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Placodermi (from Greek πλάξ 'plate' and δέρμα 'skin', literally '
plate Plate may refer to: Cooking * Plate (dishware), a broad, mainly flat vessel commonly used to serve food * Plates, tableware, dishes or dishware used for setting a table, serving food and dining * Plate, the content of such a plate (for example: ...
-skinned') is a class of armoured
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 fis ...
, known from
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
s, which lived from the
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
to the end of the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
period. Their
head A head is the part of an organism which usually includes the ears, brain, forehead, cheeks, chin, eyes, nose, and mouth, each of which aid in various sensory functions such as sight, hearing, smell, and taste. Some very simple animals may not ...
and
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the crea ...
were covered by articulated armoured plates and the rest of the body was scaled or naked, depending on the
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
. Placoderms were among the first
jawed fish 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 ...
; their jaws likely evolved from the first of their
gill arch Branchial arches, or gill arches, are a series of bony "loops" present in fish, which support the gills. As gills are the primitive condition of vertebrates, all vertebrate embryos develop pharyngeal arches, though the eventual fate of these a ...
es. Placoderms are thought to be
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
, consisting of several distinct
outgroups In sociology and social psychology, an in-group is a social group to which a person psychologically identifies as being a member. By contrast, an out-group is a social group with which an individual does not identify. People may for example i ...
or
sister taxa In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to all living jawed
vertebrate Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
s, 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 fin Pelvic fins or ventral fins are paired fins located on the ventral surface of fish. The paired pelvic fins are homologous to the hindlimbs of tetrapods. Structure and function Structure In actinopterygians, the pelvic fin consists of two e ...
s, the precursor to
hindlimb A hindlimb or back limb is one of the paired articulated appendages (limbs) attached on the caudal ( posterior) end of a terrestrial tetrapod vertebrate's torso.http://www.merriam-webster.com/medical/hind%20limb, Merriam Webster Dictionary-Hind ...
s in tetrapods, as well as true
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tea ...
. 380-million-year-old fossils of three other genera, ''
Incisoscutum ''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 ...
'', '' Materpiscis'' and '' Austroptyctodus'', represent the oldest known examples of live birth. The first identifiable placoderms appear in the
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
during the late Llandovery epoch of the early
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
. The various groups of placoderms were diverse and abundant during the Devonian, but became extinct at the end-Devonian
Hangenberg event The Hangenberg event, also known as the Hangenberg crisis or end-Devonian extinction, is a mass extinction that occurred at the end of the Famennian stage, the last stage in the Devonian Period (roughly 358.9 ± 0.4 million years ago). It is usuall ...
358.9 million years ago


