A New Basal Hadrosaurid (Dinosauria: Ornithischia) From the Latest Cretaceous Kita-ama Formation in Japan: the Rise of Hadrosaurs

doi:10.21203/rs.3.rs-225217/v1

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A New Basal Hadrosaurid (Dinosauria: Ornithischia) From the Latest Cretaceous Kita-ama Formation in Japan: the Rise of Hadrosaurs

https://doi.org/10.21203/rs.3.rs-225217/v1

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published 27 Apr, 2021

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Here we describe a partial hadrosaurid skeleton from the marine Maastrichtian Kita-ama Formation of the Izumi Group in Japan as a new taxon, Yamatosaurus izanagii gen. et sp. nov. Our phylogenetic analysis demonstrates it belongs to the Hadrosauridae, composed of Hadrosaurus foulkii + (Yamatosaurus izanagii + (Saurolophinae + Lambeosaurinae). Yamatosaurus izanagii is unique in having only one functional tooth/row in the middle of the dentary dental battery and complete absence of the branched ridges on the occlusal surfaces. The coracoid lacks a biceps tubercle as in non-hadrosaurid hadrosauroids, suggesting its presence is a key feature for the clade of Saurolophinae and Lambeosaurinae. The evolutionary rate analysis further supports that shoulder and forelimb features, which are likely to be involved in locomotion, are important for the early evolution of Hadrosauridae. Our biogeographic analyses show that the basal Hadrosauridae was widely distributed in Asia and Appalachia, and the clade of Saurolophinae and Lambeosaurinae originated in Asia. The contemporaneous occurrence of basal (Yamatosaurus izanagii) and derived (Kamuysaurus japonicus) hadrosaurids during the Maastrichtian in Japan is the first record in Asia. Because of a long geographical distance between these localities, they may not co-exist but show some level of provinciality.

Paleozoology

Evolutionary Biology

Taxonomy

hadrosaurid

skeleton

Maastrichtian Kita-ama Formation

phylogenetic analysis

biogeographic analysis

Hadrosauroids are successful herbivorous ornithischian dinosaurs during the Cretaceous. Fossil records of basal hadrosauroids have been discovered mainly from the Lower Cretaceous deposits of Europe, eastern Asia, and Africa, whereas derived forms are commonly known from the Upper Cretaceous rocks of all continents except Australia and the Indian subcontinent. A large clade of derived hadrosauroids, flourished in post-Campanian, called differently in previous studies such as Euhadrosauria ^1–3, Hadrosauridae ^4,5, and Saurolophidae ⁶, consists of two major clades with solid-crested (Hadrosaurinae ⁷ or Saurolophinae ⁶) and hollow-crested (Lambeosaurinae) skulls ^2,6. Hadrosauridae is defined as the most recent common ancestor of Edmontosaurus regalis, Saurolophus osborni, Lambeosaurus lambei, and Hadrosaurus foulkii and all of its descendants ⁶. The members of hadrosaurids were successful in both abundance ^8,9 and cosmopolitan distribution including the Arctic ^10–12. The high diversification of hadrosaurids is attributed to their efficient oral processing system, established by specialized dentitions with complex grinding surface ¹³ together with cranial kinetics ^14,15. Social behaviors, expressed by unique supracranial crests, are also likely to have contributed to the success of the derived hadrosauroids ^16–18. In addition to the functional adaptations, Kobayashi, et al. ¹⁹ recently proposed that inhabitation in marine-influenced environments by ancestral hadrosaurids also played an important role in the early evolution of hadrosaurids. However, because the members of sister clades of Hadrosauridae vary among previous studies (e.g., Telmatosaurus transsylvanicus, Plesiohadros djadokhtaensis, Eotrachodon orientalis, and Lophorhothon atopus), the rise of hadrosaurids has been obscure.

