Psittacosaurus

*Psittacosaurus mongolensis*

* Timothy Bradley Artwork
* Robert F Walters Artwork

***Psittacosaurus*** (/ˌsɪtəkəˈsɔrəs/ sit-ə-kə-sor-əs; "parrot lizard") is a genus of extinct ceratopsian dinosaur from the Early Cretaceous of what is now Asia, existing between 126 and 101 million years ago. It is notable for being the most species-rich dinosaur genus. Up to 11 species are known, from across Mongolia, Siberia, China, and possibly Thailand. The species of *Psittacosaurus* were obligate bipeds at adulthood, with a high skull and a robust beak. One individual was found preserved with long quills on the tail, similar to those of *Tianyulong*, yet scales of varying sizes and shapes across the rest of the animal. *Psittacosaurus* probably had complex behaviours, based on the proportions and relative size of the brain. It may have been active for short periods of time during the day and night, and had good smell and well developed vision. It was up to 2.5 meters (8 feet) long and weighed up to 50 kilograms (110 lbs).

*Psittacosaurus* is one of the most completely known dinosaur genera. Fossils of over 400 individuals have been collected so far, including many complete skeletons. Most different age classes are represented, from hatchling through to adult, which has allowed several detailed studies of *Psittacosaurus* growth rates and reproductive biology. The abundance of this dinosaur in the fossil record has led to the labeling of Lower Cretaceous sediments of east Asia the *Psittacosaurus biochron*.

Studies is 2016 show that *P. sp.* had countershading, with brown on the top half of its body, and orange on the bottom, as seen in the picture above. This is likely present to help the animal camouflage itself in its forest habitat.

## Description

The species of *Psittacosaurus* vary in size and specific features of the skull and skeleton, but share the same overall body shape. The best-known—*P. mongoliensis*—can reach 2 metres (6.5 ft) in length. The maximum adult body weight was most likely over 20 kilogrammes (44 lb) in *P. mongoliensis*. Several species approach *P. mongoliensis* in size (*P. lujiatunensis*, *P. neimongoliensis*, *P. xinjiangensis*), while others are somewhat smaller (*P. sinensis*, *P. meileyingensis*). The smallest known species, *P. ordosensis*, is 30% smaller than *P. mongoliensis*. The largest are *P. lujiatunensis* and *P. sibiricus*, although neither is significantly larger than *P. mongoliensis*. *Psittacosaurus* postcranial skeletons are more typical of a 'generic' bipedal ornithischian. There are only four digits on the manus ('hand'), as opposed to the five found in most other ornithischians (including all other ceratopsians), while the four-toed hindfoot is very similar to many other small ornithischians.

The skull of *Psittacosaurus* is highly modified compared to other ornithischian dinosaurs of its time. Extremely tall in height and short in length, the skull has an almost round profile in some species. The portion in front of the orbit (eye socket) is only 40% of total skull length, shorter than any other known ornithischian. The lower jaws of psittacosaurs are characterised by a bulbous vertical ridge down the centre of each tooth. Both upper and lower jaws sport a pronounced beak, formed from the rostral and predentary bones, respectively. The bony core of the beak may have been sheathed in keratin to provide a sharp cutting surface for cropping plant material. As the generic name suggests, the short skull and beak superficially resemble those of modern parrots. *Psittacosaurus* skulls share several adaptations with more derived ceratopsians, such as the unique rostral bone at the tip of the upper jaw, and the flared jugal (cheek) bones. There is still no sign of the bony neck frill or prominent facial horns which would develop in later ceratopsians. Bony horns protrude from the skull of *P. sibiricus*, but these are thought to be an example of convergent evolution.

### Soft tissue and coloration

The integument, or body covering, of *Psittacosaurus* is known from a Chinese specimen, which most likely comes from the Yixian Formation of Liaoning Province, China. The specimen, which is not yet assigned to any particular species, was likely illegally exported from China and was donated to the Senckenberg Museum in Germany. It was described while awaiting repatriation; previous repatriation attempts were unsuccessful.

