Duncostes

## Taxonomy[edit]

Restoration and size comparison of *D. terrelli*
*Dunkleosteus* was named in 1956 to honour David Dunkle, who died aged 72, then curator of vertebrate paleontology at the Cleveland Museum of Natural History. The type species *D. terrelli* was originally described in 1873 as a species of *Dinichthys*. *Dunkleosteus* is an arthrodire originally placed in the family Dinichthyidae, which is composed mostly of large, carnivorous fish like *Gorgonichthys*. Anderson (2009) suggests, because of its primitive jaw structure, *Dunkleosteus* should be placed outside the family Dinichthyidae, perhaps close to the base of the clade Pachyosteomorpha, near *Eastmanosteus*. Carr and Hlavin (2010) resurrect Dunkleosteidae and place *Dunkleosteus*, *Eastmanosteus*, and a few other genera from Dinichthyidae within it. Dinichthyidae, in turn, is made into a monospecific family.

### Species[edit]

*D. marsaisi* skull
At least ten different species of *Dunkleosteus* have been described so far.

The type species, *D. terrelli*, is the largest, best-known species of the genus, measuring 8.79 m (28.8 ft) 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) 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.

*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*.

*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 35 centimetres (1.15 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*. H. Schultze regards *D. marsaisi* as a member of *Eastmanosteus*.

*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.

*D. missouriensis* is known from fragments from Frasnian Missouri. Dunkle and Lane regard them as being very similar to *D. terrelli*.

*D. Newberry* is known primarily from a 28 centimetres (11 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 Newberry*".

*D. amblyodoratus* is known from some fragmentary remains from Late Devonian strata of Kettle Point, Canada. 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. Although it is known only from fragments, it is estimated to have been about 6 metres (19.7 ft) long in life.

*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.

## Description[edit]

Life restoration of *D. marsaisi*
The largest species, *D. terrelli*, is estimated to have grown up to 8.79 m (28.8 ft) in length and 4 t (4.4 short tons) in weight, making it one of the largest placoderms to have existed. Like other placoderms, *Dunkleosteus* had a two-part bony, armoured exterior, which may have made it a relatively slow but powerful swimmer. Instead of teeth, *Dunkleosteus* possessed two pairs of sharp bony plates which formed a beak-like structure. *Dunkleosteus*, together with most other placoderms, may have also been among the first vertebrates to internalize egg fertilization, as seen in some modern sharks.
A skull diagram of *Dunkleosteus*
Mainly the armored frontal sections of specimens have been fossilized, and consequently, the appearance of the other portions of the fish is mostly unknown. In fact, only about 5% of *Dunkleosteus* specimens have more than a quarter of their skeleton preserved. Because of this, many reconstructions of the hindquarters are often based on fossils of smaller arthrodires, such as *Coccosteus*, which have preserved hind sections. However, an exceptionally preserved specimen of *D. terrelli* preserves ceratotrichia in a pectoral fin, 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 body shape of *D. terrelli*. This new reconstruction gives D. terrelli a much more shark-like profile, including a strong anterior lobe on its tail, in contrast to reconstructions based on other placoderms.

The most famous specimens of *Dunkleosteus* are displayed at the Cleveland Museum of Natural History, and others are displayed at the American Museum of Natural History, National Museum of Natural History, State Museum of Pennsylvania, Harrisburg and in the Queensland Museum in Brisbane, Queensland.[*citation needed*]

### Diet[edit]

Life restoration of *D. terrelli*
*Dunkleosteus terrelli* possessed a four-bar li