last1=Carr |first1=Robert K. |date=2010 |title=Paleoecology of Dunkleosteus terrelli (Placodermi: Arthrodira). |url=https://www.researchgate.net/publication/235924093 |journal=Kirtlandia |volume=57}}</ref> |- | Long || 2010 || {{convert|4.5|–|8|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Long |first1=John A. |title=The Rise of Fishes: 500 Million Years of Evolution |date=2010 |publisher=Johns Hopkins University Press |location=Baltimore |pages=88–90 |edition=2nd}}</ref> |- | Sallan and Galimberti || 2015 || {{convert|8|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Sallan |first1=Lauren |last2=Galimberti |first2=Andrew K. |title=Body-size reduction in vertebrates following the end-Devonian mass extinction |journal=Science |date=13 November 2015 |volume=350 |issue=6262 |pages=812–815 |doi=10.1126/science.aac7373|pmid=26564854 |bibcode=2015Sci...350..812S |s2cid=206640186 }}</ref> |- | Ferrón ''et al.'' || 2017 || {{convert|6.88|m|ft}} (average adult, CMNH 5768) <br> {{convert|8.79|m|ft}} (largest individual, CMNH 5936) || '''Upper jaw perimeter''' || <ref name="Ferron2017" /> |- | Long ''et al.'' || 2019 || {{convert|6|–|8|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Long |first1=John A. |last2=Choo |first2=Brian |last3=Clement |first3=Alice |contribution=The Evolution of Fishes through Geological Time |title=Evolution and Development of Fishes |editor1-last=Johanson |editor1-first=Zerina|editor2-last=Underwood |editor2-first=Charlie|editor3-last=Richter|editor3-first=Martha| date=31 December 2018 |pages=3–29 |doi=10.1017/9781316832172.002|s2cid=134217082 }}</ref> |- | Johanson ''et al.'' || 2019 || {{convert|3|m|ft}} (CMNH 50322) <br> {{convert|7.1|m|ft}} (extrapolated to CMNH 5768 by Engelman 2023 assuming similar head-trunk proportions) || Methods and measurements not stated || <ref>{{cite journal |last1=Johanson |first1=Zerina |last2=Trinajstic |first2=Kate |last3=Cumbaa |first3=Stephen |last4=Ryan |first4=Michael |title=Fusion in the vertebral column of the pachyosteomorph arthrodire Dunkleosteus terrelli ('Placodermi') |journal=Palaeontologia Electronica |date=2019 |volume=22.2.20A |pages=1–13 |doi=10.26879/872|s2cid=162173408 }}</ref> |- | Engelman || 2023 || {{convert|3.41|m|ft}} (average adult, CMNH 5768) <br> {{convert|4.1|m|ft|1}} (largest individual, CMNH 5936) || '''Orbit-opercular length (head length minus snout)''' || <ref name=":0" /> |- | Engelman || 2023 || {{convert|3.41|m|ft}} (average adult, CMNH 5768) || '''Skull length in ''Coccosteus''''' || <ref name=":0" /> |- | Engelman || 2023 || {{convert|5.23|m|ft}} (average adult, CMNH 5768) || '''Infragnathal length in ''Coccosteus''''' (source considers this estimate unreliable due to ''Dunkleosteus'' having a relatively larger mouth than ''Coccosteus'') || <ref name=":0" /> |- | Engelman || 2023 || {{convert|3.47|m|ft}} (average adult, CMNH 5768) || '''Entering angle of body''' || <ref name=":0" /> |- | Engelman || 2023 || {{convert|3.88|m|ft}} (average adult, CMNH 5768) || '''Length of posteroventrolateral plate''' || <ref name=":0" /> |- | Engelman || 2023 || {{convert|3.40|m|ft}} (average adult, CMNH 5768) || '''Inferred location of pelvic girdle''' || <ref name=":0" /> |- |} ==Paleobiology== ===Diet=== [[File:Dunkleosteus terrelli CMNH 5936.png|thumb|left|Lower jaw of ''D. terrelli'' specimen CMNH 5936]] ''Dunkleosteus terrelli'' possessed a [[four-bar linkage]] mechanism for [[Fish jaw|jaw]] opening that incorporated connections between the skull, the thoracic shield, the lower jaw and the jaw muscles joined by movable joints.<ref name=Anderson&Westneat2009>{{cite journal |last=Anderson |first=P.S.L. |author2=Westneat, M. |title=A biomechanical model of feeding kinematics for ''Dunkleosteus terrelli'' (Arthrodira, Placodermi) |journal=Paleobiology |date=2009 |volume=35 |issue=2 |pages=251–269 |doi=10.1666/08011.1 |bibcode=2009Pbio...35..251A |s2cid=86203770 |url=http://projects.fieldmuseum.org/sites/default/files/DunkPaleoBio.pdf |access-date=October 29, 2017 |archive-date=November 7, 2017 |archive-url=https://web.archive.org/web/20171107014026/http://projects.fieldmuseum.org/sites/default/files/DunkPaleoBio.pdf |url-status=dead }}</ref><ref name=Anderson&Westneat2007>{{Cite journal |last=Anderson |first=P.S.L. |author2=Westneat, M. |title=Feeding mechanics and bite force modelling of the skull of Dunkleosteus terrelli, an ancient apex predator |journal=Biology Letters |volume=3 |issue=1 |pages=76–79 |date=2007 |doi=10.1098/rsbl.2006.0569 |pmid=17443970 |pmc=2373817}}</ref> This mechanism allowed ''D. terrelli'' to both achieve a high speed of jaw opening, opening their jaws in 20 milliseconds and completing the whole process in 50–60 milliseconds (comparable to modern fishes that use [[Aquatic feeding mechanisms#Suction feeding|suction feeding]] to assist in prey capture;<ref name=Anderson&Westneat2007/>) and producing high bite forces when closing the jaw, estimated at {{convert|4414|N|kg-f lb-f|0|abbr=on}} at the tip and {{convert|5363|N|kg-f lb-f|0|abbr=on}} at the blade edge,<ref name=Anderson&Westneat2007/> or even up to {{cvt|6170|N|kg-f lb-f|0}} and {{cvt|7495|N|kg-f lb-f|0}} respectively.<ref name=Anderson&Westneat2009/> The pressures generated in those regions were high enough to puncture or cut through [[cuticle]] or [[dermal bone|dermal]] armor,<ref name=Anderson&Westneat2007/> suggesting that ''D. terrelli'' was adapted to prey on free-swimming, armored prey such as [[ammonite]]s and other placoderms.<ref name=Anderson&Westneat2009/> In addition, teeth of a [[Chondrichthyes|chondrichthyan]] thought to belong to ''[[Orodus]]'' (''Orodus'' spp.) were found in association with ''Dunkleosteus'' remains, suggesting that these were probably stomach contents regurgitated from the animal. ''Orodus'' is thought to be tachypelagic, or a fast-swimming [[pelagic fish]]. Thus, ''Dunkleosteus'' might have been fast enough to catch these fast organisms, and not a slow swimmer like originally thought.<ref name="Ferron2017" /> Fossils of ''Dunkleosteus'' are frequently found with [[Bolus (digestion)|bolus]]es of fish bones, semidigested and partially eaten remains of other fish.<ref name="Virtual Fossil">{{cite web|url= http://www.fossilmuseum.net/Fossil_Galleries/Fish_Devonian/Dunkleosteous/Dunkleosteus.htm |title=Dunkleosteus Placodermi Devonian Armored Fish from Morocco |work=Fossil Archives |publisher=The Virtual Fossil Museum |access-date=April 26, 2009}}</ref> As a result, the [[fossil]] record indicates it may have routinely regurgitated prey bones rather than digest them. Mature individuals probably inhabited deep sea locations, like other placoderms, living in shallow waters during adolescence.<ref>{{Cite web|url=http://bioweb.uwlax.edu/bio203/f2013/peters_sadi/habitat.htm|title=UWL Website}}</ref> A specimen of ''Dunkleosteus'' (CMNH 5302), and ''[[Titanichthys]]'' (CMNH 9889), show damage said to be puncture damage from the bony fangs of other ''Dunkleosteus''.<ref name="Anderson&Westneat2009" /> ===Reproduction=== ''Dunkleosteus'', together with most other placoderms, may have also been among the first [[vertebrate]]s to [[internal fertilization|internalize egg fertilization]], as seen in some modern sharks.<ref name="Ahlberg">{{cite journal |last1=Ahlberg |first1=Per |first2=Kate |last2=Trinajstic |first3=Zerina |last3=Johanson |first4=John |last4=Long |s2cid=205217467 |year=2009 |title=Pelvic claspers confirm chondrichthyan-like internal fertilization in arthrodires |journal=Nature |volume=460 |issue=7257 |pages=888–889 |doi=10.1038/nature08176|pmid=19597477 |bibcode=2009Natur.460..888A }}</ref> Some other placoderms have been found with evidence that they may have been [[viviparity|viviparous]], including what appears to have been an [[umbilical cord]].<ref>{{ cite journal | last = Long | first = J. A. |author2=Trinajstic, K. |author3=Young, G. C. |author4=Senden, T. | year = 2008 | title = Live birth in the Devonian period | journal = Nature | volume = 453 | issue = 7195 | pages = 650&ndash;652 | doi = 10.1038/nature06966 | pmid = 18509443 | bibcode = 2008Natur.453..650L | s2cid = 205213348 }}</ref> ===Growth=== {{multiple image |align=right |perrow=2 |total_width=380 |image1=Dunkleosteus juvenile fossil - Cleveland Museum of Natural History (34421618320).jpg |caption1=''D. terrelli'' juvenile specimen CMNH 7424 |image2=Dunkleosteus terrelli - Cleveland Museum of Natural History - 2014-12-26 (21137540331).jpg |caption2=''D. terrelli'' adult specimen CMNH 5768 }} [[morphology (biology)|Morphological]] studies on the lower jaws of juveniles of ''D. terrelli'' reveal they were proportionally as robust as those of adults, indicating they already could produce high bite forces and likely were able to shear into resistant prey tissue similar to adults, albeit on a smaller scale. This pattern is in direct contrast to the condition common in [[tetrapod]]s in which the jaws of juveniles are more [[gracile]] than in adults.<ref name="Snively_etal_2009">{{cite journal |last=Snively |first=E. |author2=Anderson, P.S.L. |author3=Ryan, M.J. |date=2009 |title=Functional and ontogenetic implications of bite stress in arthrodire placoderms |url=https://www.biodiversitylibrary.org/page/51811342#page/60/mode/1up |journal=Kirtlandia |volume=57}}</ref> ==See also== * [[List of placoderms]] ==References== {{notelist}} {{reflist}} ==Further reading== * {{cite journal|first=Philip S. L.|last=Anderson|s2cid=86583150|year=2008|title=Shape Variation Between Arthrodire Morphotypes Indicates Possible Feeding Niches|journal=Journal of Vertebrate Paleontology|volume=28|number=4|pages=961–969|doi=10.1671/0272-4634-28.4.961|bibcode=2008JVPal..28..961A }} ==External links== {{Commons category|Dunkleosteus}} {{Wikispecies|Dunkleosteus}} * [http://www.ucmp.berkeley.edu/vertebrates/basalfish/placodermi.html ''Introduction to the Placodermi: Extinct Armored Fishes with Jaws''. Waggoner, Ben (2000). Retrieved Aug 1, 2005] * [http://www.nbcnews.com/id/15940709 MSNBC: Prehistoric fish packed a mean bite] * [http://news.bbc.co.uk/1/hi/sci/tech/6195188.stm BBC: Ancient 'Jaws' had monster bite] {{Arthrodira|E.}} {{Taxonbar|from=Q131039|from2=Q911597|from3=Q911599|from4=Q911605|from5=Q911606|from6=Q911611|from7=Q911612|from8=Q911616|from9=Q911618|from10=Q911593|from11=Q25357547}} [[Category:Dunkleosteidae]] [[Category:Apex predators]] [[Category:Placoderms of Africa]] [[Category:Fossils of Morocco]] [[Category:Placoderms of Europe]] [[Category:Fossils of Belgium]] [[Category:Placoderms of North America]] [[Category:Fossils of Canada]] [[Category:Paleontology in Missouri]] [[Category:Paleontology in Pennsylvania]] [[Category:Paleontology in Tennessee]] [[Category:Late Devonian first appearances]] [[Category:Late Devonian animals]] [[Category:Famennian extinctions]] [[Category:Fossil taxa described in 1956]] [[Category:Taxa named by John Strong Newberry]]'
