Rhenanida | |
---|---|
Gemuendina stuertzi | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | † Placodermi |
Order: | † Rhenanida Broili, 1930 |
Genera | |
†? Ohioaspis Contents |
Rhenanida ("Rhine (fish)") is an order of scaly placoderms. Unlike most other placoderms, the rhenanids' armor was made up of a mosaic of unfused scales and tubercles. The patterns and components of this "mosaic" correspond to the plates of armor in other, more advanced placoderms, suggesting that the ancestral placoderm had armor made of unfused components, as well.
All rhenanids were flattened, ray-like, bottom-dwelling predators that lived in marine environments.
The rhenanids were once presumed to be the most primitive, or at least the closest to the ancestral placoderm, as their armor was made up of a mosaic of tubercles, as opposed to the solidified plates of "advanced" placoderms, such as antiarchs and arthrodires. However, comparing the skull anatomies of Jagorina pandora with those of antiarchs, the rhenanids are considered to be the sister group of the antiarchs (together with their respective Acanthothoracid relatives).[ citation needed ]
The fossil record of Rhenanida is very sparse, with most fossils being isolated tubercles and skull fragments that are identified as being similar to Gemuendina stuertzi , the most well-known rhenanid, known from several specimens from the Hunsruck slates. Given the rhenanids' worldwide distribution, this paupacity probably did not reflect a scarcity of living individuals (when the order was alive), but reflects the fact that rhenanid armor disintegrated into isolated fragments, and scattered soon after the owner's demise. Most fossils of rhenanids are from the Early Devonian, primarily in the United States and Germany. The recently discovered Nefudina qalibahensis is known from Northeastern Saudi Arabia. Asterosteus stenocephalus is known from Mid Devonian Ohio. Another species of rhenanid was Bolivosteus chacomensis , of the Lower to Middle Devonian Malvinokaffric Fauna of Western Gondwana, in what is now Bolivia, South America. The youngest rhenanid, Jagorina pandora is known from Upper Devonian Germany.
There are five recognized species of rhenanids, in five genera, Asterosteus stenocephalus, Nefudina qalibahensis, Gemuendina stuertzi, Jagorina pandora, and Bolivosteus chacomensis. They are all placed within the family Asterosteidae, erected by Woodward in 1891: other families attributed to Rhenanida, i.e., Gemuendinidae and Jagorinidae, are considered synonyms. [1]
A sixth genus, Ohioaspis , is of questionable status, as the first specimens were ichthyoliths that were originally described as being tubercles from a new species of Asterosteus. Later examinations of these tubercles have led to the formation of two camps of experts, one of which that believe the three recognized species of Ohioaspis were rhenanids, while the other suggests that they were actually some sort of ostracoderm agnathans.
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.
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.
Petalichthyida is an extinct order of small, flattened placoderm fish. They are typified by their splayed pectoral fins, exaggerated lateral spines, flattened bodies, and numerous tubercles that decorated all of the plates and scales of their armor. They reached a peak in diversity during the Early Devonian and were found throughout the world, particularly in Europe, North America, Asia, South America, and Australia. The petalichthids Lunaspis and Wijdeaspis are among the best known. The earliest and most primitive known petalichthyid is Diandongpetalichthys, which is from earliest Devonian-aged strata of Yunnan. The presence of Diandongpetalichthys, along with other primitive petalichthyids including Neopetalichthys and Quasipetalichthys, and more advanced petalichthyids, suggest that the order may have arisen in China, possibly during the late Silurian.
Gemuendina stuertzi is a placoderm of the order Rhenanida, of the seas of Early Devonian Germany. In life, Gemuendina resembled a scaly ray with an upturned head, or a large-finned stargazer. G. stuertzi is often invoked as an example of convergent evolution- with its flat body and huge, wing-like pectoral fins it has a strong, albeit superficial similarity to rays. Unlike rays, however, both Gemuendina`s eyes and nostrils were placed atop the head, facing upward. Furthermore, G. stuertzi's upturned mouth would have enabled it to suction prey that swam overhead, rather than swallow sediment or suction prey out of the substrate like modern rays.
Antiarchi is an order of heavily armored placoderms. The antiarchs form the second-most successful group of placoderms after the arthrodires in terms of numbers of species and range of environments. The order's name was coined by Edward Drinker Cope, who, when examining some fossils that he thought were armored tunicates related to Chelyosoma, mistakenly thought that the orbital fenestra was the opening for the mouth, or oral siphon, and that the opening for the anal siphon was on the other side of the body, as opposed to having both oral and anal siphons together at one end.