Among hadrosauroid remains from the Late Cretaceous in Japan, the best-preserved specimen is Kamuysaurus japonicus from the Maastrichtian Hakobuchi Formation, Yezo Group in Hokkaido Prefecture. Although other specimens are fragmentary and isolated elements such as teeth, limb elements, and vertebrae, it is noteworthy that hadrosauroid remains have been discovered from all four major islands (Hokkaido, Honshu, Shikoku, and Kyushu) of Japan ²⁰. In addition to these specimens, an amateur fossil collector, Mr. Shingo Kishimoto, discovered a partial hadrosauroid specimen in May of 2004. This new discovery includes a dentary, a surangular, isolated dentary teeth, cervical vertebrae, a caudal vertebra, cervical ribs, and a coracoid (MNHAH D1-033516). This specimen was recovered from the Maastrichtian horizon of the Kita-ama Formation, Izumi Group, in Sumoto City of the Awaji Island in Hyogo Prefecture (Fig. 1A-B). MNHAH D1-033516 was initially identified as a lambeosaurine hadrosaurid ²¹, but it has been suggested that a further study is needed for its taxonomic assignment ¹⁹. Here we provide the first description of MNHAH D1-033516 and test its taxonomic status. The new hadrosaurid provides insight into how coastal habit impacted the early divergence of Hadrosauridae and the coexistence of the Far East coastal hadrosauroids during the early Maastrichtian.

Institutional abbreviations:

AMNH: American Museum of Natural History, New York, USA; CMN: Canadian Museum of Nature, Ottawa, Ontario, Canada; MNHAH: Museum of Nature and Human Activities, Hyogo, Sanda, Hyogo, Japan; MPC: Mongolian Paleontological Center, Ulaanbaatar, Mongolia; MOR: Museum of the Rockies, Montana, USA; NMMNH: New Mexico Museum of Natural History, Albuquerque, New Mexico, USA; ROM: Royal Ontario Museum, Toronto, Ontario, Canada.

Geological setting

The new specimen was recovered from blocks of dark grey mudstones of the upper part of the Kita-ama Formation of the Izumi Group, exposed at a cement industry quarry (Loc. Aw14, Morozumi ²²) in Yura Town, Sumoto City of the Awaji Island in Hyogo Prefecture. The upper part of the Kita-ama Formation, composed of inter-arc basin deposits including conglomerate, sandstone, and mudstone of the distal turbidite faces (Fig. 1C-E), is rich in invertebrate fossils; the ammonoid Nostoceras hetonaiense, the nautilid Eutrephoceras sp., bivalves (e.g., Apiotrigonia (Microtrigonia) amanoi, Eriphyla japonica, and Inoceramus (Endocostea) shikotanensis), and gastropods (e.g., Gigantocapulus problematicus), decapods (e.g., Archaeopus ezoensis, Hinecaris simplex, and Ahazianassa masanorii) ^22–24. Fragmentary materials of vertebrates also have been reported from this quarry such as fishes, a turtle (Mesodermochelys undulatus), a mosasaur, and a hesperornithiform ^23,25−27 (Tanaka, et al. ²⁷ erroneously listed pterosaur and plesiosaur occurrences from the locality). Plant fossils from the quarry include angiosperms (Platanus sp. and Ulmus sp.) and cycad (Zamiophyllum sp.) ²³. The exposed beds at the quarry are within subchron 32.1r ²⁸, which is currently dated as 71.94–71.69 Ma (early Maastrichtian) ²⁹, as well as the Nostoceras hetonaiense Zone ²², which is the same as the ammonite zone that includes Kamuysaurus japonicus. This ammonite zone is currently dated as 72.4–70.6 Ma (based on the dates of the underlying Didymoceras awajiense Zone ³⁰ and overlying Gaudryceras izumiense Zone ³¹), which is in concordance with the magnetostratigraphic data.

Systematic paleontology

Dinosauria Owen ³²

Ornithischia Seeley ³³

Ornithopoda Marsh ³⁴

Hadrosauridae Cope ³⁵

Yamatosaurus izanagii gen. et sp. nov.

Nomenclatural Acts. This published work and the nomenclatural acts it contains have been registered in ZooBank, the proposed online registration system for the International Code of Zoological Nomenclature. The ZooBank life science identifiers (LSID) can be resolved and the associated information viewed by appending the LSID to the prefix http://zoobank.org/. The LSID for this publication is: urn:lsid:zoobank.org:act: 1C728683-C3EA-4688-9420-096D13EB6227.

