Baryonyx

Holotype and naming where Baryonyx was found. In January 1983, the plumber and amateur fossil collector William J. Walker explored the Smokejack Clay Pit, a clay pit in the Weald Clay Formation near Ockley in Surrey, England. He found a rock wherein he discovered a large claw, but after piecing it together at home, he realised the tip of the claw was missing. Walker returned to the same spot in the pit some weeks later, and found the missing part after searching for an hour. He also found a and part of a . Walker's son-in-law later brought the claw to the Natural History Museum of London, where it was examined by the palaeontologists Alan J. Charig and Angela C. Milner, who identified it as belonging to a theropod dinosaur. Most of the bones collected were encased in siltstone nodules surrounded by fine sand and silt, with the rest lying in clay. The bones were and scattered over a area, but most were not far from their natural positions. The position of some bones was disturbed by a bulldozer, and some were broken by mechanical equipment before they were collected.). The dinosaur had been presented earlier the same year during a lecture at a conference about dinosaur systematics in Drumheller, Canada. Due to ongoing work on the bones (70 per cent had been prepared at the time), they called their article preliminary and promised a more detailed description at a later date. Baryonyx was the first large Early Cretaceous theropod found anywhere in the world by that time. In 1997, Charig and Milner published a monograph describing the holotype skeleton in detail. Assigned specimens Fossils from other parts of the UK and Iberia, mostly isolated teeth, have subsequently been attributed to Baryonyx or similar animals. A maxilla fragment from La Rioja, Spain, was attributed to Baryonyx by the palaeontologists Luis I. Viera and José Angel Torres in 1995 (although the palaeontologist Thomas R. Holtz and colleagues raised the possibility that it could have belonged to Suchomimus in 2004). In 1999, a postorbital, , tooth, vertebral remains, (hand bones), and a phalanx from the Salas de los Infantes deposit in Burgos Province, Spain, were attributed to an immature Baryonyx (though some of these elements are unknown in the holotype) by the palaeontologist Carolina Fuentes Vidarte and colleagues. Dinosaur tracks near Burgos have also been suggested to belong to Baryonyx or a similar theropod. ''. In 2011, a specimen (catalogued as ML1190 in Museu da Lourinhã) from the Papo Seco Formation in Boca do Chapim, Portugal, with a fragmentary dentary, teeth, vertebrae, ribs, hip bones, a scapula, and a phalanx bone, was attributed to Baryonyx by the palaeontologist Octávio Mateus and colleagues, the most complete Iberian remains of the animal. The skeletal elements of this specimen are also represented in the more complete holotype (which was of similar size), except for the mid-neck vertebrae. This specimen was made the basis of the new genus Iberospinus by Mateus and Darío Estraviz-López in 2022. Multiple studies found that additional spinosaurid remains from Iberia may belong to taxa other than Baryonyx, such as Vallibonavenatrix and Protathlitis, or may be indeterminate. A 2024 article by the palaeontologist Erik Isasmendi and colleagues reviewed the spinosaurid fossil record of Iberia and concluded that no specimens from there could be assigned to Baryonyx. They moved a specimen formerly assigned to Baryonyx from La Rioja to the new genus Riojavenatrix. In 2021, the palaeontologist Chris T. Barker and colleagues described two new spinosaurid genera from the Wessex Formation of the Isle of Wight, Ceratosuchops and Riparovenator (the latter named R. milnerae honouring Milner for her contributions to spinosaurid research), and stated that spinosaurid material from there that had previously been attributed to the contemporary Baryonyx could have belonged to other taxa instead. These specimens had previously been assigned to Baryonyx in a 2017 conference abstract. Barker and colleagues stated that the recognition of the Wessex Formation specimens as new genera renders the presence of Baryonyx there ambiguous, and most of the previously assigned isolated material from the Wealden Supergroup is therefore indeterminate. Possible synonyms In 2003, Milner noted that some teeth at the Natural History Museum previously identified as belonging to the genera Suchosaurus (the first named spinosaurid) and Megalosaurus probably belonged to Baryonyx. The type species of Suchosaurus, S. cultridens, was named by the biologist Richard Owen in 1841, based on teeth discovered by the geologist Gideon A. Mantell in Tilgate Forest, Sussex. Owen originally thought the