Characteristics

Many placoderms, particularly the Rhenanida, Petalichthyida,
Phyllolepida Phyllolepida ("leaf scales") 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. ...
, and
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 coin ...
, were bottom-dwellers. In particular, the antiarchs, with their highly modified, jointed bony pectoral fins, were highly successful inhabitants of Middle-Late Devonian freshwater and shallow marine habitats, with the Middle to Late Devonian
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomencla ...
, ''
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, ''Bot ...
'', known from over 100 valid species. The vast majority of placoderms were
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...
s, many of which lived at or near the substrate. Many, primarily the
Arthrodires 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 ec ...
, were active,
nekton Nekton or necton (from the ) refers to the actively swimming aquatic organisms in a body of water. The term was proposed by German biologist Ernst Haeckel to differentiate between the active swimmers in a body of water, and the passive organisms ...
ic predators that dwelled in the middle to upper portions of the water column. A study of the arthrodire '' Compagopiscis'' published in 2012 concluded that placoderms (at least this particular genus) likely possessed true teeth contrary to some early studies. The teeth had well defined pulp cavities and were made of both bone and
dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
. However, the tooth and jaw development were not as closely integrated as in modern gnathostomes. These teeth were likely homologous to the teeth of other gnathostomes. One of the largest known arthrodires, ''
Dunkleosteus ''Dunkleosteus'' is an extinct genus of large armored, jawed fishes that existed during the Late Devonian period, about 382–358 million years ago. It consists of ten species, some of which are among the largest placoderms to have ever lived: ...
terrelli '', was long, and is presumed to have had a large distribution, as its remains have been found in Europe, North America and possibly Morocco. Some paleontologists regard it as the world's first
vertebrate Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
"superpredator", preying upon other predators. Other, smaller arthrodires, such as '' Fallacosteus'' and ''
Rolfosteus ''Rolfosteus'' is an extinct monospecific genus of arthrodire placoderm from the Early Frasnian stage of the Late Devonian period, found at the Gogo Formation of Western Australia.DENNIS, KIM, and R. S. Miles. "Eubrachythoracid arthrodires with ...
'', both of the
Gogo Formation 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 ...
of Western Australia, had streamlined, bullet-shaped head armor, strongly supporting the idea that many, if not most, arthrodires were active swimmers, rather than passive ambush-hunters whose armor practically anchored them to the sea floor. Some placoderms were herbivorous, such as the Middle to Late Devonian arthrodire ''
Holonema ''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. M ...
'', and some were planktivores, such as the gigantic, long arthrodire, ''
Titanichthys ''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. Many of the species approached ''Dunkleosteus'' in size and build. Unlike it ...
''. Extraordinary evidence of internal fertilization in a placoderm was afforded by the discovery in the Gogo Formation, near
Fitzroy Crossing Fitzroy or FitzRoy may refer to: People As a given name *Several members of the Somerset family (Dukes of Beaufort) have this as a middle-name: **FitzRoy Somerset, 1st Baron Raglan (1788–1855) ** Henry Charles FitzRoy Somerset, 8th Duke of Beau ...
,
Kimberley Kimberly or Kimberley may refer to: Places and historical events Australia * Kimberley (Western Australia) ** Roman Catholic Diocese of Kimberley * Kimberley Warm Springs, Tasmania * Kimberley, Tasmania a small town * County of Kimberley, ...
, Western Australia, of a small female placoderm, about in length, which died in the process of giving birth to a 6 cm ( in) offspring and was fossilized with the umbilical cord intact. The fossil, named ''
Materpiscis attenboroughi ''Materpiscis'' (Latin for ''mother fish'') is a genus of ptyctodontid placoderm from the Late Devonian located at the Gogo Formation of Western Australia. Known from only one specimen, it is unique in having an unborn embryo present inside th ...
'' (after scientist
David Attenborough Sir David Frederick Attenborough (; born 8 May 1926) is an English broadcaster, biologist, natural historian and author. He is best known for writing and presenting, in conjunction with the BBC Natural History Unit, the nine natural histor ...
), had eggs which were fertilized internally, the mother providing nourishment to the embryo and giving birth to live young. With this discovery, the placoderm became the oldest vertebrate known to have given birth to live young ("
viviparous Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the m ...
"), pushing the date of first viviparity back some 200 million years earlier than had been previously known. Specimens of the arthrodire ''
Incisoscutum ''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 ...
ritchei'', also from the Gogo Formation, have been found with embryos inside them indicating this group also had live bearing ability. The males reproduced by inserting a long
clasper In biology, a clasper is a male anatomical structure found in some groups of animals, used in mating. Male cartilaginous fish have claspers formed from the posterior portion of their pelvic fin which serve to channel semen into the female's ...
into the female. Elongated basipterygia are also found on the phyllolepid placoderms, such as '' Austrophyllolepis'' and '' Cowralepis'', both from the Middle Devonian of Australia, suggesting that the basiptergia were used in copulation. The placoderm claspers are not homologous with the claspers in
cartilaginous fishes Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
. The similarities between the structures has been revealed to be an example of convergent evolution. While the claspers in cartilaginous fishes are specialized parts of their paired pelvic fins that have been modified for copulation due to changes in the hox genes hoxd13, the origin of the mating organs in placoderms most likely relied on different sets of hox genes and were structures that developed further down the body as an extra and independent pair of appendages, but which during development turned into body parts used for reproduction only. Because they were not attached to the pelvic fins, as are the claspers in fish like sharks, they were much more flexible and could probably be rotated forward.