Etymology. “Yamato” refers to the ancient name for a region of the Japanese archipelago, including the western half of the main island (Honshu Island), Shikoku Island, and the northern half of Kyushu Island, ruled by the Yamato Kingdom from the third to the seventh century. “Sauros” means reptiles. The specific name, “izanagi”, refers to a deity in Japanese mythology, which created eight countries of Yamato with another deity, Izanami, based on the oldest history book in Japan, called “Kojiki (Records of Ancient Matters)”, published in 712 CE (Common Era). The first country created was the Awaji Island, followed by the Shikoku, Oki, Kyushu, Iki, Tsushima, Sado, and Honshu islands.

Holotype. MNHAH D1-033516, a right dentary, a right surangular, twelve isolated dentary teeth, four cervical vertebrae, a distal caudal vertebra, three cervical ribs, and a coracoid (Figs. 1F, 2–6). This specimen is stored in the Museum of Nature and Human Activities, Hyogo, Sanda City, Hyogo Prefecture, Japan.

Locality and horizon. Loc. Aw14 (Morozumi ²²) of Yura Town, Sumoto City of the Awaji Island, Hyogo Prefecture, Japan; the early Maastrichtian (71.94–71.69 Ma) ²⁹ Kita-ama Formation of the Izumi Group.

Diagnosis. The hadrosaurid with unique characters in having the presence of only one functional tooth/row in the middle of the dentary dental battery and the complete absence of the “branched ridges” on the dentary tooth occlusal surfaces. It is also unique in the combination of the following characters: the small angle between the medial surface of dentary symphysis and lateral surface of the dentary and ventrally facing ventral surface of the surangular.

Description. The right dentary is nearly complete, missing its posterior end and the coronoid process. The lateral surface of the main body of the dentary bears multiple neurovascular foramina. A large foramen is positioned at the level of the seventh dentary tooth, and several smaller foramina are present posteriorly. In lateral view, the anterior portion of the dentary is downturned, and an angle between the ventral edge of the anterior portion and a horizontal plane, or tooth row, is 20 degrees (Fig. 2A). In medial view (Fig. 2B), the long axis of the tooth row is parallel to the ventral margin of the middle region of the dentary main body as in hadrosaurids except for Brachylophosaurini ³⁶. A thin alveolar parapet, covering the medial surface of the dental battery, is dorsoventrally taller posteriorly than anteriorly. The nutrient foramina are organized in a shallow arch ventral to the dental battery. The Meckelian groove extends along the ventral margin of the posterior two-thirds of the dentary. The ventral edge of the dentary main body below the coronoid process is weakly bowed. In dorsal view, the occlusal surface of the dental battery is straight and parallel to the lateral surface of the dentary main body (Fig. 2C).

The dentary diastema is approximately 99 mm. The length is 30% of the length between the anterior end of the dental battery and the posterior margin of the coronoid process (330 mm). Although the coronoid process is not preserved, the latter length was estimated from the base of the coronoid process. The dorsal margin of the dentary, where the lateral ramus of the predentary contacts, is strongly concave in lateral view and is angled by 110 degrees from a horizontal plane (Fig. 2B).

The symphysial process of the dentary curves medially at nearly a right angle (Fig. 2E) as in non-hadrosaurid hadrosauroids (e.g., Plesiohadros djadokhtaensis; MPC-D 100/745) and some members of Lambeosaurini, including Olorotitan arharensis ³⁷ and Parasaurolophus tubicen (NMMNH P-25100). The maximum mediolateral width of the dentary symphysial region in dorsal view is 81 mm, which is approximately twice as wide as the minimum breadth of the dentary posterior to the dentary symphysial region (Fig. 2C). The medial margin of the symphysial process is largely divergent from the lateral surface of the main body of the dentary by an angle of 20 degrees as in Eotrachodon orientalis ³⁸ but unlike a nearly parallel alignment in hadrosaurids such as Edmontosaurus regalis ³⁹ and Parasaurolophus tubicen (NMMNH P-25100). The medial surface of the symphysial process bears a shallow groove for contact with its counterpart (Fig. 2B). The ventral surface of the symphysial process has a large neurovascular foramen (Fig. 2D).

