| Nipponosaurus | |
|---|---|
 | |
| Restored skeleton | |
Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Reptilia |
| Clade: | Dinosauria |
| Clade: | †Ornithischia |
| Clade: | †Ornithopoda |
| Family: | †Hadrosauridae |
| Subfamily: | †Lambeosaurinae |
| Genus: | †Nipponosaurus Nagao,1936 |
| Type species | |
| †Nipponosaurus sachalinensis | |
Nipponosaurus (meaning "Japaneselizard") is alambeosaurinehadrosaur from sediments of theYezo Group, in Sinegorsk on the island ofSakhalin, which was part of Japan at the time of the species' classification. Thetype and only species isN. sachalinensis, known only from a singlejuvenile specimen discovered in 1934 and named in 1936, by Takumi Nagao, with further material of the same individual found in 1937.[1][2][3] Since then, the taxon has been largely ignored, and its validity has been doubted, with synonymy with other Asian hadrosaurs or status as anomen dubium being suggested. Redescriptions from 2004 and 2017, however, have supported recognition as a distinct species.[2][4][5] Dating the only specimen has been difficult, but based on associated mollusc taxa, the species likely lived sometime in the upperSantonian or lowerCampanian, around 80 million years ago.[4][5]
Theholotype (UHR 6590,University of Hokkaidō Registration) was discovered in November 1934 during the construction of ahospital for the Kawakamicolliery of theMitsui Mining Company onKarafuto Prefecture (nowSinegorsk,Sakhalin, Russia).[4] It was disarticulated when discovered.N. sachalinensis was named and described in 1936 by ProfessorTakumi Nagao of the Imperial University of Hokkaido. The generic name refers toNippon, the Japanese name ofJapan; it was the firstdinosaur named based on a find made on Japanese territory – thoughSouth Sakhalin was annexed by theSoviet Union in 1945. Thespecific name refers to Sakhalin.[1]
Although being relatively complete at the time of discovery, the specimen was missing much of theskull and limbs. A second expedition was organised in the summer of 1937, in which limb material pertaining to the holotype was recovered. The next year, Nagao authored a second article on the species describing these additional remains.[3][4]
Nipponosaurus was described as a member of the Trachodontidae; howeverTrachodon is now considered an invalidnomen dubium.[6] The family is now known as theHadrosauridae. Among its relatives, Nagao thought it was possibly closest to the taxaCheneosaurus andTetragonosaurus, even suggesting that his species might later prove to becongeneric with one of them.[1] However, these were later determined to be juvenile forms ofHypacrosaurus,Corythosaurus andLambeosaurus, and not distinct species.[4][7]
For the remainder of the 20th century, thetaxon was scarcely mentioned.[4] While they were still considered valid species, the relationship with the supposed "cheneosaurs"Cheneosaurus andProcheneosaurus was accepted;[8] after this it was rarely assigned past the Lambeosaurinae. Ahumerus from a pit nearHashima Island, Japan was assigned toNipponosaurus, variously referred to asTrachodon, in 1967.[9] No further studies have acknowledged the specimen, and the referral has been ignored, with the species still considered to be represented by a single specimen.[4] In 1977, a review of Asian dinosaur palaeontology mentioned it as a possible close relative ofMandschurosaurus without comment.[10] Later in 1989, a study noted that features of its metacarpals could indicate a basal position within the Hadrosauridae.[4][11] Near the turn of the century, in 1994, a review of Japanese dinosaurs noted the incomplete nature of many Asian hadrosaurs could mean some of them, includingNipponosaurus, may in fact be representatives of the same species.[5]
More extensive research has been conducted in the 21st century. In 2004, the species was redescribed, with focus on its growth stage. They concluded it was a valid taxon, and that the holotype was immature.[4] More recently, a 2017 study provided further descriptions of the specimen, and conducted a microscopic examination on sections of the left femur, a rib, and an isolatedchevron, which re-affirmed the age of the holotype.[2]
The immature holotype specimen measures roughly 4 metres (13 feet) in length. When it was originally described, it was considered to be an adult, because of its co-ossified sacral vertebra. Future authors doubted this, merely citing its small size. In the 2004 redescription, it was examined more closely and several characteristics were found that identified it as an juvenile individual.[4] Later, in 2017, another team re-evaluated the specimen, and determined through investigating the fossils at a microscopic level (in a process calledhistology) that this was correct, although they doubted the significance of some of the characters identified in the former study.[2]
