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Amurosaurus
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| Amurosaurus Temporal range: Late Cretaceous,
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|---|---|
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| Replica skeleton, Brussels | |
Scientific classification 
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| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Reptilia |
| Clade: | Dinosauria |
| Clade: | †Ornithischia |
| Clade: | †Ornithopoda |
| Family: | †Hadrosauridae |
| Clade: | †Euhadrosauria |
| Subfamily: | †Lambeosaurinae |
| Genus: | †Amurosaurus Bolotsky & Kurzanov, 1991 |
| Species: | †A. riabinini
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| Binomial name | |
| †Amurosaurus riabinini Bolotsky & Kurzanov, 1991
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| Synonyms | |
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Amurosaurus (/əˌmʊərəˈsɔːrəs/; "Amur lizard") is a genus of lambeosaurine hadrosaurid dinosaur found in Late Cretaceous (70 to 66 million years ago) deposits of what is now eastern Asia.[1]
Discovery and naming
[edit]
Russian paleontologists Yuri Bolotsky and Sergei Kurzanov first described and named this dinosaur in 1991. The generic name is derived from the Amur River and the Greek word sauros ("lizard"). The Amur (called Heilongjiang or "Black Dragon River" in Chinese) forms the border of Russia and China, and is near where this dinosaur's remains were found. There is one known species (A. riabinini), named in honor of the late Russian paleontologist Anatoly Riabinin, who conducted the first Russian expeditions to recover dinosaur remains in the Amur region in 1916 and 1917.[2][3]
All fossils of Amurosaurus have been recovered from a single bonebed locality, discovered in 1984 within the city limits of Blagoveschensk in the Amur Oblast of far eastern Russia. This bonebed is found in the Udurchukan Formation, the oldest geologic formation in the Tsagayan Group of far eastern Russia and northeastern China. This formation is thought to belong to the Maastrichtian stage of the Late Cretaceous Period, and was deposited about 68 million years ago, or just prior to the Lancian faunal stage of North America. The sediments were laid down in the floodplain of a river, which transported the fossils, but only a short distance, judging by the randomly assorted, disarticulated, but well-preserved bones within the bonebed, including fragile skull elements. Only a small section of the bonebed has been uncovered, but 90% of the remains found so far belong to lambeosaurines like Amurosaurus, mostly juveniles, with the rest belonging to other taxa, such as the hadrosaurine Kerberosaurus. Theropod teeth are also abundant, and there are many toothmarks on the bones, made by predators or scavengers.[4] This bonebed containing many specimens was unearthed in 2008.[5]
The holotype, or original specimen, consists of only a maxilla (upper jaw bone), and a dentary (lower jaw bone), both from the left side of the same individual. However, most of the other bones of the skull and skeleton have also been preserved in the bonebed, albeit of many different individuals. This other material was described more recently, making Amurosaurus the most abundant and completely known Russian dinosaur.[3]
A pathological ulna of a specimen referred to the species, preserved with a hypertrophied and swollen distal region and with the distal articular surface engulfed within a large overgrowth of newly formed bone, was described by Bertozzo et al. (2023), who interpreted the bone as still healing prior to the animal's death, with the misalignment of the fracture and the resulting malunion of the two fragments of the bone probably causing the animal to limp and walk on three limbs.[6]
Sahaliyania
[edit]
Sahaliyania (from "black" in Manchu, a reference to the Amur/Heilongjiang River), is a junior synonym of Amurosaurus. Its remains were found in a bonebed in the Maastrichtian-age Yuliangze Formation, alongside rarer remains of the hadrosaurine hadrosaurid (flat-headed duckbill) Wulagasaurus. Sahaliyania was named by Pascal Godefroit and colleagues in 2008. The type and only species is S. elunchunorum, named in honor of the Elunchun people.[7]
