Baryonyx

In 1997, Charig and Milner noted that two fragmentary spinosaurid snouts from the

Snout of the holotype specimen, from the left and below

Holotype dentary from the left

The skull of Baryonyx is incompletely known, and much of the middle and hind portions are not preserved. The full length of the skull is estimated to have been 91–95 centimetres (36–37 inches) long, based on comparison with that of the related genus Suchomimus (which was 20% larger).

Baryonyx had a rudimentary secondary palate, similar to crocodiles but unlike most theropod dinosaurs.^[44] A rugose (roughly wrinkled) surface suggests the presence of a horny pad in the roof of the mouth. The nasal bones were fused, which distinguished Baryonyx from other spinosaurids, and a sagittal crest was present above the eyes, on the upper mid-line of the nasals. This crest was triangular, narrow, and sharp in its front part, and was distinct from those of other spinosaurids in ending hind wards in a cross-shaped process. The lacrimal bone in front of the eye appears to have formed a horn core similar to those seen, for example, in Allosaurus, and was distinct from other spinosaurids in being solid and almost triangular. The occiput was narrow, with the paroccipital processes pointing outwards horizontally, and the basipterygoid processes were lengthened, descending far below the basioccipital (the lowermost bone of the occiput).^[3][42][45] Sereno and colleagues suggested that some of Baryonyx's cranial bones had been misidentified by Charig and Milner, resulting in the occiput being reconstructed as too deep, and that the skull was instead probably as low, long and narrow as that of Suchomimus.^[45] The front 14 cm (5.5 in) of the dentary in the mandible sloped upwards towards the curve of the snout. The dentary was very long and shallow, with a prominent Meckelian groove on the inner side. The mandibular symphysis, where the two halves of the lower jaw connected at the front, was particularly short. The rest of the lower jaw was fragile; the hind third was much thinner than the front, with a blade-like appearance. The front part of the dentary curved outwards to accommodate the large front teeth, and this area formed the mandibular part of the rosette. The dentary–like the snout—had many foramina.^[3][38]

Most of the teeth found with the holotype specimen were not in articulation with the skull; a few remained in the upper jaw, and only small replacement teeth were still borne by the lower jaw. The teeth had the shape of recurved cones, where slightly flattened from sideways, and their curvature was almost uniform. The roots were very long, and tapered towards their extremity.^[3] The carinae (sharp front and back edges) of the teeth were finely serrated with denticles on the front and back, and extended all along the crown. There were around six to eight denticles per mm (0.039 in), a much larger number than in large-bodied theropods like Torvosaurus and Tyrannosaurus. Some of the teeth were fluted, with six to eight ridges along the length of their inner sides and fine-grained enamel (outermost layer of teeth), while others bore no flutes; their presence is probably related to position or ontogeny (development during growth).^[3][38] The inner side of each tooth row had a bony wall. The number of teeth was large compared to most other theropods, with six to seven teeth in each premaxilla and thirty-two in each dentary. Based on the closer packing and smaller size of the dentary teeth compared to those in the corresponding length of the premaxilla, the difference between the number of teeth in the upper and lower jaws appears to have been more pronounced than in other theropods.^[3] The terminal rosette in the upper jaw of the holotype had thirteen dental alveoli (tooth sockets), six on the left and seven on the right side, showing tooth count asymmetry. The first four upper teeth were large (with the second and third the largest), while the fourth and fifth progressively decreased in size.^[3] The diameter of the largest was twice that of the smallest. The first four alveoli of the dentary (corresponding to the tip of the upper jaw) were the largest, with the rest more regular in size. Small subtriangular interdental plates were present between the alveoli.^[3][38]

Postcranial skeleton

Three cervical vertebrae from the neck of the holotype in left side view, the third also shown from the front (left), and reconstructed necks of the spinosaurids Sigilmassasaurus (A) and Baryonyx (B), showing their curvature (right)