Evolution and extinction

It was thought for a time that placoderms became extinct due to competition from the first
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...
and early
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimor ...
s, given a combination of the supposed inherent superiority of bony fish and the presumed sluggishness of placoderms. With more accurate summaries of prehistoric organisms, it is now thought that they systematically died out as marine and freshwater ecologies suffered from the environmental catastrophes of the
Late Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wher ...
and
end-Devonian extinction The Hangenberg event, also known as the Hangenberg crisis or end-Devonian extinction, is a mass extinction that occurred at the end of the Famennian stage, the last stage in the Devonian Period (roughly 358.9 ± 0.4 million years ago). It is usuall ...
s.


Fossil record

The earliest identifiable placoderm fossils are of Chinese origin and date to the early
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
. At that time, they were already differentiated into antiarchs and
arthrodires 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 ec ...
, as well as other, more primitive, groups. Earlier fossils of basal Placodermi have not yet been discovered. The Silurian fossil record of the placoderms is both literally and figuratively fragmented. Until the discovery of ''
Silurolepis ''Silurolepis platydorsalis'' is an extinct species of Silurian-aged "maxillate" placoderm that has been described from (mostly) articulated remains. Although it has been known for several years, it was finally described by Zhang, et al., in 2 ...
'' (and then, the discoveries of ''
Entelognathus ''Entelognathus primordialis'' (“primordial complete jaw”) is a maxillate placoderm from the late Silurian (Ludlow epoch) of Qujing, Yunnan, 419 million years ago. A team led by Min Zhu of the Academy of Sciences' Institute of Vertebrate Pa ...
'' and ''
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 ...
''), Silurian-aged placoderm specimens consisted of fragments. Some of them have been tentatively identified as antiarch or arthrodire due to histological similarities; and many of them have not yet been formally described or even named. The most commonly cited example of a Silurian placoderm, '' Wangolepis'' of Silurian China and possibly Vietnam, is known only from a few fragments that currently defy attempts to place them in any of the recognized placoderm orders. So far, only three officially described Silurian placoderms are known from more than scraps: * the basal antiarch ''
Silurolepis ''Silurolepis platydorsalis'' is an extinct species of Silurian-aged "maxillate" placoderm that has been described from (mostly) articulated remains. Although it has been known for several years, it was finally described by Zhang, et al., in 2 ...
'', from the
Ludlow epoch In the geological timescale, the Ludlow Epoch (from 427.4 ± 0.5 million years ago to 423.0 ± 2.3 million years ago) occurred during the Silurian Period, after the end of the Homerian Age. It is named for the town of Ludlow in Shropshire ...
of
Yunnan Yunnan , () is a landlocked province in the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is Kunming. The province borders the ...
, China, known from an almost complete thoracic armor * ''
Entelognathus ''Entelognathus primordialis'' (“primordial complete jaw”) is a maxillate placoderm from the late Silurian (Ludlow epoch) of Qujing, Yunnan, 419 million years ago. A team led by Min Zhu of the Academy of Sciences' Institute of Vertebrate Pa ...
'', a placoderm ''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertaint ...
'' that combines features of primitive arthrodires with jaw anatomy otherwise only seen in
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...
and
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct therapsid ...
s. * ''
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 ...
'', a close relative of ''Entelognathus'' that further links ''Entelognathus'' as a transitional form between placoderms and other stem-gnathostomes and crown-group gnathostomes. The first officially described and oldest Silurian placoderm is an antiarch, ''
Shimenolepis ''Shimenolepis granifera'' is an extinct yunnanolepid placoderm from the Late Llandovery of Li County, Hunan, China. It was the first described Silurian placoderm, and is the earliest known placoderm, known from distinctively ordered plates. D ...
'', which is known from distinctively ornamented plates from the late
Llandovery Llandovery (; cy, Llanymddyfri ) is a market town and community in Carmarthenshire, Wales. It lies on the River Tywi and at the junction of the A40 and A483 roads, about north-east of Carmarthen, north of Swansea and west of Brecon. Hist ...
of
Hunan Hunan (, ; ) is a landlocked province of the People's Republic of China, part of the South Central China region. Located in the middle reaches of the Yangtze watershed, it borders the province-level divisions of Hubei to the north, Jiangxi to ...
, China. ''Shimenolepis'' plates are very similar to the early Devonian yunnanolepid ''
Zhanjilepis ''Zhanjilepis aspartilis'' is an extinct yunnanolepid placoderm from Pridoli and Lochkovian rocks of Hunan, China. ''Zhanjilepis'' is known only from distinctively ordered plates. Description Plates ''Zhanjilepis'' plates are very similar t ...
'', also known from distinctively ornamented plates. Paleontologists and placoderm specialists suspect that the scarcity of placoderms in the Silurian fossil record is due to placoderms' living in environments unconducive to fossil preservation, rather than a genuine scarcity. This hypothesis helps to explain the placoderms' seemingly instantaneous appearance and diversity at the very beginning of the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
. During the Devonian, placoderms went on to inhabit and dominate almost all known aquatic ecosystems, both
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does inclu ...
and
saltwater Saline water (more commonly known as salt water) is water that contains a high concentration of dissolved salts (mainly sodium chloride). On the United States Geological Survey (USGS) salinity scale, saline water is saltier than brackish water ...
. But this diversity ultimately suffered many casualties during the extinction event at the
Frasnian The Frasnian is one of two faunal stages in the Late Devonian Period. It lasted from million years ago to million years ago. It was preceded by the Givetian Stage and followed by the Famennian Stage. Major reef-building was under way during t ...
Famennian The Famennian is the latter of two faunal stages in the Late Devonian Epoch. The most recent estimate for its duration estimates that it lasted from around 371.1 million years ago to 359.3 million years ago. An earlier 2012 estimate, still used b ...
boundary, the Late Devonian extinctions. The remaining species then died out during the end-Devonian extinction; not a single placoderm species has been confirmed to have survived into the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferous ...
.


