Deinosuchus

Life restoration of D. rugosus

Life restoration of D. hatcheri

Despite its large size, the overall appearance of Deinosuchus was not considerably different from that of modern crocodilians. Deinosuchus had an alligator-like, broad snout, with a slightly bulbous tip.[2] Each premaxilla contained four teeth, with the pair nearest to the tip of the snout being significantly smaller than the other two.[3] Each maxilla (the main tooth-bearing bone in the upper jaw) contained 21 or 22 teeth.[4] The tooth count for each dentary (tooth-bearing bone in the lower jaw) was at least 22.[3] All the teeth were very thick and robust; those close to the rear of the jaws were short, rounded, and blunt. They appear to have been adapted for crushing, rather than piercing.[5] When the mouth was closed, only the fourth tooth of the lower jaw would have been visible.[3] The skull of Deinosuchus itself was of a unique shape not seen in any other living or extinct crocodilians; the skull was broad, but inflated at the front around the nares. Two holes in the premaxilla in front of the nares are present in this genus and are unique autapomorphies not seen in other crocodilians, but nothing is known at present regarding their function.[6][7]

Modern saltwater crocodiles (Crocodylus porosus) have the strongest recorded bite of any living animal, with a maximum force of 16,414 N (1,673.8 kgf; 3,690 lbf) for a 4.59 meters (15.1 ft), 531 kilograms (1,171 lb) specimen.[8] The bite force of Deinosuchus has been estimated to be 18,000 N (1,835 kgf; 4,047 lbf)[2] to 102,803 N (10,483 kgf; 23,111 lbf).[8]

Deinosuchus had a secondary bony palate, which would have permitted it to breathe through its nostrils while the rest of the head remained submerged underwater.[9] The vertebrae were articulated in a procoelous manner, meaning they had a concave hollow on the front end and a convex bulge on the rear; these would have fit together to produce a ball and socket joint.[10][11] The secondary palate and procoelous vertebrae are advanced features also found in modern eusuchian crocodilians.[9][12]

The osteoderms (scutes) covering the back of Deinosuchus were unusually large, heavy, and deeply pitted; some were of a roughly hemispherical shape.[13][14] Deep pits and grooves on these osteoderms served as attachment points for connective tissue.[14] Together, the osteoderms and connective tissue would have served as load-bearing reinforcement to support the massive body of Deinosuchus out of water.[4][14] These deeply pitted osteoderms have been used to suggest that, despite its bulk, Deinosuchus could probably have walked on land much like modern-day crocodiles.[4][9]

D. riograndensis compared to other large crocodyliforms

The large size of Deinosuchus has generally been recognized despite the fragmentary nature of the fossils assigned to it. However, estimates of how large it really was have varied considerably over the years. The original estimate from 1954 for the type specimen of the then-named "Phobosuchus riograndensis" were based on a skull of 1.5 meters (4.9 ft) and a lower jaw of 1.8 meters (5.9 ft) long, reconstructed with similar proportions to the Cuban crocodile giving a total estimated length of 15 meters (49 ft).[10] However, this reconstruction is currently considered to be inaccurate.[4] Using more complete remains, it was estimated in 1999 that the size attained by specimens of Deinosuchus varied from 8 to 10 meters (26 to 33 ft) with weights from 2.5 to 5 metric tons (2.8 to 5.5 short tons).[15] This was later corroborated when it was noted that most known specimens of D. rugosus usually had skulls of about 1 meter (3.3 ft) with estimated total lengths of 8 meters (26 ft) and weights of 2.3 metric tons (2.5 short tons). A reasonably well-preserved skull specimen discovered in Texas indicated the animal's head measured about 1.31 meters (4.3 ft), and its body length was estimated at 9.8 meters (32 ft). However, the largest fragmentary remains of D. riograndensis were 1.5 times the size of those of the average D. rugosus and it was determined that the largest individuals of this species may have been up to 12 meters (39 ft) in length and perhaps weighed as much as 8.5 metric tons (9.4 short tons).[4]

A particularly large mandibular fragment from a D. riograndensis specimen was estimated to have come from an individual with a skull length of 147.5 centimeters (4.84 ft). This length was used in conjunction with a regression equation relating skull length to total length in the American alligator to estimate a total length of 10.6 meters (35 ft) for this particular specimen.[16] This is only slightly lower than previous estimates for the species. Deinosuchus has often been described as the largest crocodyliform of all time. However, other crocodyliforms such as Purussaurus, Gryposuchus, Rhamphosuchus, Euthecodon, and Sarcosuchus may have equaled or exceeded it in size.[17]

A Deinosuchus jaw fragment, exhibited at the North Carolina Museum of Natural Sciences: Fossils of this large alligatoroid have been discovered in 10 U.S. states and northern Mexico.