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'@@ -1,379 +1,1 @@ -{{Short description|Genus of extinct fishes}} -{{Automatic taxobox -| fossil_range = [[Late Devonian]] ([[Frasnian]] to [[Famennian]]), {{Fossil range|382|358}} -| image = Dunkleosteus terrelli (fossil fish) (Cleveland Shale Member, Ohio Shale, Upper Devonian; Rocky River Valley, Cleveland, Ohio, USA) 21 (34001200911).jpg -| image_caption = Partially reconstructed ''D. terrelli'' skull (specimen CMNH 5768), [[Cleveland Museum of Natural History]] -| display_parents = 7 -| taxon = Dunkleosteus -| authority = [[Jean-Pierre Lehman|Lehman]], 1956 -| type_species = †'''''Dunkleosteus terrelli''''' -| type_species_authority = [[John Strong Newberry|Newberry]], 1873 -| subdivision_ranks = [[Species]] -| subdivision = {{collapsible list| -''D. terrelli'' <small>(Newberry, 1873 [originally ''[[Dinichthys]]''])</small><br /> -''D. ? belgicus'' <small>(Newberry) 1873</small><br /> -''D. denisoni'' <small>(Kulczycki) 1957</small><br /> -''D. marsaisi'' <small>Lehmann, 1956</small><br /> -''D. magnificus'' <small>(Hussakof & Bryant) 1919</small><br /> -''D. missouriensis'' <small>(Branson) 1914</small><br /> -''D. newberryi'' <small>(Clarke) 1885</small><br /> -''D. amblyodoratus'' <small>Carr & Hlavin, 2010</small><br /> -''D. raveri'' <small>Carr & Hlavin, 2010</small><br /> -''D. tuderensis'' <small>Lebedev et. al. 2023</small><br /> -}} -}} - -'''''Dunkleosteus''''' is an [[extinct]] [[genus]] of large [[arthrodira|arthrodire]] fish that existed during the [[Late Devonian]] period, about 382–358 [[million years ago]]. It consists of ten species, some of which are among the largest [[placoderm]]s to have ever lived: ''D. terrelli'', ''D. belgicus'', ''D. denisoni'', ''D. marsaisi'', ''D. magnificus'', ''D. missouriensis'', ''D. newberryi'', ''D. amblyodoratus'', ''D. raveri'', and ''D. tuderensis,'' and the largest and most well known species is ''D. terrelli''. Since body shape is not known, various methods of estimation put the living total length of the largest known specimen between {{convert|4.1|to|10|m|ft|abbr=on}} long and weigh around {{convert|1|-|4|MT|ST|abbr=on}}.<ref name=":0" /> ''Dunkleosteus'' could quickly open and close its jaw, like modern-day [[Aquatic feeding mechanisms#Suction feeding|suction feeders]], and had a bite force of {{convert|4414|-|6170|N|kg-f lb-f|0|abbr=on}} at the tip and {{convert|5363|-|7495|N|kg-f lb-f|0|abbr=on}} at the blade edge.<ref name=Anderson&Westneat2007/><ref name=Anderson&Westneat2009/> Numerous [[fossil]]s of the various species have been found in [[North America]], [[Poland]], [[Belgium]], and [[Morocco]]. ''Dunkleosteus'' was a [[pelagic fish]] inhabiting open waters, and an [[apex predator]] of its ecosystem. - -==Discovery== -''Dunkleosteus'' fossils were first discovered in 1867 by Jay Terrell, a hotel owner and amateur paleontologist who collected fossils in the cliffs along [[Lake Erie]] near his home of [[Sheffield Lake, Ohio]] (due west of [[Cleveland]]). Terrell donated his fossils to [[John Strong Newberry]] and the Ohio Geological Survey, who in 1873 described all the material as belonging to a single new [[genus]] and [[species]]: ''[[Dinichthys|Dinichthys herzeri]]''. However, with later fossil discoveries by 1875 it became apparent multiple large fish species were present in the [[Ohio Shale]]. ''[[Dinichthys|Dinichthys herzeri]]'' came from the lowermost layer, the [[Huron Shale]], whereas most of the fossils were coming from the younger [[Cleveland Shale]] and represented a distinct species.<ref name=":1" /> Newberry named this more common species "''Dinichthys''" ''terrelli'', after Terrell. Most of Terrell's original collection does not survive, having been destroyed by a fire in [[Elyria, Ohio|Elyria]] in 1873.<ref name=":1" /><ref>{{Cite journal |last=Claypole |first=E. W. |date=1893 |title=The three great fossil placoderms of Ohio |url=https://www.biodiversitylibrary.org/item/240810#page/107/mode/1up |journal=American Geologist |volume=12 |pages=89–99}}</ref> - -The largest collection of ''Dunkleosteus'' fossils in the world is housed at the [[Cleveland Museum of Natural History]],<ref>{{cite web |url=https://www.cmnh.org/dunk |title=Dunkleosteus terrelli: Fierce prehistoric predator |author=<!--Not stated--> |website=Cleveland Museum of Natural History |access-date=July 21, 2022 |archive-date=May 19, 2021 |archive-url=https://web.archive.org/web/20210519150010/https://www.cmnh.org/dunk |url-status=dead }}</ref> with smaller collections (in descending order of size) held at the [[American Museum of Natural History]],<ref>{{cite web |url=https://www.amnh.org/exhibitions/permanent/vertebrate-origins/dunkleosteus |title=Dunkleosteus |author=<!--Not stated--> |website=American Museum of Natural History |access-date=July 21, 2022}}</ref> [[Smithsonian National Museum of Natural History]],<ref name="Smithsonian">{{cite web |title=Collections Catalog of the Department of Paleobiology of the Smithsonian National Museum of Natural History |url=https://collections.nmnh.si.edu/search/paleo/ |access-date=12 August 2022 |publisher=Smithsonian Institute}}</ref> [[Yale Peabody Museum]],<ref>{{cite web |title=Collections Database of the Yale Peabody Museum |url=https://collections.peabody.yale.edu/search/Search/Results?sort=relevance&join=AND&lookfor0%5B%5D=dunkleosteus&type0%5B%5D=AllFields&lookfor0%5B%5D=&type0%5B%5D=AllFields&lookfor0%5B%5D=&type0%5B%5D=AllFields&bool0%5B%5D=AND&lookfor1%5B%5D=VP&type1%5B%5D=CatalogNumber&lookfor1%5B%5D=VPPU&type1%5B%5D=CatalogNumber&bool1%5B%5D=OR&filter%5B%5D=%7Ecollection%3A%22Vertebrate+Paleontology%22&limit=5&daterange%5B%5D=collecting_year_first&collecting_year_firstfrom=&collecting_year_firstto= |publisher=Yale Peabody Museum |access-date=12 August 2022}}</ref> the [[Natural History Museum in London]], and the [[Cincinnati Museum Center]]. Specimens of ''Dunkleosteus'' are on display in many museums throughout the world (see table below), most of which are casts of the same specimen: CMNH 5768, the largest well-preserved individual of ''D. terrelli''.<ref name="Ferron2017" /> The original CMNH 5768 is on display in the [[Cleveland Museum of Natural History]]. - -==Taxonomy== -''Dunkleosteus'' was named in 1956 to honour [[David Dunkle]] (1911–1982), former curator of [[vertebrate paleontology]] at the [[Cleveland Museum of Natural History]]. The genus name ''Dunkleosteus'' combines [[David Dunkle]]'s surname with the Greek word {{wikt-lang|grc|ὀστέον}} ({{transliteration|grc|ostéon}} 'bone'), literally meaning 'Dunkle's-bone'. The [[type species]] ''D. terrelli'' was originally described in 1873 as a species of ''[[Dinichthys]]'', its specific epithet chosen in honor of Jay Terrell, the fossil's discoverer.<ref>"Dunkleosteus terrelli: Fierce prehistoric predator" page at ''Cleveland Museum of Natural History''. https://www.cmnh.org/dunk {{Webarchive|url=https://web.archive.org/web/20210519150010/https://www.cmnh.org/dunk |date=2021-05-19 }}</ref> - -Originally thought to be a member of the genus ''[[Dinichthys]]'', ''Dunkleosteus'' was later recognized as belonging to its own genus in 1956. It was thought to be closely related to ''Dinichthys'', and they were grouped together in the [[family (biology)|family]] [[Dinichthyidae]]. However, in the 2010 Carr & Hlavin [[phylogenetic]] study, ''Dunkleosteus'' and ''Dinichthys'' were found to belong to two separate [[clade]]s. Carr & Hlavin resurrected the family [[Dunkleosteidae]] and placed ''Dunkleosteus'', ''[[Eastmanosteus]]'', and a few other genera from Dinichthyidae within it.<ref name="Carr+2010">{{cite journal| author =Carr R. K., Hlavin V. J.| title =Two new species of Dunkleosteus Lehman, 1956, from the Ohio Shale Formation (USA, Famennian) and the Kettle Point Formation (Canada, Upper Devonian), and a cladistic analysis of the Eubrachythoraci (Placodermi, Arthrodira)| journal =Zoological Journal of the Linnean Society| volume =159| issue =1| pages =195–222| year =2010| doi =10.1111/j.1096-3642.2009.00578.x| doi-access =free}}</ref> Dinichthyidae, in turn, is left a [[Monotypic taxon|monospecific]] family.<ref name=Carr1995>{{cite journal|last=Carr|first=Robert K.|author2=William J. Hlavin|title=Dinichthyidae (Placodermi):A paleontological fiction?|journal=Geobios|date=September 2, 1995|volume=28|pages=85–87|doi=10.1016/S0016-6995(95)80092-1|bibcode=1995Geobi..28...85C }}</ref> -[[File:Dunkleosteus terrelli (fossil fish) (Cleveland Shale Member, Ohio Shale, Upper Devonian; Rocky River Valley, Cleveland, Ohio, USA) 25 (34091142396).jpg|thumb|right|Front view of ''D. terrelli'' skull]] -The [[cladogram]] below from the 2013 Zhu & Zhu study shows the placement of ''Dunkleosteus'' within [[Dunkleosteidae]] and ''[[Dinichthys]]'' within the separate clade [[Aspinothoracidi]]:<ref name=Zhu2013>{{Cite journal |author1=You-An Zhu |author2=Min Zhu |title=A redescription of Kiangyousteus yohii (Arthrodira: Eubrachythoraci) from the Middle Devonian of China, with remarks on the systematics of the Eubrachythoraci | journal=[[Zoological Journal of the Linnean Society]] |volume=169 |issue=4 |pages=798–819 |year=2013 |doi=10.1111/zoj12089 |doi-access=free }}</ref> -{{clade| style=font-size:85%;line-height:85% - |grouplabel1={{clade labels |width=10em; |label1='''Pachyosteomorphi''' |top1=65% |color1=green}} - |label1='''[[Eubrachythoraci]]''' - |1={{clade - |1={{clade - |label1='''[[Coccosteomorphi]]''' - |1={{clade - |1=''[[Coccosteus cuspidatus]]'' - |2={{clade - |1=''[[Harrytoombsia elegans]]'' - |2={{clade - |1=''[[Mcnamaraspis kaprios]]'' - |label2=[[Incisoscutoidea]] - |2={{clade - |1=''[[Compagopiscis croucheri]]'' - |2={{clade - |1=''[[Incisoscutum ritchiei]]'' - |2=''[[Incisoscutum sarahae]]'' }} - |label3=[[Camuropiscidae]] - |3={{clade - |1=''[[Latocamurus coulthardi]]'' - |2=''[[Camuropiscis laidlawi]]'' - |3={{clade - |1=''[[Rolfosteus canningensis]]'' - |2=''[[Tubonasus lennardensis]]'' - |3=''[[Fallacosteus turneri]]'' }} }} }} }} }} }} - |label2='''[[Aspinothoracidi]]''' - |2={{clade - |1=''[[Dinichthys herzeri]]'' |barbegin1=green - |2={{clade - |1=''[[Hadrosteus rapax]]'' |bar1=green - |2={{clade - |1=''[[Gorgonichthys clarki]]'' |bar1=green - |2={{clade - |1=''[[Heintzichthys gouldii]]'' |bar1=green - |label2=[[Selenosteidae]] - |2={{clade - |1=''[[Stenosteus angustopectus]]'' |bar1=green - |2={{clade - |1=''[[Gymnotrachelus hydei]]'' |bar1=green - |2={{clade - |1=''[[Rhinosteus parvulus]]'' |bar1=green - |2=''[[Pachyosteus bulla]]'' |bar2=green }} }} }} }} }} }} }} }} - |label2='''[[Dunkleosteoidea]]''' - |2={{clade - |1=''[[Westralichthys uwagedensis]]'' |bar1=green - |2={{clade - |1={{clade - |1=''[[Protitanichthys rockportensis]]'' |bar1=green - |label2=[[Panxiosteidae]] - |2={{clade - |1=''[[Panxiosteus ocullus]]'' |bar1=green - |2=''[[Janiosteus timanicus]]'' |bar2=green - |3=''[[Plourdosteus canadensis]]'' |bar3=green }} }} - |label2='''[[Dunkleosteidae]]''' - |2={{clade - |1={{clade - |1=''[[Eastmanosteus calliaspis]]'' |bar1=green - |2=''[[Xiangshuiosteus wui]]'' |bar2=green }} - |2={{clade - |1=''[[Eastmanosteus pustulosus]]'' |bar1=green - |2=''[[Kiangyousteus yohii]]'' |bar2=green - |3=''[[Golshanichthys asiatica]]'' |bar3=green - |4={{clade - |1='''''Dunkleosteus amblyodoratus''''' |bar1=green - |2='''''Dunkleosteus terrelli''''' |bar2=green - |3='''''Dunkleosteus raveri''''' |barend3=green }} }} }} }} }} }} }} - -Alternatively, the subsequent 2016 Zhu ''et al.'' study using a larger [[morphology (biology)|morphological]] dataset recovered [[Panxiosteidae]] well outside of [[Dunkleosteoidea]], leaving the status of [[Dunkleosteidae]] as a clade grouping separate from Dunkleosteoidea in doubt, as shown in the cladogram below:<ref name=Zhu2016>{{Cite journal |last1=Zhu |first1=You-An |last2=Zhu |first2=Min |last3=Wang |first3=Jun-Qing |date=April 1, 2016 |title=Redescription of Yinostius major (Arthrodira: Heterostiidae) from the Lower Devonian of China, and the interrelationships of Brachythoraci |journal=[[Zoological Journal of the Linnean Society]] |volume=176 |issue=4 |pages=806–834 |doi=10.1111/zoj.12356|issn=0024-4082 |doi-access=free }}</ref> -{{clade| style=font-size:85%;line-height:85% - |label1='''[[Eubrachythoraci]]''' - |1={{clade - |label1='''[[Coccosteomorphi]]''' - |1={{clade - |label1=[[Coccosteoidea]] - |1={{clade - |label1=[[Coccosteidae]] - |1={{clade - |1={{clade - |1=''[[Millerosteus minor]]'' - |2=''[[Coccosteus cuspidatus]]'' }} - |2={{clade - |1=''[[Dickosteus threiplandi]]'' - |2=''[[Watsonosteus fletti]]'' - |3=''[[Protitanichthys rockportensis]]'' }} }} - |label2='''[[Panxiosteidae]]''' - |2={{clade - |1=''[[Plourdosteus canadensis]]'' - |2=''[[Panxiosteus ocullus]]'' - |3=''[[Janiosteus timanicus]]'' }} }} - |label2=[[Incisoscutoidea]] - |2={{clade - |1=''[[Harrytoombsia elegans]]'' - |2={{clade - |1=''[[Torosteus tuberculatus]]'' - |2=''[[Torosteus pulchellus]]'' }} - |3={{clade - |1=''[[Mcnamaraspis kaprios]]'' - |2={{clade - |1=''[[Compagopiscis croucheri]]'' - |2=''[[Trematosteus fontanellus]]'' - |label3=[[Camuropiscidae]] - |3={{clade - |1={{clade - |1=''[[Incisoscutum ritchiei]]'' - |2=''[[Incisoscutum sarahae]]'' }} - |2={{clade - |1=''[[Rolfosteus canningensis]]'' - |2={{clade - |1=''[[Tubonasus lennardensis]]'' - |2={{clade - |1=''[[Fallacosteus turneri]]'' - |2={{clade - |1=''[[Camuropiscis laidlawi]]'' - |2=''[[Latocamurus coulthardi]]'' }} }} }} }} }} }} }} }} }} - |label2='''[[Pachyosteomorphi]]''' - |2={{clade - |1=''[[Rhachiosteus pterygiatus]]'' - |2={{clade - |label1='''[[Dunkleosteoidea]]''' - |1={{clade - |1=''[[Eastmanosteus calliaspis]]'' - |2={{clade - |1={{clade - |1=''[[Eastmanosteus pustulosus]]'' - |2=''[[Kiangyousteus yohii]]'' }} - |2={{clade - |1=''[[Golshanichthys asiatica]]'' - |2={{clade - |1=''[[Westralichthys uwagedensis]]'' - |2={{clade - |1={{clade - |1='''''Dunkleosteus raveri''''' - |2='''''Dunkleosteus terrelli''''' }} - |2={{clade - |1='''''Dunkleosteus amblyodoratus''''' - |label2=[[Heterostiidae]] - |2={{clade - |1=''[[Heterosteus ingens]]'' - |2=''[[Yinostius major]]'' }} }} }} }} }} }} }} - |label2='''[[Aspinothoracidi]]''' - |2={{clade - |1=''[[Tapinosteus heintzi]]'' - |2={{clade - |1={{clade - |1=''[[Bullerichthys fascidens]]'' - |2={{clade - |1=''[[Kendrickichthys cavernosus]]'' - |2=''[[Bruntonichthys multidens]]'' }} }} - |2={{clade - |1=''[[Dinichthys herzeri]]'' - |2={{clade - |1=''[[Hadrosteus rapax]]'' - |2={{clade - |1=''[[Gorgonichthys clarki]]'' - |label2=[[Selenosteidae]] - |2={{clade - |1=''[[Heintzichthys gouldii]]'' - |2=''[[Pachyosteus bulla]]'' - |3=''[[Gymnotrachelus hydei]]'' - |4=''[[Stenosteus angustopectus]]'' - |5=''[[Brachyosteus dietrichi]]'' - |6={{clade - |1=''[[Melanosteus occitanus]]'' - |2=''[[Rhinosteus parvulus]]'' }} }} }} }} }} }} }} }} }} }} }} - -=== Species === -At least ten different species<ref name="Carr+2010"/><ref name=Denison>{{cite book|last=Denison|first=Robert|chapter=Placodermi|volume=2|title=Handbook of Paleoichthyology|year=1978|publisher=Gustav Fischer Verlag|location=Stuttgart New York|isbn=978-0-89574-027-4|pages=128}}</ref> of ''Dunkleosteus'' have been described so far. -[[File:Dunkleosteus skull steveoc.jpg|thumb|Labelled skull diagram of ''D. terrelli'']] -The [[type species]], ''D. terrelli'', is the largest, best-known species of the genus, measuring {{convert|4.1|-|10|m|ft|abbr=on}} in length. It has a rounded snout. ''D. terrelli''{{'s}} fossil remains are found in Upper Frasnian to Upper Famennian Late Devonian strata of the United States (Huron and [[Cleveland Shale]] of Ohio, the Conneaut of [[Pennsylvania]], [[Chattanooga Shale]] of Tennessee, [[Lost Burro Formation]], California, and possibly Ives [[breccia]] of Texas<ref name=Denison/>) and [[Europe]]. - -''D. belgicus'' (?) is known from fragments described from the [[Famennian]] of [[Belgium]]. The median dorsal plate is characteristic of the genus, but, a plate that was described as a suborbital is anterolateral.<ref name=Denison/> Lelièvre (1982) considers this taxon a [[nomen dubium]] and suggests the material may actually pertain to ''[[Ardennosteus]]''.<ref>{{Cite journal |last=Lelièvre |first=Hervé |date=1982 |title=Ardennosteus ubaghsi n.g., n. sp. Brachythoraci primitif (vertébré, placoderme) du Famennien d'Esneux (Belgique) |url=https://popups.uliege.be/0037-9395/index.php?id=3220 |journal=Annales de la Société géologique de Belgique |volume=105 |issue=1}}</ref> - -''D. denisoni'' is known from a small median dorsal plate, typical in appearance for ''Dunkleosteus'', but much smaller than normal. It is comparable in skull structure to ''D. marsaisi''.<ref name=Denison/> - -[[File:Dunkleosteus marsaisi 45.JPG|thumb|''D. marsaisi'' skull]] -''D. marsaisi'' refers to the ''Dunkleosteus'' fossils from the Lower Famennian Late Devonian strata of the [[Atlas Mountains]] in [[Morocco]]. It differs in size, the known skulls averaging a length of {{convert|35|cm|ft}} and in form to ''D. terrelli''. In ''D. marsaisi'', the snout is narrower, and a postpineal fenestra may be present. Many researchers and authorities consider it a synonym of ''D. terrelli''.<ref>{{cite journal|last=Murray|first=A.M.|title=The Palaeozoic, Mesozoic and Early Cenozoic fishes of Africa|journal=Fish and Fisheries|year=2000|volume=1|issue=2|pages=111–145|doi=10.1046/j.1467-2979.2000.00015.x}}</ref> H. Schultze regards ''D. marsaisi'' as a member of ''[[Eastmanosteus]]''.<ref name=Denison/><ref name=Schultz>{{cite journal|last=Schultz|first=H|title=Large Upper Devonian arthrodires from Iran|journal=Fieldiana Geology|year=1973|volume=23|pages=53–78|doi=10.5962/bhl.title.5270|doi-access=free}}</ref> - -''D. magnificus'' is a large placoderm from the Frasnian [[Rhinestreet Shale]] of New York. It was originally described as ''[[Dinichthys]] magnificus'' by Hussakof and Bryant in 1919, then as "'' Dinichthys mirabilis''" by Heintz in 1932. Dunkle and Lane moved it to ''Dunkleosteus'' in 1971.<ref name=Denison/> - -''D. missouriensis'' is known from fragments from Frasnian [[Missouri]]. Dunkle and Lane regard them as being very similar to ''D. terrelli''.<ref name=Denison/> - -''D. newberryi'' is known primarily from a {{convert|28|cm|in}} long infragnathal with a prominent anterior cusp, found in the Frasnian portion of the [[Genesee Group]] of New York, and originally described as ''Dinichthys newberryi''.<ref name=Denison/> - -''D. amblyodoratus'' is known from some fragmentary remains from Late Devonian strata of [[Kettle Point Formation]], Ontario. The species name means 'blunt spear' and refers to the way the [[nuchal]] and [[paranuchal]] plates in the back of the head form the shape of a blunted spearhead.<ref name="Carr+2010"/> - -''D. raveri'' is a small species, possibly 1 meter long, known from an uncrushed skull roof found in a carbonate concretion from near the bottom of the Huron Shale, of the Famennian [[Ohio Shale]] strata. Besides its small size, it had comparatively large eyes. Because ''D. raveri'' was found in the strata directly below the strata where the remains of ''D. terrelli'' are found, ''D. raveri'' may have given rise to ''D. terrelli''. The species name commemorates Clarence Raver of [[Wakeman, Ohio]], who discovered the concretion where the [[holotype]] was found.<ref name="Carr+2010"/> - -''D. tuderensis'' is known from an infragnathal found in the lower-middle [[Famennian]]-aged [[Bilovo Formation]] of the [[Tver Region]] in northwest Russia. The specific name refers to the Maliy Tuder River as the holotyoe was found on its bank.<ref name=":3">{{Cite journal |last=Lebedev |first=Oleg A. |last2=Engelman |first2=Russell K. |last3=Skutschas |first3=Pavel P. |last4=Johanson |first4=Zerina |last5=Smith |first5=Moya M. |last6=Kolchanov |first6=Veniamin V. |last7=Trinajstic |first7=Kate |last8=Linkevich |first8=Valeriy V. |date=May 2023 |title=Structure, Growth and Histology of Gnathal Elements in Dunkleosteus (Arthrodira, Placodermi), with a Description of a New Species from the Famennian (Upper Devonian) of the Tver Region (North-Western Russia) |url=https://www.mdpi.com/1424-2818/15/5/648 |journal=Diversity |language=en |volume=15 |issue=5 |pages=648 |doi=10.3390/d15050648 |issn=1424-2818}}</ref> - -==Description== -===Size and anatomy=== -[[File:Dunkleosteus terrelli size.png|left|thumb|Comparison of multiple size estimates for ''D. terrelli'']] -Mainly the armored frontal sections of specimens have been [[fossilized]], and consequently, the appearance of the other portions of the fish is mostly unknown.<ref>{{cite AV media |url=https://www.youtube.com/watch?v=lEEhm4rzxEg |title=Prehistoric Monsters Revealed |date=2008 |last=Dash |first=Sean |publisher=Workaholic Productions / History Channel |location=United States |access-date=December 18, 2015}}{{cbignore}}{{Dead YouTube link|date=February 2022}}</ref> In fact, only about 5% of ''Dunkleosteus'' specimens have more than a quarter of their skeleton preserved.<ref>Carr, R, & G.L. Jackson. 2008. [https://www.researchgate.net/publication/213769842_THE_VERTEBRATE_FAUNA_OF_THE_CLEVELAND_MEMBER_FAMENNIAN_OF_THE_OHIO_SHALE The Vertebrates fauna of the Cleveland member (Famennian) of the Ohio Shale. Society of Vertebrates Paleontology]. 1–17.</ref> Because of this, many reconstructions of the hindquarters are often based on fossils of smaller [[arthrodire]]s, such as ''[[Coccosteus]]'', which have preserved hind sections,<ref name=":0" /> leading to widely varying size estimates.<ref name=":0" /> - -''Dunkleosteus'' is one of the largest known placoderms, with its maximum size being variably estimated as anywhere from {{convert|4.1|–|10|m|ft}} by different researchers.<ref name="Anderson&Westneat2007" /><ref name="Anderson&Westneat2009" /><ref name="Ferron2017" /><ref>{{cite journal|last=Williams|first=Nigel|year=2007|title=Force feeding|journal=Current Biology |volume=17 |pages=R3 |doi=10.1016/j.cub.2006.11.057|s2cid=36585467 }}</ref><ref name=":0">{{Cite journal |last=Engelman |first=Russell K. |date=2023|title=A Devonian Fish Tale: A New Method of Body Length Estimation Suggests Much Smaller Sizes for Dunkleosteus terrelli (Placodermi: Arthrodira) |journal=Diversity |language=en |volume=15 |issue=3 |pages=318 |doi=10.3390/d15030318 |issn=1424-2818 |doi-access=free }}</ref> However, most cited length estimates are speculative and lack quantitative or statistical backing.<ref name="Ferron2017" /><ref name=":0" /> Most studies that estimate the length of ''Dunkleosteus'' do not provide information as to how these estimates were calculated, the measurements used to scale them, or which specimens were examined. Most are implied to be based on either CMNH 5768 (the largest complete armor of ''Dunkleosteus'') or CMNH 5936 (the largest known jaw fragment). -{{multiple image -|align=right -|perrow=1 -|total_width=240 - -|image1=Dunkleosteus terrelli 2017 reconstruction.png -|caption1=[[Paleoart|Life reconstruction]] of ''D. terrelli'', as presented by Ferrón ''et al.'' 2017<ref name="Ferron2017"/> - -|image2=Dunkleosteus terrelli 2023 reconstruction.png -|caption2=Life reconstruction of ''D. terrelli'', as presented by Engelman 2023<ref name=":0"/> -}} - -===Revaluation of physiology=== -Most of the studies with well-defined methods produce lengths of {{convert|5|m|ft}} or less for ''Dunkleosteus terrelli'',<ref name=":0" /> with the exception of Ferrón ''et al.'' (2017), which produces larger estimates of {{convert|6.88|–|8.79|m|ft}} based on upper jaw perimeter of modern sharks.<ref name="Ferron2017" /> However, arthrodires have proportionally larger mouths than modern sharks, making the lengths estimated by Ferrón ''et al.'' (2017) unreliable.<ref name=":2">{{cite journal |last=Engelman |first=Russell |date=2023-04-10 |title=Giant, swimming mouths: oral dimensions of extant sharks do not accurately predict body size in ''Dunkleosteus terrelli'' (Placodermi: Arthrodira) |url=https://peerj.com/articles/15131 |journal=PeerJ |language=en |volume=11 |pages=e15131 |doi=10.7717/peerj.15131 |pmid=37065696 |pmc=10100833 }}</ref> Upper jaw perimeter overestimates the size of complete arthrodires like ''[[Coccosteus]]'' and the estimates of Ferrón ''et al.'' (2017) result in ''Dunkleosteus'' having an extremely small head and hyper-elongate trunk relative to the known dimensions of the fossils.<ref name=":2" /> If the reconstruction presented in Ferrón ''et al.'' (2017) is scaled to the known dimensions of CMNH 5768, it produces a length of {{convert|3.77|m|ft}}.<ref name=":2" /> - -Carr (2010) estimated a {{convert|4.6|m|ft}} long adult individual of ''Dunkleosteus terrelli'' to have weighed {{convert|665|kg|lb}}, assuming a shark-like body plan and a similar length-weight relationship.<ref name="Carr 20102">{{cite journal |last1=Carr |first1=Robert K. |date=2010 |title=Paleoecology of Dunkleosteus terrelli (Placodermi: Arthrodira). |url=https://www.researchgate.net/publication/235924093 |journal=Kirtlandia |volume=57}}</ref> Engelman (2023), using an [[ellipsoid]] [[Volume|volumetric]] method, estimated weights of {{convert|950|–|1200|kg|lb}} for typical ({{convert|3.41|m|ft}} long) adult ''Dunkleosteus'', and weights of {{convert|1490|–|1760|kg|lb}} for the largest ({{convert|4.1|m|ft|1}} in this study) individual.<ref name=":0" /> The higher weights by Engelman (2023) are mostly a result of the fact that arthrodires tend to have relatively deeper and wider bodies compared to sharks.