Incisoscutum Temporal range: Late Devonian: Frasnian, | |
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Incisoscutum specimen WAM 03.3.28 | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | † Placodermi |
Order: | † Arthrodira |
Suborder: | † Brachythoraci |
Clade: | † Eubrachythoraci |
Clade: | † Coccosteomorphi |
Superfamily: | † Incisoscutoidea |
Family: | † Incisoscutidae |
Genus: | † Incisoscutum Dennis & Miles, 1981 |
Species | |
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Synonyms | |
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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, [1] 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.
The genus is important in the study of early vertebrates as well-preserved fossilized embryos have been found in female specimens [2] and ossified pelvic claspers found in males. [3] This shows that viviparity and internal fertilization was common amongst these primitive jawed vertebrates, which are outside the crown group Gnathostomata.
In a study of fossil remains, comparison of the ontogeny of fourteen dermal plates from Compagopiscis croucheri and the more derived species Incisoscutum ritchiei suggested that lengthwise growth occurs earlier in the ontogeny than growth in width, and that dissociated allometric heterochrony has been an important mechanism in the evolution of the arthrodires, which include placoderms. [4]
Three-dimensional uncrushed Incisioscutum fossils with remarkable soft tissue preservation were discovered within the Western Australian Gogo Formation, a warm, shallow-sea reef facies of Frasnian age. [5] I. ritchiei was a small placoderm with an estimated body length of 30.3 cm (11.9 in). [6]
Incisoscutum's body size and morphology are similar to Torosteus and Compagopiscis , suggesting a possible pelagic lifestyle, although they were on different trophic levels of their ecosystem. [7] Jaw morphology suggests that Incisoscutum was durophagous, and individual fossils have been found in numbers, suggesting possible schooling behavior. [7]
When Incisoscutum ritchiei was first described, bony plates of smaller arthrodires were discovered within the body cavity of two specimens. Due to their disorganized arrangement, posterior position behind the trunk shield, apparent gastric etching and the fact that one of the small arthodires was posterior facing relative to the adult, these bony plates were thought to represent the stomach contents of the adult Incisoscutum. [1] [2]
However, spurred by the discovery of preserved embryos inside the ptyctodonts Materpiscis attenboroughi [8] and Austroptyctodus gardineri , the ‘last meals’ of the Incisoscutum have now been reinterpreted as embryos, 5 cm in length, within pregnant adult females. [2]
The apparent gastric etching is now thought to be an early stage of ossification. The disarrangement is thought to be due to scattering as a result of the body cavity opening postmortem; worm casts throughout the body and surrounding matrix reveal that the fish carcass remained in an open environment after death. Furthermore, the lack of any taxa associated with the completely preserved delicate plates means that the arthrodires can now be excluded as stomach contents and interpreted as embryos. [2]
The discovery of embryos within Incisoscutum is of evolutionary importance as these fossils reveal that arthrodires had advanced reproductive biology and were able to give birth to live young. The Incisoscutum fossils show the oldest record of viviparity in any vertebrate, therefore internal fertilization and viviparity must have evolved in the vertebrates at least 380 million years ago and outside the gnathostomata. [2]
Viviparity in Incisoscutum and the ptyctodontids show that these placoderms were the first K-strategists in relation to breeding. They invested in rearing a smaller amount of eggs, rather than a huge spawn as their ancestors must have done. It is therefore likely that the warm reef environment of the Devonian Gogo Formation was stable and predictable, having a degree of ecological balance (for example hiding places for pregnant placoderms). This would have allowed the placoderms to invest more time and energy in producing and nurturing fewer, but more developed, offspring. [9]
Sexually dimorphic pelvic claspers have been found in male and female Incisoscutum fossil specimens. [2] [3] In males (WAM 03.3.28) [3] the completely ossified clasper is a slender rod attached to a square basal plate that articulates directly with the pelvic girdle. This contrasts to modern sharks where the clasper articulates with a basipterygial cartilage element. The tip of the distal end has a small cap of dermal bone with small pits and denticles for clinging on to the female. The proximal part of the clasper expands into a plate with four foramina, two larger above two smaller. It is thought that this anatomy corresponds to the core of an erectile element as in extant sharks. [3]
Female Incisoscutum specimens differ from these male claspers and instead fossils have been found with a broad based pelvic plate that articulates with a posteriorly directed basipterygium, similar to modern sharks. The distal end has an articulation for an additional cartilaginous segment or series. [2]
Therefore, the difference in pelvic claspers between genders suggests that sexual dimorphism was already present in the arthrodires in the Devonian. Pelvic claspers have also been discovered in pyctodontid fossils [8] suggesting homology. It is therefore suggested that pelvic claspers may characterize all of the pytodontids and arthrodires. [3] As males and females have never been found in the same locality, it is possible that Incisoscutum males and females inhabited different areas throughout most their life cycles, only coming together to mate, a behavior comparable to some extant sharks. [9]
Incisoscutum is a member of the superfamily Incisoscutoidea, which belongs to the clade Coccosteomorphi, one of the two major clades within Eubrachythoraci. [10] [11] Incisoscutum was initially placed in the family Incisoscutidae. However, Incisoscutidae is currently a monotypic family, with the genus Incisoscutum as the only member, and thus the family name Incisoscutidae is not widely used. [10] Alternatively, Incisoscutum could possibly be considered a member of the closely related family Camuropiscidae, as shown in the cladogram below: [11]
Eubrachythoraci |
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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.
