Troodon

Troodon (or Troödon in older sources) is a genus of relatively small, bird-like dinosaur from the Late Cretaceous period (75-65 mya). Discovered in 1855, it was among the first dinosaurs found in North America.

The name Troodon is Greek for "wounding tooth", referring to the dinosaur's teeth, which are different from those of most other theropods. The teeth bear prominent, apically oriented serrations. These "wounding" serrations, however, are morphometrically more similar to those of herbivorous reptiles, and suggest a possibly omnivorous diet.

Description
Troodon were small dinosaurs, up to 2.4 metres (7.9 ft) in length,[5] and up to 50 kilograms (110 lb) in mass.[6] The largest specimens are comparable in size to Deinonychus and Unenlagia.[7] They had very long, slender hind limbs, suggesting that these animals were able to run quickly. They had large, retractable sickle-shaped claws on the second toes, which were raised off of the ground when running.

Their eyes were very large (perhaps suggesting a partially nocturnal lifestyle), and slightly forward facing, giving Troodon some degree of depth perception. Their light skulls contained a capsule similar to those found in ostrich dinosaurs.

Brain and inner ear
Troodon had some of the largest known brains of any dinosaur group, relative to their body mass (comparable to modern birds).[9] Troodon's cerebrum-to-brain-volume ratio was 31.5% to 63% of the way from a nonavian reptile proportion to a truly avian one.[10] Troodon had bony crista supporting their tympanic membranes that were ossified at least in their top and bottom regions. The rest of the crista were either cartilaginous or too delicate to be preserved. The metotic strut of Troodon was enlarged from side-to-side, similar to Dromaeosaurus and primitive birds like Archaeopteryx and Hesperornis.

History and classification
The name was originally spelled Troödon (with a diaeresis) by Joseph Leidy in 1856, which was officially amended to its current status by Sauvage in 1876. The type specimen of Troodon has caused problems with classification, as the entire genus is based only on a single tooth from the Judith River Formation. Troodon has historically been a highly unstable classification and has been the subject of numerous conflicting synonymies with similar theropod specimens.

The Troodon tooth was originally classified as a "lacertilian" (lizard) by Leidy, but re-assigned as a megalosaurid dinosaur by Nopcsa in 1901 (Megalosauridae having historically been a wastebin taxon for most carnivorous dinosaurs). In 1924, Gilmore suggested that the tooth belonged to the herbivorous pachycephalosaur Stegoceras, and that Stegoceras was in fact a junior synonym of Troodon (the similarity of troodontid teeth to those of herbivorous dinosaurs continues to lead many paleontologists to believe that these animals were omnivores). The classification of Troodon as a pachycephalosaur was followed for many years, during which the family Pachycephalosauridae was known as Troodontidae. In 1945, Charles Mortram Sternberg rejected the possibility that Troodon was a pachycephalosaur due to its stronger similarity to the teeth of other carnivorous dinosaurs. With Troodon now classified as a theropod, the family Troodontidae could no longer be used for the dome-headed dinosaurs, so Sternberg named a new family for them, Pachycephalosauridae.[12]

The first specimen currently assigned to Troodon that was not a tooth was named Stenonychosaurus by Sternberg in 1932, based on a foot, fragments of a hand, and some tail vertebrae from the Dinosaur Park Formation of Alberta. A remarkable feature of these remains was the enlarged claw on the second toe, which is now recognized as characteristic of early paravians. Sternberg initially classified Stenonychosaurus as a member of the family Coeluridae. Later, in 1951, Sternberg speculated that since Stenonychosaurus had a "very peculiar pes" and Troodon "equally unusual teeth", they may be closely related. Unfortunately, no comparable specimens were available at that time to test the idea.

A more complete skeleton of Stenonychosaurus was described by Dale Russell in 1969 from the Dinosaur Park Formation, which eventually formed the scientific foundation for a famous life-sized sculpture of Stenonychosaurus accompanied by its fictional, humanoid descendant, the "dinosauroid".[13] Stenonychosaurus became a well-known theropod in the 1980s, when the feet and braincase were described in more detail. Along with Saurornithoides, it formed the family Saurornithoididae. Based on differences in tooth structure, and the extremely fragmentary nature of the original Troodon formosus specimens, saurornithoidids were thought to be close relatives while Troodon was considered a dubious possible relative of the family. Phil Currie, reviewing the pertinant specimens in 1987, showed that supposed differences in tooth and jaw structure among troodontids and saurornithoidids were based on age and position of the tooth in the jaw, rather than a difference in species. He reclassified Stenonychosaurus inequalis as well as Polyodontosaurus grandis and Pectinodon bakkeri as junior synonyms of Troodon formosus. Currie also made Saurornithoididae a junior synonym of Troodontidae.[14] In 1988, Gregory S. Paul went farther and included Saurornithoides mongoliensis in the genus Troodon as T. mongoliensis,[15] but this reclassification, along with many other unilateral synonymizations of well known genera, was not adopted by other researchers. Currie's classification of all North American troodontid material in the single species Troodon formosus became widely adopted by other paleontologists, and all of the specimens once called Stenonychosaurus were referred to as Troodon in the scientific literature through the early 21st century.

However, the concept that all Late Cretaceous North American troodontids belong to one species began to be questioned soon after Currie's 1987 paper was published, including by Currie himself. Currie and colleagues (1990) noted that, while they believed the Judith River troodontids were all T. formosus, troodontid fossils from other formations, such as the Hell Creek Formation and Lance Formation, might belong to different species. In 1991, George Olshevsky assigned the Lance formation fossils, which had first been named Pectinodon bakkeri but later synonymized with Troodon formosus to the species Troodon bakkeri, and several other researchers (including Currie) have reverted to keeping the Dinosaur Park Formation fossils separate as Troodon inequalis (formerly Stenonychosaurus inequalis).[16]

In 2011, Zanno and colleagues reviewed the convoluted history of troodontid classification in Late Cretaceous North America. They followed Longrich (2008) in treating Pectinodon bakkeri as a valid genus, and noted that it is likely the numerous Late Cretaceous specimens currently assigned to Troodon formosus almost certainly represent numerous new species, but that a more thorough review of the specimens is required. Because the holotype of T. formosus is a single tooth, this may render Troodon a nomen dubium.

Taxonomy
Troodon is considered to be one of the most derived members of its family. Along with Zanabazar, Saurornithoides and Talos it forms a clade of specialized troodontids.

Below is a cladogram of Troodontidae by Zanno et al.