The base of the coronoid process is laterally expanded as in hadrosaurids and some non-hadrosaurid hadrosauroids such as Plesiohadros djadokhtaensis, Nanningosaurus dashiensis, and Adynomosaurus arcanus ^40–42 and is separated from the dental battery (Fig. 2E). The posteromedial surface of the coronoid process is largely excavated, forming the mandibular adductor fossa.

The dorsolateral portion of the right surangular is preserved (Fig. 3). The preserved region of the anterodorsal process is thin and bears a well-defined ridge for the attachment of m. adductor mandibulae externus superficialis ⁴³. The ridge is posteriorly continuous with the dorsolateral flange, which is gently arcuate in dorsal view. The lateroventral surface of the surangular bears no foramen as in derived hadrosauroids including Bactrosaurus johnsoni ^44,45. The dorsal surface of the anterior half of the surangular is concave to form the mandibular adductor fossa, whereas the posterior half forms a glenoid fossa that receives the mandibular condyles of the quadrate. The ventromedial portion of the surangular is lost before collection thus the morphology of the caudal process and the retroarticular process is unavailable.

At least 34 tooth positions are present, and each tooth position bears a maximum of four teeth in the right dentary, and at least twelve isolated dentary teeth from the left side are preserved. Tooth crowns are diamond-shaped and bear a prominent straight primary ridge in the middle teeth, while the primary ridge is sinuous in mesial and distal teeth (Figs. 2B, 4A, 4H). The secondary ridges are faint, wrinkled, and in some cases branched as in Eotrachodon orientalis ³⁸ and Hypacrosaurus altispinus ⁴⁶ but unlike the straight secondary ridges of Corythosaurus casuarius (ROM 868) and Lambeosaurus lambei (CMN 361, CMN 2869). The largest tooth crown, located in the middle portion of the dental battery, is approximately 41.94 mm high and 14.66 mm wide. The tooth crowns become shorter and narrower mesially and distally (Fig. 2B). The height/width ratios of the dentary tooth crowns are slightly less than three on average although vary largely by position. The marginal denticles along the mesial/distal edges of the coronal half of the tooth crown are proportionately smaller than those of Eotrachodon orientalis ³⁸ and Lambeosaurus lambei (CMN 351) and resemble those of Hypacrosaurus altispinus in size ⁴⁶. The marginal denticles of Yamatosaurus izanagii are formed of multiple small papillae (Fig. 4B-C), which are unorganized unlike in Eotrachodon orientalis ³⁸ and Corythosaurus casuarius (AMNH FARB 8527), but resembling Gryposaurus notabilis (AMNH FARB 8526). The angle between the crown and root of the dentary teeth is 132 degrees (Fig. 4B).

An occlusal surface of an isolated tooth changes from a flat pentagonal surface (Fig. 4D-E) to a concaved oval-shaped surface (Fig. 4F-G) as it wears. At most two teeth are functional per tooth position throughout the dental battery unlike hadrosaurids (e.g., Edmontosaurus regalis, CMN 2289; Parasaurolophus tubicen, NMMNH P-25100), which have more than two functional teeth in the middle of the dental battery. Only one functional tooth in 9th, 11th, 14th, 16th, 19th, 21st, and 23rd tooth positions (Fig. 4I-J), which is unique to Yamatosaurus izanagii. The occlusal surface of the dental buttery is steeply inclined as in Prosaurolophus maximus (CMN 2277, CMN 2280), the distal area of the dental battery of Brachylophosaurus canadensis (CMN 8893), and the mesial area of the dental battery of Gryposaurus (Gryposaurus notabilis, ROM 873; Gryposaurus latidens, AMNH FARB 5465). The occlusal surface of Yamatosaurus izanagii bears low ridges that are associated with cementum and longitudinal giant tubules that fill the pulp cavity ^13,47. The occlusal topography is much lower than in Corythosaurus ¹³ but resembles that of Brachylophosaurus canadensis (CMN 8893). The strong inclination and the low topography of the occlusal surfaces may suggest the pulp cavity of teeth of Yamatosaurus izanagii is filled mainly with transverse giant tubules, which are less wear-resistant than longitudinal giant tubules ¹³.