History of study

The earliest studies of placoderms were published by
Louis Agassiz Jean Louis Rodolphe Agassiz ( ; ) FRS (For) FRSE (May 28, 1807 – December 14, 1873) was a Swiss-born American biologist and geologist who is recognized as a scholar of Earth's natural history. Spending his early life in Switzerland, he rec ...
, in his five volumes on fossil fishes, 1833–1843. In those days, placoderms were thought to be shelled jawless fish akin to
ostracoderms Ostracoderms () are the armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic (excluding jawed fishes) (may also be polyphyletic if anaspids are closer to cyclostomes) an ...
. Some naturalists even suggested that they were shelled invertebrates or even
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked t ...
-like vertebrates. In the late 1920s, Dr.
Erik Stensiö Prof Erik Helge Osvald Stensiö HFRSE (2 October 1891 – 11 January 1984) was a Swedish paleozoologist. He later took his new surname from his place of origin and is occasionally referred to with both names (as Erik Andersson Stensiö, Erik ...
, at the
Swedish Museum of Natural History The Swedish Museum of Natural History ( sv, Naturhistoriska riksmuseet, literally, the National Museum of Natural History), in Stockholm, is one of two major museums of natural history in Sweden, the other one being located in Gothenburg. The ...
in
Stockholm Stockholm () is the capital and largest city of Sweden as well as the largest urban area in Scandinavia. Approximately 980,000 people live in the municipality, with 1.6 million in the urban area, and 2.4 million in the metropolita ...
, established the details of placoderm anatomy and identified them as true jawed fishes related to
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimor ...
s. He took fossil specimens with well-preserved skulls and ground them away, one tenth of a millimeter at a time. After each layer had been removed, he made an imprint of the next surface in
wax Waxes are a diverse class of organic compounds that are lipophilic, malleable solids near ambient temperatures. They include higher alkanes and lipids, typically with melting points above about 40 °C (104 °F), melting to give low ...
. Once the specimens had been completely ground away (and so destroyed), he made enlarged, three-dimensional models of the skulls to examine the anatomical details more thoroughly. Many other placoderm specialists thought that Stensiö was trying to shoehorn placoderms into a relationship with
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimor ...
s; however, as more fossils were found, placoderms were accepted as a sister group of chondrichthyians. Much later, the exquisitely preserved placoderm fossils from Gogo reef changed the picture again. They showed that placoderms shared anatomical features not only with chondrichthyians but with other
gnathostome 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 ...
groups as well. For example, Gogo placoderms show separate bones for the nasal capsules as in gnathostomes; in both sharks and bony fish those bones are incorporated into the braincase. Placoderms also share certain anatomical features only with the jawless
osteostracan The class Osteostraci (meaning "bony shells") is an extinct taxon of bony-armored jawless fish, termed "ostracoderms", that lived in what is now North America, Europe and Russia from the Middle Silurian to Late Devonian. Anatomically speaking, ...
s; because of this, the theory that placoderms are the sister group of chondrichthyians has been replaced by the theory that placoderms are a group of stem gnathostomes.


