Titanichthys Temporal range: | |
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Rendered reconstruction of Titanichthys clarkii | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | † Placodermi |
Order: | † Arthrodira |
Suborder: | † Brachythoraci |
Superfamily: | † Dinichthyloidea |
Family: | † Titanichthyidae Dean, 1901 |
Genus: | † Titanichthys Newberry, 1885 |
Type species | |
†Titanichthys agassizi Newberry, 1885 | |
Species | |
| |
Synonyms | |
Brontichthys |
Titanichthys is an extinct genus of giant, aberrant marine placoderm from shallow seas of the Late Devonian of Morocco, Eastern North America, and possibly Europe. [1] Many of the species approached Dunkleosteus in size and build. Unlike its relative, however, the various species of Titanichthys had small, ineffective-looking mouth-plates that lacked a sharp cutting edge. It is assumed that Titanichthys was a filter feeder that used its capacious mouth to swallow or inhale schools of small, anchovy-like fish, or possibly krill-like zooplankton, and that the mouth-plates retained the prey while allowing the water to escape as it closed its mouth. A study has since confirmed this assumption as its jaws are functionally closer to that of filter feeders like baleen whales and basking sharks, and it appears to have developed from benthic durophagists that became pelagic suspension feeders. This would make it the first (known) large-sized vertebrate filter feeder. [2] Titanichthys was estimated to have reached a length of 7–7.6 m (23–25 ft), [3] [4] [5] but Engelman (2023) suggested that Titanichthys was comparable in size to Dunkleosteus, likely measuring about or just over 4.1 metres (13.5 ft) in length. [6]
Titanichthys is thought to have been a basal aspinothoracid, closest related to Bungartius and Tafilalichthys . [7]
The genus shows a great diversity in the Famennian-aged Cleveland Shale, though species are also found in similarly aged strata in Morocco and possibly the Holy Cross Mountains in Poland. [1]
This is the type species, from the Cleveland Shale. Its infrognathals are strongly recurved medially, and is elongated with a spatula-like process at the anterior end. The headshield averages about 60 cm (24 in) in length. [1]
This Cleveland Shale species is based on an infragnathal bone more than 36 cm (14 in) in length. May possibly be a synonym of T. agassizi. [8]
This Cleveland Shale species has infragnathals that are not as recurved as T. agassizi's. The cranial roof is comparatively narrower and more rounded. It is the largest known species in the genus, and possibly one of the largest Devonian vertebrates known. The head is about 90 cm (35 in) in length. [8]
This Cleveland Shale species is known from a badly preserved, incomplete infragnathal. It was originally described by Claypole as "Brontichthys clarki" in 1894. [8] As "Brontichthys" is a junior synonym of Titanichthys, it should not be confused with another, similarly-named arthrodire, Bruntonichthys of Dunkleosteidae.
This Cleveland Shale species has an infragnathal as large as that of T. clarkii, though T. rectus' infragnathal is much straighter, and does not have a spatula-like process on its anterior end. [8]
This species placement within the genus is in doubt. It is based on incomplete nuchal and central plates found in Upper Famennian-aged marine strata of the Holy Cross Mountains in Poland. [8]
This species is found in Lower Famennian-aged marine strata of Tafilalet, Southern Morocco. The fossil material of this species strongly suggests it is as large as the Cleveland Shale' T. clarkii. [8] The average combined length of the head and trunk shields for T. termieri is estimated to be 200 cm (79 in) [8] When the first fossils of T. termieri were found by geologist Henri Termier, the specimens were originally placed within the genus Gorgonichthys - that is, after Termier was able to convince his colleagues that the bone scraps were of a placoderm, and not a dinosaur. [9]
Placoderms are vertebrate animals of the class Placodermi, an extinct group of prehistoric fish known from Paleozoic fossils during the Silurian and the Devonian periods. While their endoskeletons are mainly cartilaginous, their head and thorax were covered by articulated armoured plates, and the rest of the body was scaled or naked depending on the species.
Dunkleosteus is an extinct genus of large arthrodire ("jointed-neck") fish that existed during the Late Devonian period, about 382–358 million years ago. It was a pelagic fish inhabiting open waters, and one of the first apex predators of any ecosystem.
Hyneria is a genus of large prehistoric predatory lobe-finned fish which lived in fresh water during the Famennian stage of the Devonian period.
Groenlandaspis is an extinct genus of arthrodire from the Late Devonian. Fossils of the different species are found in late Devonian strata in all continents except eastern Asia. The generic name commemorates the fact that the first specimens of the type species were found in Greenland.