teeth to have belonged to a crocodile; he was yet to name the group Dinosauria, which happened the following year. A second species, S. girardi, was named by the palaeontologist Henri Émile Sauvage in 1897, based on jaw fragments and a tooth from Boca do Chapim, Portugal. In 2007, the palaeontologist Éric Buffetaut considered the teeth of S. girardi very similar to those of Baryonyx (and S. cultridens) except for the stronger development of the flutes (or "ribs"; lengthwise ridges), suggesting that the remains belonged to the same genus. Buffetaut agreed with Milner that the teeth of S. cultridens were almost identical to those of B. walkeri, but with a ribbier surface. The former taxon might be a senior synonym of the latter (since it was published first), depending on whether the differences were within a taxon or between different ones. According to Buffetaut, since the holotype specimen of S. cultridens is a single tooth and that of B. walkeri is a skeleton, it would be more practical to retain the newer name. In 2011, Mateus and colleagues agreed that Suchosaurus was closely related to Baryonyx, but considered both species in the former genus nomina dubia (dubious names) since their holotype specimens were not considered diagnostic (lacking distinguishing features) and could not be definitely equated with other taxa. Barker and colleagues agreed with this in 2023. Milner concurred that the material from Niger was indistinguishable from Baryonyx in 2003. Later studies have kept Baryonyx and Suchomimus separate, whereas Cristatusaurus has been proposed to be either a nomen dubium or possibly distinct from both. Barker and colleagues found Suchomimus to be closer related to the British genera Riparovenator and Ceratosuchops than to Baryonyx in 2021. ==Description==

s (Baryonyx in yellow) compared with a human Baryonyx is estimated to have been between long, in hip height, and to have weighed between . The fact that elements of the skull and vertebral column of the B. walkeri holotype specimen (NHM R9951) do not appear to have co-ossified (fused) suggests that the individual was not fully grown, and the mature animal may have been much larger (as is the case for some other spinosaurids). On the other hand, the specimen's fused sternum indicates that it may have been mature. The cervical vertebrae of the neck tapered towards the head and became progressively longer front to back. Their (the processes that connected the vertebrae) were flat, and their (processes to which neck muscles attached) were well developed. The (the second neck vertebra) was small relative to the size of the skull and had a well-developed . The of the cervical vertebrae were not always sutured to the (the bodies of the vertebrae), and the there were low and thin. The 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 (openings) in the neural arches and (hollow depressions) in the centra (primarily near the ). 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 or front of the tail. , Paris 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 and muscle attachment and the lower for articulation with the radius and ulna. The radius was short, stout and straight, and less than half the length of the humerus, while the ulna was a little longer. The ulna had a powerful and an expanded lower end. The hands had three fingers; the first finger bore a large claw measuring about along its curve in the holotype specimen. The claw would have been lengthened by a keratin (horny) sheath in life. Apart from its size, the claw's proportions were fairly typical of a theropod, i.e. it was bilaterally symmetric, slightly compressed, smoothly rounded, and sharply pointed. A groove for the sheath ran along the length of the claw. The other claws of the hand were much smaller. The (main hip bone) of the pelvis had a prominent , an anterior process that was slender and vertically expanded, and a posterior process that was long and straight. The ilium also had a prominent and a deep grove that faced downwards. The (the socket for the femur) was long from front to back. The (lower and rearmost hip bone) had a well developed at the upper part. The margin of the blade at the lower end was turned outward, and the pubic foot was not expanded. The femur lacked a groove on the fibular condyle, and, uniquely among spinosaurids, the fibula had a very shallow fibular (depression). ==Classification==