Stensioella heintzi is an enigmatic placoderm of arcane affinity. It is only known from the Lower Devonian Hunsrück slate of Germany. The genus is named after Erik Stensiö, the species name honours Anatol Heintz.
Pseudopetalichthyida is an order of lightly armored placoderms known only from rare fossils in Lower Devonian strata in Hunsrück, Germany. Like Stensioella heintzi, and the Rhenanida, the Pseudopetalichthids had armor made up of a mosaic of tubercles. Like Stensioella heintzi, the Pseudopetalichthids' placement within Placodermi is suspect. However, due to a gross lack of whole, uncrushed, articulated specimens, there are no other groups that the Pseudopetalichthids could be, for a lack of a better word, pigeonholed into.
Phyllolepida is an extinct taxon of flattened placoderms found throughout the world, with fossils being found in Devonian strata. Like other flattened placoderms, the phyllolepids were bottom-dwelling predators that ambushed prey. Unlike other flattened placoderms, the phyllolepids were inhabitants of freshwater environments.
Acanthothoraci is an extinct group of chimaera-like placoderms closely related to the rhenanid placoderms. Superficially, the acanthoracids resembled scaly chimaeras and (relatively) heavily armored ptyctodonts. They were distinguished from chimaeras by their large scales and plates, a pair of large spines that emanate from their chests, tooth-like beak plates, and the typical bone-enhanced placoderm eyeball. They were distinguished from other placoderms by differences in skull anatomy and by patterns on the skull plates and thoracic plates that are unique to this order.
The ptyctodontids ("folded-teeth") are placoderms of the order Ptyctodontida, containing the family Ptyctodontidae. With their big heads, big eyes, reduced armor and long bodies, the ptyctodontids bore a superficial resemblance to modern day chimaeras (Holocephali). Their armor was reduced to a pattern of small plates around the head and neck. Like the extinct and related acanthothoracids, and the living and unrelated holocephalians, most of the ptyctodontids are thought to have lived near the sea bottom and preyed on shellfish.
Jagorina pandora is a rhenanid placoderm of Upper Frasnian Germany. As with other rhenanids, it was a flattened, skate-like fish protected in an armor made up of unfused tubercles, and preyed on other, smaller fish by ambushing them from the sea bottom.
Lunaspis is an extinct genus of armor-plated petalichthyid placoderm fish that lived in shallow marine environments of the Early Devonian period, from approximately 409.1 to 402.5 million year ago. Fossils have been found in Germany, China and Australia. There are three different identified species of within the genus Lunaspis: L. broilii, L. heroldi, and L. prumiensis.
Phyllolepis is the type genus of Phyllolepida, an extinct taxon of arthrodire placoderm fish from the middle to late Devonian. The species of Phyllolepis, themselves, are restricted to the Famennian-aged freshwater strata of the Late Devonian, around 360 million years ago. Fossils of this genus have been found primarily in Europe and North America. The end of the Devonian saw them disappear in a mass extinction.
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.
Asterosteus is a genus of rhenanid placoderm from the Eifelian. The type species, A. stenocephalus, is known from an incomplete skull from Middle Devonian strata in Ohio. What may be a second species is described from fragments decorated with star-shaped tubercules from the Eifelian-aged Gebze beds of Turkey.
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.
Microbrachiidae is an extinct family of tiny, advanced antiarch placoderms closely related to the bothriolepidids.
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.
Romundina is a small, heavily armored extinct genus of acanthothoracid placoderms which lived in shallow marine environments in the early Devonian (Lochkovian). The name Romundina honors Canadian geologist and paleontologist Dr. Rómundur (Raymond) Thorsteinsson of Calgary, Alberta, Canada. Romundina are believed to have lived on Earth between 400 and 419 million years ago. The closest known relative to Romundina is the acanthothoracid Radotina. The type and only described species is R. stellina.
Qilinyu is a genus of early placoderm from the late Silurian of China. It contains a single species, Qilinyu rostrata, from the Xiaoxiang fauna of the Kuanti Formation. Along with its contemporary Entelognathus, Qilinyu is an unusual placoderm showing some traits more similar to bony fish, such as dermal jaw bones and lobe-like fins. It can be characterized by adaptations for a benthic lifestyle, with the mouth and nostrils on the underside of the head, similar to the unrelated antiarch placoderms. The shape of the skull has been described as "dolphin-like", with a domed cranium and a short projecting rostrum.