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Coelophysis

Coelophysis ( traditionally;   or , as heard more commonly in recent decades ) is a genus of coelophysid theropod dinosaur that lived approximately 215 to 208.5 million years ago during the Late Triassic period from the middle to late Norian age in what is now the southwestern United States. Megapnosaurus was once considered to be a species within this genus, but this interpretation has been challenged since 2017 and the genus Megapnosaurus is now considered valid.

Coelophysis was a small, slenderly-built, ground-dwelling, bipedal carnivore that could grow up to 3 m long. It is one of the earliest known dinosaur genera. Scattered material representing similar animals has been found worldwide in some Late Triassic and Early Jurassic formations.

The type species C. bauri, originally given to the genus Coelurus by Edward Drinker Cope in 1887, was described by the latter in 1889. The names Longosaurus and Rioarribasaurus are synonymous with Coelophysis. Coelophysis is one of the most specimen-rich dinosaur genera.

History of discovery
The type species of Coelophysis was originally named as a species of Coelurus. Edward Drinker Cope first named Coelophysis in 1889 to name a new genus, outside of Coelurus and Tanystropheus, which C. bauri was previously classified in, for C. bauri, C. willistoni, and C. longicollis. An amateur fossil collector working for Cope, David Baldwin, had found the first remains of the dinosaur in 1881 in the Chinle Formation in northwestern New Mexico. Early in 1887, Cope referred the specimens collected to two new species, C. bauri and C. longicollis of the genus Coelurus. Later on in 1887, Cope reassigned the material to a yet another genus, Tanystropheus. Two years later, Cope corrected his classification after realizing differences in the vertebrae and named Coelophysis, with C. bauri as the type species, which was named for Georg Baur, a comparative anatomist whose ideas were similar to Cope's. The name Coelophysis comes from the Greek words κοῖλος/koilos (meaning 'hollow') and φύσις/physis (meaning 'form'), together "hollow form", which is a reference to its hollow vertebrae. However, the first finds were too poorly preserved to give a complete picture of the new dinosaur.

In 1947, a substantial "graveyard" of Coelophysis fossils was found by George Whitaker, the assistant of Edwin H. Colbert, in New Mexico, at Ghost Ranch, close to the original find. American Museum of Natural History paleontologist Edwin H. Colbert conducted a comprehensive study of all the fossils found up to that date and assigned them to Coelophysis. The Ghost Ranch specimens were so numerous, including many well-preserved and fully articulated specimens, that one of them has since become the diagnostic, or type specimen, for the entire genus, replacing the original, poorly preserved specimen.

In the early 1990s, there was debate over the diagnostic characteristics of the first specimens collected, compared to the material excavated at the Ghost Ranch Coelophysis quarry. Some paleontologists were of the opinion that the original specimens were not diagnostic beyond themselves and, therefore, that the name C. bauri could not be applied to any additional specimens. They therefore applied a different name, Rioarribasaurus, to the Ghost Ranch quarry specimens.

Since the numerous well-preserved Ghost Ranch specimens were used as Coelophysis in most of the scientific literature, the use of Rioarribasaurus would have been very inconvenient for researchers. So, a petition was given to have the type specimen of Coelophysis transferred from the poorly preserved original specimen to one of the well-preserved Ghost Ranch specimens. This would make Rioarribasaurus a definite synonym of Coelophysis, specifically a junior objective synonym. In the end, the International Commission on Zoological Nomenclature (ICZN) voted to make one of the Ghost Ranch samples the actual type specimen for Coelophysis and dispose of the name Rioarribasaurus altogether (declaring it a nomen rejectum, or "rejected name"), thus resolving the confusion. The name Coelophysis became a nomen conservandum ("conserved name").

In a situation affecting many dinosaur taxa, some more recently discovered fossils were originally classified as new genera, but may be species of Coelophysis. For example, Prof. Mignon Talbot's 1911 discovery, which she named Podokesaurus holyokensis, has long been considered to be related to Coelophysis and some modern scientists consider Podokesaurus a synonym of Coelophysis. Another specimen from the Portland Formation of the Hartford Basin, now at the Boston Museum of Science, has also been referred to Coelophysis. The specimen consists of sandstone casts of a pubis, tibia, three ribs, and a possible vertebra that probably originated in a quarry in Middletown, Connecticut. However, both the type specimen of Podokesaurus and the Middletown specimen are typically considered indeterminate theropods today.

Sullivan & Lucas (1999) referred one specimen from Cope's original material of Coelophysis (AMNH 2706) to what they thought was a newly discovered theropod, Eucoelophysis. However, subsequent studies have shown that Eucoelophysis was misidentified and is actually a silesaurid, a type of non-dinosaurian ornithodiran closely related to Silesaurus.

Formerly assigned species
"Syntarsus" rhodesiensis was first described by Raath (1969) and assigned to Podokesauridae. The taxon "Podokesauridae", was abandoned because its type specimen was destroyed in a fire and can no longer be compared to new finds. Over the years, paleontologists assigned the genus to Ceratosauridae (Welles, 1984), Procompsognathidae (Parrish and Carpenter, 1986), and Ceratosauria (Gauthier, 1986). In 2004, "Syntarsus" was found to be synonymous with Coelophysis by Tykoski and Rowe (2004). Ezcurra and Novas (2007) and Ezcurra (2007) also concluded that "Syntarsus" was synonymous with Coelophysis. In a phylogenetic analysis by Ezcurra (2017), Megapnosaurus was recovered in a clade with Segisaurus and Camposaurus, supporting the generic distinction of Megapnosaurus. This was supported by Barta and colleagues in 2018, noting that Coelophysis still bears the vestigial 5th metacarpal, a feature absent in Megapnosaurus.

The genus Syntarsus was named by Raath in 1969 for the type species Syntarsus rhodesiensis from Africa and later applied to the North American Syntarsus kayentakatae. It was renamed by American entomologist Dr. Michael Ivie (Montana State University of Bozeman), Polish Australian Dr. Adam Ślipiński, and Polish Dr. Piotr Węgrzynowicz (Muzeum Ewolucji Instytutu Zoologii PAN of Warsaw), the three scientists who discovered that the genus name Syntarsus was already taken by a colydiine beetle described in 1869. Many paleontologists did not like the naming of Megapnosaurus, partially because taxonomists are generally expected to allow original authors of a name to correct any mistakes in their work. Raath was aware of the homonymy between the dinosaur Syntarsus and beetle Syntarsus, but the group who published Megapnosaurus have claimed that they believed Raath was deceased and unable to correct his mistake, so they proceeded accordingly. Mortimer (2012) pointed out that "Paleontologists might have reacted more positively if the replacement name (Megapnosaurus) hadn't been facetious, translating to "big dead lizard". Yates (2005) analyzed Coelophysis and Megapnosaurus and concluded that the two genera are almost identical, suggesting that Megapnosaurus was possibly synonymous with Coelophysis. In 2004, Raath co-authored two papers in which he argued that Megapnosaurus (formerly Syntarsus) was a junior synonym of Coelophysis. Megapnosaurus was regarded by Paul (1988) and Downs (2000) as being congeneric with Coelophysis. In 1993, Paul then suggested that Coelophysis should be placed in Megapnosaurus (then known as Syntarsus) to get around the above-mentioned taxonomic confusion. Downs (2000) examined Camposaurus and concluded that it was a junior synonym of Coelophysis because of its similarity to some of the Coelophysis Ghost Ranch specimens. However, a reassessment of the Camposaurus holotype by Martin Ezcurra and Stephen Brusatte, published in 2011, revealed a pair of autapomorphies in the holotype, indicating that C. arizonensis was not a synonym of C. bauri, although it was a close relative of M. rhodesiensis. Barta et al. (2018) concluded that C. bauri differed from M. rhodesiensis in that it bears its 5th metacarpal and several features in the musculature of the limbs according to Griffin (2018).

Description
Coelophysis is known from a number of complete fossil skeletons of the species C. bauri. This lightly built dinosaur measured up to 3 m long and was more than a meter tall at the hips. Gregory S. Paul (1988) estimated the weight of the gracile form at 15 kg and the weight of the robust form at 20 kg, but later presented a higher estimate of 25 kg. Coelophysis was a bipedal, carnivorous, theropod dinosaur and a fast, agile runner. Despite its basal position within Theropoda, the bauplan of Coelophysis differed from those of other basal theropods, such as Herrerasaurus, showing more derived traits common in theropods that superseded it. The torso of Coelophysis conforms to the basic theropod bauplan, but the pectoral girdle displays some special characteristics. C. bauri had a furcula (wishbone), the earliest known example in a dinosaur. Coelophysis also preserves the ancestral condition of possessing four digits on the hand (manus). It had only three functional digits, with the fourth being embedded in the flesh of the hand.

