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Ichthyovenator
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Ichthyovenator
Ichthyovenator is a genus of spinosaurid dinosaur that lived in what is now Laos, sometime between 120 and 113 million years ago, during the Aptian stage of the Early Cretaceous period. It is known from fossils collected from the Grès supérieurs Formation of the Savannakhet Basin, the first of which were found in 2010, consisting of a partial skeleton without the skull or limbs. This specimen became the holotype of the new genus and species Ichthyovenator laosensis, and was described by palaeontologist Ronan Allain and colleagues in 2012. The generic name, meaning "fish hunter", refers to its assumed piscivorous lifestyle, while the specific name alludes to the country of Laos. In 2014, it was announced that more remains from the dig site had been recovered; these fossils included teeth, more vertebrae (backbones) and a pubic bone from the same individual.
The holotype specimen is estimated to have been between 8.5 to 10.5 metres (28 to 34 feet) long and to have weighed 2.4 tonnes (2.6 short tons; 2.4 long tons). Ichthyovenator's teeth were straight and conical, and its neck resembled that of the closely related genus Sigilmassasaurus (which may be synonymous with Spinosaurus). Like others in its family, Ichthyovenator had tall neural spines that formed a sail on its back. Unlike other known spinosaurids, Ichthyovenator's sail had a sinusoidal (wave-like) shape that curved downwards over the hips and divided into two separate sails. The pelvic girdle was reduced; the ilium—the uppermost body of the pelvis—was proportionately longer than both the pubis and ischium than in other known theropod dinosaurs. Ichthyovenator was initially thought to belong to the subfamily Baryonychinae but more recent analyses place it as a primitive member of the Spinosaurinae.
As a spinosaur, Ichthyovenator would have had a long, shallow snout and robust forelimbs. Its diet likely mainly consisted of aquatic prey, hence its etymology. Spinosaurids are also known to have eaten small dinosaurs and pterosaurs in addition to fish. Ichthyovenator's conspicuous sail might have been used for sexual display or species recognition. Fossil evidence suggests spinosaurids, especially spinosaurines, were adapted for semiaquatic lifestyles. The vertebral spines of Ichthyovenator's tail were unusually tall, suggesting—as in today's crocodilians—the tail may have aided in swimming. Ichthyovenator lived alongside sauropod and ornithopod dinosaurs, as well as bivalves, fish and turtles.
The first fossils of Ichthyovenator were found in 2010 at Ban Kalum in the Grès supérieurs Formation of the Savannakhet Basin in Savannakhet Province, Laos. These fossilized bones were recovered from a red sandstone layer within a surface area of less than 2 square metres (22 sq ft). Designated under the specimen numbers MDS BK10-01 to 15, they consist of a partly articulated, well-preserved skeleton lacking the skull and limbs, and including the third-to-last dorsal (back) vertebra, the neural spine of the last dorsal vertebra, five partial sacral (hip) vertebrae, the first two caudal (tail) vertebrae, both ilia (main hip bones), a right pubis (pubic bone), both ischia (lower and rearmost hip bones) and a posterior dorsal rib. The twelfth dorsal spine is bent sideways when viewed front-to-back due to taphonomic distortion. The centra (vertebral bodies) of the sacrals are largely incomplete due to erosion, but preserved all of their accompanying spines with their upper edges intact. At the time of Ichthyovenator's description, excavations at the site were still ongoing.
After undergoing preparation in 2011, the skeleton was used as the basis, or holotype, for the type species Ichthyovenator laosensis, which was named and described in 2012 by palaeontologists Ronan Allain, Tiengkham Xeisanavong, Philippe Richir, and Bounsou Khentavong. The generic name is derived from the Old Greek word ἰχθύς (ichthys), "fish", and the Latin word venator, "hunter", in reference to its likely piscivorous (fish-eating) lifestyle. The specific name refers to its provenance from Laos. Ichthyovenator is the third named spinosaurid dinosaur from Asia after the Thai genus Siamosaurus in 1986 and the Chinese species "Sinopliosaurus" fusuiensis in 2009. The latter may represent the same animal as Siamosaurus. In 2014, Allain published a conference paper on Ichthyovenator; the abstract indicated additional remains from the original individual had been found after excavations continued in 2012. These remains include three teeth, the left pubis, and many vertebrae, including a nearly complete neck, the first dorsal vertebra, and seven more caudal vertebrae. Some of these additional vertebrae were compared with those of other spinosaurids in a 2015 paper by German palaeontologist Serjoscha Evers and colleagues, in which they noted similarities with the vertebrae of the African spinosaurid Sigilmassasaurus.