Most of the body was covered in scales. Larger scales were arranged in irregular patterns, with numerous smaller scales occupying the spaces between them, similarly to skin impressions known from other ceratopsians, such as *Chasmosaurus*. A series of what appear to be hollow, tubular bristle-like structures, approximately 16 centimetres (6.3 in) long, were also preserved, arranged in a row down the dorsal (upper) surface of the tail. These were confirmed by the authors, as well as an independent scientist, to not represent plant material. The bristle-like integumentary structures extend into the skin nearly to the vertebrae, and were likely circular or tubular before being preserved. Under ultraviolet light, they gave off the same fluorescence as scales, providing the possibility they were keratinized. The study stated that, "at present, there is no convincing evidence which shows these structures to be homologous to the structurally different integumentary filaments of theropod dinosaurs". However, they found that all other feather-like integument from the Yixian Formation could be identified as feathers.

In 2008, another study was published describing the integument and dermis of *Psittacosaurus* sp., from a different specimen. The skin remains could be observed by a natural cross-section to compare them to modern animals, showing that dinosaurian dermal layers evolved in parallel to those in many other large vertebrates. The collagen tissue fibres in *Psittacosaurus* are complex, virtually identical to all other vertebrates in structure but having an exceptional thickness of about forty layers. As the sections of dermis were collected from the abdomen, where the scales were eroded, the tissue may have assisted with the musculature of the stomach and intestines and offered protection against predators.

### Species characteristics

Skulls of *P. mongoliensis* are flat on top, especially over the back of the skull, with a triangular depression, the antorbital fossa, on the outside surface of the maxilla (an upper jaw bone). A flange is present on the lower edge of the dentary (the tooth-bearing bone of the lower jaw), although it is not as prominent as in *P. meileyingensis* or *P. major* (=*P. lujiatunensis*). *P. mongoliensis* is among the largest known species. The skull of the type specimen, which is probably a juvenile, is 15.2 centimetres (6 in) long, and the associated femur is 16.2 centimetres (6.4 in) in length. Other specimens are larger, with the largest documented femur measuring about 21 centimetres (8.25 in) long.

*P. sinensis* is readily distinguished from all other species by numerous features of the skull. Adult skulls are smaller than those of *P. mongoliensis* and have less teeth. Uniquely, the premaxillary bone contacts the jugal (cheek) bone on the outside of the skull. The jugals flare out sideways, forming 'horns' proportionally wider than in any other known Psittacosaurus species except *P. sibiricus* and *P. lujiatunensis*. Because of the flared cheeks, the skull is actually wider than it is long. A smaller 'horn' is present behind the eye, at the contact of the jugal and postorbital bones, a feature also seen in *P. sibiricus*. The mandible (lower jaw) lacks the hollow opening, or fenestra, seen in other species, and the entire lower jaw is bowed outwards, giving the animal the appearance of an underbite. The skull of an adult *P. sinensis* can reach 11.5 centimeters (4.5 in) in length.

*P. sibiricus* is the largest-known species of *Psittacosaurus*. The skull of the type specimen is 20.7 centimetres long (8.25 in), and the femur is 22.3 cm (8.75 in) in length. It is also distinguished by its neck frill, which is longer than any other species, at 15 to 18% of skull length. A very striking feature of *P. sibiricus* is the number of 'horns' around the eyes, with three prominences on each postorbital, and one in front of each eye, on the palpebral bones. Similar horns found on the postorbital of *P. sinensis* are not as pronounced but may be homologous. The jugal has extremely prominent 'horns' and may contact the premaxilla, both features also seen in the possibly related *P. sinensis*. There is a flange on the dentary of the lower jaw, similar to *P. mongoliensis*, *P. meileyingensis*, and *P. sattayaraki*. It can be told apart from the other species of *Psittacosaurus* by a combination of 32 anatomical features, including six that are unique to the species. Most of these are skull details, but one unusual feature is the presence of 23 vertebrae between the skull and pelvis, unlike the 21 or 22 in the other species where the vertebrae are known.