Coelophysis had narrow hips, arms adapted for grasping prey, and narrow feet. Its neck and tail were long and slender. The pelvis and hindlimbs of C. bauri are also slight variations on the theropod body plan. It has the open acetabulum and straight ankle hinge that define Dinosauria. The leg ended in a three-toed foot (pes) with a raised dewclaw (hallux). The tail had an unusual structure within its interlocking prezygapophysis of its vertebrae, which formed a semi-rigid lattice, apparently to stop the tail from moving up and down.

Coelophysis had a long and narrow head (approximately 270 mm), with large, forward-facing eyes that afforded it stereoscopic vision and, as a result, excellent depth perception. Rinehart et al. (2004) described the complete sclerotic ring found for a juvenile Coelophysis bauri (specimen NMMNH P-4200) and compared it to data on the sclerotic rings of reptiles (including birds), concluding that Coelophysis was a diurnal, visually oriented predator. The study found that the vision of Coelophysis was superior to most lizards' vision and ranked with that of modern birds of prey. The eyes of Coelophysis appear to be the closest to those of eagles and hawks, with a high power of accommodation. The data also suggested poor night vision, which would mean this dinosaur had a round pupil rather than a split pupil.

Coelophysis had an elongated snout with large fenestrae that helped to reduce skull weight, while narrow struts of bones preserved the structural integrity of the skull. The neck had a pronounced sigmoid curve. The braincase is known in Coelophysis bauri, but little data could be derived because the skull was crushed. Unlike some other theropods, the cranial ornamentation of Coelophysis was not located at the top of its skull. Low, laterally raised bony ridges were present on the dorsolateral margin of the nasal and lacrimal bones in the skull, directly above the antorbital fenestra.

Distinguishing anatomical features
A diagnosis is a statement of the anatomical features of an organism (or group) that collectively distinguish it from all other organisms. Some, but not all, of the features in a diagnosis are also autapomorphies. An autapomorphy is a distinctive anatomical feature that is unique to a given organism or group.

According to Ezcurra (2007) and Bristowe and Raath (2004), Coelophysis can be distinguished based on the absence of an offset rostral process of the maxilla, the quadrate being strongly caudally, a small external mandibular fenestra (which is 9–10% of the mandibular length ), and the anteroposterior length of the ventral lacrimal process is greater than 30% of its height.

Several paleontologists consider Coelophysis bauri to be the same dinosaur as Megapnosaurus rhodesiensis (formerly Syntarsus). However, this has been refuted by many paleontologists. Downs (2000) concluded that C. bauri differs from C. rhodesiensis in cervical length, proximal and distal leg proportions, and proximal caudal vertebral anatomy. Tykoski and Rowe (2004) concluded that C. bauri differs from M. rhodesiensis in that it lacks a pit at the base of the nasal process of the premaxilla. Bristowe and Raath (2004) concluded that C. bauri differs from M. rhodesiensis in having a longer maxillary tooth row. Barta et al. (2018) concluded that C. bauri differed from M. rhodesiensis in that it bears its 5th metacarpal. Griffin (2018) concluded that C. bauri differs from M. rhodesiensis in several differences in the musculature of the limbs.

Classification

Theropod cladogram based on the phylogenetic analyses conducted by Ezcurra et al. (2020).

Coelophysis is a distinct taxonomic unit (genus), composed of the single species C. bauri. Two additional originally described species, C. longicollis and C. willistoni, are now considered dubious and undiagnostic. M. rhodesiensis was referred to Coelophysis for several years, but it is likely its own genus and is known from the early Jurassic of southern Africa. A third possible species is Coelophysis kayentakatae, previously referred to the genus Megapnosaurus, from the Kayenta Formation of the southwestern US. In recent phylogenetic analyses, "Syntarsus" kayentakatae has been shown to be distantly related to Coelophysis and Megapnosaurus, suggesting that it belongs to its own genus.

Feeding
The teeth of Coelophysis were typical of predatory dinosaurs, as they were blade-like, recurved, sharp, jagged, and finely serrated on both the anterior and posterior edges. Its dentition shows that it was carnivorous, probably preying on the small, lizard-like animals that were discovered with it. It may also have hunted in packs to tackle larger prey. Coelophysis bauri has approximately 26 teeth on the maxillary bone of the upper jaw and 27 teeth on the dentary bone of the lower jaw. Kenneth Carpenter (2002) examined the bio-mechanics of theropod arms and attempted to evaluate their usefulness in predation. He concluded that the arm of Coelophysis was flexible and had a good range of motion, but its bone structure suggested that it was comparatively weak. The "weak" arms and small teeth in this genus suggested that Coelophysis preyed upon animals that were substantially smaller than itself. Rinehart et al. agreed that Coelophysis was a "hunter of small, fast-moving prey". Carpenter also identified three distinct models of theropod arm use and noted that Coelophysis was a "combination grasper-clutcher", as compared to other dinosaurs that were "clutchers" or "long armed graspers".

It has been suggested that C. bauri was a cannibal, based on supposed juvenile specimens found "within" the abdominal cavities of some Ghost Ranch specimens. However, Robert J. Gay showed in 2002 that these specimens were misinterpreted. Several specimens of "juvenile coelophysids" were actually small crurotarsan reptiles, such as Hesperosuchus. Gay's position was lent support in a 2006 study by Nesbitt et al. In 2009, new evidence of cannibalism came to light when additional preparation of previously excavated matrix revealed regurgitate material in and around the mouth of Coelophysis specimen NMMNH P-44551. This material included tooth and jaw bone fragments that Rinehart et al. considered "morphologically identical" to a juvenile Coelophysis.

In 2010, Gay examined the bones of juveniles found within the thoracic cavity of AMNH 7224 and calculated that the total volume of these bones was 17 times greater than the maximum estimated stomach volume of the Coelophysis specimen. Gay observed that the total volume would be even greater when considering that there would have been flesh on these bones. This analysis also noted the absence of tooth marks on the bones as would be expected in defleshing and the absence of expected pitting by stomach acids. Finally, Gay demonstrated that the alleged cannibalized juvenile bones were deposited stratigraphically below the larger animal that had supposedly cannibalized them. Taken together, these data suggested that the Coelophysis specimen AMNH 7224 was not a cannibal and that the bones of the juvenile and adult specimens were found in their final position as a result of "coincidental superposition of different sized individuals.

Pack behavior
The discovery of over 1,000 specimens of Coelophysis at the Whitaker quarry at Ghost Ranch has suggested gregarious behavior to researchers like Schwartz and Gillette. There is a tendency to see this massive congregation of animals as evidence for huge packs of Coelophysis roaming the land. No direct evidence for flocking exists because the deposits only indicate that large numbers of Coelophysis, along with various other Triassic animals, were buried together. Some of the evidence from the taphonomy of the site indicates that these animals may have been gathered together to feed or drink from a depleted water hole or to feed on a spawning run of fish, being later buried in a catastrophic flash flood or a drought.

With 30 specimens of C. rhodesiensis found together in Zimbabwe, some palaeontologists have suggested that Coelophysis was indeed gregarious. Again, there is no direct evidence of flocking in this case and it has also been suggested that these individuals were also victims of flash flooding as it appears to have been commonplace during this period.

Growth and sexual dimorphism
Rinehart (2009) assessed the ontogenic growth of this genus using data gathered from the length of its upper leg bone (femur) and concluded that Coelophysis juveniles grew rapidly, especially during the first year of life. Coelophysis likely reached sexual maturity between the second and third year of life and reached its full size, just above 10 feet in length, by its eighth year. This study identified four distinct growth stages: 1-year, 2-year, 4-year, and 7+ year. It was also thought that, as soon as they were hatched, they would have to fend for themselves.

Two "morphs" of Coelophysis have been identified. One is a more gracile form, as in specimen AMNH 7223, and the other is a slightly more robust form, as in specimens AMNH 7224 and NMMNH P-42200. Skeletal proportions were different between these two forms. The gracile form has a longer skull, a longer neck, shorter arms, and has sacral neural spines that are fused. The robust form has a shorter skull, a shorter neck, longer arms, and unfused sacral neural spines. Historically, many arguments have been made that this represents some sort of dimorphism in the population of Coelophysis, probably sexual dimorphism. Raath agreed that dimorphism in Coelophysis is evidenced by the size and structure of the arm. Rinehart et al. studied 15 individuals, and agreed that two morphs were present, even in juvenile specimens, and suggested that sexual dimorphism was present early in life, prior to sexual maturity. Rinehart concluded that the gracile form was female and the robust form was male based on differences in the sacral vertebrae of the gracile form, which allowed for greater flexibility for egg laying. Further support for this position was provided by an analysis showing that each morph comprised 50% of the population, as would be expected in a 50/50 sex ratio.