Sahaliyania was based on GMH W453, a partial skull. Godefroit and colleagues assigned numerous other fossils from the bonebed to their new genus, representing much of the skull, pectoral girdle, upper arm, and pelvis. It can be distinguished from other hadrosaurids by a variety of anatomical details. Godefroit and colleagues performed a phylogenetic analysis that places Sahaliyania as a lambeosaurine of uncertain relationships.[7] A 2022 article reassessed Sahaliyania and considered it a junior synonym of Amurosaurus.[8]
Description
[edit]
Amurosaurus is characterized by many autapomorphies, or unique features, of the skull, as well as the sigmoidal shape of the ulna (the lower arm bone) when viewed from the front or side. Most other known lambeosaurines have hollow crests on the top of their skulls, and although the bones that would make up such a crest are unknown for this dinosaur, the bones on the roof of the skull are modified to support one, so it can be assumed that Amurosaurus was crested as well. It has been estimated at 8 metres (26 ft) in length and about 3 metric tons (3.0 long tons; 3.3 short tons) in weight.[9] As a hadrosaurid, Amurosaurus would have been herbivorous.[10]
Classification
[edit]
The following cladogram on the left shows the results of a phylogenetic analysis of lambeosaur relationships from a 2013 study by Albert Prieto-Márquez and colleagues, showing Amurosaurus as a derived member of the tribe Lambeosaurini.[11] The cladogram on the right depicts the results from a 2014 study by Yuri Bolotsky and colleagues; this study incorporated novel skull material of Amurosaurus which they found to indicate a more primitive spot among lambeosaurs.[12]
The 2020 osteological description of the material previously constituting the distinct genus Sahaliyania found the latter genus to be a synonym of Amurosaurus, allowing its newly restudied material to contribute to more robust tests of the classification of Amurosaurus. With this new data, Amurosaurus was found to be a derived member of Lambeosaurinae closely related to Lambeosaurus, something that had been recovered in some previous studies. The cladogram from this study is reproduced below:[13]
See also
[edit]References
[edit]- ^ Godefroit, P., Lauters, P., Van Itterbeeck, J., Bolotsky, Y. and Bolotsky, I.Y. (2011). "Recent advances on study of hadrosaurid dinosaurs in Heilongjiang (Amur) River area between China and Russia." Global Geology, 2011(3).
- ^ Bolotsky, Y.L. & Kurzanov, S.K. 1991. [The hadrosaurs of the Amur Region.] In: [Geology of the Pacific Ocean Border]. Blagoveschensk: Amur KNII. 94-103. [In Russian]
- ^ a b Godefroit, P., Bolotsky, Y.L., & Van Itterbeeck, J. 2004. The lambeosaurine dinosaur Amurosaurus riabinini, from the Maastrichtian of Far Eastern Russia. Acta Palaeontologica Polonica 49(4): 585–618. Available online as PDF Archived December 24, 2005, at the Wayback Machine
- ^ Lauters, P.; Bolotsky, Y. L.; Van Itterbeeck, J.; Godefroit, P. (2008-03-01). "Taphonomy and Age Profile of a Latest Cretaceous Dinosaur Bone Bed in Far Eastern Russia". PALAIOS. 23 (3): 153–162. Bibcode:2008Palai..23..153L. doi:10.2110/palo.2006.p06-031r. ISSN 0883-1351. S2CID 128911990.
- ^ Lauters, Pascaline; Bolotsky, Yuri L.; Van Itterbeeck, Jimmy; Godefroit, Pascal (March 1, 2008). "Taphonomy and Age Profile of a Latest Cretaceous Dinosaur Bone Bed in Far Eastern Russia". PALAIOS. 23 (3). SEPM Society for Sedimentary Geology: 153–162. Bibcode:2008Palai..23..153L. doi:10.2110/palo.2006.p06-031r. S2CID 128911990.
- ^ Bertozzo F, Bolotsky I, Bolotsky YL, Poberezhskiy A, Ruffell A, Godefroit P, Murphy E (2023). "A pathological ulna of Amurosaurus riabinini from the Upper Cretaceous of Far Eastern Russia". Historical Biology: An International Journal of Paleobiology. 35 (2): 268–275. Bibcode:2023HBio...35..268B. doi:10.1080/08912963.2022.2034805. S2CID 247003496.
- ^ a b Godefroit, Pascal; Hai Shulin; Yu Tingxiang; Lauters, Pascaline (2008). "New hadrosaurid dinosaurs from the uppermost Cretaceous of north−eastern China" (PDF). Acta Palaeontologica Polonica. 53 (1): 47–74. doi:10.4202/app.2008.0103.