*P. xinjiangensis* is distinguished by a prominent jugal 'horn' that is flattened on the front end, as well as some features of the teeth. The ilium, one of the three bones of the pelvis, also bears a characteristically long bony process behind the acetabulum (hip socket). An adult femur has a published length of about 16 centimetres (6.3 in). *P. meileyingensis* has the shortest snout and neck frill of any species, making the skull nearly circular in profile. The orbit (eye socket) is roughly triangular, and there is a prominent flange on the lower edge of the dentary, a feature also seen in specimens of *P. lujiatunensis*, and to a lesser degree in *P. mongoliensis*, *P. sattayaraki*, and *P. sibiricus*. The complete type skull, probably adult, is 13.7 centimetres (5.5 in) long. The dentary of *P. sattayaraki* has a flange similar to that found in *P. mongoliensis*, *P. sibiricus*, *P. lujiatunensis* and *P. meileyingensis*, although it is less pronounced than in those species. The material appears to be roughly the same size as *P. sinensis*. The frontal bone of *P. neimongoliensis* is distinctly narrow compared to that of other species, resulting in a narrower skull overall. The ischium bone of the pelvis is also longer than the femur, which differs from other species in which these bones are known. The type specimen has a skull length of 13.2 centimetres (5.2 in) and a femoral length of 13 centimetres (5.1 in), but is not fully grown. An adult *P. neimongoliensis* was probably smaller than *P. mongoliensis*, with a proportionately longer skull and tail. *P. ordosensis* can be distinguished by numerous features of the jugals, which have very prominent 'horns'. It is also the smallest known species. One adult skull measures only 9.5 centimeters (3.75 in) in length.

The type skull of *P. lujiatunensis* measures 19 cm (7.5 in) in length, while the largest-known skull is 20.5 centimetres (8 in) long, so this species was similar in size to *P. mongoliensis* and *P. sibiricus*. There is a fossa in front of the eye, as in *P. mongoliensis*. The jugal bones flare outwards widely, making the skull wider than it is long, as seen in *P. sinensis*. Widely flared jugals are also found in *P. sibiricus*. Overall, this species is thought to exhibit several primitive characteristics compared to other species of *Psittacosaurus*, which is consistent with its greater geological age. *P. gobiensis* was small-bodied (one metre (3 ft 3 in) long) and differs from other species of *Psittacosaurus* by "significant, but structurally minor, details." These include the presence of a pyramidal horn on the postorbital, a depression on the postorbital-jugal contact, and enamel thickness. *P. mongoliensis* was a contemporary.

## History of discovery

*Psittacosaurus* was first described as a genus in 1923, by Henry Fairfield Osborn. He named the type species *P. mongoliensis*, for the location of its discovery in Mongolia, placing it in the new family Psittacosauridae. Remains of this dinosaur were first discovered the year before, on the third American Museum of Natural History expedition to the Gobi Desert of Mongolia, when one of the expedition's drivers, Wong, found the type specimen (AMNH 6254), which preserves a nearly complete skull, as well as a post cranial skeleton lacking sections of the limbs. This same expedition turned up the remains of many other famous Mongolian dinosaurs, including *Protoceratops*, *Oviraptor*, and *Velociraptor*. Many later expeditions by various combinations of Mongolian, Russian, Chinese, American, Polish, Japanese, and Canadian paleontologists also recovered specimens from throughout Mongolia and northern China. In these areas, *Psittacosaurus mongoliensis* fossils are found in most sedimentary strata dating to the Aptian to Albian stages of the Early Cretaceous Period, or approximately 125 to 100 mya. Fossil remains of over 75 individuals have been recovered, including nearly 20 complete skeletons with skulls. Individuals of all ages are known, from hatchlings less than 13 centimetres (5.1 in) long, to very old adults reaching nearly 2 metres (6 ft 7 in) in length.

When describing *Psittacosaurus mongoliensis* in 1923, Osborn also gave the name *Protiguanodon mongoliense* to another skeleton found nearby, believing it to represent an ancestor of the ornithopod *Iguanodon*, in the new subfamily Protiguanodontinae. *Protiguanodon mongoliense*, AMNH 6523, measured 1.35 m (4 ft 5 in) long, and was known from much of the skeleton, although at the time of description the neck vertebrae were still covered by matrix. Osborn diagnosed his taxa on the basis of features of the teeth and snout. However, modern taxonomists find these features insignificant, instead placing *Protiguanodon mongoliense* within *Psittacosaurus mongoliensis*. When the skeleton was prepared further, it became clear that it was nearly identical to *Psittacosaurus mongoliensis*. In 1958, Chinese paleontologist Yang Zhongjian (better known as C. C. Young) renamed the skeleton *Psittacosaurus protiguanodonensis*. Today the specimen is generally referred to as the species *Psittacosaurus mongoliensis* and the names *Protiguanodon mongoliense* and *Psittacosaurus protiguanodonensis* are considered junior synonyms of the name *Psittacosaurus mongoliensis*, which was coined first.