However, more recent research has found that C. bauri and C. rhodesiensis had highly variable growth between individuals, with some specimens being larger in their immature phase than smaller adults were when completely mature. This indicates that the supposed presence of distinct morphs is simply the result of individual variation. This highly variable growth was likely ancestral to dinosaurs but later lost and may have given such early dinosaurs an evolutionary advantage in surviving harsh environmental challenges.

Reproduction
Through the compilation and analysis of a database of nearly three dozen reptiles (including birds) and comparison with existing data about the anatomy of Coelophysis, Rinehart et al. (2009) drew the following conclusions. It was estimated that average egg of Coelophysis was 31–33.5 millimeters across its minor diameter and that each female would lay between 24 and 26 eggs in each clutch. The evidence suggested that some parental care was necessary to nurture the relatively small hatchlings during the first year of life, where they would reach 1.5 meters in length by the end of their first growth stage. Coelophysis bauri invested as much energy in reproduction as other extinct reptiles of its approximate size.

Paleopathology
In a 2001 study conducted by Bruce Rothschild and other paleontologists, 14-foot bones referred to Coelophysis were examined for signs of stress fracture, but none were found.

In C. rhodesiensis, healed fractures of the tibia and metatarsus have been observed, but are very rare. "[T]he supporting butresses of the second sacral rib" in one specimen of Syntarsus rhodesiensis showed signs of fluctuating asymmetry. Fluctuating asymmetry results from developmental disturbances and is more common in populations under stress and can therefore be informative about the quality of conditions a dinosaur lived under.

Ichnology
Edwin H. Colbert has suggested that the theropod footprints referred to the ichnogenus Grallator, located in the Connecticut River Valley across Connecticut and Massachusetts, may have been made by Coelophysis. The footprints are from the Late Triassic to Early Jurassic aged Newark Supergroup. They clearly show digits II, III, and IV, but not I or V. That condition is strange for footprints of their age. The digits I and V were presumed to be stubby and ineffective, not touching the ground when the dinosaur was walking or running. They have been thought to be from an unidentified, primitive saurischian similar to Coelophysis by David B. Weishampel and L. Young more recently. Skeletal remains resembling Coelophysis have also been found in the valley, supporting the idea that a species similar to Coelophysis is responsible for the footprints.

Paleoenvironment
Specimens of Coelophysis have been recovered from the Chinle Formation of New Mexico and Arizona, more famously at the Ghost Ranch (Whitaker) quarry in the Rock Point member among other quarries in the underlying Petrified Forest member, the sediments of which have been dated to approximately 212 million years ago, making them part of the middle Norian stage of the Late Triassic, but Thomas Holtz Jr. interpreted that it was during the Rhaetian stage from approximately 204 to 201.6 million years ago.

C. rhodesiensis has been recovered in the Upper Elliott Formation in the Cape and Free State provinces of South Africa, as well as the Chitake River bonebed quarry at the Forest Sandstone Formation in Zimbabwe.

Ghost Ranch was located close to the equator over 200 million years ago, and had a warm, monsoon-like climate with heavy seasonal precipitation. Hayden Quarry, a new excavation site at Ghost Ranch, New Mexico, has yielded a diverse collection of fossil material that included the first evidence of dinosaurs and less-advanced dinosauromorphs from the same time period. The discovery indicates that the two groups lived together during the early Triassic period 235 million years ago.

Therrien and Fastovsky (2001) examined the paleoenvironment of Coelophysis and other early theropods from Petrified Forest National Park in Arizona and determined that this genus lived in an environment that consisted of floodplains marked by distinct dry and wet seasons. There was a great deal of competition during drier times when animals struggled for water in riverbeds that were drying up.

In the upper sections of the Chinle Formation where Coelophysis is found, dinosaurs were rare. So far, only Chindesaurus and Daemonosaurus are known, the terrestrial fauna being dominated instead by other reptiles like the rhynchocephalian Whitakersaurus, the pseudosuchian Revueltosaurus, the aetosaurs Desmatosuchus, Typothorax and Heliocanthus, the crocodilomorph Hesperosuchus, the "rauisuchians" Shuvosaurus, Effigia, and Vivaron, along with other rare components like the dinosauriform Eucoelophysis and the amniote Kraterokheirodon. In the waterways there are the phytosaur Machaeroprosopus, the archosauromorph Vancleavea, the amphibians Apachesaurus and Koskinonodon, and the fishes Reticulodus, Arganodus, and Lasalichthyes.

Taphonomy
The multitude of specimens deposited so closely together at Ghost Ranch was probably the result of a flash flood that swept away a large number of Coelophysis and buried them quickly and simultaneously. In fact, it seems that such flooding was commonplace during this period of the Earth's history and, indeed, the Petrified Forest of nearby Arizona is the result of a preserved log jam of tree trunks that were caught in one such flood. Whitaker quarry at Ghost Ranch is considered a monotaxic site because it features multiple individuals of a single taxon. The quality of preservation and the ontogenic (age) range of the specimens helped make Coelophysis one of the best known of all genera. In 2009, Rinehart et al. noted that in one case the Coelophysis specimens were "washed into a topographic low containing a small pond, where they probably drowned and were buried by a sheet flood event from a nearby river."

The 30 specimens of C. rhodesiensis found together in Zimbabwe was also probably the result of a flash flood that swept away a large number of Coelophysis and buried them quickly and simultaneously as well.

Cultural significance
Coelophysis was the second dinosaur in space, following Maiasaura (STS-51-F). A Coelophysis skull from the Carnegie Museum of Natural History was aboard the Space Shuttle Endeavour mission STS-89 when it left the atmosphere on 22 January 1998. It was also taken onto the space station Mir before being returned to Earth.

Being over 100 years old, Coelophysis is one of the best-known dinosaurs in literature. It was designated as the official state fossil of New Mexico in 1981 and is now the logo of the New Mexico Museum of Natural History.

title: Coelophysis


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Hadrosauridae

Hadrosaurids, or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod family, which includes genera such as Edmontosaurus and Parasaurolophus, was a common group of herbivores during the Late Cretaceous Period. Hadrosaurids are descendants of the Late Jurassic/Early Cretaceous iguanodontian dinosaurs and had a similar body layout. Hadrosaurs were among the most dominant herbivores during the Late Cretaceous in Asia and North America, and during the close of the Cretaceous several lineages dispersed into Europe, Africa, and South America.

Like other ornithischians, hadrosaurids had a predentary bone and a pubic bone which was positioned backwards in the pelvis. Unlike more primitive iguanodonts, the teeth of hadrosaurids are stacked into complex structures known as dental batteries, which acted as effective grinding surfaces. Hadrosauridae is divided into two principal subfamilies: the lambeosaurines (Lambeosaurinae), which had hollow cranial crests or tubes; and the saurolophines (Saurolophinae), identified as hadrosaurines (Hadrosaurinae) in most pre-2010 works, which lacked hollow cranial crests (solid crests were present in some forms). Saurolophines tended to be bulkier than lambeosaurines. Lambeosaurines included the aralosaurins, tsintaosaurins, lambeosaurins and parasaurolophins, while saurolophines included the brachylophosaurins, kritosaurins, saurolophins and edmontosaurins.

Hadrosaurids were facultative bipeds, with the young of some species walking mostly on two legs and the adults walking mostly on four.

History of discovery
Ferdinand Vandeveer Hayden, during expeditions near the Judith River in 1854 through 1856, discovered the very first dinosaur fossils recognized from North America. These specimens were obtained by Joseph Leidy, who described and named them in 1856; two of the several species named were Trachodon mirabilis of the Judith River Formation and Thespesius occidentalis of the "Great Lignite Formation". The former was based on a collection of teeth whilst the latter on two  and a. Although most of the Trachodon teeth turned out to belong to ceratopsids, the holotype and remains of T. occidentalis would come to be recognized as the first recognized hadrosaur specimens. Around the same time in Philadelphia, on the other side of the continent, geologist William Parker Foulke was informed of numerous large bones accidentally uncovered by farmer John E. Hopkins some twenty years earlier. Foulke obtained permission to investigate the now scattered fossils in 1858, and these specimens as well were given to Leidy. They were described in the same year as Hadrosaurus foulkii, giving a slightly better picture of the form of a hadrosaur. Leidy provided additional description in a 1865 paper. Among his 1858 work Leidy briefly suggested that the animal was likely amphibious in nature; this school of thought about hadrosaurs would come to be dominant for over a century to come.