In 2016, Gregory S. Paul estimated Ichthyovenator to have been approximately 8.5 metres (28 ft) long and to have weighed 2 tonnes (2.2 short tons; 2.0 long tons). The same year, Rubén Molina-Pérez and Asier Larramendi gave an estimate of 10.5 m (34 ft) in length, 2.95 m (9 ft 8 in) tall at the hips, and 2.4 t (2.6 short tons; 2.4 long tons) in weight.
Ichthyovenator's teeth were conical, straight, and bore no serrations. The front edges of the maxillary and dentary teeth were evident on the base of the tooth crown. The front articulating surfaces of Ichthyovenator's rear cervical and front dorsal vertebrae were one-and-a-half times wider than they were high and wider than the length of their centra. They also bore robust front tubercles (processes for skeletal muscle attachment) and lacked interzygapophyseal laminae (bony plates), which resulted in their spinopre- and spinopostzygapophyseal fossae (depressions) having open undersides. The first dorsal vertebra had extensive transverse processes (wing-like projections that articulate with the ribs), as well as deep excavations at the front and back of its base that were filled by air sacs in life. The parapophyses (processes that articulated with the capitulum of the ribs) increased in height from the rear cervicals to the first dorsal; its underside remained in contact with the front lower edge of the centrum. This is unlike the condition in most theropods, in which the parapophyses shifted towards the top of the vertebra during the transition from cervical to dorsal vertebrae. All of these features were also present in Sigilmassasaurus. Ichthyovenator's mid-cervical vertebrae had elongate, somewhat wider-than-tall centra that became progressively shorter towards the rear of the neck, as well as well-developed keels on their bottom surfaces, traits that were shared with the spinosaurids Baryonyx, Suchomimus, Sigilmassasaurus, and Vallibonavenatrix. The cervical neural spines of Ichthyovenator were taller than in Sigilmassasaurus and Baryonyx but shared the blade-like shape with those two taxa at the mid-cervicals. The holotype dorsal rib, which was found near the twelfth dorsal vertebra, had a head typical of the ribs of other moderate-to-large-sized theropods. The rib shaft formed a half-circle. The rib's lower end was slightly expanded both sideways and to the front and back. This condition, which differs from the tapered, pointed tips seen in the ribs of other theropods, suggests the rearmost dorsal ribs articulated with the complex of the sternum (breast bone).
Like many other spinosaurids, Ichthyovenator had a sail on its back and hips that was formed by the elongated neural spines of its vertebrae. Uniquely among known members of the family, Ichthyovenator's sail was divided in two over the hips and had a sinusoid (wave-like) curvature. This is in contrast to the related genera Spinosaurus and Suchomimus—both of which had continuous sails that rose to a peak before sloping down again—and Baryonyx, which showed a less-developed sail, having much lower dorsal spines. Ichthyovenator's preserved dorsal and sacral spinal column, which is over 1 m (3 ft 3 in) long, shows a very high spine on the twelfth dorsal vertebra representing a crest that rose from the back and a lower rounded sail that extended from the sacral vertebrae of the hips; the latter sail's apex was located above the third and fourth sacrals. The 54.6-centimetre (21.5 in) high spine of the twelfth dorsal vertebra widened towards the top, giving it a trapezoidal shape—similar to the condition seen in a neural spine from Vallibonavenatrix—whereas the spines of other known spinosaurids were roughly rectangular. It also lacked the forward or backward inclination of Spinosaurus's neural spines. Its front corner formed a 3-centimetre (1.2 in) narrow-process, pointing upwards. The spine of the thirteenth dorsal vertebra has been only partly preserved; its upper and lower ends had broken off from taphonomic causes. From its general shape, the describers inferred however, it was about as long as the preceding spine. This would imply the back edge of the front sail would form a rectangular corner, because the spine of the much lower first sacral vertebra is about 21 cm (8.3 in), creating a sudden hiatus in the sail's profile. The spine of the second sacral vertebra steeply curved upwards again, joining the 39 cm (15 in) and 48 cm (19 in) high, broad, fan-shaped spines of the third and fourth sacrals. The 40 cm (16 in) tall spine of the fifth sacral gradually descended. Unlike in the sails of most other theropods, including that of Suchomimus, the sacral spines were not fused and had no extensive contacts. The second and third sacral centra were fused; the suture connecting them was still visible. Only the rearmost two dorsal vertebrae are preserved, so to what extent the dorsal sail continued towards the front of the animal's back is unknown.