- ^ Xing, Hai; Gu, Wei; Hai, Shulin; Yu, Tingxiang; Han, Dong; Zhang, Yuguang; Zhang, Shujun (2022). "Osteological and taxonomic reassessments of Sahaliyania elunchunorum (Dinosauria, Hadrosauridae) from the Upper Cretaceous Yuliangzi Formation, northeast China". Journal of Vertebrate Paleontology. 41 (6) e2085111. doi:10.1080/02724634.2021.2085111. S2CID 250463301.
- ^ Paul, G.S., 2010, The Princeton Field Guide to Dinosaurs, Princeton University Press p. 308
- ^ Horner, John R.; Weishampel, David B.; Forster, Catherine A (2004). "Hadrosauridae". In Weishampel, David B.; Dodson, Peter; Osmólska, Halszka (eds.). The Dinosauria (2nd ed.). Berkeley: University of California Press. pp. 438–463. ISBN 978-0-520-24209-8.
- ^ Prieto-Márquez, A.; Dalla Vecchia, F. M.; Gaete, R.; Galobart, À. (2013). "Diversity, Relationships, and Biogeography of the Lambeosaurine Dinosaurs from the European Archipelago, with Description of the New Aralosaurin Canardia garonnensis" (PDF). PLOS ONE. 8 (7) e69835. Bibcode:2013PLoSO...869835P. doi:10.1371/journal.pone.0069835. PMC 3724916. PMID 23922815.
- ^ Bolotsky, Y. L., et al. "Hadrosaurs from the Far East: Historical perspective and new Amurosaurus material from Blagoveschensk (Amur region, Russia)." Hadrosaurs. 2014. 315-331.
- ^ Xing, Hai; Gu, Wei; Hai, Shulin; Yu, Tingxiang; Han, Dong; Zhang, Yuguang; Zhang, Shujun (2022). "Osteological and taxonomic reassessments of Sahaliyania elunchunorum (Dinosauria, Hadrosauridae) from the Upper Cretaceous Yuliangzi Formation, northeast China". Journal of Vertebrate Paleontology. 41 (6) e2085111. doi:10.1080/02724634.2021.2085111. S2CID 250463301.
Amurosaurus
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Discovery and naming
Etymology
The genus name Amurosaurus is derived from the Amur River, which marks the border between Russia and China in the region where the fossils were discovered, combined with the Greek word sauros, meaning "lizard" or "reptile".[5][6] The specific epithet riabinini honors the Russian paleontologist Anatoly Riabinin (1874–1942), who pioneered studies of Upper Cretaceous vertebrates, including early dinosaur discoveries, in the Amur region.[5][6][7] The fossils of A. riabinini were found near Blagoveshchensk, further tying the name to this specific site.[6]History of discovery
The discovery of Amurosaurus riabinini began in 1984 when paleontologist Yuri L. Bolotsky, working with the Amur Complex Integrated Research Institute (Amur KNII) of the Far Eastern Branch of the Soviet Academy of Sciences, identified a rich bonebed during fieldwork near Blagoveshchensk in the Amur Region of Far Eastern Russia.[5] This site, located at the base of Cretaceous sediments along the upper part of Nagornaia Street west of the city, revealed a continuous accumulation of disarticulated dinosaur bones spanning approximately 200 square meters.[5] Excavations yielded several hundred skeletal elements, with more than 90% belonging to lambeosaurine hadrosaurids, predominantly juveniles based on the sizes of centra, neural arches, and other postcranial remains, though some adult material was also present.[5] In 1991, Bolotsky and Sergei K. Kurzanov formally named the taxon Amurosaurus riabinini in a brief description published in Russian, designating the holotype as specimen AEHM 1/12—an associated left maxilla and dentary from the bonebed.[5] The fossils were recovered from the Udurchukan Formation, dated to the late Maastrichtian stage of the Late Cretaceous, approximately 68 million years ago, based on the Wodehouseia spinata–Aquilapollenites subtilis palynozone.[5] This initial publication focused on the diagnostic jaw elements, establishing A. riabinini as a new lambeosaurine genus from the region.[5] Subsequent studies expanded on the original material. In 2004, Pascal Godefroit, Bolotsky, and Johan Van Itterbeeck provided a comprehensive redescription incorporating additional specimens from the same bonebed, confirming its lambeosaurine affinities and detailing the taphonomic context of the assemblage.[5] Further reassessment occurred in 2022 with the description of a pathological ulna (AEHM 1/1037) from the Udurchukan Formation, offering new insights into individual variation and pathology within the species.[8]Description
Size and general anatomy