Further discoveries such as "Hadrosaurus minor" and "Ornithotarsus immanis" would come from the East, and Edward Drinker Cope led an expedition to the Judith River Formation where Trachodon was found. Upon the fragments discovered he named seven new species in two genera, as well as assigning material to Hadrosaurus. Cope had studied the jaws of hadrosaurs and come to the conclusion that the teeth were fragile and could have been dislodged incredibly easily. As such, he supposed the animals must have fed largely on soft water plants; he presented this idea to the Philadelphia Academy in 1883, and this idea would come to be very influential on future study. Research would continue in the Judith River area for years to come, but the formation never yielded much more than fragmentary remains, and Cope's species as well as Trachodon itself would in time be seen as of doubtful validity. The Eastern states, too, would never yield particularly informative specimens. Instead, other sites in the American West would come to provide many very complete specimens that would form the backbone of hadrosaur research. One such specimen was the very complete AMNH 5060 (belonging to Edmontosaurus annectens), recovered in 1908 by the fossil collector Charles Hazelius Sternberg and his three sons in Converse County, Wyoming. It was described by Henry Osborn in 1912, who dubbed it the "Dinosaur mummy". This specimen's skin was almost completely preserved in the form of impressions. The skin around its hands, thought to represent webbing, was seen as further bolstering the idea that hadrosaurs were very aquatic animals.

Cope had planned to write a monograph about the group Ornithopoda, but never made much progress towards it before his death. This unrealized endeavor would come to be the inspiration for Richard Swann Lull and Nelda Wright to work on a similar project decades later. Eventually they realized the whole of Ornithopoda was too broad of a scope, until eventually it was narrowed down to specifically North American hadrosaurs. Their monograph, Hadrosaurian Dinosaurs of North America, was published in 1942, and looked back at the whole of understanding about the family. It was designed as a definitive work, covering all aspects of their biology and evolution, and as part of it every known species was re-evaluated and many of them redescribed. They agreed with prior authors on the semi-aquatic nature of hadrosaurs, but re-evaluated Cope's idea of weak jaws and found quite the opposite. The teeth were rooted in strong batteries and would be continuously replaced to prevent them getting worn down. Such a system seemed incredibly overbuilt for the job of eating soft Mesozoic plants, and this fact confused the authors. Though they still proposed a diet of water plants, they considered it likely this would be supplemented by occasional forrays into browsing on land plants.

Twenty years later, in 1964, another very important work would be published, this time by John H. Ostrom. It challenged the idea that hadrosaurs were semi-aquatic animals, which had been held since the work of Leidy back in the 1850s. This new approach was backed using evidence of the environment and climate they lived in, co-existing flora and fauna, physical anatomy, and preserved stomach contents from mummies. Based on evaluation of all this data, Ostrom found the idea that hadrosaurs were adapted for aquatic life incredibly lacking, and instead proposed they were capable terrestrial animals that browsed on plants such as conifers. He remained uncertain, however, as to the purpose of the paddle-like hand Osborn had described, as well as their long and somewhat paddle-like tails. Thus he agreed with the idea that hadrosaurs would have taken refuge from predators in water. Numerous important studies would follow this; Ostrom's student Peter Dodson published a paper about lambeosaur skull anatomy that made enormous changes to hadrosaur taxonomy in 1975, and Michael K. Brett-Surman conducted a full revision of the group as part of his Graduate studies through the 1970s and 1980s. John R. Horner would also begin to leave his impact on the field, including with the naming of Maiasaura in 1979.

Hadrosaur research experienced a surge in the decade of the 2000s, similar to the research of other dinosaurs. In response to this, the Royal Ontario Museum and the Royal Tyrrell Museum collaborated to arrange the International Hadrosaur Symposium, a professional meeting about ongoing hadrosaur research that was held at the latter institution on September 22 and 23 in 2011. Over fifty presentations were made at the event, thirty-six of which were later incorporated into a book, titled Hadrosaurs, published in 2015. The volume was brought together primarily by palaeontologists David A. Eberth and David C. Evans, and featured an afterword from John R. Horner, all of whom also contributed to one or more of the studies published therein. The first chapter of the volume was a study by David B. Weishampel about the rate of ornithopod research over history, and the interest in different aspects of it over that history, using the 2004 volume The Dinosauria as the source of data on the amount of works published in each decade. Various periods of high and low activity were found, but the twenty-first century was found to overwhelmingly be the most prolific time, with over two-hundred papers published. The advent of the internet was cited as a likely catalyst for this boom. Hadrosaur research experienced high levels of diversity within the decade, with previously uncommon subjects such as growth, phylogeny, and biogeography experiencing more attention, though the functional morphology of hadrosaurids was found to have declined in study since the Dinosaur Renaissance.

Distribution
Hadrosaurids likely originated in North America, before shortly dispersing into Asia. During the late Campanian-Maastrichtian, a saurolophine hadrosaurid migrated into South America from North America, giving rise to the clade Austrokritosauria, which is closely related to the tribe Kritosaurini. During the late early Maastrichtian, several lineages of Lambeosaurinae from Asia migrated into the European Ibero-Armorican Island (what is now France and Spain), including Arenysaurini and Tsintaosaurini. One of these lineages later dispersed from Europe into North Africa, as evidenced by Ajnabia, a member of Arenysaurini.

Classification
The family Hadrosauridae was first used by Edward Drinker Cope in 1869, then containing only Hadrosaurus. Since its creation, a major division has been recognized in the group between the hollow-crested subfamily Lambeosaurinae and the subfamily Saurolophinae, historically known as Hadrosaurinae. Both of these have been robustly supported in all recent literature. Phylogenetic analysis has increased the resolution of hadrosaurid relationships considerably, leading to the widespread usage of tribes (a taxonomic unit below subfamily) to describe the finer relationships within each group of hadrosaurids.

Lambeosaurines have also been traditionally split into Parasaurolophini and Lambeosaurini. These terms entered the formal literature in Evans and Reisz's 2007 redescription of Lambeosaurus magnicristatus. Lambeosaurini is defined as all taxa more closely related Lambeosaurus lambei than to Parasaurolophus walkeri, and Parasaurolophini as all those taxa closer to P. walkeri than to L. lambei. In recent years Tsintaosaurini and Aralosaurini have also emerged.

The use of the term Hadrosaurinae was questioned in a comprehensive study of hadrosaurid relationships by Albert Prieto-Márquez in 2010. Prieto-Márquez noted that, though the name Hadrosaurinae had been used for the clade of mostly crestless hadrosaurids by nearly all previous studies, its type species, Hadrosaurus foulkii, has almost always been excluded from the clade that bears its name, in violation of the rules for naming animals set out by the ICZN. Prieto-Márquez defined Hadrosaurinae as just the lineage containing H. foulkii, and used the name Saurolophinae instead for the traditional grouping.

Phylogeny
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. 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.

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.

The following cladogram is from Ramírez-Velasco (2022), including most recently named taxa.

Anatomy
The most recognizable aspect of hadrosaur anatomy is the flattened and laterally stretched rostral bones, which gives the distinct duck-bill look. Some members of the hadrosaurs also had massive crests on their heads, probably for display and/or to make noises. In some genera, including Edmontosaurus, the whole front of the skull was flat and broadened out to form a beak, which was ideal for clipping leaves and twigs from the forests of Asia, Europe and North America. However, the back of the mouth contained thousands of teeth suitable for grinding food before it was swallowed. This has been hypothesized to have been a crucial factor in the success of this group in the Cretaceous compared to the sauropods.

Skin impressions of multiple hadrosaurs have been found. From these impressions, the hadrosaurs were determined to be scaled, and not feathered like some dinosaurs of other groups.

Hadrosaurs, much like sauropods, are noted for having their manus united in a fleshy, often nail-less pad.

The two major divisions of hadrosaurids are differentiated by their cranial ornamentation. While members of the Lambeosaurinae subfamily have hollow crests that differ depending on species, members of the Saurolophinae (Hadrosaurinae) subfamily have solid crests or none at all. Lambeosaurine crests had air chambers that may have produced a distinct sound and meant that their crests could have been used for both an audio and visual display.

Diet
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. 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. However, Purnell said these conclusions were less secure than the more conclusive evidence regarding the motion of teeth while chewing.

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. 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. As a result of that finding, Tweet concluded in September 2008 that the animal was likely a browser, not a grazer. 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.