<ref name=":0" /> - -An exceptionally preserved specimen of ''D. terrelli'' preserves a [[Fish fin|pectoral fin]] outline with [[ceratotrichia]], implying that the fin morphology of placoderms was much more variable than previously thought, and was heavily influenced by locomotory requirements. This knowledge, coupled with the knowledge that fish morphology is more heavily influenced by feeding niche than phylogeny, allowed a 2017 study to infer the [[Fish fin|caudal fin]] shape of ''D. terrelli'', reconstructing this fin with a strong ventral lobe, a high [[aspect ratio]], narrow [[caudal peduncle]], in contrast to previous reconstructions based on the [[Fish locomotion|anguilliform]] caudal fin of coccosteomorph placoderms.<ref name="Ferron2017">{{Cite journal|last1=Ferrón|first1=Humberto G.|last2=Martínez-Pérez|first2=Carlos|last3=Botella|first3=Héctor|date=2017|title=Ecomorphological inferences in early vertebrates: reconstructing Dunkleosteus terrelli (Arthrodira, Placodermi) caudal fin from palaeoecological data|journal=PeerJ|language=en|volume=5|pages=e4081|doi=10.7717/peerj.4081|pmid=29230354|pmc=5723140|issn=2167-8359}}</ref> - -====Length estimations of ''D. terrelli''==== -{| class="wikitable sortable" style="margin-left: auto; margin-right: auto; border: none;" -|+ Length estimates of ''Dunkleosteus terrelli'' (modified from Table 7 of Engelman 2023<ref name=":0" />) -! Study (author) -! Year -! Length -! Method -! Reference -|- -| Newberry || 1875 || {{convert|4.5|–|5.5|m|ft}} || '''Extrapolated from ''Coccosteus cuspidatus'', measurements and specimen used unclear''' || <ref name=":1">{{cite book |last1=Newberry |first1=John S. |url=https://www.biodiversitylibrary.org/item/116994#page/15/mode/1up |title=Report of the Geological Survey of Ohio. Volume II. Geology and Paleontology |date=1875 |publisher=Nevins and Myers, State Printers |volume=2 |location=Columbus |page=24 |contribution=Descriptions of fossil fishes}}</ref> -|- -| Newberry || 1889 || {{convert|4.5|m|ft}} || Unstated (implied extrapolation from ''Coccosteus'') || <ref>{{cite journal |last1=Newberry |first1=John S. |date=1889 |title=Paleozoic fishes of North America |url=https://www.biodiversitylibrary.org/item/51025#page/5/mode/1up |journal=Monographs of the U.S. Geological Survey |volume=16 |page=24}}</ref> -|- -| Dean || 1895 || {{convert|3|m|ft}} || Methods, measurements, and specimens unstated || <ref>{{cite book |last1=Dean |first1=Bashford |title=Fishes, Living and Fossil: An Outline of Their Forms and Probable Relationships |date=1895 |publisher=Macmillan and Company |location=London |page=130}}</ref> -|- -| Hussakof || 1905 || {{convert|1.67|m|ft}} (AMNH FF 195) <br> {{convert|3.79|m|ft}} (extrapolated to CMNH 5768 by Engelman 2023<ref name=":0"/> assuming similar head-trunk proportions) || '''Entering angle of body''' || <ref>{{cite journal |last1=Hussakof |first1=Louis |title=Notes on the Devonian "placoderm" ''Dinichthys intermedius'' Newb. |journal=Bulletin of the American Museum of Natural History |date=1905 |volume=21 |pages=27–36}}</ref> -|- -| Anonymous|| 1923 || {{convert|7.6|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Anonymous |title=Cleveland shale fishes |journal=Bulletin of the Cleveland Museum of Natural History |date=1923 |volume=9 |pages=36}}</ref> -|- -| Hyde || 1926 || {{convert|4.5|–|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Hyde |first1=Jesse E. |title=Collecting fossil fishes from the Cleveland Shale |journal=Natural History |date=1926 |volume=26 |pages=497–504}}</ref> -|- -| Romer || 1966 || {{convert|9|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Romer |first1=Alfred S. |title=Vertebrate Paleontology |date=1966 |publisher=University of Chicago Press |location=Chicago |page=49 |edition=3rd}}</ref> -|- -| Colbert || 1969 || {{convert|9|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Colbert |first1=Edwin H. |title=Evolution of the Vertebrates: A History of Backboned Animals Through Time |date=1969 |publisher=John Wiley and Sons |location=Hoboken |page=36 |edition=2nd}}</ref> -|- -| Denison || 1978 || {{convert|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref name=Denison /> -|- -| Williams || 1992 || {{convert|5|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Williams |first1=Michael E. |title=Jaws: The Early Years |journal=Explorer |date=1992 |volume=34 |pages=4–8}}</ref> -|- -| Janvier || 2003 || {{convert|6|–|7|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Janvier |first1=P. |title=Early Vertebrates |date=2003 |publisher=Clarendon Press |location=Oxford |page=12}}</ref> -|- -| Young || 2003 || {{convert|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Young |first1=Gavin C. |title=Did placoderm fish have teeth? |journal=Journal of Vertebrate Paleontology |date=24 December 2003 |volume=23 |issue=4 |pages=987–990 |doi=10.1671/31|bibcode=2003JVPal..23..987Y |s2cid=85572061 }}</ref> -|- -| Anderson and Westneat || 2007 || {{convert|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref name="Anderson&Westneat2007" /> -|- -| Anderson and Westneat || 2009 || {{convert|10|m|ft}} || Methods, measurements, and specimens used not stated || <ref name="Anderson&Westneat2009" /> -|- -| Carr || 2010 || {{convert|4.5|–|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref name="Carr 2010">{{cite journal |last1=Carr |first1=Robert K. |date=2010 |title=Paleoecology of Dunkleosteus terrelli (Placodermi: Arthrodira). |url=https://www.researchgate.net/publication/235924093 |journal=Kirtlandia |volume=57}}</ref> -|- -| Long || 2010 || {{convert|4.5|–|8|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Long |first1=John A. |title=The Rise of Fishes: 500 Million Years of Evolution |date=2010 |publisher=Johns Hopkins University Press |location=Baltimore |pages=88–90 |edition=2nd}}</ref> -|- -| Sallan and Galimberti || 2015 || {{convert|8|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Sallan |first1=Lauren |last2=Galimberti |first2=Andrew K. |title=Body-size reduction in vertebrates following the end-Devonian mass extinction |journal=Science |date=13 November 2015 |volume=350 |issue=6262 |pages=812–815 |doi=10.1126/science.aac7373|pmid=26564854 |bibcode=2015Sci...350..812S |s2cid=206640186 }}</ref> -|- -| Ferrón ''et al.'' || 2017 || {{convert|6.88|m|ft}} (average adult, CMNH 5768) <br> {{convert|8.79|m|ft}} (largest individual, CMNH 5936) || '''Upper jaw perimeter''' || <ref name="Ferron2017" /> -|- -| Long ''et al.'' || 2019 || {{convert|6|–|8|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Long |first1=John A. |last2=Choo |first2=Brian |last3=Clement |first3=Alice |contribution=The Evolution of Fishes through Geological Time |title=Evolution and Development of Fishes |editor1-last=Johanson |editor1-first=Zerina|editor2-last=Underwood |editor2-first=Charlie|editor3-last=Richter|editor3-first=Martha| date=31 December 2018 |pages=3–29 |doi=10.1017/9781316832172.002|s2cid=134217082 }}</ref> -|- -| Johanson ''et al.'' || 2019 || {{convert|3|m|ft}} (CMNH 50322) <br> {{convert|7.1|m|ft}} (extrapolated to CMNH 5768 by Engelman 2023 assuming similar head-trunk proportions) || Methods and measurements not stated || <ref>{{cite journal |last1=Johanson |first1=Zerina |last2=Trinajstic |first2=Kate |last3=Cumbaa |first3=Stephen |last4=Ryan |first4=Michael |title=Fusion in the vertebral column of the pachyosteomorph arthrodire Dunkleosteus terrelli ('Placodermi') |journal=Palaeontologia Electronica |date=2019 |volume=22.2.20A |pages=1–13 |doi=10.26879/872|s2cid=162173408 }}</ref> -|- -| Engelman || 2023 || {{convert|3.41|m|ft}} (average adult, CMNH 5768) <br> {{convert|4.1|m|ft|1}} (largest individual, CMNH 5936) || '''Orbit-opercular length (head length minus snout)''' || <ref name=":0" /> -|- -| Engelman || 2023 || {{convert|3.41|m|ft}} (average adult, CMNH 5768) || '''Skull length in ''Coccosteus''''' || <ref name=":0" /> -|- -| Engelman || 2023 || {{convert|5.23|m|ft}} (average adult, CMNH 5768) || '''Infragnathal length in ''Coccosteus''''' (source considers this estimate unreliable due to ''Dunkleosteus'' having a relatively larger mouth than ''Coccosteus'') || <ref name=":0" /> -|- -| Engelman || 2023 || {{convert|3.47|m|ft}} (average adult, CMNH 5768) || '''Entering angle of body''' || <ref name=":0" /> -|- -| Engelman || 2023 || {{convert|3.88|m|ft}} (average adult, CMNH 5768) || '''Length of posteroventrolateral plate''' || <ref name=":0" /> -|- -| Engelman || 2023 || {{convert|3.40|m|ft}} (average adult, CMNH 5768) || '''Inferred location of pelvic girdle''' || <ref name=":0" /> -|- -|} - -==Paleobiology== -===Diet=== -[[File:Dunkleosteus terrelli CMNH 5936.png|thumb|left|Lower jaw of ''D. terrelli'' specimen CMNH 5936]] -''Dunkleosteus terrelli'' possessed a [[four-bar linkage]] mechanism for [[Fish jaw|jaw]] opening that incorporated connections between the skull, the thoracic shield, the lower jaw and the jaw muscles joined by movable joints.<ref name=Anderson&Westneat2009>{{cite journal |last=Anderson |first=P.S.L. |author2=Westneat, M. |title=A biomechanical model of feeding kinematics for ''Dunkleosteus terrelli'' (Arthrodira, Placodermi) |journal=Paleobiology |date=2009 |volume=35 |issue=2 |pages=251–269 |doi=10.1666/08011.1 |bibcode=2009Pbio...35..251A |s2cid=86203770 |url=http://projects.fieldmuseum.org/sites/default/files/DunkPaleoBio.pdf |access-date=October 29, 2017 |archive-date=November 7, 2017 |archive-url=https://web.archive.org/web/20171107014026/http://projects.fieldmuseum.org/sites/default/files/DunkPaleoBio.pdf |url-status=dead }}</ref><ref name=Anderson&Westneat2007>{{Cite journal |last=Anderson |first=P.S.L. |author2=Westneat, M. |title=Feeding mechanics and bite force modelling of the skull of Dunkleosteus terrelli, an ancient apex predator |journal=Biology Letters |volume=3 |issue=1 |pages=76–79 |date=2007 |doi=10.1098/rsbl.2006.0569 |pmid=17443970 |pmc=2373817}}</ref> This mechanism allowed ''D. terrelli'' to both achieve a high speed of jaw opening, opening their jaws in 20 milliseconds and completing the whole process in 50–60 milliseconds (comparable to modern fishes that use [[Aquatic feeding mechanisms#Suction feeding|suction feeding]] to assist in prey capture;<ref name=Anderson&Westneat2007/>) and producing high bite forces when closing the jaw, estimated at {{convert|4414|N|kg-f lb-f|0|abbr=on}} at the tip and {{convert|5363|N|kg-f lb-f|0|abbr=on}} at the blade edge,<ref name=Anderson&Westneat2007/> or even up to {{cvt|6170|N|kg-f lb-f|0}} and {{cvt|7495|N|kg-f lb-f|0}} respectively.<ref name=Anderson&Westneat2009/> The pressures generated in those regions were high enough to puncture or cut through [[cuticle]] or [[dermal bone|dermal]] armor,<ref name=Anderson&Westneat2007/> suggesting that ''D. terrelli'' was adapted to prey on free-swimming, armored prey such as [[ammonite]]s and other placoderms.<ref name=Anderson&Westneat2009/> - -In addition, teeth of a [[Chondrichthyes|chondrichthyan]] thought to belong to ''[[Orodus]]'' (''Orodus'' spp.) were found in association with ''Dunkleosteus'' remains, suggesting that these were probably stomach contents regurgitated from the animal. ''Orodus'' is thought to be tachypelagic, or a fast-swimming [[pelagic fish]]. Thus, ''Dunkleosteus'' might have been fast enough to catch these fast organisms, and not a slow swimmer like originally thought.<ref name="Ferron2017" /> Fossils of ''Dunkleosteus'' are frequently found with [[Bolus (digestion)|bolus]]es of fish bones, semidigested and partially eaten remains of other fish.<ref name="Virtual Fossil">{{cite web|url= http://www.fossilmuseum.net/Fossil_Galleries/Fish_Devonian/Dunkleosteous/Dunkleosteus.htm |title=Dunkleosteus Placodermi Devonian Armored Fish from Morocco |work=Fossil Archives |publisher=The Virtual Fossil Museum |access-date=April 26, 2009}}</ref> As a result, the [[fossil]] record indicates it may have routinely regurgitated prey bones rather than digest them. Mature individuals probably inhabited deep sea locations, like other placoderms, living in shallow waters during adolescence.<ref>{{Cite web|url=http://bioweb.uwlax.edu/bio203/f2013/peters_sadi/habitat.htm|title=UWL Website}}</ref> - -A specimen of ''Dunkleosteus'' (CMNH 5302), and ''[[Titanichthys]]'' (CMNH 9889), show damage said to be puncture damage from the bony fangs of other ''Dunkleosteus''.<ref name="Anderson&Westneat2009" /> - -===Reproduction=== -''Dunkleosteus'', together with most other placoderms, may have also been among the first [[vertebrate]]s to [[internal fertilization|internalize egg fertilization]], as seen in some modern sharks.