Dunkleosteus is an extinct genus of large arthrodire ("jointed-neck") fish that existed during the Late Devonian period, about 382–358 million years ago.
[1] Dunkleosteus consists of ten species, some of which are among the largest placoderms ("plate-skinned") 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.
Since body shape is not known, various methods of estimation put the living total length of the largest known specimen of D. terrelli between 4.1 to 10 m (13 to 33 ft) long and weigh around 1–4 t (1.1–4.4 short tons).
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 herzeri.
Dunkleosteus was named by Jean-Pierre Lehman in 1956 to honour David Dunkle (1911–1984), former curator of vertebrate paleontology at the Cleveland Museum of Natural History.
[15] The cladogram below from the study of Zhu & Zhu (2013) shows the placement of Dunkleosteus within Dunkleosteidae and Dinichthys within the separate clade Aspinothoracidi:[16] Coccosteus cuspidatus Harrytoombsia elegans Mcnamaraspis kaprios Compagopiscis croucheri Incisoscutum ritchiei Incisoscutum sarahae Latocamurus coulthardi Camuropiscis laidlawi Rolfosteus canningensis Tubonasus lennardensis Fallacosteus turneri Dinichthys herzeri Hadrosteus rapax Gorgonichthys clarki Heintzichthys gouldii Stenosteus angustopectus Gymnotrachelus hydei Rhinosteus parvulus Pachyosteus bulla Westralichthys uwagedensis Protitanichthys rockportensis Panxiosteus ocullus Janiosteus timanicus Plourdosteus canadensis Eastmanosteus calliaspis Xiangshuiosteus wui Eastmanosteus pustulosus Kiangyousteus yohii Golshanichthys asiatica Dunkleosteus amblyodoratus Dunkleosteus terrelli Dunkleosteus raveri Alternatively, the subsequent 2016 Zhu et al. study using a larger 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:[17] Millerosteus minor Coccosteus cuspidatus Dickosteus threiplandi Watsonosteus fletti Protitanichthys rockportensis Plourdosteus canadensis Panxiosteus ocullus Janiosteus timanicus Harrytoombsia elegans Torosteus tuberculatus Torosteus pulchellus Mcnamaraspis kaprios Compagopiscis croucheri Trematosteus fontanellus Incisoscutum ritchiei Incisoscutum sarahae Rolfosteus canningensis Tubonasus lennardensis Fallacosteus turneri Camuropiscis laidlawi Latocamurus coulthardi Rhachiosteus pterygiatus Eastmanosteus calliaspis Eastmanosteus pustulosus Kiangyousteus yohii Golshanichthys asiatica Westralichthys uwagedensis Dunkleosteus raveri Dunkleosteus terrelli Dunkleosteus amblyodoratus Heterosteus ingens Yinostius major Tapinosteus heintzi Bullerichthys fascidens Kendrickichthys cavernosus Bruntonichthys multidens Dinichthys herzeri Hadrosteus rapax Gorgonichthys clarki Heintzichthys gouldii Pachyosteus bulla Gymnotrachelus hydei Stenosteus angustopectus Brachyosteus dietrichi Melanosteus occitanus Rhinosteus parvulus At least ten different species[14][18] of Dunkleosteus have been described so far.
[2] D. terrelli's fossil remains are found in Upper Frasnian to Upper Famennian Late Devonian strata of the United States (Huron, Chagrin, and Cleveland Shales of Ohio, the Conneaut and Chadakoin Formations of Pennsylvania, the Chattanooga Shale of Tennessee, the Lost Burro Formation of California, and possibly the Ives breccia of Texas[18]) and Europe.
[18] Lelièvre (1982) considers this taxon a nomen dubium ("doubtful name") and suggests the material may actually pertain to Ardennosteus.
[18] D. marsaisi refers to the Dunkleosteus fossils from the Lower Famennian Late Devonian strata of the Atlas Mountains in Morocco.
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.
[14] D. tuderensis is known from an infragnathal found in the lower-middle Famennian-aged Bilovo Formation of the Tver Region in northwest Russia.
[19] In total, of the ten or so species listed above only four are agreed upon as valid species of Dunkleosteus by all researchers: D. terrelli (which may or may not include Dunkleosteus material from Morocco), D. raveri, D. tuderensis, and possibly D. amblyodoratus (which is known from limited material that appears distinct but is difficult to compare with other dunkleosteids).
[19][25] Dunkleosteus was covered in dermal bone forming armor plates across its skull and front half of its trunk.
[29] The plates of Dunkleosteus had both a hard cortical and a marrow-filled cancellous layer, unlike most teleost fishes and more similar to tetrapod bones.
[32] Because of this, many reconstructions of the hindquarters are often based on fossils of smaller arthrodires, such as Coccosteus, which have preserved hind sections,[2] leading to widely varying size estimates.
[2] Dunkleosteus terrelli is one of the largest known placoderms, with its maximum size being variably estimated as anywhere from 4.1–10 metres (13–33 ft) by different researchers.
Additionally, these reconstructions often require Dunkleosteus to lack many features consistent across the body plans of other arthrodires like Coccosteus and Amazichthys.
[12] However, arthrodires have proportionally larger mouths than modern sharks, making the lengths estimated by Ferrón et al. (2017) unreliable.
[4] 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.
[4] Carr (2010) estimated a 4.6 metres (15 ft) long adult individual of Dunkleosteus terrelli to have weighed 665 kilograms (1,466 lb), assuming a shark-like body plan and a similar length-weight relationship.
[2] 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.
[37][3] This, along with a ridge on the inside of the trunk armor suggesting an unusually well-developed attachment for the horizontal septum, suggests Dunkleosteus may have had an anteriorly stiffened spine and specialized connective tissues to transmit force generated by the anterior trunk muscles to the tail fin, similar to thunniform vertebrates like lamnids and tunas.
[3] However, because these specimens were excavated from cliff faces, they were probably found in close to the armor, suggesting these fins were associated with the end of the ventral shield as in other arthrodires.
Mature individuals probably inhabited deep sea locations, like other placoderms, living in shallow waters during adolescence.