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Placodermi
Placodermi (from Greek πλάξ 'plate' and δέρμα 'skin', literally ' plate-skinned') 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. Placoderms are thought to be paraphyletic, consisting of several distinct outgroups or sister taxa to all living jawed vertebrates, which originated among their ranks. In contrast, one 2016 analysis concluded that placodermi are likely monophyletic, though these analyses have been further dismissed with more transitional taxa between placoderms and modern gnathosthomes, solidifying their paraphyletic status. Placoderms were also the first fish to develop pelvic fins, the precursor to hindlimbs in tetrapods, as well as tr ...
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Prehistoric Fish
The evolution of fish began about 530 million years ago during the Cambrian explosion. It was during this time that the early chordates developed the skull and the vertebral column, leading to the first craniates and vertebrates. The first fish lineages belong to the Agnatha, or jawless fish. Early examples include ''Haikouichthys''. During the late Cambrian, eel-like jawless fish called the conodonts, and small mostly armoured fish known as ostracoderms, first appeared. Most jawless fish are now extinct; but the extant lampreys may approximate ancient pre-jawed fish. Lampreys belong to the Cyclostomata, which includes the extant hagfish, and this group may have split early on from other agnathans. The earliest jawed vertebrates probably developed during the late Ordovician period. They are first represented in the fossil record from the Silurian by two groups of fish: the armoured fish known as placoderms, which evolved from the ostracoderms; and the Acanthodii (or spi ...
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Gnathostomata
Gnathostomata (; from Greek: (') "jaw" + (') "mouth") are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. In addition to opposing jaws, living gnathostomes have true teeth (a characteristic which has subsequently been lost in some), paired appendages (pectoral and pelvic fins, arms, legs, wings, etc.), the elastomeric protein of elastin, and a horizontal semicircular canal of the inner ear, along with physiological and cellular anatomical characters such as the myelin sheaths of neurons, and an adaptive immune system that has the discrete lymphoid organs of spleen and thymus, and uses V(D)J recombination to create antigen recognition sites, rather than using genetic recombination in the variable lymphocyte receptor gene. It is now assumed that Gnathostomata evolved from ancestors that already possessed a pair of both pectoral and pelvic fins. Until recently these ancesto ...
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Pseudopetalichthyida
Pseudopetalichthyida is an extinct 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. On the other hand, according to anatomical studies done on the crushed specimens that have been found, those experts who do regard the Pseudopetalichthyida as placoderms consider them to be a group more advanced than the Ptyctodonts. And as such, pro-placoderm experts consider Pseudopetalichthyida to be the sister group of the Arthrodires + Phyllolepida + Antiarchi trichotomy and the Acanthothoraci + Rhenanida dichotomy. The best known ...
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Stensioella
''Stensioella heintzi'' is an enigmatic placoderm of arcane affinity. It is only known from the Lower Devonian Hunsrück slate of Germany. Anatomy ''Stensioella heintzi'' has an elongated body, a whip-like tail, and long, wing-like pectoral fins. In life, the animal would have looked vaguely like an elongated ratfish. Like the sympatric '' Gemuendina'', ''S. heintzi'' had armor made up of a complex mosaic of small, scale-like tubercles. Taxonomy ''Stensioella'' is tentatively placed within Placodermi as being among the most basal of all placoderms, as from what can be discerned from the only whole specimen found, the shoulder joints of its armor appear to be very similar to other placoderms. Despite this detail, coupled with superficial similarities in skull plates, and gross, superficial similarities between its tubercles, and the tubercles of the rhenanids, some paleontologists believe that there are very few concrete reasons for ''S. heintzis placement in Placodermi. T ...
