Marsaisi, D. Magnificus, D. Missouriensis, D. Newberryi, D. Amblyodoratus, and D. Raveri Some

Dunkleosteus was a genus of large predatory placoderm fish that lived during the Late Devonian period, around 358-382 million years ago. It contained 10 species and was named after David Dunkle. The largest species, D. terrelli, could grow up to 6 meters long and weigh over 1 ton, making it one of the largest placoderms. Dunkleosteus had a powerful bite enabled by a unique jaw mechanism, allowing it to prey on other armored fish and ammonites.

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Marsaisi, D. Magnificus, D. Missouriensis, D. Newberryi, D. Amblyodoratus, and D. Raveri Some

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Dunkleosteus is an extinct genus of arthrodire placoderm fish that existed during the Late

Devonian period, about 358–382 million years ago. The name Dunkleosteus combines the Greek
ὀστέον, osteon, meaning "bone", and Dunkle, in honor of David Dunkle of the Cleveland
Museum of Natural History. It consists of ten species: D. terrelli, D. belgicus, D. denisoni, D.
marsaisi, D. magnificus, D. missouriensis, D. newberryi, D. amblyodoratus, and D. raveri; some
of which are among the largest placoderms to have ever lived. The largest species, D. terrelli
grew up to 6 m (19.7 ft) long and 1 t (1.1 short tons) in weight. Few other placoderms rivaled
Dunkleosteus in size. Dunkleosteus could quickly open and close its jaw, like modern day
suction feeders, and had a bite force of 6,000 N (612 kgf; 1,349 lbf) at the tip and 7,400 N
(755 kgf; 1,664 lbf) at the blade edge. Numerous fossils of the various species have been found in
North America, Poland, Belgium, and Morocco.

Contents
 1 Taxonomy
o 1.1 Species
 2 Description
o 2.1 Diet
o 2.2 Juveniles
 3 See also
 4 References
 5 Further reading
 6 External links
Taxonomy[edit]

Restoration and size comparison of D. terreli

Dunkleosteus was named in 1956 to honour David Dunkle, 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.[1]
Dinichthyidae, in turn, is made into a monospecific family.[2]
Species[edit]
D. marsaisi skull
At least 10 different species[1][3] of Dunkleosteus have been described so far.
The type species, D. terrelli, is the largest, best-known species of the genus. 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[3]) 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
apparently an anteriolateral plate.[3]
D. denisoni is known from a small median dorsal plate, typical in appearance for Dunkleosteus,
but much smaller than normal.[3]
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.[4] H. Schultze regards D. marsaisi as a member of Eastmanosteus.[3][5]
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.[3]
D. missouriensis is known from fragments from Frasnian Missouri. Dunkle and Lane regard
them as being very similar to D. terrelli.[3]
D. newberryi 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 newberryi".[3]
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.[1]
D. raveri is a small, possibly 1-m-long species 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.[1]
Description[edit]

Life restoration of D. marsaisi

The largest species, D. terrelli, is estimated to have grown up to 6 m (19.7 ft) in length and 1 t
(1.1 short tons) in weight, making it one of the largest placoderms to have existed.[6][7][8] 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.[7] Dunkleosteus, together with most other
placoderms, may have also been among the first vertebrates to internalize egg fertilization, as
seen in some modern sharks.[9]

A skull diagram of Dunkleosteus

Mainly the armoured frontal sections of specimens have been fossilized, and consequently the
appearance of the other portions of the fish is mostly unknown.[10] Because of this, many
reconstructions of the hindquarters are often based on smaller arthrodires, such as Coccosteus,
which had 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.[11]
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 linkage mechanism for jaw opening that incorporated
connections between the skull, the thoracic shield, the lower jaw and the jaw muscles joined
together by movable joints.[7][6] 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 suction feeding to assist in prey capture;
[6]
) and producing high bite forces when closing the jaw, estimated at 6,000 N (612 kgf; 1,349 lbf)
at the tip and 7,400 N (755 kgf; 1,664 lbf) at the blade edge in the largest individuals.[7] The
pressures generated in those regions were high enough to puncture or cut through cuticle or
dermal armor[6] suggesting that D. terrelli was adapted to prey on free-swimming, armored prey
such as ammonites and other placoderms.[7] Fossils of Dunkleosteus are frequently found with
boluses of fish bones, semidigested and partially eaten remains of other fish.[12] As a result, the
fossil record indicates it may have routinely regurgitated prey bones rather than digest them. It
probably inhabited inshore waters.[citation needed]
Juveniles[edit]
Morphological studies on the lower jaws of juveniles of D. terrelli reveal they were
proportionally as robust as those of adults, indicating they already had the ability to 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 tetrapods in which the
jaws of juveniles are more gracile than in adults.[