Taxonomy and phylogeny

Currently, Placodermi are divided into eight recognized orders. There are two further controversial orders: One is the
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
Stensioellida, containing the enigmatic ''
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 ...
''; the other is the equally enigmatic
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 ...
. These orders are considered to be basal or primitive groups within Placodermi, though their precise placement within the class remains unsure. Fossils of both are currently known only from the Hunsruck lagerstatten.


Placoderm orders


Arthrodira

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 ec ...
("jointed neck") were the most diverse and numerically successful of the placoderm orders, occupying roles from giant
apex predator An apex predator, also known as a top predator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the highest trophic lev ...
s to
detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts commun ...
-nibbling bottom dwellers. They had a movable joint between armour surrounding the head and body. As the lower jaw moved down, the head shield moved, allowing for a larger opening. All arthrodires, save for ''Compagopiscis'', lacked teeth, and used instead the sharpened edges of a bony plate, termed a "tooth plate," as a biting surface (''Compagopiscis'' had true teeth in addition to tooth plates). The eye sockets are protected by a bony ring, a feature shared by birds and some
ichthyosaurs Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, altho ...
. Early arthrodires, such as the genus '' Arctolepis'', were well-armoured fishes with flattened bodies. The largest member of this group, ''
Dunkleosteus ''Dunkleosteus'' is an extinct genus of large armored, jawed fishes that existed during the Late Devonian period, about 382–358 million years ago. It consists of ten species, some of which are among the largest placoderms to have ever lived: ...
'', was a true "superpredator" of the latest Devonian period, reaching 3 to as much as 8 metres in length. In contrast, the long-nosed ''
Rolfosteus ''Rolfosteus'' is an extinct monospecific genus of arthrodire placoderm from the Early Frasnian stage of the Late Devonian period, found at the Gogo Formation of Western Australia.DENNIS, KIM, and R. S. Miles. "Eubrachythoracid arthrodires with ...
'' measured just 15 cm. Fossils of ''
Incisoscutum ''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 ...
'' have been found containing unborn fetuses, indicating that arthrodires gave birth to live young.


Antiarchi

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 coin ...
("opposite anus") were the second most successful order of placoderms known, after the
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 ec ...
. The order's name was coined by
Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontologist, comparative anatomist, herpetologist, and ichthyologist. Born to a wealthy Quaker family, Cope distinguished himself as a child prodigy interested ...
, who, after incorrectly identifying the first fossils as being those of an armored
tunicate A tunicate is a marine invertebrate animal, a member of the subphylum Tunicata (). It is part of the Chordata, a phylum which includes all animals with dorsal nerve cords and notochords (including vertebrates). The subphylum was at one time ca ...
, mistakenly thought the eye-hole was the mouth, and 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 armoured, to the point of literally resembling a box with eyes, with the sometimes scaled, sometimes naked rear portions often becoming
sinuous Sinuosity, sinuosity index, or sinuosity coefficient of a continuously differentiable curve having at least one inflection point is the ratio of the curvilinear length (along the curve) and the Euclidean distance ( straight line) between the ...
, particularly with later forms. The pair of
pectoral fins Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
were modified into a pair of
caliper A caliper ( British spelling also calliper, or in plurale tantum sense a pair of calipers) is a device used to measure the dimensions of an object. Many types of calipers permit reading out a measurement on a ruled scale, a dial, or a digital d ...
-like, or
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chitin, o ...
-like limbs. In primitive forms, such as ''
Yunnanolepis ''Yunnanolepis'' is an extinct genus of primitive antiarch placoderm. The fossils of the various species are found in Early to Middle Devonian strata in Southern China ( Xishancun, Lianhuashan and Xitun Formations). External links ''Yunnan ...
'', the limbs were thick and short, while in advanced forms, such as ''
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, ''Bot ...
'', the limbs were long and had elbow-like joints. The function of the limbs is still not perfectly understood, but most hypothesize that they helped their owners pull themselves across the substrate, as well as allowing their owners to bury themselves into the substrate.