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Ichthyovenator
Ichthyovenator is a genus of spinosaurid dinosaur that lived in what is now Laos, sometime between 120 and 113 million years ago, during the Aptian stage of the Early Cretaceous period. It is known from fossils collected from the Grès supérieurs Formation of the Savannakhet Basin, the first of which were found in 2010, consisting of a partial skeleton without the skull or limbs. This specimen became the holotype of the new genus and species Ichthyovenator laosensis, and was described by palaeontologist Ronan Allain and colleagues in 2012. The generic name, meaning "fish hunter", refers to its assumed piscivorous lifestyle, while the specific name alludes to the country of Laos. In 2014, it was announced that more remains from the dig site had been recovered; these fossils included teeth, more vertebrae (backbones) and a pubic bone from the same individual.
The holotype specimen is estimated to have been between 8.5 to 10.5 metres (28 to 34 feet) long and to have weighed 2.4 tonnes (2.6 short tons; 2.4 long tons). Ichthyovenator's teeth were straight and conical, and its neck resembled that of the closely related genus Sigilmassasaurus (which may be synonymous with Spinosaurus). Like others in its family, Ichthyovenator had tall neural spines that formed a sail on its back. Unlike other known spinosaurids, Ichthyovenator's sail had a sinusoidal (wave-like) shape that curved downwards over the hips and divided into two separate sails. The pelvic girdle was reduced; the ilium—the uppermost body of the pelvis—was proportionately longer than both the pubis and ischium than in other known theropod dinosaurs. Ichthyovenator was initially thought to belong to the subfamily Baryonychinae but more recent analyses place it as a primitive member of the Spinosaurinae.
As a spinosaur, Ichthyovenator would have had a long, shallow snout and robust forelimbs. Its diet likely mainly consisted of aquatic prey, hence its etymology. Spinosaurids are also known to have eaten small dinosaurs and pterosaurs in addition to fish. Ichthyovenator's conspicuous sail might have been used for sexual display or species recognition. Fossil evidence suggests spinosaurids, especially spinosaurines, were adapted for semiaquatic lifestyles. The vertebral spines of Ichthyovenator's tail were unusually tall, suggesting—as in today's crocodilians—the tail may have aided in swimming. Ichthyovenator lived alongside sauropod and ornithopod dinosaurs, as well as bivalves, fish and turtles.
The first fossils of Ichthyovenator were found in 2010 at Ban Kalum in the Grès supérieurs Formation of the Savannakhet Basin in Savannakhet Province, Laos. These fossilized bones were recovered from a red sandstone layer within a surface area of less than 2 square metres (22 sq ft). Designated under the specimen numbers MDS BK10-01 to 15, they consist of a partly articulated, well-preserved skeleton lacking the skull and limbs, and including the third-to-last dorsal (back) vertebra, the neural spine of the last dorsal vertebra, five partial sacral (hip) vertebrae, the first two caudal (tail) vertebrae, both ilia (main hip bones), a right pubis (pubic bone), both ischia (lower and rearmost hip bones) and a posterior dorsal rib. The twelfth dorsal spine is bent sideways when viewed front-to-back due to taphonomic distortion. The centra (vertebral bodies) of the sacrals are largely incomplete due to erosion, but preserved all of their accompanying spines with their upper edges intact. At the time of Ichthyovenator's description, excavations at the site were still ongoing.
After undergoing preparation in 2011, the skeleton was used as the basis, or holotype, for the type species Ichthyovenator laosensis, which was named and described in 2012 by palaeontologists Ronan Allain, Tiengkham Xeisanavong, Philippe Richir, and Bounsou Khentavong. The generic name is derived from the Old Greek word ἰχθύς (ichthys), "fish", and the Latin word venator, "hunter", in reference to its likely piscivorous (fish-eating) lifestyle. The specific name refers to its provenance from Laos. Ichthyovenator is the third named spinosaurid dinosaur from Asia after the Thai genus Siamosaurus in 1986 and the Chinese species "Sinopliosaurus" fusuiensis in 2009. The latter may represent the same animal as Siamosaurus. In 2014, Allain published a conference paper on Ichthyovenator; the abstract indicated additional remains from the original individual had been found after excavations continued in 2012. These remains include three teeth, the left pubis, and many vertebrae, including a nearly complete neck, the first dorsal vertebra, and seven more caudal vertebrae. Some of these additional vertebrae were compared with those of other spinosaurids in a 2015 paper by German palaeontologist Serjoscha Evers and colleagues, in which they noted similarities with the vertebrae of the African spinosaurid Sigilmassasaurus.