At least four cervical vertebrae are preserved, and three of them are fairly complete (Fig. 5A-O). Although the exact position of these vertebrae is ambiguous, two are identified as anterior (fourth or fifth and fifth or sixth) and one as a middle (seventh to ninth) cervical vertebra based on comparisons with Gobihadros mongoliensis (MPC-D 100/746). The anterior cervical vertebrae are nearly complete other than postzygapophyses, whereas the middle cervical vertebra is missing the ventral half of its centrum. The anterior centra are strongly opisthocoelous. The anterior convexity and the posterior concavity are the most pronounced in the anteriormost cervical vertebra. The lateroventral surface of the centrum is deeply excavated and bears several foramina. The ventral surfaces of the centra are slightly convex and do not form a distinct keel. The neural canal is slightly wider than its height in the anteriormost cervical vertebra and becomes wider posteriorly. The transverse process is the shortest in the anteriormost cervical vertebra and becomes longer posteriorly. The dorsal surfaces of the prezygapophyses are slightly inclined mediodorsally. The diapophyses become longer posteriorly relative to the articular facets of the prezygapophyses. The neural spine is faint and lacks significant dorsal projection in the anteriormost cervical vertebrae and becomes larger posteriorly. Both sides of the base of the neural spine are shallowly depressed in the anteriormost cervical vertebra. The depressions are shallower and smaller in posterior cervicals. The postzygapophyses are less than three times as long as the anteroposterior length of the neural arch unlike hadrosaurids, including Maiasaura peeblesorum (ROM 44770) and Hypacrosaurus stebingeri (MOR 549) but resemble non-hadrosaurid hadrosauroids such as Gilmoreosaurus mongoliensis (AMNH FARB 6551). The postzygapophyseal articular surface is anteroposteriorly longer than mediolaterally wide in the anterior cervical vertebrae, whereas the articulation surface is subcircular in the middle cervical vertebra.

A distal caudal vertebra is nearly complete, missing a right prezygapophysis and a distal portion of the neural spine (Fig. 5V-Z). The centrum is amphiplatyan and is hexagonal in anterior and posterior aspects. The lateral surfaces of the centrum bear a ridge in the middle, whereas the ventral surface is gently convex. The transverse process is absent as in the 38th and more posterior caudal vertebrae of a lambeosaurine indet. (CMN 8330). The neural canal is a mediolaterally wide ellipsoid. The rod-shaped prezygapophysis lacks a distinct articular facet. The postzygapophysis is reduced and indistinguishable from the neural spine.

One right and two left cervical ribs are preserved (Fig. 5P-U). The capitulum is longer and more massive than the tuberculum in all preserved cervical ribs. The lateral crest is dorsoventrally flat and well-developed in the anterior-most preserved cervical rib and gradually diminishes posteriorly.

The right coracoid is complete (Fig. 6). The articular facet for the scapula is shorter than the glenoid and angled 114 degrees from the glenoid in the external view. The coracoid foramen is ellipsoid and does not intersect the scapulocoracoid contact. The anterior margin of the coracoid is straight. The biceps tubercle is absent. The ventral process is only half as long as its base as in non-hadrosaurid hadrosauroids (e.g., Gilmoreosaurus mongoliensis, AMNH FARB 30722), yet the ventral process is recurved caudoventrally as in hadrosaurids (e.g., Brachylophosaurus canadensis, MOR 1071 8-10-99-541; Hypacrosaurus altispinus, AMNH FARB 5272).

The ontogenetic stage of Yamatosaurus izanagii cannot be examined histologically due to the absence of any weight-bearing limb bones. However, the size of the preserved dentary (Supplementary Table S1) is comparable with those of large, presumably adult hadrosaurids including Kamuysaurus japonicus (526 mm) ¹⁹ and Hypacrosaurus stebingeri (500 mm) ⁴⁸. In addition, the neurocentral sutures of preserved cervical vertebrae are closed (Fig. 5), suggesting the type specimen of Yamatosaurus izanagii was at least close to maturity at the time of its death ^49,50.