Coccosteus is an extinct genus of arthrodire placoderm from the Devonian period. Its fossils have been found throughout Europe and North America. The majority of these have been found in freshwater sediments, though such a large range suggests that they may have been able to enter saltwater. It was a small placoderm, with Coccosteus cuspidatus measuring 29.6–39.4 cm (11.7–15.5 in) long.
Dinichthys is an extinct monospecific genus of large marine arthrodire placoderm from the Late Devonian measuring around 3 metres (9.8 ft) long. Fossils were recovered from the Ohio Shale Formation along the Olentangy River in Delaware County, Ohio.
Dunkleosteidae is an extinct family of arthrodire placoderms that lived during the Devonian period. The gigantic apex predator Dunkleosteus terrelli is the best known member of this group.
Holonema is an extinct genus of relatively large, barrel-shaped arthrodire placoderms that were found in oceans throughout the world from the Mid to Late Devonian, when the last species perished in the Frasnian-Fammian extinction event. Most species of the genus are known from fragments of their armor, but the Gogo Reef species, H. westolli, is known from whole, articulated specimens.
Gorgonichthys is extinct monospecific genus of large arthrodire placoderm. Fossils are found in the Upper Famennian Cleveland Shales of Late Devonian in Ohio. The type species is Gorgonichthys clarki.
Holdenius is an extinct genus of arthrodire placoderm fish which lived during the Late Devonian period.
Plourdosteus is an extinct genus of placoderm arthrodire which was relatively widespread in Euramerica during the Givetian to Frasnian ages of the Devonian. It was a small placoderm, with P. canadensis specimen MNHM 2-177 measuring 37.5 cm (14.8 in) long.
Stenosteus is an extinct monospecific genus of medium-sized selenosteid arthrodire placoderms of the Late Devonian period known from the Upper Famennian Cleveland Shale of Ohio. Estimated skull lengths range from 6 to 9 centimeters Most fossils of Stenosteus have been scraps of armor and portions of tooth-plates suggestive of Selenosteus. In 1996, enough material of a new species, S. angustopectus, was recovered to allow a reconstruction of armor that resembles that of Selenosteus.
Gymnotrachelus is an extinct monospecific genus of large selenosteid arthrodire placoderm of the Late Devonian known from the Late Famennian Cleveland Shale of Ohio. The type species Gymnotrachelus hydei was originally reconstructed as physically resembling Selenosteus, with slightly smaller orbits. Later specimens led to a reappraisal, and now G. hydei is thought to have a more gar-like or barracuda-like build.
Diplognathus is a genus of arthrodire placoderm from the Late Famennian Cleveland Shale of Late Devonian Ohio, known only from incomplete fragments of jaws and skulls. What fragments are known suggest that the living animals were large-eyed piscivores with weak, but widely gaping jaws. D. mirabilis is thought to be fairly large, with infragnathals up to 45 centimeters in length. The second species, D. larfargei, was much smaller, with inferognathals averaging about 4 centimeters in length.
Homostiidae is a family of flattened arthrodire placoderms from the Early to Middle Devonian. Fossils appear in various strata in Europe, Russia, Morocco, Australia, Canada and Greenland.
Heterosteus is an extinct genus of heterosteid placoderm of the Middle Devonian known from remains discovered in Europe and Greenland. According to Denison, 1978, Heterosteus might have been planktivorous, along with Homosteus, and Titanichthys.
Africanaspis is an extinct genus of groenlandaspidid placoderm known from two species, Africanaspis doryssa, named in 1997 from fossils discovered in South Africa and Africanaspis edmountaini, named from fossils described from South Africa during 2017. A. edmountaini is only known from juvenile specimens. Both species are known from the Witpoort Formation. A. doryssa reached 23 cm (9.1 in) long.
Millerosteus is an extinct genus of coccosteid arthrodire placoderm from the Early Givetian stage of the Middle Devonian period. Fossils are found in Orkney and Caithness, Scotland. It was a small placoderm with an body length of 14 cm (5.5 in). Millerosteus is one of the few arthrodires known from specimens preserving the entire skeleton.
Watsonosteus is an extinct genus of coccosteid arthrodire placoderm from the Late Givetian stage of the Middle Devonian period. Fossils are found in the Orkney Islands, Scotland. It was a small placoderm with a total body length of 57 cm (22 in), with the largest individuals reaching lengths of 1 m (39 in). It is one of the few arthrodires for which complete body fossils are known.
Amazichthys is an extinct genus of selenosteid arthrodire from the Middle Famennian of the Late Devonian of the Anti-Atlas Mountains of Morocco. It contains a single species, Amazichthys trinajsticae. It is one of a few example of placoderm known from whole body shape, including cartilaginous axial and fin elements.
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