Brindabellaspida

Brindabellaspida (" Brindabella's shield") was a long-snouted placoderm from the
Early Devonian The Early Devonian is the first of three epochs comprising the Devonian period, corresponding to the Lower Devonian series. It lasted from and began with the Lochkovian Stage , which was followed by the Pragian from and then by the Emsian, wh ...
. When it was first discovered in 1980, it was originally regarded as a weejasperaspid acanthothoracid due to anatomical similarities with the other species found at the same locality. According to Philippe Janvier, anatomical similarities in the brain of '' Brindabellaspis stensioi'' and the brain of a
jawless fish Agnatha (, Ancient Greek 'without jaws') is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both present (cyclostomes) and extinct (conodonts and ostracoderms) species. Among recent animals, cyclostome ...
suggest it is a basal placoderm closest to the ancestral placoderm. Various Early to Middle Devonian placoderm ''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertaint ...
'' have also been inserted in the order.


Phyllolepida

Phyllolepida Phyllolepida ("leaf scales") 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. ...
("leaf scales") were flattened placoderms found throughout the world. Like other flattened placoderms they were bottom-dwelling predators that ambushed prey. Unlike other flattened placoderms, they were freshwater fish. Their armour was made of whole plates, rather than the numerous tubercles and scales of Petalichthyida. The eyes were on the sides of the head, unlike visual bottom-dwelling predators, such as stargazers or
flatfish A flatfish is a member of the ray-finned demersal fish order Pleuronectiformes, also called the Heterosomata, sometimes classified as a suborder of Perciformes. In many species, both eyes lie on one side of the head, one or the other migratin ...
, which have eyes on the top of their head. The orbits for the eyes were extremely small, suggesting the eyes were vestigial and that the phyllolepids may have been blind.


Ptyctodontida

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 ch ...
("folded teeth") were lightly armoured placoderms with big heads, big eyes and long bodies. They have a strong but superficial resemblance to modern day
chimaera Chimaeras are cartilaginous fish in the order Chimaeriformes , known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively. A ...
s. Their armour was reduced to a pattern of small plates around the head and neck. Like the extinct and related
acanthothoracids 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. ...
, and the living and unrelated holocephalians, most of the ptyctodontids are thought to have lived near the sea bottom and preyed on
shellfish Shellfish is a colloquial and fisheries term for exoskeleton-bearing aquatic invertebrates used as food, including various species of molluscs, crustaceans, and echinoderms. Although most kinds of shellfish are harvested from saltwater enviro ...
. On account of their lack of armour, some paleontologists have suggested that the Ptyctodontida were not placoderms, but holocephalians or the ancestors of holocephalians. Anatomical examinations of whole fossil specimens have shown that the similarities between these two groups are superficial. The major differences were that holocephalians have
shagreen Shagreen is a type of rawhide consisting of rough untanned skin, historically from a horse's or onager's back, or from shark or ray. Etymology The word derives from the French ''chagrin'' and is related to Italian ''zigrino'' and Venetia ...
on their skin, while ptyctodontids do not; the armoured plates and scales of holocephalians are made of
dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
, while those of ptyctodontids are made of bone; the craniums of holocephalians are similar to sharks, while those of ptyctodontids are similar to those of other placoderms; and, most importantly, that holocephalians have true teeth, while ptyctodonts have beak-like tooth plates. Ptyctodontids were
sexually dimorphic Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most anim ...
, with the males having pelvic
clasper In biology, a clasper is a male anatomical structure found in some groups of animals, used in mating. Male cartilaginous fish have claspers formed from the posterior portion of their pelvic fin which serve to channel semen into the female's ...
s and possibly claspers on the head as well.