<ref name="Ahlberg">{{cite journal |last1=Ahlberg |first1=Per |first2=Kate |last2=Trinajstic |first3=Zerina |last3=Johanson |first4=John |last4=Long |s2cid=205217467 |year=2009 |title=Pelvic claspers confirm chondrichthyan-like internal fertilization in arthrodires |journal=Nature |volume=460 |issue=7257 |pages=888–889 |doi=10.1038/nature08176|pmid=19597477 |bibcode=2009Natur.460..888A }}</ref> Some other placoderms have been found with evidence that they may have been [[viviparity|viviparous]], including what appears to have been an [[umbilical cord]].<ref>{{ cite journal | last = Long | first = J. A. |author2=Trinajstic, K. |author3=Young, G. C. |author4=Senden, T. | year = 2008 | title = Live birth in the Devonian period | journal = Nature | volume = 453 | issue = 7195 | pages = 650&ndash;652 | doi = 10.1038/nature06966 | pmid = 18509443 | bibcode = 2008Natur.453..650L | s2cid = 205213348 }}</ref> - -===Growth=== -{{multiple image -|align=right -|perrow=2 -|total_width=380 - -|image1=Dunkleosteus juvenile fossil - Cleveland Museum of Natural History (34421618320).jpg -|caption1=''D. terrelli'' juvenile specimen CMNH 7424 - -|image2=Dunkleosteus terrelli - Cleveland Museum of Natural History - 2014-12-26 (21137540331).jpg -|caption2=''D. terrelli'' adult specimen CMNH 5768 - -}} -[[morphology (biology)|Morphological]] studies on the lower jaws of juveniles of ''D. terrelli'' reveal they were proportionally as robust as those of adults, indicating they already could produce high bite forces and likely were able to shear into resistant prey tissue similar to adults, albeit on a smaller scale. This pattern is in direct contrast to the condition common in [[tetrapod]]s in which the jaws of juveniles are more [[gracile]] than in adults.<ref name="Snively_etal_2009">{{cite journal |last=Snively |first=E. |author2=Anderson, P.S.L. |author3=Ryan, M.J. |date=2009 |title=Functional and ontogenetic implications of bite stress in arthrodire placoderms |url=https://www.biodiversitylibrary.org/page/51811342#page/60/mode/1up |journal=Kirtlandia |volume=57}}</ref> - -==See also== -* [[List of placoderms]] - -==References== -{{notelist}} -{{reflist}} - -==Further reading== -* {{cite journal|first=Philip S. L.|last=Anderson|s2cid=86583150|year=2008|title=Shape Variation Between Arthrodire Morphotypes Indicates Possible Feeding Niches|journal=Journal of Vertebrate Paleontology|volume=28|number=4|pages=961–969|doi=10.1671/0272-4634-28.4.961|bibcode=2008JVPal..28..961A }} - -==External links== -{{Commons category|Dunkleosteus}} -{{Wikispecies|Dunkleosteus}} -* [http://www.ucmp.berkeley.edu/vertebrates/basalfish/placodermi.html ''Introduction to the Placodermi: Extinct Armored Fishes with Jaws''. Waggoner, Ben (2000). Retrieved Aug 1, 2005] -* [http://www.nbcnews.com/id/15940709 MSNBC: Prehistoric fish packed a mean bite] -* [http://news.bbc.co.uk/1/hi/sci/tech/6195188.stm BBC: Ancient 'Jaws' had monster bite] - -{{Arthrodira|E.}} -{{Taxonbar|from=Q131039|from2=Q911597|from3=Q911599|from4=Q911605|from5=Q911606|from6=Q911611|from7=Q911612|from8=Q911616|from9=Q911618|from10=Q911593|from11=Q25357547}} - -[[Category:Dunkleosteidae]] -[[Category:Apex predators]] -[[Category:Placoderms of Africa]] -[[Category:Fossils of Morocco]] -[[Category:Placoderms of Europe]] -[[Category:Fossils of Belgium]] -[[Category:Placoderms of North America]] -[[Category:Fossils of Canada]] -[[Category:Paleontology in Missouri]] -[[Category:Paleontology in Pennsylvania]] -[[Category:Paleontology in Tennessee]] -[[Category:Late Devonian first appearances]] -[[Category:Late Devonian animals]] -[[Category:Famennian extinctions]] -[[Category:Fossil taxa described in 1956]] -[[Category:Taxa named by John Strong Newberry]] +Sksksks '
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[ 0 => '{{Short description|Genus of extinct fishes}}', 1 => '{{Automatic taxobox', 2 => '| fossil_range = [[Late Devonian]] ([[Frasnian]] to [[Famennian]]), {{Fossil range|382|358}}', 3 => '| image = Dunkleosteus terrelli (fossil fish) (Cleveland Shale Member, Ohio Shale, Upper Devonian; Rocky River Valley, Cleveland, Ohio, USA) 21 (34001200911).jpg', 4 => '| image_caption = Partially reconstructed ''D. terrelli'' skull (specimen CMNH 5768), [[Cleveland Museum of Natural History]]', 5 => '| display_parents = 7', 6 => '| taxon = Dunkleosteus', 7 => '| authority = [[Jean-Pierre Lehman|Lehman]], 1956', 8 => '| type_species = †'''''Dunkleosteus terrelli'''''', 9 => '| type_species_authority = [[John Strong Newberry|Newberry]], 1873', 10 => '| subdivision_ranks = [[Species]]', 11 => '| subdivision = {{collapsible list|', 12 => '''D. terrelli'' <small>(Newberry, 1873 [originally ''[[Dinichthys]]''])</small><br />', 13 => '''D. ? belgicus'' <small>(Newberry) 1873</small><br />', 14 => '''D. denisoni'' <small>(Kulczycki) 1957</small><br />', 15 => '''D. marsaisi'' <small>Lehmann, 1956</small><br />', 16 => '''D. magnificus'' <small>(Hussakof & Bryant) 1919</small><br />', 17 => '''D. missouriensis'' <small>(Branson) 1914</small><br />', 18 => '''D. newberryi'' <small>(Clarke) 1885</small><br />', 19 => '''D. amblyodoratus'' <small>Carr & Hlavin, 2010</small><br />', 20 => '''D. raveri'' <small>Carr & Hlavin, 2010</small><br />', 21 => '''D. tuderensis'' <small>Lebedev et. al. 2023</small><br />', 22 => '}}', 23 => '}}', 24 => '', 25 => ''''''Dunkleosteus''''' is an [[extinct]] [[genus]] of large [[arthrodira|arthrodire]] fish that existed during the [[Late Devonian]] period, about 382–358 [[million years ago]]. It consists of ten species, some of which are among the largest [[placoderm]]s to have ever lived: ''D. terrelli'', ''D. belgicus'', ''D. denisoni'', ''D. marsaisi'', ''D. magnificus'', ''D. missouriensis'', ''D. newberryi'', ''D. amblyodoratus'', ''D. raveri'', and ''D. tuderensis,'' and the largest and most well known species is ''D. terrelli''. Since body shape is not known, various methods of estimation put the living total length of the largest known specimen between {{convert|4.1|to|10|m|ft|abbr=on}} long and weigh around {{convert|1|-|4|MT|ST|abbr=on}}.<ref name=":0" /> ''Dunkleosteus'' could quickly open and close its jaw, like modern-day [[Aquatic feeding mechanisms#Suction feeding|suction feeders]], and had a bite force of {{convert|4414|-|6170|N|kg-f lb-f|0|abbr=on}} at the tip and {{convert|5363|-|7495|N|kg-f lb-f|0|abbr=on}} at the blade edge.<ref name=Anderson&Westneat2007/><ref name=Anderson&Westneat2009/> Numerous [[fossil]]s of the various species have been found in [[North America]], [[Poland]], [[Belgium]], and [[Morocco]]. ''Dunkleosteus'' was a [[pelagic fish]] inhabiting open waters, and an [[apex predator]] of its ecosystem.', 26 => '', 27 => '==Discovery==', 28 => '''Dunkleosteus'' fossils were first discovered in 1867 by Jay Terrell, a hotel owner and amateur paleontologist who collected fossils in the cliffs along [[Lake Erie]] near his home of [[Sheffield Lake, Ohio]] (due west of [[Cleveland]]). Terrell donated his fossils to [[John Strong Newberry]] and the Ohio Geological Survey, who in 1873 described all the material as belonging to a single new [[genus]] and [[species]]: ''[[Dinichthys|Dinichthys herzeri]]''. However, with later fossil discoveries by 1875 it became apparent multiple large fish species were present in the [[Ohio Shale]]. ''[[Dinichthys|Dinichthys herzeri]]'' came from the lowermost layer, the [[Huron Shale]], whereas most of the fossils were coming from the younger [[Cleveland Shale]] and represented a distinct species.<ref name=":1" /> Newberry named this more common species "''Dinichthys''" ''terrelli'', after Terrell. Most of Terrell's original collection does not survive, having been destroyed by a fire in [[Elyria, Ohio|Elyria]] in 1873.<ref name=":1" /><ref>{{Cite journal |last=Claypole |first=E. W. |date=1893 |title=The three great fossil placoderms of Ohio |url=https://www.biodiversitylibrary.org/item/240810#page/107/mode/1up |journal=American Geologist |volume=12 |pages=89–99}}</ref>', 29 => '', 30 => 'The largest collection of ''Dunkleosteus'' fossils in the world is housed at the [[Cleveland Museum of Natural History]],<ref>{{cite web |url=https://www.cmnh.org/dunk |title=Dunkleosteus terrelli: Fierce prehistoric predator |author=<!--Not stated--> |website=Cleveland Museum of Natural History |access-date=July 21, 2022 |archive-date=May 19, 2021 |archive-url=https://web.archive.org/web/20210519150010/https://www.cmnh.org/dunk |url-status=dead }}</ref> with smaller collections (in descending order of size) held at the [[American Museum of Natural History]],<ref>{{cite web |url=https://www.amnh.org/exhibitions/permanent/vertebrate-origins/dunkleosteus |title=Dunkleosteus |author=<!--Not stated--> |website=American Museum of Natural History |access-date=July 21, 2022}}</ref> [[Smithsonian National Museum of Natural History]],<ref name="Smithsonian">{{cite web |title=Collections Catalog of the Department of Paleobiology of the Smithsonian National Museum of Natural History |url=https://collections.nmnh.si.edu/search/paleo/ |access-date=12 August 2022 |publisher=Smithsonian Institute}}</ref> [[Yale Peabody Museum]],<ref>{{cite web |title=Collections Database of the Yale Peabody Museum |url=https://collections.peabody.yale.edu/search/Search/Results?sort=relevance&join=AND&lookfor0%5B%5D=dunkleosteus&type0%5B%5D=AllFields&lookfor0%5B%5D=&type0%5B%5D=AllFields&lookfor0%5B%5D=&type0%5B%5D=AllFields&bool0%5B%5D=AND&lookfor1%5B%5D=VP&type1%5B%5D=CatalogNumber&lookfor1%5B%5D=VPPU&type1%5B%5D=CatalogNumber&bool1%5B%5D=OR&filter%5B%5D=%7Ecollection%3A%22Vertebrate+Paleontology%22&limit=5&daterange%5B%5D=collecting_year_first&collecting_year_firstfrom=&collecting_year_firstto= |publisher=Yale Peabody Museum |access-date=12 August 2022}}</ref> the [[Natural History Museum in London]], and the [[Cincinnati Museum Center]]. Specimens of ''Dunkleosteus'' are on display in many museums throughout the world (see table below), most of which are casts of the same specimen: CMNH 5768, the largest well-preserved individual of ''D. terrelli''.<ref name="Ferron2017" /> The original CMNH 5768 is on display in the [[Cleveland Museum of Natural History]].', 31 => '', 32 => '==Taxonomy==', 33 => '''Dunkleosteus'' was named in 1956 to honour [[David Dunkle]] (1911–1982), former curator of [[vertebrate paleontology]] at the [[Cleveland Museum of Natural History]]. The genus name ''Dunkleosteus'' combines [[David Dunkle]]'s surname with the Greek word {{wikt-lang|grc|ὀστέον}} ({{transliteration|grc|ostéon}} 'bone'), literally meaning 'Dunkle's-bone'. The [[type species]] ''D. terrelli'' was originally described in 1873 as a species of ''[[Dinichthys]]'', its specific epithet chosen in honor of Jay Terrell, the fossil's discoverer.<ref>"Dunkleosteus terrelli: Fierce prehistoric predator" page at ''Cleveland Museum of Natural History''. https://www.cmnh.org/dunk {{Webarchive|url=https://web.archive.org/web/20210519150010/https://www.cmnh.org/dunk |date=2021-05-19 }}</ref>', 34 => '', 35 => 'Originally thought to be a member of the genus ''[[Dinichthys]]'', ''Dunkleosteus'' was later recognized as belonging to its own genus in 1956. It was thought to be closely related to ''Dinichthys'', and they were grouped together in the [[family (biology)|family]] [[Dinichthyidae]]. However, in the 2010 Carr & Hlavin [[phylogenetic]] study, ''Dunkleosteus'' and ''Dinichthys'' were found to belong to two separate [[clade]]s. Carr & Hlavin resurrected the family [[Dunkleosteidae]] and placed ''Dunkleosteus'', ''[[Eastmanosteus]]'', and a few other genera from Dinichthyidae within it.<ref name="Carr+2010">{{cite journal| author =Carr R. K., Hlavin V. J.| title =Two new species of Dunkleosteus Lehman, 1956, from the Ohio Shale Formation (USA, Famennian) and the Kettle Point Formation (Canada, Upper Devonian), and a cladistic analysis of the Eubrachythoraci (Placodermi, Arthrodira)| journal =Zoological Journal of the Linnean Society| volume =159| issue =1| pages =195–222| year =2010| doi =10.1111/j.1096-3642.2009.00578.x| doi-access =free}}</ref> Dinichthyidae, in turn, is left a [[Monotypic taxon|monospecific]] family.<ref name=Carr1995>{{cite journal|last=Carr|first=Robert K.