In 1931, C. C. Young named a new species of *Psittacosaurus* for a partial skull discovered in Inner Mongolia, China. The skull was named *P. osborni* after Henry Fairfield Osborn. The validity of this species is now considered equivocal. Sereno (1990) considered it a synonym of *P. mongoliensis*, which is found in nearby strata of the same age. You and Dodson (2004) listed it as valid in a table, but not in their text. In a 2010 review, Sereno again regarded *P. osborni* as a synonym of *P. mongoliensis*, but noted it was tentative because of the presence of multiple valid psittacosaur species in Inner Mongolia. Young also described the species P. tingi in the same 1931 report which contained *P. osborni*. It is based on several skull fragments. He later synonymised the two species under the name *P. osborni*. You and Dodson (2004) followed this in a table, but Sereno regarded both species as synonyms of *P. mongoliensis*; a table in the latter reported *P. tingi* as a *nomen dubium*, however. The front half of a skull from Guyang County in Inner Mongolia was described as *Psittacosaurus guyangensis* in 1983. Disarticulated postcranial remains representing multiple individuals were found at the same locality and were assigned to the species. While it differs from the type specimen of *P. mongoliensis*, it falls within the range of individual variation seen in other specimens of that species and is no longer recognised as a valid species. You and Dodson (2004) included *P. guyangensis* in a table of valid taxa, but did not include it as such in their text.

### Assigned species

Seventeen species have been referred to the genus *Psittacosaurus*, although only nine to eleven are considered valid today. This is the highest number of valid species currently assigned to any single dinosaur genus (not including birds). In contrast, most other dinosaur genera are monospecific, containing only a single known species. The difference is most likely due to artifacts of the fossilisation process. While *Psittacosaurus* is known from hundreds of fossil specimens, most other dinosaur species are known from far fewer, and many are represented by only a single specimen. With a very high sample size, the diversity of *Psittacosaurus* can be analysed more completely than that of most dinosaur genera, resulting in the recognition of more species. Most extant animal genera are represented by multiple species, suggesting that this may have been the case for extinct dinosaur genera as well, although most of these species may not have been preserved. In addition, most dinosaurs are known solely from bones and can only be evaluated from a morphological standpoint, whereas extant species often have very similar skeletal morphology but differ in other ways which would not normally be preserved in the fossil record, such as behaviour, or colouration. Therefore, actual species diversity may be much higher than currently recognised in this and other dinosaur genera. As some species are known only from skull material, species of *Psittacosaurus* are primarily distinguished by features of the skull and teeth. Several species can be recognised by features of the pelvis as well.

#### *P. sinensis*

In the 1950s, a new Chinese species of *Psittacosaurus* was found in the Aptian-Albian Qingshan Formation of Shandong Province, southeast of Beijing. C. C. Young called it *P. sinensis* to differentiate it from *P. mongoliensis*, which had originally been found in Mongolia. Fossils of more than twenty individuals have since been recovered, including several complete skulls and skeletons, making this the most well-known species after *P. mongoliensis*. Chinese paleontologist Zhao Xijin named a new species after his mentor, C. C. Young, in 1962. However, the type specimen of *P. youngi* (a partial skeleton and skull) was discovered in the same rocks as *P. sinensis* and appears to be very similar, so *P. youngi* is generally considered a junior synonym of that better-known species. As with *P. guyangensis* and *P. osborni*, You and Dodson (2004) listed it as valid in a table, but not in their text.

#### *P. xinjiangensis*

In 1988, Zhao and American paleontologist Paul Sereno described *P. xinjiangensis*, named after the Xinjiang Autonomous Region in which it was discovered. Several individuals of different ages were discovered in the early 1970s by Chinese paleontologists and described by Sereno and Zhao, although the holotype and most complete skeleton belonged to a juvenile. An adult skeleton was later discovered at a different locality in Xinjiang. These specimens come from the upper part of the Tugulu Group, which is regarded as Aptian-Albian in age.