Initially thought to have lacked the sigmoid curve typical of theropods,^[3] the neck of Baryonyx does appear to have formed an S shape, though straighter than in other theropods.^[46] The cervical vertebrae of the neck tapered towards the head and became progressively longer front to back. Their zygapophyses (the processes that connected the vertebrae) were flat, and their epipophyses (processes to which neck muscles attached) were well developed. The axis (the second neck vertebra) was small relative to the size of the skull and had a well-developed hyposphene. The neural arches of the cervical vertebrae were not always sutured to the centra (the bodies of the vertebrae), and the neural spines there were low and thin. The cervical ribs were short, similar to those of crocodiles, and possibly overlapped each other somewhat. The centra of the dorsal vertebrae of the back were similar in size. Like in other theropods, the skeleton of Baryonyx showed skeletal pneumaticity, reducing its weight through fenestrae (openings) in the neural arches and pleurocoels (hollow depressions) in the centra (primarily near the transverse processes). From front to back, the neural spines of the dorsal vertebrae changed from short and stout to tall and broad. One isolated dorsal neural spine was moderately elongated and slender, indicating that Baryonyx may have had a hump or ridge along the centre of its back (though incipiently developed compared to those of other spinosaurids). Baryonyx was unique among spinosaurids in having a marked constriction from side to side in a vertebra that either belonged to the sacrum or front of the tail.^[3][6][45]

The coracoid tapered hind-wards when viewed in profile, and, uniquely among spinosaurids, connected with the scapula in a peg-and-notch articulation. The scapulae were robust and the bones of the forelimb were short in relation to the animal's size, but broad and sturdy. The humerus was short and stout, with its ends broadly expanded and flattened—the upper side for the

In their original description, Charig and Milner

Barker and colleagues found support for a Baryonychinae-Spinosaurinae split in 2021, and the following cladogram shows the position of Baryonyx within Spinosauridae according to their study:^[26]

	Megalosauridae
Spinosauridae	Vallibonavenatrix Baryonychinae ML1190 (cf. Baryonyx sp.) Baryonyx Ceratosuchopsini Suchomimus Riparovenator Ceratosuchops Spinosaurinae Camarillasaurus Ichthyovenator Irritator Spinosaurini Sigilmassasaurus "Spinosaurus B" MSNM-V4047 FSAC-KK11888 Spinosaurus holotype

Evolution

Spinosaurids appear to have been widespread from the

megalosaurid relatives. They also suggested that the spinosaurines and baryonychines diverged before the Barremian age of the Early Cretaceous.^[45]

Several theories have been proposed about the

stepping stone between Europe and Africa, which is supported by the presence of baryonychines in Iberia. The direction of the dispersal between Europe and Africa is still unknown,^[30] and subsequent discoveries of spinosaurid remains in Asia and possibly Australia indicate that it may have been complex.^[18]

Candeiro and colleagues suggested in 2017 that spinosaurids of northern Gondwana were replaced by other predators, such as

abelisauroids, since no definite spinosaurid fossils are known from after the Cenomanian anywhere in the world. They attributed the disappearance of spinosaurids and other shifts in the fauna of Gondwana to changes in the environment, perhaps caused by transgressions in sea level.^[40] Malafaia and colleagues stated in 2020 that Baryonyx remains the oldest unquestionable spinosaurid, while acknowledging that older remains had also been tentatively assigned to the group.^[23] Barker and colleagues found support for a European origin for spinosaurids in 2021, with an expansion to Asia and Gondwana during the first half of the Early Cretaceous. In contrast to Sereno, these authors suggested there had been at least two dispersal events from Europe to Africa, leading to Suchomimus and the African part of Spinosaurinae.^[26]

Palaeobiology

Diet and feeding

In 1986, Charig and Milner suggested that its elongated snout with many finely serrated teeth indicated that Baryonyx was

storks.^[3][1] In 1987, the biologist Andrew Kitchener disputed the piscivorous behaviour of Baryonyx and suggested that it would have been a scavenger, using its long neck to feed on the ground, its claws to break into a carcass, and its long snout (with nostrils far back for breathing) for investigating the body cavity.^[55] Kitchener argued that Baryonyx's jaws and teeth were too weak to kill other dinosaurs and too heavy to catch fish, with too many adaptations for piscivory.^[55] According to the palaeontologist Robin E. H. Reid, a scavenged carcass would have been broken up by its predator and large animals capable of doing so—such as grizzly bears—are also capable of catching fish (at least in shallow water).^[56]

iguanodontid

found within the rib cage of the B. walkeri holotype

In 1997, Charig and Milner demonstrated direct dietary evidence in the stomach region of the B. walkeri holotype. It contained the first evidence of piscivory in a theropod dinosaur, acid-etched scales and teeth of the common fish

macro-predator like Allosaurus. They suggested that Baryonyx mainly used its forelimbs and large claws to catch, kill and tear apart larger prey. An apparent gastrolith (gizzard stone) was also found with the specimen.^[3] The German palaeontologist Oliver Wings suggested in 2007 that the low number of stones found in theropods like Baryonyx and Allosaurus could have been ingested by accident.^[58] In 2004, a pterosaur neck vertebra from Brazil with a spinosaurid tooth embedded in it reported by Buffetaut and colleagues confirmed that the latter were not exclusively piscivorous.^[59]

crocodilians

that were compared to those of spinosaurs in a 2013 study (right)