|author2=William J. Hlavin|title=Dinichthyidae (Placodermi):A paleontological fiction?|journal=Geobios|date=September 2, 1995|volume=28|pages=85–87|doi=10.1016/S0016-6995(95)80092-1|bibcode=1995Geobi..28...85C }}</ref>', 36 => '[[File:Dunkleosteus terrelli (fossil fish) (Cleveland Shale Member, Ohio Shale, Upper Devonian; Rocky River Valley, Cleveland, Ohio, USA) 25 (34091142396).jpg|thumb|right|Front view of ''D. terrelli'' skull]] ', 37 => 'The [[cladogram]] below from the 2013 Zhu & Zhu study shows the placement of ''Dunkleosteus'' within [[Dunkleosteidae]] and ''[[Dinichthys]]'' within the separate clade [[Aspinothoracidi]]:<ref name=Zhu2013>{{Cite journal |author1=You-An Zhu |author2=Min Zhu |title=A redescription of Kiangyousteus yohii (Arthrodira: Eubrachythoraci) from the Middle Devonian of China, with remarks on the systematics of the Eubrachythoraci | journal=[[Zoological Journal of the Linnean Society]] |volume=169 |issue=4 |pages=798–819 |year=2013 |doi=10.1111/zoj12089 |doi-access=free }}</ref>', 38 => '{{clade| style=font-size:85%;line-height:85%', 39 => ' |grouplabel1={{clade labels |width=10em; |label1='''Pachyosteomorphi''' |top1=65% |color1=green}}', 40 => ' |label1='''[[Eubrachythoraci]]'''', 41 => ' |1={{clade', 42 => ' |1={{clade', 43 => ' |label1='''[[Coccosteomorphi]]'''', 44 => ' |1={{clade', 45 => ' |1=''[[Coccosteus cuspidatus]]''', 46 => ' |2={{clade', 47 => ' |1=''[[Harrytoombsia elegans]]''', 48 => ' |2={{clade', 49 => ' |1=''[[Mcnamaraspis kaprios]]''', 50 => ' |label2=[[Incisoscutoidea]]', 51 => ' |2={{clade', 52 => ' |1=''[[Compagopiscis croucheri]]''', 53 => ' |2={{clade', 54 => ' |1=''[[Incisoscutum ritchiei]]''', 55 => ' |2=''[[Incisoscutum sarahae]]'' }}', 56 => ' |label3=[[Camuropiscidae]]', 57 => ' |3={{clade', 58 => ' |1=''[[Latocamurus coulthardi]]''', 59 => ' |2=''[[Camuropiscis laidlawi]]''', 60 => ' |3={{clade', 61 => ' |1=''[[Rolfosteus canningensis]]''', 62 => ' |2=''[[Tubonasus lennardensis]]''', 63 => ' |3=''[[Fallacosteus turneri]]'' }} }} }} }} }} }}', 64 => ' |label2='''[[Aspinothoracidi]]'''', 65 => ' |2={{clade', 66 => ' |1=''[[Dinichthys herzeri]]'' |barbegin1=green', 67 => ' |2={{clade', 68 => ' |1=''[[Hadrosteus rapax]]'' |bar1=green', 69 => ' |2={{clade', 70 => ' |1=''[[Gorgonichthys clarki]]'' |bar1=green', 71 => ' |2={{clade', 72 => ' |1=''[[Heintzichthys gouldii]]'' |bar1=green', 73 => ' |label2=[[Selenosteidae]]', 74 => ' |2={{clade', 75 => ' |1=''[[Stenosteus angustopectus]]'' |bar1=green', 76 => ' |2={{clade', 77 => ' |1=''[[Gymnotrachelus hydei]]'' |bar1=green', 78 => ' |2={{clade', 79 => ' |1=''[[Rhinosteus parvulus]]'' |bar1=green', 80 => ' |2=''[[Pachyosteus bulla]]'' |bar2=green }} }} }} }} }} }} }} }}', 81 => ' |label2='''[[Dunkleosteoidea]]'''', 82 => ' |2={{clade', 83 => ' |1=''[[Westralichthys uwagedensis]]'' |bar1=green', 84 => ' |2={{clade', 85 => ' |1={{clade', 86 => ' |1=''[[Protitanichthys rockportensis]]'' |bar1=green', 87 => ' |label2=[[Panxiosteidae]]', 88 => ' |2={{clade', 89 => ' |1=''[[Panxiosteus ocullus]]'' |bar1=green', 90 => ' |2=''[[Janiosteus timanicus]]'' |bar2=green', 91 => ' |3=''[[Plourdosteus canadensis]]'' |bar3=green }} }}', 92 => ' |label2='''[[Dunkleosteidae]]'''', 93 => ' |2={{clade', 94 => ' |1={{clade', 95 => ' |1=''[[Eastmanosteus calliaspis]]'' |bar1=green', 96 => ' |2=''[[Xiangshuiosteus wui]]'' |bar2=green }}', 97 => ' |2={{clade', 98 => ' |1=''[[Eastmanosteus pustulosus]]'' |bar1=green', 99 => ' |2=''[[Kiangyousteus yohii]]'' |bar2=green', 100 => ' |3=''[[Golshanichthys asiatica]]'' |bar3=green', 101 => ' |4={{clade', 102 => ' |1='''''Dunkleosteus amblyodoratus''''' |bar1=green', 103 => ' |2='''''Dunkleosteus terrelli''''' |bar2=green', 104 => ' |3='''''Dunkleosteus raveri''''' |barend3=green }} }} }} }} }} }} }}', 105 => '', 106 => 'Alternatively, the subsequent 2016 Zhu ''et al.'' study using a larger [[morphology (biology)|morphological]] dataset recovered [[Panxiosteidae]] well outside of [[Dunkleosteoidea]], leaving the status of [[Dunkleosteidae]] as a clade grouping separate from Dunkleosteoidea in doubt, as shown in the cladogram below:<ref name=Zhu2016>{{Cite journal |last1=Zhu |first1=You-An |last2=Zhu |first2=Min |last3=Wang |first3=Jun-Qing |date=April 1, 2016 |title=Redescription of Yinostius major (Arthrodira: Heterostiidae) from the Lower Devonian of China, and the interrelationships of Brachythoraci |journal=[[Zoological Journal of the Linnean Society]] |volume=176 |issue=4 |pages=806–834 |doi=10.1111/zoj.12356|issn=0024-4082 |doi-access=free }}</ref>', 107 => '{{clade| style=font-size:85%;line-height:85%', 108 => ' |label1='''[[Eubrachythoraci]]'''', 109 => ' |1={{clade', 110 => ' |label1='''[[Coccosteomorphi]]'''', 111 => ' |1={{clade', 112 => ' |label1=[[Coccosteoidea]]', 113 => ' |1={{clade', 114 => ' |label1=[[Coccosteidae]]', 115 => ' |1={{clade', 116 => ' |1={{clade', 117 => ' |1=''[[Millerosteus minor]]''', 118 => ' |2=''[[Coccosteus cuspidatus]]'' }}', 119 => ' |2={{clade', 120 => ' |1=''[[Dickosteus threiplandi]]''', 121 => ' |2=''[[Watsonosteus fletti]]''', 122 => ' |3=''[[Protitanichthys rockportensis]]'' }} }}', 123 => ' |label2='''[[Panxiosteidae]]'''', 124 => ' |2={{clade', 125 => ' |1=''[[Plourdosteus canadensis]]''', 126 => ' |2=''[[Panxiosteus ocullus]]''', 127 => ' |3=''[[Janiosteus timanicus]]'' }} }}', 128 => ' |label2=[[Incisoscutoidea]]', 129 => ' |2={{clade', 130 => ' |1=''[[Harrytoombsia elegans]]''', 131 => ' |2={{clade', 132 => ' |1=''[[Torosteus tuberculatus]]''', 133 => ' |2=''[[Torosteus pulchellus]]'' }}', 134 => ' |3={{clade', 135 => ' |1=''[[Mcnamaraspis kaprios]]''', 136 => ' |2={{clade', 137 => ' |1=''[[Compagopiscis croucheri]]''', 138 => ' |2=''[[Trematosteus fontanellus]]''', 139 => ' |label3=[[Camuropiscidae]]', 140 => ' |3={{clade', 141 => ' |1={{clade', 142 => ' |1=''[[Incisoscutum ritchiei]]''', 143 => ' |2=''[[Incisoscutum sarahae]]'' }}', 144 => ' |2={{clade', 145 => ' |1=''[[Rolfosteus canningensis]]''', 146 => ' |2={{clade', 147 => ' |1=''[[Tubonasus lennardensis]]''', 148 => ' |2={{clade', 149 => ' |1=''[[Fallacosteus turneri]]''', 150 => ' |2={{clade', 151 => ' |1=''[[Camuropiscis laidlawi]]''', 152 => ' |2=''[[Latocamurus coulthardi]]'' }} }} }} }} }} }} }} }} }}', 153 => ' |label2='''[[Pachyosteomorphi]]'''', 154 => ' |2={{clade', 155 => ' |1=''[[Rhachiosteus pterygiatus]]''', 156 => ' |2={{clade', 157 => ' |label1='''[[Dunkleosteoidea]]'''', 158 => ' |1={{clade', 159 => ' |1=''[[Eastmanosteus calliaspis]]''', 160 => ' |2={{clade', 161 => ' |1={{clade', 162 => ' |1=''[[Eastmanosteus pustulosus]]''', 163 => ' |2=''[[Kiangyousteus yohii]]'' }}', 164 => ' |2={{clade', 165 => ' |1=''[[Golshanichthys asiatica]]''', 166 => ' |2={{clade', 167 => ' |1=''[[Westralichthys uwagedensis]]''', 168 => ' |2={{clade', 169 => ' |1={{clade', 170 => ' |1='''''Dunkleosteus raveri'''''', 171 => ' |2='''''Dunkleosteus terrelli''''' }}', 172 => ' |2={{clade', 173 => ' |1='''''Dunkleosteus amblyodoratus'''''', 174 => ' |label2=[[Heterostiidae]]', 175 => ' |2={{clade', 176 => ' |1=''[[Heterosteus ingens]]''', 177 => ' |2=''[[Yinostius major]]'' }} }} }} }} }} }} }}', 178 => ' |label2='''[[Aspinothoracidi]]'''', 179 => ' |2={{clade', 180 => ' |1=''[[Tapinosteus heintzi]]''', 181 => ' |2={{clade', 182 => ' |1={{clade', 183 => ' |1=''[[Bullerichthys fascidens]]''', 184 => ' |2={{clade', 185 => ' |1=''[[Kendrickichthys cavernosus]]''', 186 => ' |2=''[[Bruntonichthys multidens]]'' }} }}', 187 => ' |2={{clade', 188 => ' |1=''[[Dinichthys herzeri]]''', 189 => ' |2={{clade', 190 => ' |1=''[[Hadrosteus rapax]]''', 191 => ' |2={{clade', 192 => ' |1=''[[Gorgonichthys clarki]]''', 193 => ' |label2=[[Selenosteidae]]', 194 => ' |2={{clade', 195 => ' |1=''[[Heintzichthys gouldii]]''', 196 => ' |2=''[[Pachyosteus bulla]]''', 197 => ' |3=''[[Gymnotrachelus hydei]]''', 198 => ' |4=''[[Stenosteus angustopectus]]''', 199 => ' |5=''[[Brachyosteus dietrichi]]''', 200 => ' |6={{clade', 201 => ' |1=''[[Melanosteus occitanus]]''', 202 => ' |2=''[[Rhinosteus parvulus]]'' }} }} }} }} }} }} }} }} }} }} }}', 203 => '', 204 => '=== Species ===', 205 => 'At least ten different species<ref name="Carr+2010"/><ref name=Denison>{{cite book|last=Denison|first=Robert|chapter=Placodermi|volume=2|title=Handbook of Paleoichthyology|year=1978|publisher=Gustav Fischer Verlag|location=Stuttgart New York|isbn=978-0-89574-027-4|pages=128}}</ref> of ''Dunkleosteus'' have been described so far.', 206 => '[[File:Dunkleosteus skull steveoc.jpg|thumb|Labelled skull diagram of ''D. terrelli'']]', 207 => 'The [[type species]], ''D. terrelli'', is the largest, best-known species of the genus, measuring {{convert|4.1|-|10|m|ft|abbr=on}} in length. It has a rounded snout. ''D. terrelli''{{'s}} fossil remains are found in Upper Frasnian to Upper Famennian Late Devonian strata of the United States (Huron and [[Cleveland Shale]] of Ohio, the Conneaut of [[Pennsylvania]], [[Chattanooga Shale]] of Tennessee, [[Lost Burro Formation]], California, and possibly Ives [[breccia]] of Texas<ref name=Denison/>) and [[Europe]].', 208 => '', 209 => '''D. belgicus'' (?) is known from fragments described from the [[Famennian]] of [[Belgium]]. The median dorsal plate is characteristic of the genus, but, a plate that was described as a suborbital is anterolateral.<ref name=Denison/> Lelièvre (1982) considers this taxon a [[nomen dubium]] and suggests the material may actually pertain to ''[[Ardennosteus]]''.<ref>{{Cite journal |last=Lelièvre |first=Hervé |date=1982 |title=Ardennosteus ubaghsi n.g., n. sp. Brachythoraci primitif (vertébré, placoderme) du Famennien d'Esneux (Belgique) |url=https://popups.uliege.be/0037-9395/index.php?id=3220 |journal=Annales de la Société géologique de Belgique |volume=105 |issue=1}}</ref>', 210 => '', 211 => '''D. denisoni'' is known from a small median dorsal plate, typical in appearance for ''Dunkleosteus'', but much smaller than normal. It is comparable in skull structure to ''D. marsaisi''.<ref name=Denison/>', 212 => '', 213 => '[[File:Dunkleosteus marsaisi 45.JPG|thumb|''D. marsaisi'' skull]]', 214 => '''D. marsaisi'' refers to the ''Dunkleosteus'' fossils from the Lower Famennian Late Devonian strata of the [[Atlas Mountains]] in [[Morocco]]. It differs in size, the known skulls averaging a length of {{convert|35|cm|ft}} and in form to ''D. terrelli''. In ''D. marsaisi'', the snout is narrower, and a postpineal fenestra may be present. Many researchers and authorities consider it a synonym of ''D. terrelli''.<ref>{{cite journal|last=Murray|first=A.M.|title=The Palaeozoic, Mesozoic and Early Cenozoic fishes of Africa|journal=Fish and Fisheries|year=2000|volume=1|issue=2|pages=111–145|doi=10.1046/j.1467-2979.2000.00015.x}}</ref> H. Schultze regards ''D. marsaisi'' as a member of ''[[Eastmanosteus]]''.<ref name=Denison/><ref name=Schultz>{{cite journal|last=Schultz|first=H|title=Large Upper Devonian arthrodires from Iran|journal=Fieldiana Geology|year=1973|volume=23|pages=53–78|doi=10.5962/bhl.title.5270|doi-access=free}}</ref>', 215 => '', 216 => '''D. magnificus'' is a large placoderm from the Frasnian [[Rhinestreet Shale]] of New York. It was originally described as ''[[Dinichthys]] magnificus'' by Hussakof and Bryant in 1919, then as "'' Dinichthys mirabilis''" by Heintz in 1932. Dunkle and Lane moved it to ''Dunkleosteus'' in 1971.<ref name=Denison/>', 217 => '', 218 => '''D. missouriensis'' is known from fragments from Frasnian [[Missouri]]. Dunkle and Lane regard them as being very similar to ''D. terrelli''.<ref name=Denison/>', 219 => '', 220 => '''D. newberryi'' is known primarily from a {{convert|28|cm|in}} long infragnathal with a prominent anterior cusp, found in the Frasnian portion of the [[Genesee Group]] of New York, and originally described as ''Dinichthys newberryi''.<ref name=Denison/>', 221 => '', 222 => '''D. amblyodoratus'' is known from some fragmentary remains from Late Devonian strata of [[Kettle Point Formation]], Ontario. The species name means 'blunt spear' and refers to the way the [[nuchal]] and [[paranuchal]] plates in the back of the head form the shape of a blunted spearhead.