Amurosaurus riabinini was a large lambeosaurine hadrosaurid, with estimates indicating a body length of approximately 9 meters and a mass of around 4 metric tons for the largest individuals, based on scaling from the biggest humerus recovered from the Blagoveschensk bonebed.[9] These dimensions place it among the larger members of its subfamily, comparable to other Late Cretaceous Asian hadrosaurs. Alternative reconstructions suggest slightly smaller sizes of 7–8 meters in length and 2.5–3 metric tons in mass, derived from comparative skeletal proportions with North American relatives. Hip height likely reached up to 3 meters, inferred from limb bone robusticity and body proportions in similarly sized lambeosaurines.[10] Like other mature hadrosaurids, Amurosaurus exhibited facultative locomotion, capable of both bipedal and quadrupedal gaits, supported by robust fore- and hindlimbs adapted for efficient terrestrial movement across varied terrains.[11] The limb bones, including humeri and femora, show increased robusticity in larger specimens, facilitating weight-bearing during quadrupedal stance while allowing bipedal agility for foraging or evasion.[12] As a herbivore, it possessed a complex dental battery in the maxilla and dentary, comprising hundreds of tightly packed teeth that continuously replaced and sheared tough plant material, enabling efficient grinding of fibrous vegetation.[5] The fossil assemblage from the Blagoveschensk locality, dominated by Amurosaurus remains, primarily consists of late juveniles and small subadults, with adult specimens rare and representing less than 20% of identifiable elements based on long bone size distributions.[13] This age profile suggests ontogenetic changes in behavior or habitat, with hatchlings and early juveniles possibly segregated in aquatic or riparian environments before integrating into adult groups at about half-grown size, as inferred from the absence of very small individuals.[13] No direct evidence of sexual dimorphism is apparent in the skeletal record, though the predominance of immature individuals may mask subtle differences in adult morphology.[12]Skull and postcranial features
The skull of Amurosaurus riabinini is characterized by a low, hollow supracranial crest formed primarily by the nasal bones, prefrontals, and frontals, resembling the crest morphology observed in Lambeosaurus.[5] The premaxilla is elongated and narrow, lacking a premaxillary foramen, which aligns with lambeosaurine synapomorphies.[5] The maxilla exhibits a nearly symmetrical profile in lateral view, with an elevated body approximately twice as long as high and a high, triangular dorsal process whose apex lies about 20% posterior to the midpoint.[5] A distinctive sagittal crest on the parietal is particularly elevated caudally, forming a high, triangular, and deeply excavated process on the occipital aspect of the skull.[5] The dental structure includes complex jaw batteries, with the holotype maxilla bearing 39 vertical tooth columns and the dentary featuring approximately 37 columns, each containing 4–5 teeth, resulting in hundreds of teeth per battery across both jaws.[5] Maxillary teeth are diamond-shaped and straight, adorned with a prominent median carina of enamel and minimal denticulations, while dentary teeth are broader (with a height-to-width ratio of about 3.5) and display more developed lateral denticles along with occasional secondary ridges; enamel wear patterns on these teeth show angled facets typical of hadrosaurid occlusion.[5] Postcranially, the ulna is robust and displays a characteristic sigmoidal curvature in both cranial and lateral views, with the proximal portion convex medially and the distal portion convex laterally, potentially aiding in forelimb support or manipulation.[5] The femur is sturdy, featuring a globular head on a short neck, a craniocaudally extended greater trochanter, and a prominent, triangular fourth trochanter at midshaft.[5] The tibia has an enlarged proximal head with a laterally deflected, wing-like cnemial crest, a straight ovoid shaft, and a mediolaterally broadened distal end.[5] Vertebrae include opisthocoelous cervical centra that are wider than tall or long, heart-shaped and opisthocoelous dorsal centra, and amphiplatyan caudal centra that are subrectangular; neural spines vary regionally, with hook-like forms in the cervicals, subrectangular dorsal spines about three times higher than long and variably oriented, and long, slender, posteriorly inclined caudal spines bearing rough surfaces suggestive of cartilage caps in life.[5] Inferences from related lambeosaurine taxa suggest the hollow crest may have been covered by a keratinous sheath or soft tissue extension, though direct evidence for Amurosaurus is limited to the bony structure.[5] A depressed area on the parietal likely served as an attachment site for the ligamentum nuchae, supporting neck musculature.[5]Classification