In their original description, Charig and Milner Buffetaut also supported this relationship in 1989. In 1990, Charig and Milner dismissed the spinosaurid affinities of Baryonyx, since they did not find their remains similar enough. In 1997, they agreed that Baryonychidae and Spinosauridae were related, but disagreed that the former name should become a synonym of the latter, because the completeness of Baryonyx compared to Spinosaurus made it a better type genus for a family, and because they did not find the similarities between the two significant enough. They also united the spinosaurids and their closest relatives in the superfamily Spinosauroidea, but in 2010, the palaeontologist Roger Benson considered this a junior synonym of Megalosauroidea (an older name). In a 2007 conference abstract, the palaeontologist Denver W. Fowler suggested that since Suchosaurus is the first named genus in its group, the clade names Spinosauroidea, Spinosauridae, and Baryonychinae should be replaced by Suchosauroidea, Suchosauridae, and Suchosaurinae, regardless of whether or not the name Baryonyx is retained. A 2017 study by the palaeontologists Marcos A. F. Sales and Cesar L. Schultz found that the clade Baryonychinae was not well supported, since serrated teeth may be an ancestral trait among spinosaurids. (C, D), and Baryonyx'' (E) 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: They shared features such as long, narrow, crocodile-like skulls; sub-circular teeth, with fine to no serrations; the terminal rosette of the snout; and a secondary palate that made them more resistant to torsion. In contrast, the primitive and typical condition for theropods was a tall, narrow snout with blade-like (ziphodont) teeth with serrated carinae. Sereno and colleagues proposed in 1998 that the large thumb-claw and robust forelimbs of spinosaurids evolved in the Middle Jurassic, before the elongation of the skull and other adaptations related to fish-eating, since the former features are shared with their megalosaurid relatives. They also suggested that the spinosaurines and baryonychines diverged before the Barremian age of the Early Cretaceous. Milner suggested in 2003 that spinosaurids originated in Laurasia during the Jurassic, and dispersed via the Iberian land bridge into Gondwana, where they radiated. In 2007, Buffetaut pointed out that palaeogeographical studies had demonstrated that Iberia was near northern Africa during the Early Cretaceous, which he found to confirm Milner's idea that the Iberian region was a 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, and subsequent discoveries of spinosaurid remains in Asia and possibly Australia indicate that it may have been complex. 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. 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. 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. ==Palaeobiology==

Diet and feeding In 1986, Charig and Milner suggested that its elongated snout with many finely serrated teeth indicated that Baryonyx was piscivorous (fish-eating), speculating that it crouched on a riverbank and used its claw to gaff fish out of the water (similar to the modern grizzly bear). pointed out that the spinosaurid snouts from Niger were similar to those of the modern gharial and suggested a behaviour similar to herons or storks. 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). 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 Scheenstia mantelli (then classified in the genus Lepidotes), and abraded or etched bones of a young iguanodontid. They also presented circumstantial evidence for piscivory, such as crocodile-like adaptations for catching and swallowing prey: long, narrow jaws with their "terminal rosette", similar to those of a gharial, and the downturned tip and notch of the snout. In their view, these adaptations suggested that Baryonyx would have caught small to medium-sized fish in the manner of a crocodilian: gripping them with the notch of the snout (giving the teeth a "stabbing function"), tilting the head backwards, and swallowing them headfirst. 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. A 2005 beam-theory study by the palaeontologist François Therrien and colleagues was unable to reconstruct force profiles of Baryonyx, but found that the related Suchomimus would have used the front part of its jaws to capture prey, and suggested that the jaws of spinosaurids were adapted for hunting smaller terrestrial prey in addition to fish. They envisaged that spinosaurids could have captured smaller prey with the rosette of teeth at the front of the jaws, and finished it by shaking it. Larger prey would instead have been captured and killed with their forelimbs instead of their bite, since their skulls would not be able to resist the bending stress. They also agreed that the conical teeth of spinosaurids were well-developed for impaling and holding prey, with their shape enabling them to withstand bending loads from all directions. A 2007 finite element analysis of 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. 