Holonema | |
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H. westolli | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | † Placodermi |
Order: | † Arthrodira |
Suborder: | † Brachythoraci |
Family: | † Holonematidae |
Genus: | † Holonema Newberry, 1889 |
Type species | |
Pterichthys rugosum Claypole, 1883 | |
Species | |
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Synonyms | |
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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.
Holonema is thought to have been a benthic-dwelling fish, based on its cambered body shape, which performs better hydrodynamically when closer to the sea bed. [1] Although previously thought to have fed in the bottom seafloor mud, more recent morphological studies of the jaw and toothplates suggest a more planktivorous lifestyle. [2] Holonema fossil individuals have small stones in the trunk shield, and pieces of feldspar in the abdominal region, suggesting Holonema used these as gastroliths for processing food. [3]
H. rugosum is the type species. It was originally described as a species of the antiarch genus, Pterichthys . Fossils are found in Frasnian strata of New York and Pennsylvania, and Givetian strata of Wisconsin and Michigan. It is a fairly large species, though the type specimen is of an 18 centimeter long median dorsal plate belonging to a small individual. Fossils of H. horridum and Glyptaspis eastmani may be this species. Fossils from Frasnian Iran (H. cf. rugosum) may also belong to this species. [4]
A large species from Givetian-aged strata in Novaya Zemlya, Russia, known only from two plates from one individual's trunk shield. [4]
H. bruehni is a recently discovered species from the Middle Eifelian-aged Brandenberg stratum of Sauerland, Northwestern Germany. Juveniles had a dermal ornamentation of irregularly spaced tubercles, while the dermal surface of adults' armor were smooth. [5]
Known from fragments from the Givetian-aged "Traverse Group" stratum in Michigan. The median dorsal plates range up to 20 cm in length. The median dorsal plate is fairly broad at the posterior end, but narrows as one approaches the anterior end. [4]
Known from fragments from the Givetian section of the Jefferson formation in Idaho. The median dorsal plate is estimated to be about 28 cm in length, and is very narrow compared to those of other species. [4]
Found in the Givetian-aged Burtnieki beds of Estonia. It is distinguished from other species in that the spinal and subpectoral margins of the anterior-ventral plates meet at almost a right angle. [4]
Found in the Eifelian-aged Aruküla beds of Estonia. The median dorsal plate has a small, horn-like process, and the trunkshield is somewhat circular when viewed from the top or bottom. [4]
From the Givetian-aged Bressay flags of the Shetland Islands. Imperfectly known. Denison 1978 suggests that it may not belong to Holonema. [4]
This species is known primarily from the Frasnian-aged Shelon beds of the Baltic, and the Donetz Basin of Russia. Specimens and fragments from the Frasnian-aged Holy Cross Mountain of Poland, and the Givetian sections of the Fiskekløfta formation in Spitzbergen. It is a fairly large species with a moderately narrow median dorsal plate. The median dorsal plate averages about 30 cm in length. [4]
This is the best-known species of the genus, known from numerous, often articulated specimens from the Frasnian-aged Gogo Reef. The species is very similar to H. rugosum, but the trunk shield is comparatively more narrow. This species reached around 60.6 cm (23.9 in) long. [6]
Placodermi is a class of armoured prehistoric fish, known from fossils, which lived from the Silurian to the end of the Devonian period. Their head and thorax were covered by articulated armoured plates and the rest of the body was scaled or naked, depending on the species. Placoderms were among the first jawed fish; their jaws likely evolved from the first of their gill arches.
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.
Ctenurella is an extinct genus of ptyctodont placoderm from the Late Devonian of Germany. The first fossils were found in the Strunde valley in the Paffrather Kalkmulde.
Bothriolepis was a widespread, abundant and diverse genus of antiarch placoderms that lived during the Middle to Late Devonian period of the Paleozoic Era. Historically, Bothriolepis resided in an array of paleo-environments spread across every paleocontinent, including near shore marine and freshwater settings. Most species of Bothriolepis were characterized as relatively small, benthic, freshwater detritivores, averaging around 30 centimetres (12 in) in length. However, the largest species, B. rex, had an estimated bodylength of 170 centimetres (67 in). Although expansive with over 60 species found worldwide, comparatively Bothriolepis is not unusually more diverse than most modern bottom dwelling species around today.
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.
Onychodus is a genus of prehistoric lobe-finned fish which lived during the Devonian period. It is one of the best known of the group of onychodontiform fishes. Scattered fossil bones of Onychodus were first discovered in 1857, in North America, and described by John Strong Newberry. Other species were found in Australia, England, Norway and Germany showing that it had a widespread range.