Synonymy
Amurosaurus riabinini was initially named and briefly described in 1991 based on disarticulated hadrosaurid bones from the Udurchukan Formation in far eastern Russia, with early comparisons noting similarities to the North American lambeosaurine Corythosaurus due to shared cranial features such as the overall skull shape and crest morphology.[5] In 2008, a new lambeosaurine genus and species, Sahaliyania elunchunorum, was erected by Godefroit et al. for a partial skull (including the braincase, quadrates, and basicranium) collected from the Upper Cretaceous Yuliangze Formation in Heilongjiang Province, northeastern China, near the Russian border.[14] This taxon was distinguished from Amurosaurus primarily by purported differences in the basisphenoid and other cranial elements, positioning it as a derived lambeosaurine closely related to corythosaurins and parasaurolophins.[14] A comprehensive reassessment in 2022 by Xing et al. reexamined the holotype of S. elunchunorum and compared it directly with Amurosaurus material, finding that the previously cited diagnostic differences were either misinterpretations due to preservation artifacts or shared traits, such as the configuration of the alar process on the basisphenoid and the overall proportions of the postorbital and squamosal.[15] Geographic proximity between the type localities—spanning the Amur River region—and stratigraphic correlation of the Yuliangze and Udurchukan formations further supported synonymy, leading to the conclusion that Sahaliyania elunchunorum represents a junior synonym of Amurosaurus riabinini.[15] This revision was bolstered by shared lambeosaurine autapomorphies, including the elongate crest and specific dental battery features.[15] The synonymization has significant implications for specimen referral, allowing additional Chinese hadrosaurid fossils from the Yuliangze Formation—previously assigned to Sahaliyania or left indeterminate—to be confidently attributed to Amurosaurus, thereby expanding its known geographic range across the Sino-Russian border and indicating a broader distribution in eastern Asia during the Maastrichtian.[15] No other genera or species have been proposed as synonyms of Amurosaurus, though early Russian hadrosaurid finds from the region were occasionally misidentified as North American forms like Corythosaurus before the recognition of endemic Asian lambeosaurines.[5]Phylogenetic position
Amurosaurus is recognized as a basal member of the Lambeosaurinae subfamily within Hadrosauridae, positioned as the sister taxon to a clade that includes Lambeosaurus, Corythosaurus, and Hypacrosaurus.[16] This placement is supported by cladistic analyses incorporating extensive cranial and postcranial character data, highlighting its evolutionary ties to other crested hadrosaurids from the Late Cretaceous of Laurasia.[16] Phylogenetic studies have consistently affirmed Amurosaurus's position through parsimony-based analyses. The original description utilized 40 characters to recover Amurosaurus as a basal lambeosaurine, sister to the combined parasaurolophin and corythosaur clades.[5] A species-level analysis of Lambeosaurinae further positioned it as the terminal Asian taxon in a sequential dispersal pattern from Asia to North America, preceding the diversification of more derived North American forms. More recent work incorporating osteological data from the related Asian lambeosaurine Sahaliyania elunchunorum reinforced this basal stance among Asian lambeosaurines, resolving Amurosaurus specifically as sister to Lambeosaurus in a single most parsimonious tree.[15] Amurosaurus shares key derived traits with other lambeosaurines, notably the presence of hollow nasal crests formed by inflated nasal and premaxillary bones, which likely functioned in vocalization and resonance.[5] These features underscore its membership in a clade that evolved endemically in Asia following the dispersal of hadrosaurids from North America across Beringia during the Campanian, with Amurosaurus representing a late-branching Asian lineage.[16] As a Maastrichtian taxon, Amurosaurus is among the latest-surviving hadrosaurids, persisting into the latest Late Cretaceous and potentially outcompeting earlier Asian forms in its far eastern Russian habitat.[15]Paleoecology