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Acanthothoraci
Acanthothoraci (''spine chests'') is an extinct group of chimaera-like placoderms who were closely related to the rhenanid placoderms. Superficially, the acanthoracids resembled scaly chimaeras, or (relatively) heavily armored ptyctodonts. They were distinguished from chimaeras by the presence of large scales and plates, a pair of large spines that emanate from their chests (thus, the order's name), tooth-like beak plates, and the typical bone-enhanced placoderm eyeball. They were distinguished from other placoderms due to differences in the anatomy of their skulls, and due to patterns on the skull plates and thoracic plates that are unique to this order. Fossil record Fossils of the Acanthothoracids are found in various deposits from the Lower Devonian throughout the world. Fossils of the Palaeacanthaspids are found in Eurasia and Canada, while the Weejasperaspids have only been found in the Taemas Wee Jasper reef, in Southeastern Australia. Ecology From what can be inferr ...
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Bothriolepis
''Bothriolepis'' (from el, βόθρος , 'trench' and el, λεπίς 'scale') 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 (organisms that obtain nutrients by consuming decomposing plant/animal material), averaging around in length. However, the largest species, ''B. rex'', had an estimated bodylength of . Although expansive with over 60 species found worldwide, comparatively ''Bothriolepis'' is not unusually more diverse than most modern bottom dwelling species around today. Classification ''Bothriolepis'' is a genus placed within the placoderm order Antiarchi. The earliest antiarch placoderms first appeared i ...
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Arthrodira
Arthrodira (Greek for "jointed neck") is an order of extinct armored, jawed fishes of the class Placodermi that flourished in the Devonian period before their sudden extinction, surviving for about 50 million years and penetrating most marine ecological niches. Arthrodires were the largest and most diverse of all groups of Placoderms. Description Arthrodire placoderms are notable for the movable joint between armor surrounding their heads and bodies. Like all placoderms, they lacked distinct teeth; instead, they used the sharpened edges of a bony plate on their jawbone as a biting surface. The eye sockets are protected by a bony ring, a feature shared by birds and some ichthyosaurs. Early arthrodires, such as the genus ''Arctolepis'', were well-armoured fishes with flattened bodies. The largest member of this group, ''Dunkleosteus'', was a true superpredator of the latest Devonian period, reaching as much as 6 m in length. In contrast, the long-nosed ''Rolfosteus'' measured just ...
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Rhenanida
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. Evolution 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. Through 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). Presence in the fossil record ...
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Petalichthyida
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 (especially in Germany), 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. Because they had compressed body forms, it is speculated they were ...
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Antiarchi
Antiarchi ("opposite anus") 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 '' Chelysoma'', mistakenly thought that the orbital fenestra (i.e., the hole in the headshield for the eyes, nose and pineal foramen) 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. The front portions of their bodies were heavily armored, to the point of literally resembling a box with eyes, with the sometimes scaled, sometimes naked rear portions often becoming sinuous, particularly with later forms. The pair of pectoral fins were modified into a pair of caliper-like, or arthropod-like li ...
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Qilinyu
''Qilinyu rostrata'' is a "maxillate" placoderm from the late Ludlow epoch of Qujing, Yunnan, 419 million years ago.Zhu, Min, et al. "A Silurian maxillate placoderm illuminates jaw evolution." Science 354.6310 (2016): 334-336/ref> Specimens and taxonomy The holotype and paratype of ''Q. rostrata'' are two exquisitely preserved specimens both featuring a domed cranium and a curved rostrum presenting a "dolphin-like profile." The researchers' cladistic diagram shows ''Q. rostrata'' as the sister taxon of ''Entelognathus'', ''Janusiscus'' and the crown gnathostomes (i.e., bony and cartilaginous fishes and their descendants). Evolutionary significance ''Qilinyu rostrata'', together with ''Entelognathus'', demonstrates additional evidence that modern gnathostomes evolved from placoderms.Long, John A. "The first jaws." Science 354.6310 (2016): 280-281. See also * ''Entelognathus ''Entelognathus primordialis'' (“primordial complete jaw”) is a maxillate placoderm from t ...
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Ptyctodontida
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. On account of their radically reduced armor, some paleontologists have suggested that the Ptyctodontida were not actually placoderms, but actual holocephalians, some primitive group of elasmobranch fish, or even were the ancestors of the holocephalians, including the chimaeras. Thorough anatomical examinations of whole fossil specimens reveal that the profound similarities between these two groups are actually very superficial. The major differences ...
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