Homosteus is a genus of flattened arthrodire placoderm from the Middle Devonian. Fossils are found primarily in Eifelian-epoch aged strata of Europe, Canada, Greenland, and Estonia. All of the species had comparatively large, flattened heads with, as suggested by the upward opening orbits, upward-pointing eyes. These adaptations suggest that the various species were benthic predators. A study on Titanichthys, in contrast, suggests that species of Homosteus may have been filter-feeders instead.
The Gogo Formation in the Kimberley region of Western Australia is a Lagerstätte that exhibits exceptional preservation of a Devonian reef community. The formation is named after Gogo Station, a cattle station where outcrops appear and fossils are often collected from, as is nearby Fossil Downs Station.
Eastmanosteus is a fossil genus of dunkleosteid placoderms. It was closely related to the giant Dunkleosteus, but differed from that genus in size, in possessing a distinctive tuberculated bone ornament, a differently shaped nuchal plate and a more zig-zagging course of the sutures of the skull roof.
Incisoscutum is an extinct genus of arthrodire placoderm from the Early Frasnian Gogo Reef, from Late Devonian Australia. The genus contains two species I. ritchiei, named after Alex Ritchie, a palaeoichthyologist and senior fellow of the Australian Museum, and I. sarahae, named after Sarah Long, daughter of its discoverer and describer, John A. Long.
Deinodus is a form genus that includes two species: the form found in the Onondaga Formation of western New York, Deinodus bennetti, and the form found in the Columbus and Limestone of central Ohio, Deinodus ohioensis. Both species are limited to the Eifelian age of the middle Devonian Period, which occurred 398-391 million years ago.
Holonematidae is an extinct family of relatively large arthrodire placoderms from the Early to Late Devonian. Almost all fossil specimens are of armor fragments, though, all have distinctive ornamentation, often of unique arrangements and patterns of tubercles, that are diagnostic of the family. The trunkshield is very elongated, giving the armor an overall "barrel" like appearance.
Rhenonema eifeliense is a large, extinct, high-crested holonematid arthrodire placoderm from Givetian-aged strata of Middle Devonian Gerolstein, Germany. It is known from some fragments of armor, including an anterior-lateral plate estimated to be around 24 centimetres (9.4 in) long, and a portion of a median dorsal plate with a very tall crest running along the median line of the dorsal surface. The ornamentation is very similar to that of Holonema, but the concentrically arranged ridges are much coarser in Rhenonema. The holotype was originally described by Kayser, in 1880, as a species of Dinichthys, but was then redescribed in 1964 by Obruchev as a holonematid.
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.
Aspidichthys is a genus of large, distinctively tuberculated arthrodire placoderm of uncertain affinities from Upper Devonian marine strata in the Eastern United States and Europe.
Oxyosteus is a genus of trout-sized, highly compressed arthrodire placoderms from the Late Devonian of Europe: The two described species are restricted to the Late Frasnian-aged Kellwasserkalk Fauna of Bad Wildungen, while a median dorsal plate of an unnamed species is known from the Middle Frasnian Holy Cross Mountains of Poland.
Microbrachius is an extinct genus of tiny, advanced antiarch placoderms closely related to the bothriolepids. Specimens range in age from the Lower Devonian Late Emsian Stage to the Middle Devonian Upper Givetian Stage. They are characterized by having large heads with short thoracic armor of an average length of 2-4 cm. There are patterns of small, but noticeable tubercles on the armor, with the arrangement varying from species to species. Specimens of Microbrachius have been found in Scotland, Belarus, Estonia, and China.
Heightingtonaspis is an extinct genus of primitive arthrodire placoderm from Canada, England and Wales with two species previously allied with the genus Kujdanowiaspis.
Kimbryanodus is a genus of extinct ptyctodontid placoderm fish from the Frasnian of Australia.These placoderms can be told apart from others due to the large eyes, crushing tooth plates, long bodies, reduced armor, and a superficial resemblance to holocephalid fish. The group is so far the only Placoderms known with sexually dimorphic features. The fossils occur as small three dimensional isolated plates. Because of these new specimens the Ptyctodontid grouping got a taxonomic classification, it found that the genus Rhamphodopsis to be the most basal taxa. They are divided by having the more basal taxa having a median dorsal spine, a simple spinal plate, and a simple V-shaped overlap of the anterior lateral and the anterior dorsolateral plates.
Compagopiscis is an extinct genus of placoderm known from the Gogo Formation. It lived in the Upper Devonian of Western Australia. The genus is monotypic, with its only species being Compagopiscis croucheri.