<ref name="Carr+2010"/>', 223 => '', 224 => '''D. raveri'' is a small species, possibly 1 meter long, known from an uncrushed skull roof found in a carbonate concretion from near the bottom of the Huron Shale, of the Famennian [[Ohio Shale]] strata. Besides its small size, it had comparatively large eyes. Because ''D. raveri'' was found in the strata directly below the strata where the remains of ''D. terrelli'' are found, ''D. raveri'' may have given rise to ''D. terrelli''. The species name commemorates Clarence Raver of [[Wakeman, Ohio]], who discovered the concretion where the [[holotype]] was found.<ref name="Carr+2010"/>', 225 => '', 226 => '''D. tuderensis'' is known from an infragnathal found in the lower-middle [[Famennian]]-aged [[Bilovo Formation]] of the [[Tver Region]] in northwest Russia. The specific name refers to the Maliy Tuder River as the holotyoe was found on its bank.<ref name=":3">{{Cite journal |last=Lebedev |first=Oleg A. |last2=Engelman |first2=Russell K. |last3=Skutschas |first3=Pavel P. |last4=Johanson |first4=Zerina |last5=Smith |first5=Moya M. |last6=Kolchanov |first6=Veniamin V. |last7=Trinajstic |first7=Kate |last8=Linkevich |first8=Valeriy V. |date=May 2023 |title=Structure, Growth and Histology of Gnathal Elements in Dunkleosteus (Arthrodira, Placodermi), with a Description of a New Species from the Famennian (Upper Devonian) of the Tver Region (North-Western Russia) |url=https://www.mdpi.com/1424-2818/15/5/648 |journal=Diversity |language=en |volume=15 |issue=5 |pages=648 |doi=10.3390/d15050648 |issn=1424-2818}}</ref>', 227 => '', 228 => '==Description==', 229 => '===Size and anatomy===', 230 => '[[File:Dunkleosteus terrelli size.png|left|thumb|Comparison of multiple size estimates for ''D. terrelli'']]', 231 => 'Mainly the armored frontal sections of specimens have been [[fossilized]], and consequently, the appearance of the other portions of the fish is mostly unknown.<ref>{{cite AV media |url=https://www.youtube.com/watch?v=lEEhm4rzxEg |title=Prehistoric Monsters Revealed |date=2008 |last=Dash |first=Sean |publisher=Workaholic Productions / History Channel |location=United States |access-date=December 18, 2015}}{{cbignore}}{{Dead YouTube link|date=February 2022}}</ref> In fact, only about 5% of ''Dunkleosteus'' specimens have more than a quarter of their skeleton preserved.<ref>Carr, R, & G.L. Jackson. 2008. [https://www.researchgate.net/publication/213769842_THE_VERTEBRATE_FAUNA_OF_THE_CLEVELAND_MEMBER_FAMENNIAN_OF_THE_OHIO_SHALE The Vertebrates fauna of the Cleveland member (Famennian) of the Ohio Shale. Society of Vertebrates Paleontology]. 1–17.</ref> Because of this, many reconstructions of the hindquarters are often based on fossils of smaller [[arthrodire]]s, such as ''[[Coccosteus]]'', which have preserved hind sections,<ref name=":0" /> leading to widely varying size estimates.<ref name=":0" />', 232 => '', 233 => '''Dunkleosteus'' is one of the largest known placoderms, with its maximum size being variably estimated as anywhere from {{convert|4.1|–|10|m|ft}} by different researchers.<ref name="Anderson&Westneat2007" /><ref name="Anderson&Westneat2009" /><ref name="Ferron2017" /><ref>{{cite journal|last=Williams|first=Nigel|year=2007|title=Force feeding|journal=Current Biology |volume=17 |pages=R3 |doi=10.1016/j.cub.2006.11.057|s2cid=36585467 }}</ref><ref name=":0">{{Cite journal |last=Engelman |first=Russell K. |date=2023|title=A Devonian Fish Tale: A New Method of Body Length Estimation Suggests Much Smaller Sizes for Dunkleosteus terrelli (Placodermi: Arthrodira) |journal=Diversity |language=en |volume=15 |issue=3 |pages=318 |doi=10.3390/d15030318 |issn=1424-2818 |doi-access=free }}</ref> However, most cited length estimates are speculative and lack quantitative or statistical backing.<ref name="Ferron2017" /><ref name=":0" /> Most studies that estimate the length of ''Dunkleosteus'' do not provide information as to how these estimates were calculated, the measurements used to scale them, or which specimens were examined. Most are implied to be based on either CMNH 5768 (the largest complete armor of ''Dunkleosteus'') or CMNH 5936 (the largest known jaw fragment).', 234 => '{{multiple image', 235 => '|align=right', 236 => '|perrow=1', 237 => '|total_width=240', 238 => '', 239 => '|image1=Dunkleosteus terrelli 2017 reconstruction.png', 240 => '|caption1=[[Paleoart|Life reconstruction]] of ''D. terrelli'', as presented by Ferrón ''et al.'' 2017<ref name="Ferron2017"/>', 241 => '', 242 => '|image2=Dunkleosteus terrelli 2023 reconstruction.png', 243 => '|caption2=Life reconstruction of ''D. terrelli'', as presented by Engelman 2023<ref name=":0"/>', 244 => '}}', 245 => '', 246 => '===Revaluation of physiology===', 247 => 'Most of the studies with well-defined methods produce lengths of {{convert|5|m|ft}} or less for ''Dunkleosteus terrelli'',<ref name=":0" /> with the exception of Ferrón ''et al.'' (2017), which produces larger estimates of {{convert|6.88|–|8.79|m|ft}} based on upper jaw perimeter of modern sharks.<ref name="Ferron2017" /> However, arthrodires have proportionally larger mouths than modern sharks, making the lengths estimated by Ferrón ''et al.'' (2017) unreliable.<ref name=":2">{{cite journal |last=Engelman |first=Russell |date=2023-04-10 |title=Giant, swimming mouths: oral dimensions of extant sharks do not accurately predict body size in ''Dunkleosteus terrelli'' (Placodermi: Arthrodira) |url=https://peerj.com/articles/15131 |journal=PeerJ |language=en |volume=11 |pages=e15131 |doi=10.7717/peerj.15131 |pmid=37065696 |pmc=10100833 }}</ref> Upper jaw perimeter overestimates the size of complete arthrodires like ''[[Coccosteus]]'' and the estimates of Ferrón ''et al.'' (2017) result in ''Dunkleosteus'' having an extremely small head and hyper-elongate trunk relative to the known dimensions of the fossils.<ref name=":2" /> If the reconstruction presented in Ferrón ''et al.'' (2017) is scaled to the known dimensions of CMNH 5768, it produces a length of {{convert|3.77|m|ft}}.<ref name=":2" />', 248 => '', 249 => 'Carr (2010) estimated a {{convert|4.6|m|ft}} long adult individual of ''Dunkleosteus terrelli'' to have weighed {{convert|665|kg|lb}}, assuming a shark-like body plan and a similar length-weight relationship.<ref name="Carr 20102">{{cite journal |last1=Carr |first1=Robert K. |date=2010 |title=Paleoecology of Dunkleosteus terrelli (Placodermi: Arthrodira). |url=https://www.researchgate.net/publication/235924093 |journal=Kirtlandia |volume=57}}</ref> Engelman (2023), using an [[ellipsoid]] [[Volume|volumetric]] method, estimated weights of {{convert|950|–|1200|kg|lb}} for typical ({{convert|3.41|m|ft}} long) adult ''Dunkleosteus'', and weights of {{convert|1490|–|1760|kg|lb}} for the largest ({{convert|4.1|m|ft|1}} in this study) individual.<ref name=":0" /> The higher weights by Engelman (2023) are mostly a result of the fact that arthrodires tend to have relatively deeper and wider bodies compared to sharks.<ref name=":0" />', 250 => '', 251 => 'An exceptionally preserved specimen of ''D. terrelli'' preserves a [[Fish fin|pectoral fin]] outline with [[ceratotrichia]], implying that the fin morphology of placoderms was much more variable than previously thought, and was heavily influenced by locomotory requirements. This knowledge, coupled with the knowledge that fish morphology is more heavily influenced by feeding niche than phylogeny, allowed a 2017 study to infer the [[Fish fin|caudal fin]] shape of ''D. terrelli'', reconstructing this fin with a strong ventral lobe, a high [[aspect ratio]], narrow [[caudal peduncle]], in contrast to previous reconstructions based on the [[Fish locomotion|anguilliform]] caudal fin of coccosteomorph placoderms.<ref name="Ferron2017">{{Cite journal|last1=Ferrón|first1=Humberto G.|last2=Martínez-Pérez|first2=Carlos|last3=Botella|first3=Héctor|date=2017|title=Ecomorphological inferences in early vertebrates: reconstructing Dunkleosteus terrelli (Arthrodira, Placodermi) caudal fin from palaeoecological data|journal=PeerJ|language=en|volume=5|pages=e4081|doi=10.7717/peerj.4081|pmid=29230354|pmc=5723140|issn=2167-8359}}</ref>', 252 => '', 253 => '====Length estimations of ''D. terrelli''====', 254 => '{| class="wikitable sortable" style="margin-left: auto; margin-right: auto; border: none;"', 255 => '|+ Length estimates of ''Dunkleosteus terrelli'' (modified from Table 7 of Engelman 2023<ref name=":0" />)', 256 => '! Study (author)', 257 => '! Year', 258 => '! Length', 259 => '! Method', 260 => '! Reference', 261 => '|-', 262 => '| Newberry || 1875 || {{convert|4.5|–|5.5|m|ft}} || '''Extrapolated from ''Coccosteus cuspidatus'', measurements and specimen used unclear''' || <ref name=":1">{{cite book |last1=Newberry |first1=John S. |url=https://www.biodiversitylibrary.org/item/116994#page/15/mode/1up |title=Report of the Geological Survey of Ohio. Volume II. Geology and Paleontology |date=1875 |publisher=Nevins and Myers, State Printers |volume=2 |location=Columbus |page=24 |contribution=Descriptions of fossil fishes}}</ref>', 263 => '|-', 264 => '| Newberry || 1889 || {{convert|4.5|m|ft}} || Unstated (implied extrapolation from ''Coccosteus'') || <ref>{{cite journal |last1=Newberry |first1=John S. |date=1889 |title=Paleozoic fishes of North America |url=https://www.biodiversitylibrary.org/item/51025#page/5/mode/1up |journal=Monographs of the U.S. Geological Survey |volume=16 |page=24}}</ref>', 265 => '|-', 266 => '| Dean || 1895 || {{convert|3|m|ft}} || Methods, measurements, and specimens unstated || <ref>{{cite book |last1=Dean |first1=Bashford |title=Fishes, Living and Fossil: An Outline of Their Forms and Probable Relationships |date=1895 |publisher=Macmillan and Company |location=London |page=130}}</ref>', 267 => '|-', 268 => '| Hussakof || 1905 || {{convert|1.67|m|ft}} (AMNH FF 195) <br> {{convert|3.79|m|ft}} (extrapolated to CMNH 5768 by Engelman 2023<ref name=":0"/> assuming similar head-trunk proportions) || '''Entering angle of body''' || <ref>{{cite journal |last1=Hussakof |first1=Louis |title=Notes on the Devonian "placoderm" ''Dinichthys intermedius'' Newb. |journal=Bulletin of the American Museum of Natural History |date=1905 |volume=21 |pages=27–36}}</ref>', 269 => '|-', 270 => '| Anonymous|| 1923 || {{convert|7.6|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Anonymous |title=Cleveland shale fishes |journal=Bulletin of the Cleveland Museum of Natural History |date=1923 |volume=9 |pages=36}}</ref>', 271 => '|-', 272 => '| Hyde || 1926 || {{convert|4.5|–|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Hyde |first1=Jesse E. |title=Collecting fossil fishes from the Cleveland Shale |journal=Natural History |date=1926 |volume=26 |pages=497–504}}</ref>', 273 => '|-', 274 => '| Romer || 1966 || {{convert|9|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Romer |first1=Alfred S. |title=Vertebrate Paleontology |date=1966 |publisher=University of Chicago Press |location=Chicago |page=49 |edition=3rd}}</ref>', 275 => '|-', 276 => '| Colbert || 1969 || {{convert|9|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Colbert |first1=Edwin H. |title=Evolution of the Vertebrates: A History of Backboned Animals Through Time |date=1969 |publisher=John Wiley and Sons |location=Hoboken |page=36 |edition=2nd}}</ref>', 277 => '|-', 278 => '| Denison || 1978 || {{convert|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref name=Denison />', 279 => '|-', 280 => '| Williams || 1992 || {{convert|5|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Williams |first1=Michael E. |title=Jaws: The Early Years |journal=Explorer |date=1992 |volume=34 |pages=4–8}}</ref>', 281 => '|-', 282 => '| Janvier || 2003 || {{convert|6|–|7|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Janvier |first1=P. |title=Early Vertebrates |date=2003 |publisher=Clarendon Press |location=Oxford |page=12}}</ref>', 283 => '|-', 284 => '| Young || 2003 || {{convert|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Young |first1=Gavin C. |title=Did placoderm fish have teeth? |journal=Journal of Vertebrate Paleontology |date=24 December 2003 |volume=23 |issue=4 |pages=987–990 |doi=10.1671/31|bibcode=2003JVPal..23..987Y |s2cid=85572061 }}</ref>', 285 => '|-', 286 => '| Anderson and Westneat || 2007 || {{convert|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref name="Anderson&Westneat2007" />', 