Deinosuchus was present on both sides of the Western Interior Seaway.[18] Specimens have been described from 10 U.S. states: Utah, Montana, Wyoming, New Mexico, New Jersey, Georgia, Alabama, Mississippi, Texas, and North Carolina.[19] David Schwimmer has said Deinosuchus fossils have been found in South Carolina and Delaware as well, but none of them from those two states have been formally described.[18] A Deinosuchus osteoderm from the San Carlos Formation was also reported in 2006, so the giant crocodilian's range may have included parts of northern Mexico.[20] There is also a report describing a possible Deinosuchus scute from Colorado.[21] Deinosuchus fossils are most abundant in the Gulf Coastal Plain region of Georgia, near the Alabama border.[18] All known specimens of Deinosuchus were found in rocks dated to the Campanian stage of the Late Cretaceous period. The oldest examples of this genus lived approximately 82 Ma, and the youngest lived around 73 Ma.[22]

The distribution of Deinosuchus specimens indicates these giant crocodilians may have preferred estuarine environments.[18] In the Aguja Formation of Texas, where some of the largest specimens of Deinosuchus have been found, these massive predators probably inhabited brackish-water bays.[23] Although some specimens have also been found in marine deposits, it is not clear whether Deinosuchus ventured out into the ocean (like modern-day saltwater crocodiles); these remains might have been displaced after the animals died.[18] Deinosuchus has been described as a "conspicuous" component of a purportedly distinct biome occupying the southern half of Late Cretaceous North America.[24]

Deinosuchus may have preyed upon large ornithopods. Kritosaurus lived alongside the giant crocodilian in the Aguja Formation ecosystem.[25]

In 1954, Edwin H. Colbert and Roland T. Bird speculated that Deinosuchus "may very well have hunted and devoured some of the dinosaurs with which it was contemporaneous".[10] Colbert restated this hypothesis more confidently in 1961: "Certainly this crocodile must have been a predator of dinosaurs; otherwise why would it have been so overwhelmingly gigantic? It hunted in the water where the giant theropods could not go."[26][27] David R. Schwimmer proposed in 2002 that several hadrosaurid tail vertebrae found near Big Bend National Park show evidence of Deinosuchus tooth marks, strengthening the hypothesis that Deinosuchus fed on dinosaurs in at least some instances.[5] In 2003, Christopher A. Brochu agreed that Deinosuchus "probably dined on ornithopods from time to time."[28] Deinosuchus is generally thought to have employed hunting tactics similar to those of modern crocodilians, ambushing dinosaurs and other terrestrial animals at the water's edge and then submerging them until they drowned.[29] A 2014 study suggested that it would have been able to perform a "death roll", like modern crocodiles.[30]

Reconstructed skull of D. riograndensis, Museo de la Evolución de Puebla

Schwimmer and G. Dent Williams proposed in 1996 that Deinosuchus may have preyed on marine turtles.[31] Deinosuchus would probably have used the robust, flat teeth near the back of its jaws to crush the turtle shells.[5] The "side-necked" sea turtle Bothremys was especially common in the eastern habitat of Deinosuchus, and several of its shells have been found with bite marks that were most likely inflicted by the giant crocodilian.[5][31]

Schwimmer concluded in 2002 that the feeding patterns of Deinosuchus most likely varied by geographic location; the smaller Deinosuchus specimens of eastern North America would have been opportunistic feeders in an ecological niche similar to that of the modern American alligator. They would have consumed marine turtles, large fish, and smaller dinosaurs.[4] The bigger, but less common, Deinosuchus that lived in Texas and Montana might have been more specialized hunters, capturing and eating large dinosaurs.[4] Schwimmer noted no theropod dinosaurs in Deinosuchus's eastern range approached its size, indicating the massive crocodilian could have been the region's apex predator.[5]

The osteoderms of Deinosuchus, as illustrated by W.J. Holland. They are proportionately much thicker than those of modern crocodilians.

A 1999 study by Gregory M. Erickson and Christopher A. Brochu suggested the growth rate of Deinosuchus was comparable to that of modern crocodilians, but was maintained over a far longer time. Their estimates, based on growth rings in the dorsal osteoderms of various specimens, indicated each Deinosuchus might have taken over 35 years to reach full adult size, and the oldest individuals may have lived for more than 50 years. This was a completely different growth strategy than that of large dinosaurs, which reached adult size much more quickly and had shorter lifespans. According to Erickson, a full-grown Deinosuchus "must have seen several generations of dinosaurs come and go".[32]

Schwimmer noted in 2002 that Erickson and Brochu's assumptions about growth rates are only valid if the osteodermal rings reflect annual periods, as they do in modern crocodilians. According to Schwimmer, the growth ring patterns observed could have been affected by a variety of factors, including "migrations of their prey, wet-dry seasonal climate variations, or oceanic circulation and nutrient cycles". If the ring cycle were biannual rather than annual, this might indicate Deinosuchus grew faster than modern crocodilians, and had a similar maximum lifespan.[4]