A 2005

CT scanned snouts by the palaeontologist Emily J. Rayfield and colleagues indicated that the biomechanics of Baryonyx were most similar to those of the gharial and unlike those of the American alligator and more-conventional theropods, supporting a piscivorous diet for spinosaurids. Their secondary palate helped them resist bending and torsion of their tubular snouts.^[44] A 2013 beam-theory study by the palaeontologists Andrew R. Cuff and Rayfield compared the biomechanics of CT-scanned spinosaurid snouts with those of extant crocodilians, and found the snouts of Baryonyx and Spinosaurus similar in their resistance to bending and torsion. Baryonyx was found to have relatively high resistance in the snout to dorsoventral bending compared with Spinosaurus and the gharial. The authors concluded (in contrast to the 2007 study) that Baryonyx performed differently than the gharial; spinosaurids were not exclusive piscivores, and their diet was determined by their individual size.^[8]

In a 2014 conference abstract, the palaeontologist Danny Anduza and Fowler pointed out that grizzly bears do not gaff fish out of the water as was suggested for Baryonyx, and also ruled out that the dinosaur would not have darted its head like herons, since the necks of spinosaurids were not strongly S-curved, and their eyes were not well-positioned for

fishing cats. They did not find the teeth of spinosaurids suitable for dismembering prey, due to their lack of serrations, and suggested they would have swallowed prey whole (while noting they could also have used their claws for dismemberment).^[61]

A 2016 study by the palaeontologist Christophe Hendrickx and colleagues found that adult spinosaurs could displace their mandibular rami (halves of the lower jaw) sideways when the jaw was depressed, which allowed the

pike conger eels; these fish also have jaws that are compressed side to side (whereas the jaws of crocodilians are compressed from top to bottom), an elongated snout with a "terminal rosette" that bears enlarged teeth, and a notch behind the rosette with smaller teeth. Such jaws likely evolved for grabbing prey in aquatic environments with low light, and may have helped in prey detection.^[62]

braincase of Baryonyx and Ceratosuchops (left), and 3D endocast

of the brain of Baryonyx (right)

A 2023 study by Barker and colleagues based on CT scans of the braincases of Baryonyx and Ceratosuchops found that the brain anatomy of these baryonychines was similar to that of other non-

reptile encephalization quotient values imply that the cognitive capacity and behavioural sophistication of baryonychines did not deviate much from that of other basal theropods.^[63]

Motion and semi-aquatic habits

In their original description, Charig and Milner did not consider Baryonyx to be aquatic (due to its nostrils being on the sides of its snout—far from the tip—and the form of the post-cranial skeleton), but thought it was capable of swimming, like most land vertebrates.

centre of body mass. The authors found quadrupedality unlikely for Baryonyx, since the better-known legs of the closely related Suchomimus did not support this posture.^[52]

In 2017, the palaeontologists David E. Hone and Holtz hypothesized that the head crests of spinosaurids were probably used for sexual or threat display. The authors also pointed out that (like other theropods) there was no reason to believe that the forelimbs of Baryonyx were able to

pronate (crossing the radius and ulna bones of the lower arm to turn the hand), and thereby make it able to rest or walk on its palms. Resting on or using the forelimbs for locomotion may have been possible (as indicated by tracks of a resting theropod), but if this was the norm, the forelimbs would probably have showed adaptations for this. Hone and Holtz furthermore suggested that the forelimbs of spinosaurids do not seem optimal for trapping prey, but instead appear similar to the forelimbs of digging animals. They suggested that the ability to dig would have been useful when excavating nests, digging for water, or to reach some kinds of prey. Hone and Holtz also believed that spinosaurids would have waded and dipped in water rather than submerging themselves, due to their sparsity of aquatic adaptations.^[11]

carcharodontosaurids

, and spinosaurids (the latter strongly associated with coastal environments), and their global occurrences through time (right)

A 2010 study by the palaeontologist Romain Amiot and colleagues proposed that spinosaurids were semi-aquatic, based on the

carcharodontosaurids, and concluded that spinosaurids had the strongest support for association with coastal palaeoenvironments. Spinosaurids also appear to have inhabited inland environments (with their distribution there being comparable to carcharodontosaurids), which indicates they may have been more generalist than usually thought.^[65]