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 a height of 2 m when standing quadrupedally, and up to a height of 5 m bipedally.

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. Examination of hadrosaur coprolites from the Grand Staircase-Escalante indicates that shellfish such as crustaceans were also an important component of the hadrosaur diet.

It's thought that chewing behaviour may have changed throughout life in hadrosaurids. Very young specimens show simple cup shapes occlusion zones, or areas where the teeth contact one another in chewing, whereas in adulthood there is a "dual function" arrangement with two distinct areas of different tooth wear. This change during growth may have helped transition from a diet of softer plants when young to more tough and fibrous ones during adulthood. It's thought the transition between states, characterized by a more gradual transition from one wear state to another, occurred at different times in the growth of different species; in Hypacrosaurus stebingeri it did not occur until a nearly adult stage, whereas in a saurolophine specimen likely less than a year old from the Dinosaur Park Formation the transition had already begun.

Neurology
Hadrosaurs have been noted as having the most complex brains among ornithopods, and indeed among ornithischian dinosaurs as a whole. The brains of hadrosaurid dinosaurs have been studied as far back at the late 19th century, when Othniel Charles Marsh made an endocast of a specimen then referred to Claosaurus annectens; only basic remarks were possible but it was noted that the organ was proportionally small. John Ostrom would give a more informed analysis and review in 1961, pulling on data from Edmontosaurus regalis, E. annectens, and Gryposaurus notabilis (then considered a synonym of Kritosaurus). Though still obviously small, Ostrom recognized that the brains may be more significantly developed than expected, but supported the view that dinosaur brains would have only filled some of the endocranial cavity, limiting possibility of analysis. In 1977 James Hopson introduced the use of estimated encephalization quotients to the topic of dinosaur intelligence, finding Edmontosaurus to have an EQ of 1.5, above that of other ornithischians including earlier relatives like Camptosaurus and Iguanodon and similar to that of carnosaurian theropods and modern crocodilians but below that of coelurosaurian theropods. Reasonings suggested for their comparably high intelligence were the need for acute senses in the lack of defensive weapons, and more complex intraspecific behaviours as indicated by their acoustic and visual display structures.

The advent of CT scanning for use in palaeontology has allowed for more widespread application of this without the need for specimen destruction. Modern research using these methods has focused largely on hadrosaurs. In a 2009 study by palaeontologist David C. Evans and colleagues, the brains of lambeosaurine hadrosaur genera Hypacrosaurus (adult specimen ROM 702), Corythosaurus (juvenile specimen ROM 759 and subadult specimen CMN 34825), and Lambeosaurus (juvenile specimen ROM 758) were scanned and compared to each other (on a phylogenetic and ontogenetic level), related taxa, and previous predictions, the first such large-scale look into the neurology of the subfamily. Contra the early works, Evans' studies indicate that only some regions of the hadrosaur brain (the dorsal portion and much of the hindbrain) were loosely correlated to the brain wall, like modern reptiles, with the ventral and lateral regions correlating fairly closely. Also unlike modern reptiles, the brains of the juveniles did not seem to correlate any closer to the brain wall than those of adults. It was cautioned, however, that very young individuals were not included in the study. As with previous studies, EQ values were investigated, although a wider number range was given to account for uncertainty in brain and body mass. The range for the adult Hypacrosaurus was 2.3 to 3.7; the lowest end of this range was still higher than modern reptiles and most non-maniraptoran dinosaurs (nearly all having EQs below two), but fell well short of maniraptorans themselves, which had quotients higher than four. The size of the cerebral hemispheres was, for the first time, remarked upon. It was found to taking up around 43% of endocranial volume (not considering olfactory bulbs) in ROM 702. This is comparable to their size in saurolophine hadrosaurs, but far larger than in any ornithischians outside of Hadrosauriformes, and all large saurischian dinosaurs; maniraptors Conchoraptor and Archaeopteryx, an early bird, had very similar proportions. This lends further support to the idea of complex behaviours and relatively high intelligence, for non-avian dinosaurs, in hadrosaurids.

Amurosaurus, a close relative of the taxa from the 2009 study, was the subject of a 2013 paper once again looking into a cranial endocast. A nearly identical EQ range of 2.3 to 3.8 was found, and it was again noted this was higher than that of living reptiles, sauropods and other ornithischians, but different EQ estimates for theropods were cited, placing the hadrosaur numbers significantly below even more basal theropods like Ceratosaurus (with an EQ range of 3.31 to 5.07) and Allosaurus (with a range of 2.4 to 5.24, compared to only 1.6 in the 2009 study); more bird-like coelurosaurians theropods such as Troodon had stated EQs higher than seven. Additionally, the relative cerebral volume was only 30% in Amurosaurus, significantly lower than in Hypacrosaurus, closer to that of theropods like Tyrannosaurus (with 33%), though still distinctly larger than previously estimated numbers for more primitive iguanodonts like Lurdusaurus and Iguanodon (both at 19%). This demonstrated a previously unrecognized level of variation in neuro-anatomy within Hadrosauridae.

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

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.

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.

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.

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

Evidence suggests that young hadrosaurs would have walked on only their two hind legs, while adults would have walked on all four. 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. Furthermore, the animals' front limbs were shorter than their back limbs.

Daily activity patterns
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.

Pathology
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 Langerhans cell histiocytosis, hemangiomas, desmoplastic fibroma, metastatic cancer, and osteoblastomas, found in genera such as Brachylophosaurus and Edmontosaurus. Osteochondrosis is also commonly found in hadrosaurs.

license: Creative Commons - Attribution Share-Alike - https://creativecommons.org/licenses/by-sa/4.0/

title: Hadrosauridae


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Barapasaurus

Barapasaurus is a genus of basal sauropod dinosaur from Jurassic rocks of India. The only species is B. tagorei. Barapasaurus comes from the lower part of the Kota Formation, which is of Early to Middle Jurassic age. It is therefore one of the earliest known sauropods. Barapasaurus is known from approximately 300 bones from at least six individuals, so that the skeleton is almost completely known except for the anterior cervical vertebrae and the skull. This makes Barapasaurus one of the most completely known sauropods from the early Jurassic.

Discovery and naming
All known fossils come from a single locality in the vicinity of the village of Pochampally Pin Code: 442504, bordering Telangana (Pochampally Sironcha Taluka, Gadchiroli District, Vidarbha, Maharastra, in central India). The first bones were discovered in 1958, but most specimens were unearthed in 1960 and 1961. In 1975, the finds were described scientifically by palaeontologist Sohan Lal Jain and colleagues. In 2010, a more detailed osteological description was published by Bandyopadhyay and colleagues. The material is archived in the palaeontological collection of the Indian Statistical Institute (ISI), while a majority of the bones are part of a mount at the Geological Museum of the ISI.

Etymology
The name Barapasaurus ("big-legged lizard") is derived from bara meaning 'big' and pa meaning 'leg' in several Indian languages including Bengali; the Greek word sauros means 'lizard'. This name was used as a nomen nudum since a femur measuring over 1.7 m was unearthed at 1961. The specific name tagorei means 'Tagore's', which honours Bengali poet, writer, painter, and musician Rabindranath Tagore. The first year of fieldwork was carried out in the centenary year of Tagore's birth.

Fossil record
Barapasaurus tagorei is known from a large bone bed found in the Lower Kota Formation, containing approximately 300 bones from at least six individuals of various ages. No remains of the skull, other than teeth, have been found. The age of this fossil site is disputed, because no volcanic rocks whose age can be determined by radiometric dating are associated with the Kota Formation and its age can only be estimated by biostratigraphic comparisons to other rock layers, which is made difficult by the lack of reliable index fossils in the formation. The age has generally been interpreted as Early Jurassic, with the Lower Kota Formation in particular dating to the Sinemurian to Pliensbachian, roughly 184–200 million years ago. However, some studies have argued that the Kota Formation dates to the Middle Jurassic or even later.

Taphonomy
The approximately 300 bones were found together with large trunks of trees scattered over an area of 276 square meters. Although one of the specimens was found partly articulated, most bones were found disarticulated. Because there are six left femora, the total number of individuals is at least six.

Bandyopadhyay and colleagues (2002, 2010) interpret this assemblage as a herd that died due to a catastrophic event, likely a flood. This flood could have unearthed the trees and transported both trees and Barapasaurus a distance before they began to decompose. After decomposition progressed, the bones began to disarticulate. The disarticulated skull bones were removed by the water stream because they were light, leaving only the heavy postcranial bones at the site, which would explain why no skull bones were found.