#### *P. meileyingensis*

A second species described in 1988 by Sereno and Zhao, along with two Chinese colleagues, was *P. meileyingensis* from the Jiufotang Formation, near the town of Meileyingzi, Liaoning Province, northeastern China. This species is known from four fossil skulls, one associated with some skeletal material, found in 1973 by Chinese scientists. The age of the Jiufotang in Liaoning is unknown, but in the neighbouring province of Inner Mongolia, it has been dated to about 110 Ma, in the Albian stage of the Early Cretaceous.

#### *P. sattayaraki*

French paleontologist Eric Buffetaut and a Thai colleague, Varavudh Suteethorn, described a partial upper and lower jaw from the Aptian-Albian Khok Kruat Formation of Thailand in 1992, giving it the name *P. sattayaraki*. In 2000, Sereno questioned the validity of this species, citing its eroded and fragmentary nature, and noted an absence of features characteristic of the genus *Psittacosaurus*. However, in 2002 the original authors published new images of the fossil which seem to show teeth in the lower jaw that exhibit the bulbous vertical ridge characteristic of psittacosaurs. Other authors have also defended its validity, while some continue to regard it as dubious. Sereno (2010) proposed that the best assignment for the type material may be Ceratopsia *incertae sedis*.

#### *P. neimongoliensis* and *P. ordosensis?*

Two new species of *Psittacosaurus* were described by Canadian Dale Russell and Zhao in 1996. The first was named *P. neimongoliensis*, after the Mandarin Chinese name for Inner Mongolia. It is based on a nearly complete fossil skeleton, including most of the skull, found in the Early Cretaceous Ejinhoro Formation with seven other individuals. Russell and Zhao also named *P. ordosensis* in 1996, after the Ordos prefecture of the Inner Mongolia Autonomous Region. The type specimen is a nearly complete skeleton, including part of the skull. However, only the skull, lower jaw, and foot have been described. Three other specimens were referred to this species but remain undescribed. Like *P. neimongoliensis*, this species was discovered in the Eijnhoro Formation. Sereno (2010) found the species as described to be indistinguishable from *P. sinensis*, another small species, but suggested that additional study of *P. ordosensis* might reveal diagnostic features. He provisionally designated *P. ordosensis* a *nomen dubium*.

#### *P. mazongshanensis?*

Xu Xing, another Chinese paleontologist, named a new species of *Psittacosaurus* in 1997, based on a complete skull with associated vertebrae and a forelimb. This material was recovered in Gansu Province, near the border with Inner Mongolia. This species is named *P. mazongshanensis* after the nearby mountain called Mazongshan (Horse Mane Mountain) and has been described in a preliminary manner. Unfortunately, the skull was damaged while in the care of the Chinese Institute of Vertebrate Paleontology and Paleoanthropology (IVPP), and several fragments have been lost, including all of the teeth. The remains were found in the Lower Xinminbao Formation, which have not been precisely dated, although there is some evidence that they were deposited in the late Barremian through Aptian stages. Sereno suggested in 2000 that *P. mazongshanensis* was a *nomen dubium*, with no unique features that separate it from any other species of *Psittacosaurus*. However, more recent authors have noted that it can be distinguished by its proportionally long snout compared to other species of *Psittacosaurus*, as well as a prominent bony protuberance, pointing outwards and downwards, on the maxilla of the upper jaw. The maxillary protuberance is also now missing. Other features originally used to distinguish the species have been recognised as the results of the deformation of the skull after fossilisation. Sereno (2010) remained unconvinced of its validity.

#### *P. sibiricus*

Beginning in the 1950s, Russian paleontologists began excavating *Psittacosaurus* remains at a locality near the village of Shestakovo in the oblast of Kemerovo in Siberia. Two other nearby localities were explored in the 1990s, one of which produced several complete skeletons. This species was named *P. sibiricus* in 2000 in a scientific paper written by five Russian paleontologists, but credit for the name is officially given to two of those authors, Alexei Voronkevich and Alexander Averianov. The remains were not completely described until 2006. Two nearly complete, articulated skeletons and a variety of disarticulated material from other individuals of all ages are known from the Ilek Formation of Siberia, which ranges from the Aptian to Albian stages of the Early Cretaceous.