Though the quality of bone preservation is generally poor, the holotype skeleton is estimated to be 60% complete.[4] It consists of a leftmaxilla anddentary,parietal, various isolated skull elements, thirteencervical vertebrae, sixdorsal vertebrae, twosacral vertebrae, a series of 35caudal vertebrae, a leftscapula, lower portions of bothhumeri, most elements of the lower forelimbs, anischium, leftilium, and most of the hindlimbs.[2][4]
Some authors have deemed the species anomen dubium, questioning its supposed diagnostic characteristics, especially in light of its presumed immature status. The validity of the taxon has therefore been controversial. The 2004 redescription denied this, proposing three diagnostic characters.[4] In their description ofSahaliyania andWulagasaurus,Pascal Godefroit and colleagues reported based on direct observation of thetype specimen that all of these supposed unique traits were found in other hadrosaurids. The lack of certain portions of the skull key to evaluating its systematic position was also cited as a reason to doubt the validity of the genus, and the problematic nature of the skull had indeed been noted before.[5][12] Later, a 2017 study once again questioned the validity of the proposed diagnostic characters; however, upon a re-examination of the specimen, three new ones were put forward.[2]
Acladistic analysis in 2004 placedNipponosaurus sachalinensis very close to the well-known North AmericanHypacrosaurus altispinus within theLambeosaurinae. They suggested it might be congeneric with the species, but because of the incomplete nature of the holotype and the questionablemonophyly of the genusHypacrosaurus, they refrained from reassigning it.[4] However, later in 2007, a study providing a redescription ofLambeosaurus magnicristatus re-evaluated a character relating to the size of the distal ischial foot, and found this drewNipponosaurus away fromH. altispinus and into a more basal position within their Corythosaurini (now generally referred to as theLambeosaurini). They however noted the only character uniting it specifically with this group, an increased number ofcervical vertebrae, was a state poorly known in more basal Asianlambeosaurines, and cautioned its position was likely liable to change.[13] The left cladogram was recovered in the 2004 study, and the right cladogram was recovered by the 2007 study:
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More recently, another analysis, accounting for characters that vary throughontogeny, instead found it to be closely related to the European taxaArenysaurus andBlasisaurus, outside of Lambeosaurini, unlike the earlier studies. Its position outside of theclade containingParasaurolophini and Lambeosaurini was supported by the absence of twosynapomorphies of the group in its teeth. Four characters united it withArenysaurus andBlasisaurus; the end margin of thecoronoid process overlapping with the back portion of thedental battery (which was suggested to perhaps reflect a difference in feeding habits from other taxa), a shortedentulous slope, and two characters of thejugal.[2] The followingcladogram was recovered in their analysis:
The specimen was collected from UpperYezo Group, which was known as the Upper Ammonites Bed at the time of the original description. Because the area where it was discovered no longer belongs to Japan, follow up research has become difficult. Additionally, field notes from the time of discovery have been lost. This has made determining the age of the horizon problematic. However, Nagao noted fossil molluscs now calledParapuzosia japonica andSphenoceramus schmidti were also found at the locality, and these are known to hail from the lowerCampanian. An age from the upperSantonian or lower Campanian is therefore likely.[4][5]
Based on the sediments it was preserved in, theNipponosaurus specimen is thought to have been buried in a marine setting. However, this would not have been far from the shore, as indicated by the relatively complete nature of the specimen and the presence of fossilised terrestrial plants found alongside it. This indicates the species may have lived primarily on low-lying plains near the coast, a lifestyle consistent with some of its North American relatives.[5]