Description
Although a very early and unspecialised sauropod, Barapasaurus shows the building plan typical for later, more derived sauropods: the cervical vertebrae were elongated, resulting in a long neck. The trunk was short and holds columnar limbs which indicate an obligate quadrupedal posture. Even the size, which is estimated at approximately 12-14 meters long and 7 tonnes in weight, is comparable with that of later sauropods.

The vertebral column already shows many traits that are typical for later sauropods which allowed them to attain great body sizes, although in later sauropods these traits are much more developed. The central and neural spines show early hints of hollowing as a weight-saving measure. The dorsal vertebrae are stabilised with hyposphene-hypantrum articulations, accessory projections that link the vertebrae with each other. The sacrum is strengthened through an additional fourth sacral vertebra.

From the skull, only three whole teeth and three crowns are known. The largest known tooth is 5.8 cm in height. Like that of later sauropods, the teeth are spoon shaped and show wrinkled enamel. A basal trait is the coarse serration.

Classification
The relationships of this genus within the Sauropoda are debated. When first described in 1975, it was not attributed to one specific group at all, although the presence of many basal, prosauropod-like features was noted. Since 1984, Barapasaurus was united with another early sauropod, Vulcanodon, in a family called Vulcanodontidae, although this family was declared invalid by Paul Upchurch in 1995 because it was recognised as polyphyletic. Upchurch erected a clade named Eusauropoda that includes all known sauropods except some very basal forms. While Vulcanodon was classified outside the Eusauropoda, Barapasaurus was classified inside it, which means that Barapasaurus is more derived than Vulcanodon. Although Upchurch's classification was accepted by most palaeontologists, a recent study from Bandyopadhyay and colleagues came to a contrary conclusion: these palaeontologists stated that Barapasaurus was in fact more basal than Vulcanodon and removed it from Eusauropoda.

Habitat
Barapasaurus comes from clay and sandstone that belongs to the lower part of the Kota Formation. Other vertebrates of this part include another early sauropod, Kotasaurus, as well as the mammals Kotatherium, Indotherium and Indozostrodon. The upper part of the Kota Formation additionally contained a pterosaur (Campylognathoides), a turtle, two rhynchocephalians, a lepidosaur and some mammals.

title: Barapasaurus


text:
User talk:Irielion

This is an Rasta term. Irie means happy and lion stands for Him Imperial Majesty, Haile Selassi I and therefore stands for Rastafari. You can interpret the term as "Happy Man"

photo
Thanks for uploading the photo of Compay and Fidel. Alas, you neglected to indicate your source and its copyright status. As you may be aware, we have to deal with copyright issues, as a big a nuisance as they are. If the copyright status is not clarified, it may be deleted. Please add this info ASAP. -- Viajero 08:04, 9 Apr 2005 (UTC)

Image Tagging Image:Compay y castro.jpg
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title: User talk:Irielion


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Boundary Street

Boundary Street is a three-lane one-way street in Kowloon, Hong Kong. It runs in an easterly direction from its start at the intersection with Tung Chau Street in the west, and ends at its intersection with Prince Edward Road West in the east, near the former Kai Tak Airport.

History
The street previously marked the boundary between the southern part of Kowloon, ceded by the Qing dynasty to Great Britain in 1860 along with Stonecutters Island, and the northern part of Kowloon (New Kowloon) (which remained part of China until it was leased as part of the New Territories to the United Kingdom in 1898 for 99 years under the Second Convention of Peking). After the lease, the boundary was renamed from Boundary Line to Old Frontier Line.

The boundary was marked by a long line of tall bamboo fences, which effectively blocked smuggling between Chinese Kowloon and British Kowloon at that time. The barrier became obsolete when the New Territories joined the colony.

Although the street marks the historical boundary, the road did not come to exist until 1934, more than 30 years after the lease of boundary north. The road was built to accelerate the development of Kowloon Tong and identify the difference in calculation of rates between the boundary north and south.

Present
In everyday life, New Kowloon is no longer regarded as part of the New Territories, but as part of the Kowloon urban area (on both sides of Boundary Street), except for statutory and land rent proposes, where the original arrangement has remained in place.

Administratively its portion west of the East Rail line forms part of the boundary between Yau Tsim Mong and Sham Shui Po districts, until the former western coast of the peninsula, from whereupon the boundary follows the streets on the reclamation.

On 1 July 1997, sovereignty of the territories on both sides of Boundary Street was handed over from the United Kingdom to the People's Republic of China, along with the rest of Hong Kong.

Notable places along the street

 * Fa Hui Park
 * La Salle Primary School
 * Maryknoll Convent School
 * Sycamore Street Playground

title: Boundary Street


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Tanner

Tanner may refer to:

 * Tanner (occupation), the tanning of leather and hides

People

 * Tanner (given name)
 * Tanner (surname), a surname (including a list of people with the name)
 * The Tanner Sisters, also referred to as "The Harbingers of Weirdness"
 * Simon the Tanner (New Testament), a Jaffa resident where Saint Peter stayed, as told in the New Testament
 * Simon the Tanner (10th century), a Coptic Orthodox saint
 * Theodotus the Tanner (fl. late 2nd century), an early Christian writer from Byzantium

Populated places

 * In the United States
 * Tanner, Alabama, an unincorporated community
 * Tanner, Georgia, an unincorporated community
 * Tanner, Indiana, an unincorporated community
 * Tanner, Kentucky, an unincorporated community
 * Tanner, Missouri, an unincorporated community
 * Tanner, Washington, a census-designated place
 * Tanner, West Virginia, an unincorporated community
 * Tanner Township, Kidder County, North Dakota, a civil township

 * Elsewhere
 * Tanner's Settlement, Lunenburg County, Nova Scotia, Canada

Other places

 * Tanner Block, a historic building in Oswego, New York, United States
 * Tanner Farmhouse, a historic residence near Wilmer, Alabama, United States
 * Tanner High School, a school in Limestone County, Alabama, United States
 * Tanner Hill Estate, a public housing estate in Tanner Hill Road, North Point, Hong Kong
 * Tanner–Hiller Airport, a public airport in Barre, Massachusetts, United States
 * Tanner Island, the westernmost and largest of the Pickersgill Islands, off the south coast of South Georgia
 * Tanner Moor, a moorland in Austria
 * Tanner Trail, a hiking trail located on the South Rim of the Grand Canyon National Park in Arizona, United States

Popular culture

 * Tanner (film), a 1985 Swiss film
 * Tanner '88, a television series about a fictional US presidential nominee
 * Tanner, fictional video game character and protagonist of the Driver series
 * Tanner on Tanner, a 2004 comedy
 * "Mr. Tanner", a song by Harry Chapin from his 1974 album Short Stories
 * Tanner (band), an American band active in the 1990s

Other uses

 * Tanner, the sixpence, a British pre-decimal coin
 * 13668 Tanner, a main-belt asteroid discovered on 28 April 1997
 * Tanner crab (Chionoecetes bairdi), a crab species
 * Tanner Cup, a New Zealand sailing competition
 * Tanner graph, a bipartite graph used to state constraints or equations which specify error correcting codes
 * Tanner scale, a scale of physical development in children, adolescents and adults
 * USNS Tanner, a ship
 * USS Tanner (AGS-15), an attack cargo ship of the U.S. Navy
 * Prionus coriarius, the tanner beetle

title: Tanner


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Claude Closky

Claude Closky (born 22 May 1963) is a French Contemporary Artist who lives and works in Paris, France.

Reception
Closky won the "Grand prix des Arts plastiques" (1999) and the Marcel Duchamp Prize (2005) awarded by the ADIAF.

Dike Blair wrote in Artforum Magazine that "The lightness of Closky's art belies the depth of its absurdist heredity. Working in a post-modernist mode, Closky's art works combine aspects of the Situationists, Fluxus, Beckett, Tall, Daniel Buren, and Andy Warhol."

Life and career
Claude Closky has no formal training as a visual artist. He entered the ENSAD (Ecole National Supérieure des Arts Décoratifs) in 1982, but quit at the end of the first year to co-found The Ripoulin Brothers, a street artist collective, with Bla+Bla+Bla, Nina Childress, Jean Faucheur, Pierre Huyghe, Manhu, Ox, Trois carrés. In 1988, he left the group to develop his independent work, using voluntarily poor means such as drawing and printed matter.