#### *P. lujiatunensis'*

*P. lujiatunensis*, named in 2006 by Chinese paleontologist Zhou Chang-Fu and three Chinese colleagues, is one of the oldest-known species, based on four skulls from the lower beds of Yixian Formation, near the village of Lujiatun. While this bed has been dated differently by different authors, from 128 Ma in the Barremian stage, to 125 Ma in the earliest Aptian, revised dating methods have shown them to be about 123 million years old. *P. lujiatunensis* was contemporaneous with another psittacosaurid species, *Hongshanosaurus houi*, which was found in the same beds. It is potentially synonymous with H. houi; Sereno (2010), who proposed that *Hongshanosaurus* is a synonym of *Psittacosaurus*, opted to leave *P. lujiatunensis* and *H. houi* separate species due to the inadequacies of the latter's type specimen.

#### *P. major*

One nearly complete skeleton of *P. lujiatunensis* from the same lower beds of the Yixian Formation had previously been classified in its own species, *Psittacosaurus major*, named for the large size of its skull by Sereno, Zhao and two colleagues in 2007. You and colleagues described an additional specimen and concurred that it was distinct from *P. lujiatunensis*. *P. major* was originally characterised by a proportionately large skull, which was 39% of the length of its torso, compared to 30% in *P. mongoliensis*, and other features. However, a 2013 study utilising morphometric analysis showed that the supposed differences between *P. lujiatunensis* and *P. major* were due to differences in preservation and crushing. The study concluded that both represented a single species.

#### *P. houi?*

A third species of Lujiatun psittacosaur, the first to be named, was described as *Hongshanosaurus houi* in 2003. The generic name *Hongshanosaurus* was derived from the Mandarin Chinese words 紅 (hóng: "red") and 山 (shān: "hill"), as well as the Greek word sauros ("lizard"). This name refers to the ancient Hongshan culture of northeastern China, who lived in the same general area in which the fossil skull of *Hongshanosaurus* was found. The type and only named species, *H. houi*, honours Hou Lianhai, a professor at the IVPP in Beijing, who curated the specimen. Genus and species were both named by Chinese paleontologists You Hailu, Xu Xing, and Wang Xiaolin in 2003. Sereno (2010) regarded its distinct proportions as due to crushing and compression of the *Hongshanosaurus* skulls. He regarded *Hongshanosaurus* as a junior synonym of *Psittacosaurus*, and potentially the same as *P. lujiatunensis*. He did not synonymise the two species because of difficulties with the holotype skull of *H. houi*, instead considering new combination *P. houi* a *nomen dubium* within *Psittacosaurus*. Sereno's hypothesis was supported by a morphometric study in 2013, which found *P. houi* and *P. lujiatunensis* to be synonymous. While *P. houi* is the oldest available name, the researchers argued that because the type specimen of *P. lujiatunensis* was better preserved, the correct name for this species should be *P. lujiatunensis* rather than *P. houi*, which would normally have priority.

#### *P. gobiensis*

*P. gobiensis* is named for the region it was found in 2001, and first described by Sereno, Zhao and Lin in 2010. It is known from a skull and partial articulated skeleton with gastroliths. Many other specimens either cannot be determined to belong to any particular species, or have not yet been assigned to one. These specimens are generally all referred to as *Psittacosaurus* sp., although it is not assumed that they belong to the same species. More than 200 specimens of *Psittacosaurus* have been found in the Yixian Formation, which is famous for its fossils of feathered dinosaurs. The vast majority of these have not been assigned to any published species, although many are very well preserved and some have already been partially described. Nearly 100 *Psittacosaurus* skeletons were excavated in Mongolia during the summers of 2005 and 2006 by a team led by Mongolian paleontologist Bolortsetseg Minjin and American Jack Horner from the Museum of the Rockies in Montana. Although only *P. mongoliensis* has been described from Mongolia so far, these specimens are still in preparation and have not yet been assigned to a species.