Sales and Schultz agreed in 2017 that spinosaurids were semi-aquatic and partially piscivorous, based on skull features such as conical teeth, snouts that were compressed from side to side, and retracted nostrils. They interpreted the fact that

mechanoreception) when hunting fish. Olfaction may have been more useful for spinosaurids that also fed on terrestrial prey, such as baryonychines.^[39] A 2022 study by the palaeontologist Matteo Fabbri and colleagues revealed that Baryonyx possessed dense bones that would have allowed it to dive underwater. This same adaptation was revealed in the related Spinosaurus, and they are believed to have been subaqueous foragers that dived after aquatic prey, while Suchomimus was better adapted to a non-diving lifestyle by comparison according to the provided analysis. This discovery also showcases the unique and ecologically disparate lifestyles spinosaurids had, with more hollow-boned genera preferring to hunt in shallower water.^[66][67][68]

Palaeoenvironment

dinosaurs of the Wessex Formation

The Weald Clay Formation consists of sediments of

conifers.^[76][77]

Other dinosaurs from the Wessex Formation of the Isle of Wight where Baryonyx may have occurred include the theropods Riparovenator, Ceratosuchops,

ankylosaur Polacanthus.^[78][26] Barker and colleagues stated in 2021 that the identification of the two additional spinosaurids from the Wealden Supergroup, Riparovenator and Ceratosuchops, has implications for potential ecological separation within Spinosauridae if these and Baryonyx were contemporary and interacted. They cautioned that it is possible the Upper Weald Clay and Wessex Formations and the spinosaurids known from them were separated in time and distance.^[26]

It is generally thought that large predators occur with small taxonomic diversity in any area due to ecological demands, yet many Mesozoic assemblages include two or more

sympatric theropods that were comparable in size and morphology, and this also appears to have been the case for spinosaurids. Barker and colleagues suggested that high diversity within Spinosauridae in a given area may have been the result of environmental circumstances benefiting their niche. While it has been generally assumed that only identifiable anatomical traits related to resource partitioning allowed for coexistence of large theropods, Barker and colleagues noted that this does not preclude that similar and closely related taxa could coexist and overlap in ecological requirements. Possible niche partitioning could be in time (seasonal or daily), in space (between habitats in the same ecosystems), or depending on conditions, and they could also have been separated by their choice of habitat within their regions (which may have ranged in climate).^[26]

Taphonomy

Charig and Milner presented a possible scenario explaining the taphonomy (changes during decay and fossilisation) of the B. walkeri holotype specimen.^[3] The fine-grained sediments around the skeleton, and the fact that the bones were found close together (skull and forelimb elements at one end of the excavation area and the pelvis and hind-limb elements at the other), indicates that the environment was quiet at the time of deposition, and water currents did not carry the carcass far—possibly because the water was shallow. The area where the specimen died seems to have been suitable for a piscivorous animal. It may have caught fish and scavenged on the mud plain, becoming mired before it died and was buried. Since the bones are well-preserved and had no gnaw marks, the carcass appears to have been undisturbed by scavengers (suggesting that it was quickly covered by sediment).^[3]

The disarticulation of the bones may have been the result of soft-tissue decomposition. Parts of the skeleton seem to have weathered to different degrees, perhaps because water levels changed or the sediments shifted (exposing parts of the skeleton). The girdle and limb bones, the dentary, and a rib were broken before fossilisation, perhaps from trampling by large animals while buried. Most of the tail appears to have been lost before fossilisation, perhaps due to scavenging, or having rotted and floated off. The orientation of the bones indicates that the carcass lay on its back (perhaps tilted slightly to the left, with the right side upwards), which may explain why all the lower teeth had fallen out of their sockets and some upper teeth were still in place.^[3][2]

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External links

• Natural History Museum – "Baryonyx: the discovery of an amazing fish-eating dinosaur" – four minute video presented by Angela C. Milner

Wikimedia Commons has media related to Baryonyx.

Wikispecies has information related to Baryonyx.

This article was submitted to WikiJournal of Science for external academic peer review in 2018 (reviewer reports). The updated content was reintegrated into the Wikipedia page under a CC-BY-SA-3.0 license (2018). The version of record as reviewed is: Michael Bech; et al. (2019). "Baryonyx" (PDF). WikiJournal of Science. 2 (1): 3.

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