He has participated in the Biennials of Lyon (1995), Sydney (1996), Taipei (2000), València (2001), Sharjah (2005), Venice (2017), Mardin (2024). In 2000, the Mudam (Luxembourg Art Museum) commissioned him to conceive and manage its website, for which he made a magazine and a gallery dedicated to the internet. The site was launched at the Luxembourg Pavilion in the 2001, Venice Biennale. It presented specific works by Heath Bunting, François Curlet, Pierre Leguillon, Aleksandra Mir, Peter Kogler, David Shrigley, among others. Since 2005, he has taught at the École nationale supérieure des Beaux-Arts in Paris. In 2012, he curated This & There, an exhibition to celebrate the tenth year of the Pavilion, Palais de Tokyo Laboratory for Creation (Paris) which presented the work of 74 artists in 74 different spaces. He has curated in 2020 X at the Frac Pays de la Loire and in 2023 Offset at the Centre des livres d'artistes.

Work
The work of Claude Closky is mainly immaterial. Language is his model to articulate images, text, numbers, and sounds collected in our environment, or made in his studio. Although Closky is reluctant to produce objects and spectacular effects, his work still addresses issues about visibility and space appropriation.

Claude Closky's projects always find alternative ways to emancipate themselves from the formats imposed by the sites where they are exhibited. He seeks to point out the contradictions of our contemporary society and its representations, but also to question the role of art as producer of a cultural consensus and set of values. His works confront and question our environment, the conditions and benefits of artistic production, its relation to an audience.

title: Claude Closky


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History of Hong Kong (1800s–1930s)

Hong Kong (1800s–1930s) oversaw the founding of the new crown colony of Hong Kong under the British Empire. After the First Opium War, the territory was ceded by the Qing Empire to the United Kingdom of Great Britain and Ireland through Treaty of Nanjing (1842) and Convention of Peking (1860) in perpetuity. Together with additional land that was leased to the British under the Convention for the Extension of Hong Kong Territory (1898), Hong Kong became one of the first parts of East Asia to undergo industrialisation.

Beginning of trade

China was the main supplier of its native tea to the British, whose annual domestic consumption reached 30050000 lb in 1830, an average of 1.04 lb per head of population.

From the British economic standpoint, Chinese tea was a crucial item since it provided massive wealth for the taipans—foreign (especially British) businessmen in China—while the duty on tea accounted for 10% of the government's income. Some of the earliest items sold to China in exchange for tea were British clocks, watches and musical boxes known as "sing-songs". However, these were not enough to compensate for the trade imbalance and the insistence by the Chinese that they be paid in silver. Opium exports from India after 1830 provided the silver needed to balance the trade. Lin Zexu, a special Chinese commissioner appointed by the Qing Daoguang Emperor, wrote a letter to Queen Victoria in 1839 taking a stance against the acceptance of opium in trade. He confiscated more than 20,000 chests of opium already in Hong Kong, which had already been used years earlier as a transhipment point, and supervised their destruction.

Confrontation
London saw the destruction of British products as an insult and sent the first expeditionary force to the region. The First Opium War (1839–1842) began at the hands of Captain Charles Elliot of the Royal Navy and Capt. Anthony Blaxland Stransham of the Royal Marines. After a series of Chinese defeats, Hong Kong Island was occupied by the British on 20 January 1841. Sir Edward Belcher, aboard HMS Sulphur, landed in Hong Kong on 25 January 1841. Possession Street still exists to mark the event. Commodore Sir Gordon Bremer raised the Union Jack and claimed Hong Kong as a colony on 26 January 1841. He erected naval store sheds there in April 1841.

The island was first used by the British as a staging post during the war, and while the East India Company intended to establish a permanent base on the island of Zhoushan, Elliot took it upon himself to claim the island on a permanent basis. The ostensible authority for the occupation was negotiated between Captain Eliot and the Viceroy of Liangguang, the Manchu official Qishan. The Convention of Chuenpi was concluded but had not been recognised by the Qing Dynasty court at Beijing. Subsequently, Hong Kong Island was ceded to Britain in 1842 under the Treaty of Nanking, when the territory became a Crown colony.

The Opium War was ostensibly fought to liberalise trade with China. With a base in Hong Kong, British traders, opium dealers, and merchants including Jardine Matheson & Co. and Dent & Co. launched the city which would become the 'free trade' nexus of the East. American opium traders and merchant bankers such as the Russell, Perkins and Forbes families would soon join the trade.

On signature of the 1860 Convention of Peking, which marked the end of formal ended hostilities in the Second Opium War (1856–1858), Britain acquired the area south of Boundary Street on the Kowloon Peninsula rent-free under a perpetual lease. Later, in 1898, the Qing government reluctantly agreed to the Convention between Great Britain and China Respecting an Extension of Hong Kong Territory (also known as the Second Convention of Peking) that compelled China to cede a further area north of Boundary Street to the Sham Chun River along with more than two hundred nearby islands. Seen by the British government as vital to safeguard the defensive capabilities of Hong Kong, these areas became collectively known as the New Territories. The 99-year lease would expire at midnight on 30June, 1997.

Population
When the union flag was raised over Possession Point on 26 January 1841, the population of Hong Kong island was about 6,000, mostly Tanka fishermen and Hakka charcoal burners living in a number of coastal villages. In the 1850s large numbers of Chinese would emigrate from China to Hong Kong due to the Taiping Rebellion. Other events such as floods, typhoons and famine in mainland China would also play a role in establishing Hong Kong as a place to escape the mayhem. According to the census of 1865, Hong Kong had a population of 125,504, of which some 2,000 were Americans and Europeans. In 1914 despite an exodus of 60,000 Chinese fearing an attack on the colony during World War I, Hong Kong's population continued to increase from 530,000 in 1916 to 725,000 in 1925 and 1.6 million by 1941.

Segregation
The establishment of the free port made Hong Kong a major entrepôt from the start, attracting people from China and Europe alike. The society remained racially segregated and polarised due to British colonial policies and attitudes. Despite the rise of a British-educated Chinese upper class by the late 19th century, race laws such as the Peak Reservation Ordinance prevented Chinese from living in elite areas like Victoria Peak. Politically, the majority Chinese population also had little to no official governmental influence throughout much of the early years. There were, however, a small number of Chinese elites that the British governors relied on, including Sir Kai Ho and Robert Hotung. They accepted their place in the Hong Kong hierarchy, and served as main communicators and mediators between the government and the Chinese population. Sir Kai Ho was an unofficial member of the Legislative Council. Robert Hotung wanted Chinese citizens to recognise Hong Kong as the new home after the fall of China's last dynasty in 1911. As a millionaire with financial influence, he emphasised that no part of the demographics was purely indigenous.

Lifestyle

The east portion of Colonial Hong Kong was mostly dedicated to the British; filled with race courses, parade grounds, barracks, cricket and polo fields. The west portion was filled with Chinese shops, crowded markets and tea houses. The Hong Kong tea culture began in this period and evolved into yum cha. One of the most common breakfasts was congee with fish and barley. In the mid-19th century many of the merchants would sell silk, jade and consult feng shui to open shops that favoured better spiritual arrangements. Other lower ranked groups like coolies arrived with the notion that hard work would better position them for the future. Due to the commercial success of merchants, boatmen, carters and fishermen there, Hong Kong overtook China's most populous port in Canton. By 1880 Hong Kong's port would handle 27% of the mainland's export and 37% of imports. A British traveller, Isabella Bird, described Hong Kong in the 1870s as a colony filled with comforts and entertainment only a Victorian society would be able to enjoy. Other descriptions mentioned courts, hotels, post offices, shops, city hall complexes, museums, libraries and structures in impressive manner for the era. Many European businessmen went to Hong Kong to do business. They were referred to as tai-pans or "bigshot". One of the more notable Tai-pan hangout spots was the Hong Kong Club at Queen's Road.

Education
In 1861, Frederick Stewart would become the founder of Hong Kong education system bringing Western-style pedagogy to the East. Some have argued that his contribution is the key turning point between the group of Chinese that were able to modernise Hong Kong versus the group that did not in China. The education would bring Western-style finance, science, history, technology into the culture. The father of modern China, Sun Yat-sen was also educated in Hong Kong's Central School.

Law and order
In 1843 the legislative council was established. The governor of Hong Kong generally served as the British plenipotentiary in the far east in the early years. The Colonial Secretary would also assist in legal matters. A colonial police force was established in the 1840s to handle the high crime rate in Hong Kong. By China's standards, colonial Hong Kong's code of punishment was considered laughably loose and lenient. The lack of intimidation may have been the leading cause for the continual rise in crime. Po Leung Kuk became one of the first organisations established to deal with the abduction of women and prostitution crisis. Crime at sea was also common as some pirates had access to cutlasses and revolvers. Court sessions for criminal and admiralty matters were first held on 4March 1844 under the aegis of the first governor, Lieutenant General Sir Henry Pottinger and the Lieutenant-governor George D'Aguilar.