Rhenanida

Rhenanida ("
Rhine ), Surselva, Graubünden, Switzerland , source1_coordinates= , source1_elevation = , source2 = Rein Posteriur/Hinterrhein , source2_location = Paradies Glacier, Graubünden, Switzerland , source2_coordinates= , sour ...
fish") were flattened, ray-like, bottom-dwelling
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...
s with large, upturned mouths 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 armour was made of unfused components—a mosaic of tubercles—as opposed to the solidified plates of "advanced" placoderms, such as
antiarch 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 coi ...
s and
arthrodire 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 ec ...
s. However, through comparisons of skull anatomies, rhenanids are now considered to be the sister group of the antiarchs. When rhenanids die, their "mosaics" come apart, and it has been suggested that the rarity of rhenanids in the fossil record reflects postmortem disassociation, and is not an actual rarity of the species.


Acanthothoraci

Acanthothoraci ("spine chests") were a group of chimaera-like placoderms closely related to the rhenanid placoderms. Superficially, acanthoracids resembled scaly
chimaera Chimaeras are cartilaginous fish in the order Chimaeriformes , known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively. A ...
s or small, scaly arthrodires with blunt rostrums. They were distinguished from chimaeras by a pair of large spines that emanate from their chests, the presence of large scales and plates, 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. From what can be inferred from the mouthplates of fossil specimens, acanthothoracids were shellfish hunters ecologically similar to modern-day chimaeras. Competition with their relatives, the ptyctodont placoderms, may have been one of the main reasons for the acanthothoracids' extinction prior to the mid-Devonian extinction event.


Petalichthyida

Petalichthyida ("thin-plated fish") were small, flattened placoderms, typified by their splayed fins and numerous tubercles that decorated all of the plates and scales of their armour. They reached a peak in diversity during the
Early Devonian The Early Devonian is the first of three epochs comprising the Devonian period, corresponding to the Lower Devonian series. It lasted from and began with the Lochkovian Stage , which was followed by the Pragian from and then by the Emsian, wh ...
and were found throughout the world. The petalichthids '' Lunaspis'' and ''Wijdeaspis'' are among the best known. There was an independent diversification event that occurred in what is now Southern China, producing a handful of unique genera that were once placed in their own order, "Quasipetalichthyida", named after the first discovered species there, '' Quasipetalichthys haikouensis''. Soon after the petalichthids' diversification, they went into decline. Because they had compressed body forms, it is supposed they were bottom-dwellers that pursued or ambushed smaller fish. Their diet is not clear, as none of the fossil specimens found have preserved mouth parts.


Pseudopetalichthyida

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 ...
("false petalichthyids") is a group of elongated, possibly flattened fishes comprising three, poorly preserved and poorly studied genera. It is known only from rare fossils in Lower Devonian strata in
Hunsrück The Hunsrück () is a long, triangular, pronounced upland in Rhineland-Palatinate, Germany. It is bounded by the valleys of the Moselle-Saar (north-to-west), the Nahe (south), and the Rhine (east). It is continued by the Taunus mountains, past ...
, Germany. Like '' Stensioella heintzi'', and the Rhenanida, the pseudopetalichthids had armour made up of a mosaic of tubercles. Like '' Stensioella heintzi'', the pseudopetalichthids' placement within Placodermi is suspect. The matter is not easy to resolve because there are no complete, undamaged and articulated specimens. The anatomical studies done on the crushed specimens that have been found indicate that if they are placoderms, they may be a group more advanced than the ptyctodonts. As such, placoderm experts consider
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 ...
to be the sister group of the
Arthrodires 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 ec ...
+
Phyllolepida Phyllolepida ("leaf scales") 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. ...
+
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 coin ...
trichotomy and the Acanthothoraci + Rhenanida
dichotomy A dichotomy is a partition of a whole (or a set) into two parts (subsets). In other words, this couple of parts must be * jointly exhaustive: everything must belong to one part or the other, and * mutually exclusive: nothing can belong simult ...
.