287 => '|-', 288 => '| Anderson and Westneat || 2009 || {{convert|10|m|ft}} || Methods, measurements, and specimens used not stated || <ref name="Anderson&Westneat2009" />', 289 => '|-', 290 => '| Carr || 2010 || {{convert|4.5|–|6|m|ft}} || Methods, measurements, and specimens used not stated || <ref name="Carr 2010">{{cite journal |last1=Carr |first1=Robert K. |date=2010 |title=Paleoecology of Dunkleosteus terrelli (Placodermi: Arthrodira). |url=https://www.researchgate.net/publication/235924093 |journal=Kirtlandia |volume=57}}</ref>', 291 => '|-', 292 => '| Long || 2010 || {{convert|4.5|–|8|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Long |first1=John A. |title=The Rise of Fishes: 500 Million Years of Evolution |date=2010 |publisher=Johns Hopkins University Press |location=Baltimore |pages=88–90 |edition=2nd}}</ref>', 293 => '|-', 294 => '| Sallan and Galimberti || 2015 || {{convert|8|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite journal |last1=Sallan |first1=Lauren |last2=Galimberti |first2=Andrew K. |title=Body-size reduction in vertebrates following the end-Devonian mass extinction |journal=Science |date=13 November 2015 |volume=350 |issue=6262 |pages=812–815 |doi=10.1126/science.aac7373|pmid=26564854 |bibcode=2015Sci...350..812S |s2cid=206640186 }}</ref>', 295 => '|-', 296 => '| Ferrón ''et al.'' || 2017 || {{convert|6.88|m|ft}} (average adult, CMNH 5768) <br> {{convert|8.79|m|ft}} (largest individual, CMNH 5936) || '''Upper jaw perimeter''' || <ref name="Ferron2017" />', 297 => '|-', 298 => '| Long ''et al.'' || 2019 || {{convert|6|–|8|m|ft}} || Methods, measurements, and specimens used not stated || <ref>{{cite book |last1=Long |first1=John A. |last2=Choo |first2=Brian |last3=Clement |first3=Alice |contribution=The Evolution of Fishes through Geological Time |title=Evolution and Development of Fishes |editor1-last=Johanson |editor1-first=Zerina|editor2-last=Underwood |editor2-first=Charlie|editor3-last=Richter|editor3-first=Martha| date=31 December 2018 |pages=3–29 |doi=10.1017/9781316832172.002|s2cid=134217082 }}</ref>', 299 => '|-', 300 => '| Johanson ''et al.'' || 2019 || {{convert|3|m|ft}} (CMNH 50322) <br> {{convert|7.1|m|ft}} (extrapolated to CMNH 5768 by Engelman 2023 assuming similar head-trunk proportions) || Methods and measurements not stated || <ref>{{cite journal |last1=Johanson |first1=Zerina |last2=Trinajstic |first2=Kate |last3=Cumbaa |first3=Stephen |last4=Ryan |first4=Michael |title=Fusion in the vertebral column of the pachyosteomorph arthrodire Dunkleosteus terrelli ('Placodermi') |journal=Palaeontologia Electronica |date=2019 |volume=22.2.20A |pages=1–13 |doi=10.26879/872|s2cid=162173408 }}</ref>', 301 => '|-', 302 => '| Engelman || 2023 || {{convert|3.41|m|ft}} (average adult, CMNH 5768) <br> {{convert|4.1|m|ft|1}} (largest individual, CMNH 5936) || '''Orbit-opercular length (head length minus snout)''' || <ref name=":0" />', 303 => '|-', 304 => '| Engelman || 2023 || {{convert|3.41|m|ft}} (average adult, CMNH 5768) || '''Skull length in ''Coccosteus''''' || <ref name=":0" />', 305 => '|-', 306 => '| Engelman || 2023 || {{convert|5.23|m|ft}} (average adult, CMNH 5768) || '''Infragnathal length in ''Coccosteus''''' (source considers this estimate unreliable due to ''Dunkleosteus'' having a relatively larger mouth than ''Coccosteus'') || <ref name=":0" />', 307 => '|-', 308 => '| Engelman || 2023 || {{convert|3.47|m|ft}} (average adult, CMNH 5768) || '''Entering angle of body''' || <ref name=":0" />', 309 => '|-', 310 => '| Engelman || 2023 || {{convert|3.88|m|ft}} (average adult, CMNH 5768) || '''Length of posteroventrolateral plate''' || <ref name=":0" />', 311 => '|-', 312 => '| Engelman || 2023 || {{convert|3.40|m|ft}} (average adult, CMNH 5768) || '''Inferred location of pelvic girdle''' || <ref name=":0" />', 313 => '|-', 314 => '|}', 315 => '', 316 => '==Paleobiology==', 317 => '===Diet===', 318 => '[[File:Dunkleosteus terrelli CMNH 5936.png|thumb|left|Lower jaw of ''D. terrelli'' specimen CMNH 5936]] ', 319 => '''Dunkleosteus terrelli'' possessed a [[four-bar linkage]] mechanism for [[Fish jaw|jaw]] opening that incorporated connections between the skull, the thoracic shield, the lower jaw and the jaw muscles joined by movable joints.<ref name=Anderson&Westneat2009>{{cite journal |last=Anderson |first=P.S.L. |author2=Westneat, M. |title=A biomechanical model of feeding kinematics for ''Dunkleosteus terrelli'' (Arthrodira, Placodermi) |journal=Paleobiology |date=2009 |volume=35 |issue=2 |pages=251–269 |doi=10.1666/08011.1 |bibcode=2009Pbio...35..251A |s2cid=86203770 |url=http://projects.fieldmuseum.org/sites/default/files/DunkPaleoBio.pdf |access-date=October 29, 2017 |archive-date=November 7, 2017 |archive-url=https://web.archive.org/web/20171107014026/http://projects.fieldmuseum.org/sites/default/files/DunkPaleoBio.pdf |url-status=dead }}</ref><ref name=Anderson&Westneat2007>{{Cite journal |last=Anderson |first=P.S.L. |author2=Westneat, M. |title=Feeding mechanics and bite force modelling of the skull of Dunkleosteus terrelli, an ancient apex predator |journal=Biology Letters |volume=3 |issue=1 |pages=76–79 |date=2007 |doi=10.1098/rsbl.2006.0569 |pmid=17443970 |pmc=2373817}}</ref> This mechanism allowed ''D. terrelli'' to both achieve a high speed of jaw opening, opening their jaws in 20 milliseconds and completing the whole process in 50–60 milliseconds (comparable to modern fishes that use [[Aquatic feeding mechanisms#Suction feeding|suction feeding]] to assist in prey capture;<ref name=Anderson&Westneat2007/>) and producing high bite forces when closing the jaw, estimated at {{convert|4414|N|kg-f lb-f|0|abbr=on}} at the tip and {{convert|5363|N|kg-f lb-f|0|abbr=on}} at the blade edge,<ref name=Anderson&Westneat2007/> or even up to {{cvt|6170|N|kg-f lb-f|0}} and {{cvt|7495|N|kg-f lb-f|0}} respectively.<ref name=Anderson&Westneat2009/> The pressures generated in those regions were high enough to puncture or cut through [[cuticle]] or [[dermal bone|dermal]] armor,<ref name=Anderson&Westneat2007/> suggesting that ''D. terrelli'' was adapted to prey on free-swimming, armored prey such as [[ammonite]]s and other placoderms.<ref name=Anderson&Westneat2009/>', 320 => '', 321 => 'In addition, teeth of a [[Chondrichthyes|chondrichthyan]] thought to belong to ''[[Orodus]]'' (''Orodus'' spp.) were found in association with ''Dunkleosteus'' remains, suggesting that these were probably stomach contents regurgitated from the animal. ''Orodus'' is thought to be tachypelagic, or a fast-swimming [[pelagic fish]]. Thus, ''Dunkleosteus'' might have been fast enough to catch these fast organisms, and not a slow swimmer like originally thought.<ref name="Ferron2017" /> Fossils of ''Dunkleosteus'' are frequently found with [[Bolus (digestion)|bolus]]es of fish bones, semidigested and partially eaten remains of other fish.<ref name="Virtual Fossil">{{cite web|url= http://www.fossilmuseum.net/Fossil_Galleries/Fish_Devonian/Dunkleosteous/Dunkleosteus.htm |title=Dunkleosteus Placodermi Devonian Armored Fish from Morocco |work=Fossil Archives |publisher=The Virtual Fossil Museum |access-date=April 26, 2009}}</ref> As a result, the [[fossil]] record indicates it may have routinely regurgitated prey bones rather than digest them. Mature individuals probably inhabited deep sea locations, like other placoderms, living in shallow waters during adolescence.<ref>{{Cite web|url=http://bioweb.uwlax.edu/bio203/f2013/peters_sadi/habitat.htm|title=UWL Website}}</ref>', 322 => '', 323 => 'A specimen of ''Dunkleosteus'' (CMNH 5302), and ''[[Titanichthys]]'' (CMNH 9889), show damage said to be puncture damage from the bony fangs of other ''Dunkleosteus''.<ref name="Anderson&Westneat2009" />', 324 => '', 325 => '===Reproduction===', 326 => '''Dunkleosteus'', together with most other placoderms, may have also been among the first [[vertebrate]]s to [[internal fertilization|internalize egg fertilization]], as seen in some modern sharks.<ref name="Ahlberg">{{cite journal |last1=Ahlberg |first1=Per |first2=Kate |last2=Trinajstic |first3=Zerina |last3=Johanson |first4=John |last4=Long |s2cid=205217467 |year=2009 |title=Pelvic claspers confirm chondrichthyan-like internal fertilization in arthrodires |journal=Nature |volume=460 |issue=7257 |pages=888–889 |doi=10.1038/nature08176|pmid=19597477 |bibcode=2009Natur.460..888A }}</ref> Some other placoderms have been found with evidence that they may have been [[viviparity|viviparous]], including what appears to have been an [[umbilical cord]].<ref>{{ cite journal | last = Long | first = J. A. |author2=Trinajstic, K. |author3=Young, G. C. |author4=Senden, T. | year = 2008 | title = Live birth in the Devonian period | journal = Nature | volume = 453 | issue = 7195 | pages = 650&ndash;652 | doi = 10.1038/nature06966 | pmid = 18509443 | bibcode = 2008Natur.453..650L | s2cid = 205213348 }}</ref>', 327 => '', 328 => '===Growth===', 329 => '{{multiple image', 330 => '|align=right', 331 => '|perrow=2', 332 => '|total_width=380', 333 => '', 334 => '|image1=Dunkleosteus juvenile fossil - Cleveland Museum of Natural History (34421618320).jpg', 335 => '|caption1=''D. terrelli'' juvenile specimen CMNH 7424', 336 => '', 337 => '|image2=Dunkleosteus terrelli - Cleveland Museum of Natural History - 2014-12-26 (21137540331).jpg', 338 => '|caption2=''D. terrelli'' adult specimen CMNH 5768', 339 => '', 340 => '}}', 341 => '[[morphology (biology)|Morphological]] studies on the lower jaws of juveniles of ''D. terrelli'' reveal they were proportionally as robust as those of adults, indicating they already could produce high bite forces and likely were able to shear into resistant prey tissue similar to adults, albeit on a smaller scale. This pattern is in direct contrast to the condition common in [[tetrapod]]s in which the jaws of juveniles are more [[gracile]] than in adults.<ref name="Snively_etal_2009">{{cite journal |last=Snively |first=E. |author2=Anderson, P.S.L. |author3=Ryan, M.J. |date=2009 |title=Functional and ontogenetic implications of bite stress in arthrodire placoderms |url=https://www.biodiversitylibrary.org/page/51811342#page/60/mode/1up |journal=Kirtlandia |volume=57}}</ref>', 342 => '', 343 => '==See also==', 344 => '* [[List of placoderms]]', 345 => '', 346 => '==References==', 347 => '{{notelist}}', 348 => '{{reflist}}', 349 => '', 350 => '==Further reading==', 351 => '* {{cite journal|first=Philip S. L.|last=Anderson|s2cid=86583150|year=2008|title=Shape Variation Between Arthrodire Morphotypes Indicates Possible Feeding Niches|journal=Journal of Vertebrate Paleontology|volume=28|number=4|pages=961–969|doi=10.1671/0272-4634-28.4.961|bibcode=2008JVPal..28..961A }}', 352 => '', 353 => '==External links==', 354 => '{{Commons category|Dunkleosteus}}', 355 => '{{Wikispecies|Dunkleosteus}}', 356 => '* [http://www.ucmp.berkeley.edu/vertebrates/basalfish/placodermi.html ''Introduction to the Placodermi: Extinct Armored Fishes with Jaws''. Waggoner, Ben (2000). Retrieved Aug 1, 2005]', 357 => '* [http://www.nbcnews.com/id/15940709 MSNBC: Prehistoric fish packed a mean bite]', 358 => '* [http://news.bbc.co.uk/1/hi/sci/tech/6195188.stm BBC: Ancient 'Jaws' had monster bite]', 359 => '', 360 => '{{Arthrodira|E.}}', 361 => '{{Taxonbar|from=Q131039|from2=Q911597|from3=Q911599|from4=Q911605|from5=Q911606|from6=Q911611|from7=Q911612|from8=Q911616|from9=Q911618|from10=Q911593|from11=Q25357547}}', 362 => '', 363 => '[[Category:Dunkleosteidae]]', 364 => '[[Category:Apex predators]]', 365 => '[[Category:Placoderms of Africa]]', 366 => '[[Category:Fossils of Morocco]]', 367 => '[[Category:Placoderms of Europe]]', 368 => '[[Category:Fossils of Belgium]]', 369 => '[[Category:Placoderms of North America]]', 370 => '[[Category:Fossils of Canada]]', 371 => '[[Category:Paleontology in Missouri]]', 372 => '[[Category:Paleontology in Pennsylvania]]', 373 => '[[Category:Paleontology in Tennessee]]', 374 => '[[Category:Late Devonian first appearances]]', 375 => '[[Category:Late Devonian animals]]', 376 => '[[Category:Famennian extinctions]]', 377 => '[[Category:Fossil taxa described in 1956]]', 378 => '[[Category:Taxa named by John Strong Newberry]]' ]
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