Arthrodira 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.
Coccosteus is an extinct genus of arthrodire placoderm from the Devonian period. Its fossils have been found throughout Europe and North America. The majority of these have been found in freshwater sediments, though such a large range suggests that they may have been able to enter saltwater. It was a small placoderm, with Coccosteus cuspidatus measuring 29.6–39.4 cm (11.7–15.5 in) long.
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.
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.
Materpiscis 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 the mother, with remarkable preservation of a mineralised placental feeding structure. This makes Materpiscis the oldest known vertebrate to show viviparity, or giving birth to live young.
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.
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.
Bruntonichthys is an extinct monospecific genus of arthrodire placoderm from the Early Frasnian stage of the Late Devonian period. Fossils are found in the Gogo Formation of the Kimberley region of Australia. The skull is about 139 millimetres long, and had disproportionally large eye sockets. Researchers suggest it may have preyed on small mollusks.
Mcnamaraspis is an extinct monospecific genus of arthrodire placoderm that inhabited the ancient reef system of north Western Australia during the Frasnian epoch of the Late Devonian period. The type specimen was found and described by John A. Long from the Gogo Formation near Fitzroy Crossing. This fossil fish showed new anatomical features in arthrodires, like the well-preserved annular (ring-shaped) cartilages of the snout, previously inferred to be present by Erik Stensiö of Sweden. It is occasionally referred to as "The Gogo Fish" after the locale the holotype was excavated from.
Camuropiscidae is a family of mostly small, bullet or spindle-shaped extinct arthrodire placoderms from the Late Devonian. With the exception of the snub-nosed Simosteus, camuropiscid placoderms are characterized by an elongated, tubular snout. The entire family is restricted to the Frasnian Gogo Reef Formation of Australia.
Fallacosteus is an extinct monospecific genus of arthrodire placoderm from the Early Frasnian stage of the Late Devonian period, found at the Gogo Formation of Kimberley, Western Australia. As with almost all other camuropiscids, F. turneri had an elongated snout that may have enhanced its hydrodynamic streamlining.
Tubonasus is an extinct monospecific genus of long-snouted arthrodire placoderm from the Early Frasnian stage of the Late Devonian period, found at the Gogo Formation of Kimberley, Western Australia.
Xiangshuiosteus wui is an extinct monospecific genus of brachythoracid arthrodire placoderm from the Late Emsian stage of the Early Devonian epoch, discovered in Wuding County of Yunnan province, China. It has recently been reassessed as a dunkleosteid.
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.
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.
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.
Watsonosteus is an extinct genus of coccosteid arthrodire placoderm from the Late Givetian stage of the Middle Devonian period. Fossils are found in the Orkney Islands, Scotland. It was a small placoderm with a total body length of 57 cm (22 in), with the largest individuals reaching lengths of 1 m (39 in). It is one of the few arthrodires for which complete body fossils are known.
Torosteus is an extinct genus of arthrodire placoderm from the Early Frasnian stage of the Late Devonian period. Fossils are found in the Kimberley region of Australia.