Stensioellida

Stensioellida (" eintz'slittle Stensio") contains another problematic placoderm of uncertain affinity, known only from the Lower Devonian
Hunsrück The Hunsrück () is a long, triangular, pronounced upland in Rhineland-Palatinate, Germany. It is bounded by the valleys of the Moselle-Saar (north-to-west), the Nahe (south), and the Rhine (east). It is continued by the Taunus mountains, past ...
slates of Germany. ''Stensioella'' was a thin fish that, when alive, looked vaguely like an elongated
ratfish Chimaeras are cartilaginous fish in the order Chimaeriformes , known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively. A ...
, or a skinny ''
Gemuendina ''Gemuendina stuertzi'' (named for Gemünden from where it was discovered) 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 st ...
'' with thin, strap-like pectoral fins. Similar to those of the Rhenanida, its armour was a complex mosaic of small, scale-like tubercles. The shoulder joints of its armour are similar to other placoderms, and there are superficial similarities in skull plates, and even more superficial similarities between its tubercles and the tubercles of the
rhenanids 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 ar ...
. It is tentatively placed within Placodermi as a primitive placoderm, though some paleontologists believe the rationale for the placement is inadequate. The paleontologist Philippe Janvier has suggested that ''Stensioella'' is not a placoderm, but instead is a
holocephali Holocephali ("complete heads"), sometimes given the term Euchondrocephali, is a subclass of cartilaginous fish in the class Chondrichthyes. The earliest fossils are of teeth and come from the Devonian period. Little is known about these primiti ...
an. If this is true, then the holocephalians diverged from sharks before the
Chondrichthyan Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
radiation. Critics of Janvier's position say that aside from a bodyplan superficially similar to primitive
holocephali Holocephali ("complete heads"), sometimes given the term Euchondrocephali, is a subclass of cartilaginous fish in the class Chondrichthyes. The earliest fossils are of teeth and come from the Devonian period. Little is known about these primiti ...
ans, the two groups have little else in common anatomically.


Cladogram

The following
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
shows the interrelationships of placoderms according to Carr ''et al.'' (2009): However, the cladogram had changed significantly over the years, and the placoderms are now thought to be
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
, with some being more closer to the
Eugnathostomata 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 ...
than others. The updated cladogram (Zhu et al., 2016):


See also

*
Acanthodii Acanthodii or acanthodians is an extinct class of gnathostomes (jawed fishes), typically considered a paraphyletic group. They are currently considered to represent a grade of various fish lineages leading up to the extant Chondrichthyes, which ...
* List of placoderms *
Ostracoderm Ostracoderms () are the armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic (excluding jawed fishes) (may also be polyphyletic if anaspids are closer to cyclostomes) an ...
*
Chondrichthyes Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
*
Entelognathus ''Entelognathus primordialis'' (“primordial complete jaw”) is a maxillate placoderm from the late Silurian (Ludlow epoch) of Qujing, Yunnan, 419 million years ago. A team led by Min Zhu of the Academy of Sciences' Institute of Vertebrate Pa ...


Notes


References


Citations


Other references

* * Janvier, P. ''Early Vertebrates'' Oxford, New York: Oxford University Press, 1998. * * * Long, J.A. ''The Rise of Fishes: 500 Million Years of Evolution'' Baltimore: The Johns Hopkins University Press, 1996. * * * *


External links


Annetta Markussen-Brown, "Devonian Armoured Fish" 2000


*




BBC—report on ''Dunkleosteus terrelli''
{{Authority control Prehistoric fish classes Devonian fish Silurian fish Llandovery first appearances Devonian extinctions Taxa named by Frederick McCoy Paraphyletic groups