## Classification

*Psittacosaurus* is the type genus of the family Psittacosauridae, which was also named by Osborn in 1923. Psittacosaurids were basal to almost all known ceratopsians except Yinlong and perhaps the Chaoyangsauridae. While Psittacosauridae was an early branch of the ceratopsian family tree, *Psittacosaurus* itself was probably not directly ancestral to any other groups of ceratopsians. All other ceratopsians retained the fifth digit of the hand, a plesiomorphy or primitive trait, whereas all species of *Psittacosaurus* had only four digits on the hand. In addition, the antorbital fenestra, an opening in the skull between the eye socket and nostril, was lost during the evolution of Psittacosauridae, but is still found in most other ceratopsians and in fact most other archosaurs. It is considered highly unlikely that the fifth digit or antorbital fenestra would evolve a second time.

In 2014, the describers of a new taxon of basal ceratopsian published a phylogenetic analysis encompassing *Psittacosaurus*. The below cladogram is from their analysis, placing the genus as one of the most primitive ceratopsians. The authors (Farke et al.) noted that all taxa outside of Leptoceratopsidae and Coronosauria with the exception of their genus *Aquilops* are from Asia, meaning the group likely originated there.

Although many species of *Psittacosaurus* have been named, their relationships to each other have not yet been fully explored and no scientific consensus exists on the subject. Several phylogenetic analyses have been published, with the most detailed being those by Alexander Averianov and colleagues in 2006, Hai-Lu You and colleagues in 2008, and Paul Sereno in 2010. The middle one is shown below.

In 2005, Zhou and colleagues suggested that *P. lujiatunensis* is basal to all other species. This would be consistent with its earlier appearance in the fossil record.

## Paleobiology

The brain of *P. lujiatunensis* is well known; a study on the anatomy and functionality of three specimens was published in 2007. Until the study, it was generally thought the brain of *Psittacosaurus* would have been similar to other ceratopsians with low Encephalisation Quotients. Russell and Zhao (1996) believed "the small brain size of Psittacosaurs implies a very restrictive behavioural repertoire relative to that of modern mammals of similar body size." However, the 2007 study dispelled this theory when it found the brain to be more advanced. There is generally negative allometry for brain size with development in vertebrates, but this was shown not to be true in *Psittacosaurus*. The EQ score for *P. lujiatunensis* is 0.31, significantly higher than genera such as *Triceratops*. A higher EQ correlates with more complex behaviour, and various dinosaurs have high EQs, similar to birds, which range from 0.36 to 2.98. Thus, *Psittacosaurus* behaviour could have been as complex as that in *Tyrannosaurus*, whose EQ ranges from 0.30 to 0.38. Behaviours influenced by high EQs include nest-building, parental care, and bird-like sleeping, some of which have been shown to be present in *Psittacosaurus*.

The senses of *Psittacosaurus* can be inferred from the endocast. Large olfactory bulbs are present, indicating the genus had an acute sense of smell. The size of these bulbs are comparable to large predatory theropods, although they likely evolved to avoid predators instead of to seek out prey. The sclerotic rings in reptiles directly show the size of the eyeball. The rings are not well preserved in *Psittacosaurus*, with one individual preserving them likely contracted postmortem, but if they are similar to those of *Protoceratops*, *Psittacosaurus* would have had large eyes and acute vision. The curvature of the semicircular canals is related to the agility of reptiles, and the large curved canals in *Psittacosaurus* show that the genus was much more agile than later ceratopsians. Comparisons between the scleral rings of *Psittacosaurus* and modern birds and reptiles suggest that it may have been cathemeral, active throughout the day and for short intervals at night.

Ford and Martin (2010) proposed that *Psittacosaurus* was semi-aquatic, swimming with its tail like a crocodile, and paddling and kicking. They based their interpretation on evidence including: the lacustrine (lake) depositional setting of many specimens; the position of the nostrils and eyes; interpretations of the motions of the arms and legs; tails with long chevrons (and with the bristles on the tail interpreted as possibly skin-covered, forming a fin), providing a propulsive surface; and the presence of gastroliths, interpreted as ballast. They further suggested that some species of *Psittacosaurus* were more terrestrial than others.