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. 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 binocular vision. Instead, they suggested the jaws would have made sideways sweeps to catch fish, like the gharial, with the hand claws probably used to stamp down and impale large fish, whereafter they manipulated them with their jaws, in a manner similar to grizzly bears and 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). 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 pharynx (opening that connects the mouth to the oesophagus) to be widened. This jaw-articulation is similar to that seen in pterosaurs and living pelicans, and would likewise have allowed spinosaurids to swallow large prey such as fish and other animals. They also reported that some possible Portuguese Baryonyx fossils were found associated with isolated Iguanodon teeth, and listed it along with other such associations as support for opportunistic feeding behaviour in spinosaurs. 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-maniraptoriform theropods. Their neurosensory capabilities such as hearing and olfaction (sense of smell) were unexceptional, and their gaze stabilisation less developed than those of spinosaurines, so their behavioural adaptations were probably comparable to those of other large-bodied terrestrial theropods. This suggests that their transition from terrestrial hypercarnivores to semi-aquatic "generalists" during their evolution did not require substantial modification of their brain and sensory systems. This could mean that spinosaurids were either pre-adapted for detection and capture of aquatic prey, or that their transition to semi-aquatic lifestyles only required modifications to the bones associated with the mouth. Their reptile encephalization quotient values imply that the cognitive capacity and behavioural sophistication of baryonychines did not deviate much from that of other basal theropods. 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. In 2016, Sales and colleagues statistically examined the fossil distribution of spinosaurids, abelisaurids, and 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. 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 histological data indicates some spinosaurids were more terrestrial than others as reflecting ecological niche partitioning among them. As some spinosaurids have smaller nostrils than others, their olfactory abilities were presumably lesser, as in modern piscivorous animals, and they may instead have used other senses (such as vision and mechanoreception) when hunting fish. Olfaction may have been more useful for spinosaurids that also fed on terrestrial prey, such as baryonychines. ==Palaeoenvironment==

of the Wessex Formation The Weald Clay Formation consists of sediments of Hauterivian (Lower Weald Clay) to Barremian (Upper Weald Clay) age, about 130–125 million years old. The original Baryonyx specimen was found in the latter, in clay representing non-marine still water, which has been interpreted as a fluvial or mudflat environment with shallow water, lagoons, and marshes. Dinosaurs from the locality include the ornithopods Mantellisaurus, Iguanodon, and small sauropods. Other vertebrates from the Weald Clay include crocodiles, pterosaurs, lizards (such as Dorsetisaurus), amphibians, sharks (such as Hybodus), and bony fishes (including Scheenstia). Members of ten orders of insects have been identified, including Valditermes, Archisphex, and Pterinoblattina. Other invertebrates include ostracods, isopods, conchostracans, and bivalves. The plants Weichselia and the aquatic, herbaceous Bevhalstia were common. Other plants found include ferns, horsetails, club mosses, and conifers. Other dinosaurs from the Wessex Formation of the Isle of Wight where Baryonyx may have occurred include the theropods Riparovenator, Ceratosuchops, Neovenator, Eotyrannus, Aristosuchus, Thecocoelurus, Calamospondylus, and Ornithodesmus; the ornithopods Iguanodon, Hypsilophodon, and Valdosaurus; the sauropods Ornithopsis, Eucamerotus, and Chondrosteosaurus; and the ankylosaur Polacanthus. 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. (front) and Ceratosuchops (rear), spinosaurids from the Wessex Formation which may have lived alongside Baryonyx'' 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). Taphonomy Charig and Milner presented a possible scenario explaining the taphonomy (changes during decay and fossilisation) of the B. walkeri holotype specimen. 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). 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. ==References==