Hadrosauridae

Hadrosauridae was first defined as a clade, by Forster, in a 1997 abstract, as simply "Lambeosaurinae plus Hadrosaurinae and their most recent common ancestor". In 1998, Paul Sereno defined the clade Hadrosauridae as the most inclusive possible group containing Saurolophus (a well-known saurolophine) and Parasaurolophus (a well-known lambeosaurine), later emending the definition to include Hadrosaurus, the type genus of the family, which ICZN rules state must be included, despite its status as a nomen dubium. According to Horner et al. (2004), Sereno's definition would place a few other well-known hadrosaurs (such as Telmatosaurus and Bactrosaurus) outside the family, which led them to define the family to include Telmatosaurus by default. Prieto-Marquez reviewed the phylogeny of Hadrosauridae in 2010, including many taxa potentially within the family.[7]

Premaxilla of Eotrachodon, the taxon named by Prieto-Marquez et al. 2016

Below is a cladogram from Prieto-Marquez et al. 2016. This cladogram is a recent modification of the original 2010 analysis, including more characters and taxa. The resulting cladistic tree of their analysis was resolved using Maximum-Parsimony. 61 hadrosauroid species were included, characterized for 273 morphological features: 189 for cranial features and 84 for postcranial features. When characters had multiple states that formed an evolutionary scheme, they were ordered to account for the evolution of one state into the next. The final tree was run through TNT version 1.0.[10]

Skull of Lambeosaurus, the type taxon of Lambeosaurinae

Skull of Saurolophus, the type taxon of Saurolophinae

Early restoration by Charles R. Knight of hadrosaurs as semi-aquatic animals that could only chew soft water plants, a popular idea at the time.

While studying the chewing methods of hadrosaurids in 2009, the paleontologists Vincent Williams, Paul Barrett, and Mark Purnell found that hadrosaurs likely grazed on horsetails and vegetation close to the ground, rather than browsing higher-growing leaves and twigs. This conclusion was based on the evenness of scratches on hadrosaur teeth, which suggested the hadrosaur used the same series of jaw motions over and over again.[13] As a result, the study determined that the hadrosaur diet was probably made of leaves and lacked the bulkier items, such as twigs or stems, that might have required a different chewing method and created different wear patterns.[14] However, Purnell said these conclusions were less secure than the more conclusive evidence regarding the motion of teeth while chewing.[15]

The hypothesis that hadrosaurs were likely grazers rather than browsers appears to contradict previous findings from preserved stomach contents found in the fossilized guts in previous hadrosaur studies.[15] The most recent such finding before the publication of the Purnell study was conducted in 2008, when a team led by University of Colorado at Boulder graduate student Justin S. Tweet found a homogeneous accumulation of millimeter-scale leaf fragments in the gut region of a well-preserved partially grown Brachylophosaurus.[16][17] As a result of that finding, Tweet concluded in September 2008 that the animal was likely a browser, not a grazer.[17] In response to such findings, Purnell said that preserved stomach contents are questionable because they do not necessarily represent the usual diet of the animal. The issue remains a subject of debate.[18]

Edmontosaurus dentary with teeth, typical of hadrosauridae

Mallon et al. (2013) examined herbivore coexistence on the island continent of Laramidia, during the Late Cretaceous. It was concluded that hadrosaurids could reach low-growing trees and shrubs that were out of the reach of ceratopsids, ankylosaurs, and other small herbivores. Hadrosaurids were capable of feeding up to 2 m when standing quadrupedally, and up to 5 m bipedally.[19]

Coprolites (fossilized droppings) of some Late Cretaceous hadrosaurs show that the animals sometimes deliberately ate rotting wood. Wood itself is not nutritious, but decomposing wood would have contained fungi, decomposed wood material and detritus-eating invertebrates, all of which would have been nutritious.[20] Examination of hadrosaur coprolites from the Grand Staircase-Escalante indicates that shellfish such as crustaceans were also an important component of the hadrosaur diet.[21]

Neonate sized hadrosaur fossils have been documented in the scientific literature.[22] Tiny hadrosaur footprints have been discovered in the Blackhawk Formation of Utah.[22]

The head of Gryposaurus notabilis.

In a 2001 review of hadrosaur eggshell and hatchling material from Alberta's Dinosaur Park Formation, Darren Tanke and M. K. Brett-Surman concluded that hadrosaurs nested in both the ancient upland and lowlands of the formation's depositional environment. The upland nesting grounds may have been preferred by the less common hadrosaurs, like Brachylophosaurus and Parasaurolophus. However, the authors were unable to determine what specific factors shaped nesting ground choice in the formation's hadrosaurs. They suggested that behavior, diet, soil condition, and competition between dinosaur species all potentially influenced where hadrosaurs nested.[22]

Sub-centimeter fragments of pebbly-textured hadrosaur eggshell have been reported from the Dinosaur Park Formation. This eggshell is similar to the hadrosaur eggshell of Devil's Coulee in southern Alberta as well as that of the Two Medicine and Judith River Formations in Montana, United States. While present, dinosaur eggshell is very rare in the Dinosaur Park Formation and is only found in two different microfossil sites. These sites are distinguished by large numbers of pisidiid clams and other less common shelled invertebrates, like unionid clams and snails. This association is not a coincidence, as the invertebrate shells would have slowly dissolved and released enough basic calcium carbonate to protect the eggshells from naturally occurring acids that otherwise would have dissolved them and prevented fossilization.[22]

In contrast with eggshell fossils, the remains of very young hadrosaurs are somewhat common. Tanke has observed that an experienced collector could discover multiple juvenile hadrosaur specimens in a single day. The most common remains of young hadrosaurs in the Dinosaur Park Formation are dentaries, bones from limbs and feet, as well as vertebral centra. The material showed little or none of the abrasion that would have resulted from transport, meaning the fossils were buried near their point of origin. Bonebeds 23, 28, 47, and 50 are productive sources of young hadrosaur remains in the formation, especially bonebed 50. The bones of juvenile hadrosaurs and fossil eggshell fragments are not known to have been preserved in association with each other, despite both being present in the formation.[22]

The limbs of the juvenile hadrosaurs are anatomically and proportionally similar to those of adult animals.[22] However, the joints often show "predepositional erosion or concave articular surfaces",[22] which was probably due to the cartilaginous cap covering the ends of the bones.[22] The pelvis of a young hadrosaur was similar to that of an older individual.[22]

Evidence suggests that young hadrosaurs would have walked on only their two hind legs, while adults would have walked on all four.[2] As the animal aged, the front limbs became more robust in order to take on weight, while the back legs became less robust as they transitioned to walking on all four legs.[2] Furthermore, the animals’ front limbs were shorter than their back limbs.[2]

Comparisons between the scleral rings of several hadrosaur genera (Corythosaurus, Prosaurolophus, and Saurolophus) and modern birds and reptiles suggest that they may have been cathemeral, active throughout the day at short intervals.[23]

Spondyloarthropathy has been documented in the spine of a 78-million year old hadrosaurid. Other examples of pathologies in hadrosaurs include healed wounds from predators, such as those found in Edmontosaurus annectens, and tumors such as hemangiomas, desmoplastic fibroma, metastatic cancer, and osteoblastomas, found in genera such as Brachylophosaurus and Edmontosaurus.[24][25] Osteochondrosis is also commonly found in hadrosaurs.[26]

Tiny hadrosaur footprints have been discovered in the Blackhawk Formation of Utah.[22]