## Paleochronology

*Psittacosaurus* is known from hundreds of individual specimens, of which over 75 have been assigned to the type species, *P. mongoliensis*. All *Psittacosaurus* fossils discovered so far have been found in Early Cretaceous sediments in Asia, from southern Siberia to northern China, and possibly as far south as Thailand. The most common age of geologic formations bearing *Psittacosaurus* fossils is from the Late Barremian through Albian stages of the Early Cretaceous, or approximately 126 to 101 mya (million years ago). Many terrestrial sedimentary formations of this age in Mongolia and northern China have produced fossils of *Psittacosaurus*, leading to the definition of this time period in the region as the *Psittacosaurus* biochron.

The earliest known species is *P. lujiatunensis*, found in the lowest beds of the Yixian Formation. Over 200 specimens attributed to this genus have been recovered from these and other beds of the Yixian, the age of which is the subject of much debate. Although many early studies using radiometric dating put the Yixian in the Jurassic Period, tens of millions of years outside of the expected temporal range of *Psittacosaurus*, most recent work dates it to the Early Cretaceous. Using argon–argon dating, a team of Chinese scientists dated the lowest beds in the formation to about 128 mya, and the highest to approximately 122 mya. A more recent Chinese study, using uranium–lead dating, suggests that the lower beds are younger, approximately 123.2 mya, while agreeing with an age of 122 mya for the upper beds.

## JPInstitute.com Description

Psittacosaurus was discovered in Outer Mongolia in 1922 by the Roy Chapman Andrews expeditions. It was these expeditions and Andrews' adventures that inspired the Indiana Jones character. Psittacosaurus was one of the very first in the family of dinosaurs that later included Triceratops. It was very small and still walked on two legs much of the time. Its later and much larger family members would need to walk on all four legs all the time.

This is possibly the best-known genus of dinosaurs, as many well-preserved specimens have been found. It represents a possible transitional creature, resembling the ornithopod Hypsilophodon, but clearly showing the traits that would later mark the ceratopsian dinosaurs.

Named by Henry Osborn, Psittacosaurus is the earliest and most primitive in the ceratopsian family. The beak for which Psittacosaurus was named was a typical ceratopsian feature. Although the wide squamosals and the beak gave it a ceratopsian resemblance, there are no horns or frill at all. The ragged spade-shaped teeth indicate that it had a diet of course vegetation. The body itself was much lighter than Protoceratops and the smaller size of it's front limbs would indicate that it spent much of its time on two legs. The "hands" show some ability to grasp objects. Gastroliths have been found associated with some specimens.

## Dinosaur Field Guide Description

Psittocosaurus (parrot lizard") was one of the first to appear of the last great group of dinosaurs: ceratopsia, the horned dinosaurs. Psittacosaurus has two features common to all ceratopsians: an extra upper beak bone on top of the upper lip bone, and the beginnings of what will become the ceratopsian frill. Over eleven species of Psittocosaurus have been named--more than in 99 percent of all other dinosaurs. But are they all valid? Some scientists will name a new species based on very slight differences in the skull. These differences can be the result, however, of sex, age, health, and normal variation within populations. There are also stratigraphic differences. This means that over time, later populations can evolve larger featres than their ancestors. Psittacosaurus remains have been found with gastroliths, or stomach stones. Because most herbivorous dinosaurs could not chew their food. they swallowed a lot of unprocessed plants. To aid their digestion, they also swallowed small stones.

### Fun Facts

Ceratopsians are so popular that they have their own book, The Horned Dinosaurs, by Peter Dodson.

### Trivia

Psittacosaurus has horns on its cheeks, called jugal horns.

## Appearance in other media

### *Jurassic Park*

* *Psittacosaurus* appeared at *Jurassic Park: The Ride* and is located in *Ultrasaurus* lagoon. Like the version seen in *Jurassic Park III: Park Builder*, it lacks the quills found on its tail. The hooting sound of the *Psittacosaurus* is made from the hooting sounds of the white-bellied green pigeon, a bird native to Asia much like the dinosaur itself.

## Links

http://web.archive.org/web/20040806074407fw\_/http://www.jpinstitute.com/dinopedia/dinocards/dc\_psitt.html#