Geological context
The fossils of Amurosaurus riabinini are primarily known from the Udurchukan Formation, a Maastrichtian-aged unit of the Tsagayan Group exposed along the Russia-China border in the Amur River valley of the Zeya-Bureya Basin.[5] This formation consists of continental sediments, including mudstones, sandstones, conglomerates, and diamicts, deposited in an alluvial setting dominated by meandering river systems and associated floodplains.[17] The Udurchukan Formation correlates stratigraphically and biostratigraphically with the Yuliangze Formation across the border in China, extending the depositional record of similar environments in the region.[5] The age of the Udurchukan Formation is determined as late Maastrichtian (approximately 70–66 million years ago) based on palynological assemblages, including the Wodehouseia spinata–Aquilapollenites subtilis zone, which aligns with well-calibrated North American reference sections.[17] Although direct radiometric dates are lacking, indirect correlations support an age around 68 million years ago for the dinosaur-bearing horizons.[5] Sedimentary structures such as cross-bedding and channel fills indicate a dynamic fluvial-lacustrine system influenced by sediment gravity flows from nearby uplifted margins, with evidence of periodic overbank deposition in low-energy lake margins. The depositional environment reflects a warm, humid subtropical climate, inferred from diverse palynomorphs and associated paleosols suggesting seasonal precipitation and vegetation suited to moist conditions.[17] Taphonomic analysis of Amurosaurus bonebeds, particularly at sites like Blagoveshchensk and Kundur, reveals concentrations of disarticulated juvenile and subadult skeletons within diamict layers, likely resulting from catastrophic flood events that entrained and rapidly buried carcasses on alluvial plains. These deposits show minimal weathering or scavenging, with low rates of abrasion and tooth marks (<2%), pointing to quick entombment in high-energy flows that preserved attritional mortality assemblages.Associated fauna
The fossil record of the Udurchukan Formation reveals a diverse vertebrate assemblage co-occurring with Amurosaurus riabinini in the late Maastrichtian floodplain ecosystems of the Amur region, characterized by meandering rivers, wetlands, and subtropical forests.[18] Among herbivores, Amurosaurus shared its habitat with other hadrosaurids, notably the non-crested saurolophine Kerberosaurus manakini, which may indicate niche partitioning between lambeosaurines adapted for browsing in forested areas and saurolophines suited to more open or aquatic vegetation. Armored dinosaurs, including nodosaurid remains such as osteoderms and teeth, represent a minor but present component of the thyreophoran diversity, likely inhabiting similar riparian zones. Theropod dinosaurs formed the primary carnivorous guild, with tyrannosaurid teeth and postcranial elements indicating large predators capable of preying on hadrosaur herds, while ornithomimid tibiae suggest smaller, possibly omnivorous or fast-running forms that scavenged or hunted juveniles. Aquatic and semi-aquatic reptiles occupied marginal niches in the riverine environment, including lindholmemydid turtles like Amuremys planicostata and indeterminate trionychids, alongside crocodilian osteoderms and skulls, which likely ambushed prey near water bodies. Bonebeds at Blagoveshchensk, dominated by over 90% Amurosaurus remains with attritional profiles favoring late juveniles and subadults, point to gregarious herd behavior among this dominant herbivore, though direct evidence of social structures remains absent.[5] Rare tooth marks on hadrosaur bones (<2% of specimens) and a documented pathological ulna exhibiting a healed mid-shaft fracture with malunion—potentially from theropod predation or conspecific aggression—underscore biotic pressures, with juveniles particularly vulnerable in muddy terrains where robust limbs aided mobility.[8]References
- https://www.wikidata.org/wiki/Q28359184