In 2016, Gregory S. Paul estimated Ichthyovenator to have been approximately 8.5 metres (28 ft) long and to have weighed 2 tonnes (2.2 short tons; 2.0 long tons). The same year, Rubén Molina-Pérez and Asier Larramendi gave an estimate of 10.5 m (34 ft) in length, 2.95 m (9 ft 8 in) tall at the hips, and 2.4 t (2.6 short tons; 2.4 long tons) in weight.
Ichthyovenator's teeth were conical, straight, and bore no serrations. The front edges of the maxillary and dentary teeth were evident on the base of the tooth crown. The front articulating surfaces of Ichthyovenator's rear cervical and front dorsal vertebrae were one-and-a-half times wider than they were high and wider than the length of their centra. They also bore robust front tubercles (processes for skeletal muscle attachment) and lacked interzygapophyseal laminae (bony plates), which resulted in their spinopre- and spinopostzygapophyseal fossae (depressions) having open undersides. The first dorsal vertebra had extensive transverse processes (wing-like projections that articulate with the ribs), as well as deep excavations at the front and back of its base that were filled by air sacs in life. The parapophyses (processes that articulated with the capitulum of the ribs) increased in height from the rear cervicals to the first dorsal; its underside remained in contact with the front lower edge of the centrum. This is unlike the condition in most theropods, in which the parapophyses shifted towards the top of the vertebra during the transition from cervical to dorsal vertebrae. All of these features were also present in Sigilmassasaurus. Ichthyovenator's mid-cervical vertebrae had elongate, somewhat wider-than-tall centra that became progressively shorter towards the rear of the neck, as well as well-developed keels on their bottom surfaces, traits that were shared with the spinosaurids Baryonyx, Suchomimus, Sigilmassasaurus, and Vallibonavenatrix. The cervical neural spines of Ichthyovenator were taller than in Sigilmassasaurus and Baryonyx but shared the blade-like shape with those two taxa at the mid-cervicals. The holotype dorsal rib, which was found near the twelfth dorsal vertebra, had a head typical of the ribs of other moderate-to-large-sized theropods. The rib shaft formed a half-circle. The rib's lower end was slightly expanded both sideways and to the front and back. This condition, which differs from the tapered, pointed tips seen in the ribs of other theropods, suggests the rearmost dorsal ribs articulated with the complex of the sternum (breast bone).
Like many other spinosaurids, Ichthyovenator had a sail on its back and hips that was formed by the elongated neural spines of its vertebrae. Uniquely among known members of the family, Ichthyovenator's sail was divided in two over the hips and had a sinusoid (wave-like) curvature. This is in contrast to the related genera Spinosaurus and Suchomimus—both of which had continuous sails that rose to a peak before sloping down again—and Baryonyx, which showed a less-developed sail, having much lower dorsal spines. Ichthyovenator's preserved dorsal and sacral spinal column, which is over 1 m (3 ft 3 in) long, shows a very high spine on the twelfth dorsal vertebra representing a crest that rose from the back and a lower rounded sail that extended from the sacral vertebrae of the hips; the latter sail's apex was located above the third and fourth sacrals. The 54.6-centimetre (21.5 in) high spine of the twelfth dorsal vertebra widened towards the top, giving it a trapezoidal shape—similar to the condition seen in a neural spine from Vallibonavenatrix—whereas the spines of other known spinosaurids were roughly rectangular. It also lacked the forward or backward inclination of Spinosaurus's neural spines. Its front corner formed a 3-centimetre (1.2 in) narrow-process, pointing upwards. The spine of the thirteenth dorsal vertebra has been only partly preserved; its upper and lower ends had broken off from taphonomic causes. From its general shape, the describers inferred however, it was about as long as the preceding spine. This would imply the back edge of the front sail would form a rectangular corner, because the spine of the much lower first sacral vertebra is about 21 cm (8.3 in), creating a sudden hiatus in the sail's profile. The spine of the second sacral vertebra steeply curved upwards again, joining the 39 cm (15 in) and 48 cm (19 in) high, broad, fan-shaped spines of the third and fourth sacrals. The 40 cm (16 in) tall spine of the fifth sacral gradually descended. Unlike in the sails of most other theropods, including that of Suchomimus, the sacral spines were not fused and had no extensive contacts. The second and third sacral centra were fused; the suture connecting them was still visible. Only the rearmost two dorsal vertebrae are preserved, so to what extent the dorsal sail continued towards the front of the animal's back is unknown.
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