Baryonyx

***Baryonyx*** (meaning "heavy claw") is a species of large, carnivorous dinosaur discovered in clay pits just south of Dorking, England.

The species only known from a single; subadult; specimen, and because remains of its last meal were discovered fossilized in its ribcage; both remains of fish and an Iguanodon; it gave true clear indication of the spinosaurids diet as Generalist carnivores similiar to crocodilian species and large avians; different from previously thought as solely piscivores. It lived in the Barremian period of Early Cretaceous, around 125 million years ago.

## Description

*Baryonyx* is estimated to have been between 7.5 and 10 m (25 and 33 ft) long, 2.5 m (8.2 ft) in hip height, and to have weighed between 1.2 and 1.9 t (1.3 and 2.1 short tons; 1.2 and 1.9 long tons). 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.

### Skull

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). It was elongated, and the front 17 cm (6.7 in) of the premaxillae formed a long, narrow, and low snout (rostrum) with a smoothly rounded upper surface. The external nares (bony nostrils) were long, low, and placed far back from the snout tip. The front 13 cm (5.1 in) of the snout expanded into a spatulate (spoon-like), "terminal rosette", a shape similar to the rostrum of the modern gharial. The front 7 cm (2.8 in) of the lower margin of the premaxillae was downturned (or hooked), whereas that of the front portion of the maxillae was upturned. This morphology resulted in a sigmoid or S shaped margin of the lower upper tooth row, in which the teeth from the front of the maxilla were projecting forward. The snout was particularly narrow directly behind the rosette; this area received the large teeth of the mandible. The maxilla and premaxilla of *Baryonyx* fit together in a complex articulation, and the resulting gap between the upper and lower jaw is known as the subrostral notch. A downturned premaxilla and a sigmoid lower margin of the upper tooth row was also present in distantly related theropods such as *Dilophosaurus*. The snout had extensive foramina (openings), which would have been exits for blood vessels and nerves, and the maxilla appears to have housed sinuses.

*Baryonyx* had a rudimentary secondary palate, similar to crocodiles but unlike most theropod dinosaurs. 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). 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*. 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.

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. 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). 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. 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. 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.

### Postcranial Skeleton

Initially thought to have lacked the sigmoid curve typical of theropods, the neck of *Baryonyx* does appear to have formed an S shape, though straighter than in other theropods. 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.

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 deltopectoral crest 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 olecranon and an expanded lower end. The hands had three fingers; the first finger bore a large claw measuring about 31 cm (12 in) 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 ilium (main hip bone) of the pelvis had a prominent supracetabular crest, an anterior process that was slender and vertically expanded, and a posterior process that was long and straight. The ilium also had a prominent brevis shelf and a deep grove that faced downwards. The acetabulum (the socket for the femur) was long from front to back. The ischium (lower and rearmost hip bone) had a well developed obturator process at the upper part. The margin of the pubic 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 fossa (depression).