Pandemics and disasters
The Third Pandemic of bubonic plague broke out in China in the 1880s. By the spring of 1894 about 100,000 were reported dead in the mainland. In May 1894 the disease erupted into Hong Kong's overcrowded Chinese quarter of Tai Ping Shan. By the end of the month, an estimated 450 people died of the illness. At its height, the epidemic was killing 100 people per day, and it killed a total of 2,552 people that year. The disease was greatly detrimental to trade and produced a temporary exodus of 100,000 Chinese from the colony. Plague continued to be a problem in the territory for the next 30 years.

In the 1870s a typhoon hit Hong Kong one evening, reaching its height by midnight. An estimated 2,000 people lost their lives in a span of just six hours.

Transport
The growth of Hong Kong depended greatly on domestic transport of citizens and cargo across Victoria Harbour. The establishment of the Star Ferry and the Yaumati Ferry would prove to be vital. In 1843 the colony had built the first ship at a private shipyard. Some of the customers later included the Spanish government in the Philippines and the Chinese navy. The Peak Tram would begin in 1888 along with the Tramway service in 1904. The first railway line was also launched in 1910 as the Kowloon-Canton Railway. On land the rickshaws were extremely popular when they were first imported from Japan in 1874, since it was affordable and necessary for street merchants to haul goods. Sedan chairs were the preferred mode of the transport for the wealthy Europeans who lived on Victoria Peak due to the steep grade which ruled out rickshaws until the introduction of the Peak Tram. The first automobiles in Hong Kong had petrol-driven internal combustion engines and arrived between 1903 and 1905. Initially they were not well received by the public. Only around 1910 did the cars begin to gain appeal. Most of the owners were British. Buses operated by various independent companies flourished in the 1920s until the government formally issued franchises for the China Motor Bus and Kowloon Motor Bus companies in 1933. The flying boats were the first British aeroplanes to reach Hong Kong in 1928. By 1924 the Kai Tak Airport would also be found. The first flight service from Imperial Airways would become available by 1937 at a price of 288 pounds per ticket.

Hospitals and hospitality
Soon after the British occupied Hong Kong in 1841, Protestant and Catholic missionaries started to provide social service. Italian missionaries began to provide boy-only education to British and Chinese youth in 1843. "The Catholic French Sisters of St. Paul de Chartres" was one of the first orphanage and elderly home was established in 1848.

In 1870 the Tung Wah Hospital became the first official hospital in Hong Kong. It handled much of the social services and was providing free vaccinations in Hong Kong Island and Kwang Tung. After raising funds for the 1877 famine in China, a number of the hospital officials became Tung Wah elites with much authority and power representing the Chinese majority. Some of the booming hotel businesses of the era included the Victoria Hotel, New Victoria Hotel and the King Edward Hotel.

Finance
In 1864 the first large scale modern bank Hong Kong Shanghai Bank would be established turning Hong Kong into the focal point of financial affairs in Asia. Its chief manager, Sir Thomas Jackson, has a statue in Statue Square. The bank first leased Wardley House at HKD 500 a month in 1864. After raising a capital of HKD 5 million, the bank opened its door in 1865. The Association of Stockbrokers would also be established in 1891.

Resources
In December 1890 the Hongkong Electric company went into production with help from Catchick Paul Chater. It was the first step in allowing the transition of gas lamps to light bulbs. Other companies like Jardine Matheson would launch the "Hong Kong Land Investment and Agency company Ltd" accumulating a wealth as large as the entire government's total revenue. (See also China Light and Power.)

Politics

One observer summed up the decades as "politics, propaganda, panic, rumour, riot, revolution and refugees". The role of Hong Kong as a political safe haven for Chinese political refugees further cemented its status, and few serious attempts to revert its ownership were launched in the early 20th century. Both Chinese Communist and Nationalist agitators found refuge in the territory, when they did not actively participate in the turmoil in China. However, the dockworkers strikes in the 1920s and 1930s were widely attributed to the Communists by the authorities, and caused a backlash against them. A strike in 1920 was ended with a wage increase of HKD 32 cents. Ambrose King, in his controversial 1975 paper Administrative Absorption of Politics in Hong Kong, described the colonial Hong Kong's administration as "elite consensual government". In it, he claimed, any coalition of elites or forces capable of challenging the legitimacy of Hong Kong's administrative structure would be co-opted by the existing apparatus through the appointment of leading political activists, business figures and other elites to oversight committees, by granting them British honours, and by bringing them into elite institutions like Hong Kong's horse racing clubs. He called this "synarchy", an extension of John K. Fairbank's use of the word to describe the mechanisms of government under the late Qing dynasty in China. When modern China began after the fall of the last dynasty, one of the first political statements made in Hong Kong was the immediate change from long queue hairstyles to short haircuts. In 1938, Guangzhou fell to the hands of the Japanese, Hong Kong was considered a strategic military outpost for all trades in the far east. Though Winston Churchill assured that Hong Kong was an "impregnable fortress", it was taken as a reality check response since the British Army actually stretched too thin to battle on two fronts.

title: History of Hong Kong (1800s–1930s)


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Dan Turpin

Daniel "Terrible" Turpin is a character published by DC Comics. He first appeared as Brooklyn in Detective Comics #64 (June 1942), and first appeared as Dan Turpin in New Gods #5 (November 1971).

Publication history
Due to a recent retcon, Dan Turpin was made the adult version of Jack Kirby's Golden Age "kid-gang" character Brooklyn, of the Boy Commandos.

Fictional character biography
In Turpin's first appearance, he is one of the few citizens of Metropolis aware that a secretive war is taking place in the city between super-powered beings. When this war turns violent, Turpin's boss tries to take him off the case, but Turpin ignores him. Turpin leads the fight against the rampaging Kalibak, using the energy of the entire city to assist Lightray and Orion in defeating him. Turpin suffers multiple injuries but survives.

He keeps his job and becomes Lieutenant Inspector of the Metropolis Special Crimes Unit. In the third Superboy series, Turpin is sent to Hawaii to investigate whether or not they need a Special Crimes Unit of their own. While initially skeptical, a visit from the Female Furies convinces Turpin that the islands need an SCU. In current Superman titles, Turpin is extremely loyal to Maggie Sawyer before her transfer to Gotham City. His romantic feelings for her fall apart when he learns she is lesbian. Turpin has a daughter named Maisie.

He has spent a long time fighting threats in Metropolis; such as the rampaging group of mutants called the 'Underworlders'.

During the Final Crisis event, Turpin is called out of retirement to investigate the case of several missing children and the death of the New God Orion. He is later possessed by Darkseid, but freed after Batman shoots him with a Radion bullet, which is poisonous to New Gods.

Other versions

 * An alternate reality version of Dan Turpin appears in Superman: The Dark Side.
 * An alternate reality version of Dan Turpin appears in Superman's Metropolis. This version is a low-class worker and member of a revolution against Superman, who is the ruler of a dystopian Metropolis.
 * Dan Turpin appears in Elseworld's Finest.

Television

 * Dan Turpin appears in Superman: The Animated Series, voiced by Joseph Bologna. This version is the leader of the Metropolis Special Crimes Unit and is physically modeled after Jack Kirby. In the two-part episode "Apokolips... Now!", Turpin is killed by Darkseid while helping Superman fend off his invasion of Earth.
 * Dan Turpin appears in the Smallville episode "Bulletproof", portrayed by David Paetkau. This version is a rookie Metropolis police officer.

Film

 * Dan Turpin makes a non-speaking cameo appearance in Superman: Unbound.
 * Dan Turpin appears in The Death of Superman, voiced by Rick Pasqualone.

Video games
Dan Turpin appears as a character summon in Scribblenauts Unmasked: A DC Comics Adventure.

Miscellaneous

 * Dan Turpin appears in the GraphicAudio production of Superman: The Never Ending Battle.
 * Dan Turpin appears in the Smallville: Season Eleven comic-book continuation of Smallville.
 * Dan Turpin appears in the Injustice 2 prequel comic series. After Superman is defeated and his Regime disassembled, Turpin is assigned to oversee his imprisonment before being killed by Talia al Ghul's daughter Athanasia.

title: Dan Turpin


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Bahujan Samaj Party of Nepal

Bahujan Samaj Party, Nepal (Majority Society Party, Nepal) is a Nepalese political party. It is inspired by the Kanshi Ram's Bahujan Samaj Party of India and led by Madan Bahadur Pradhan. In the 1999 general elections it had 3 candidates. The party got 0.01% of the votes nationwide. Right now its national president is Ganga Prasad Mahara.

title: Bahujan Samaj Party of Nepal