Our phylogenetic analysis produced 12 most parsimonious trees (MPTs) of 1181 steps, each with a consistency index of 0.430 and a retention index of 0.834. Hadrosaurus foulkii, which has been regarded as a nomen dubium ^7,51, is recovered as a sister taxon to the clade of Yamatosaurus izanagii, Saurolophinae, and Lambeosaurinae. Since the phylogenetic position of Hadrosaurus foulkii is comparable to those proposed by Prieto-Márquez ⁶, this study follows Prieto-Márquez ⁶ in the definitions of Hadrosauridae, Lambeosaurinae, and Saurolophinae.

The analysis shows that Yamatosaurus izanagii belongs to Hadrosauridae, sharing two of five synapomorphies of the family (small denticles of dentary teeth and long ventral process of the coracoid), and is a sister taxon to the clade of Saurolophinae and Lambeosaurinae (Fig. 7). The clade of Yamatosaurus izanagii, Saurolophinae, and Lambeosaurinae is weakly supported, sharing a single synapomorphy (dentary tooth crown height-width ratio in between 2.70 and 3.30). Yamatosaurus izanagii is a sister taxon to the clade of Saurolophinae and Lambeosaurinae, which is well supported by five unambiguous characters: functional teeth up to two; symphysial region of dentary strongly curved lingually; short postzygapophysis of cervical vertebrae; straight craniodorsal margin of coracoid; and absence of biceps tubercle of the coracoid.

Three phylogenetic autapomorphies of Yamatosaurus izanagii are the crown and the root of the dentary teeth angled in between 110º and 130º, a small angle between the medial surface of dentary symphysis and lateral surface of the dentary, and ventrally facing ventral surface of the surangular. The moderately angled dentary teeth crown and root is shared with derived lambeosaurines (Parasaurolophini + Lambeosaurini) and saurolophines other than Saurolophini and Kritosaurini. The small angle between the medial surface of dentary symphysis and lateral surface of dentary is commonly seen in more primitive forms such as Eotrachodon orientalis, Gilmoreosaurus mongoliensis, Bactrosaurus johnsoni, and more primitive taxa, indicative of a reversal in Yamatosaurus izanagii. The ventrally facing surface of the surangular main body is a character for derived saurolophines (e.g., Edmontosaurus and Saurolophus) and lambeosaurines (e.g., Corythosaurus and Hypacrosaurus).

In addition to the phylogenetic autapomorphies, Yamatosaurus izanagii is unique among Eolambia caroljonesa and higher taxa in the presence of only one functional tooth/row, even in the middle of the dentary dental battery (Fig. 4I-J). The dentary occlusal surface is also unique in the complete absence of the “branched ridges” ¹³ in any of the preserved teeth. Since the ridges are formed at the plugged pulp cavity ^47,52, the absence of the “branched ridges” may indicate a unique tooth ontogeny of Yamatosaurus izanagii. Other than the dentition, Yamatosaurus izanagii dentary exhibits derived forms. It differs from the other derived non-hadrosaurid hadrosauroids, including Eotrachodon orientalis ³⁸, Zhanghenglong yangchengensis ⁷, and Plesiohadros djadokhtaensis ⁴⁰, in having the ventral deflection of the anterior dentary, the posterior elongation of the dental battery, and the anteromedial elongation of the symphysial process.

While the dentary possesses multiple derived features, the coracoid of Yamatosaurus izanagii resembles those of non-hadrosaurid hadrosauroids such as Eolambia caroljonesa and Gilmoreosaurus mongoliensis in its straight anterior margin, short ventral hook, and undeveloped biceps tubercle. Dilkes ⁵³ interpreted the biceps tubercle of hadrosaurids as the origin of m. biceps, which extends to the proximal ends of the ulna and radius. Since m. biceps functions to lift the antebrachium, the presence and enlargement of the biceps tubercles in saurolophines and lambeosaurines may represent a functional change in forelimb use. The evolutionary rate analyses on pectoral girdle + forelimb phylogenetic characters demonstrate a significantly high evolutionary rate in the branch leading to the clade of Saurolophinae and Lambeosaurinae (Fig. 7B). Although one may concern the high evolutionary rate may be an artifact by the incompleteness of Yamatosaurus izanagii as warned by Lloyd ⁵⁴, the same trend is recovered in an analysis without Yamatosaurus izanagii (Supplementary Fig. S1). Therefore, the pectoral girdle + forelimb morphology is likely to have experienced rapid morphological changes at the base of Hadrosauridae. The rapid pectoral girdle + forelimb modifications may be associated with hadrosauroid gait change. Although Maidment and Barrett ⁵⁵ suggested that facultative quadrupedality was acquired more basal to Gilmoreosaurus mongoliensis, the evolutionary rates of the pectoral girdle + forelimb may suggest the gait shift was accomplished in basal hadrosaurids. Previously, changes in jaw elements and dentition have been considered as the key features for the origin of Hadrosauridae, but this study adds that forelimb locomotion, resulting from shoulder and forelimb structural modification, is also an important feature within the origin of this family.

The biogeographic analyses resulted in higher fits in the analyses using the “starting” and the “relaxed” dispersal multiplier matrices than in the “harsh” (Supplementary Table S2). The results suggest that permitting intercontinental dispersal probabilities can explain hadrosaurid dispersal history better than considering intercontinental dispersal was nearly impossible. The results in the “starting” and the “relaxed” matrices are nearly identical to each other thus only the “starting” result is presented here (Fig. 8; see Supplementary Figs. S2 and S3 for the “relaxed” and “harsh” results). The analysis suggested that successive nodes of derived hadrosauroids from Eotrachodon orientalis to the node of Hadrosauridae were widely distributed in Asia and Appalachia, but not the intervening landmasses of Laramidia or Europe. Previous studies argued that the largely diversified clade of Saurolophinae and Lambeosaurinae (Hadrosauridae sensu Xing, et al. ⁷) originated in North America ^11,56 or Asia ⁵, but our analysis supports the Asian origin of this clade thus refining other previous broader models of ornithopod origins in Asia ⁵⁷ while also contributing to a better understanding of the origins of high latitude dinosaurs ⁵⁸. Further biogeographic resolution will be aided with additional specimens, especially from pre-Campanian Upper Cretaceous deposits, because the successive taxa of non-hadrosaurid hadrosauroids, especially from Eotrachodon orientalis to Plesiohadros djadokhtaensis, have long ghost lineages up to 20 my. Yamatosaurus izanagii shows the longest ghost lineage duration of roughly 30 my among hadrosaurids. Interestingly, given the length of the ghost lineages of the Maastrichtian non-hadrosaurid hadrosauroids Plesiohadros djadokhtaensis from Mongolia and Tanius sinensis from China, considered with the ghost lineage represented by Yamatosaurus izanagii, it may be that eastern Asia (Japan, Mongolia and China) served as a refugium of relict hadrosauroid taxa.

Hadrosauroid materials have been reported from the Upper Cretaceous deposits of Japan ²⁰, and the youngest occurrence is Kamuysaurus japonicus from the early Maastrichtian marine deposits of the Hakobuchi Formation in Mukawa Town in Hokkaido Prefecture ¹⁹, which is equivalent age to Yamatosaurus izanagii. Although paleo-latitudes of Mukawa Town and Sumoto City during the Cretaceous are estimated to be similar to the current position (approximately 1,100 km apart), it is clear that these dinosaurs were present at the same geologic time. It was suggested that the non-hadrosaurid hadrosauroid Plesiohadros djadokhtaensis in late Campanian was replaced by hadrosaurids such as Saurolophus angustirostris and Barsboldia sicinskii in Maastrichtian in Mongolia, indicating that the co-existence of hadrosaurid and non-hadrosaurid hadrosauroid is unlikely in the fluvial depositional environments ⁴⁰. The replacement of hadrosauroids by more derived members during the Maastrichtian is suggested in Laurasia ⁴⁰ but not in Europe (Telmatosaurus transsylvanicus and Tethyshadros insularis) because the European archipelago persisted as island relicts ^3,59,60. The occurrence of Yamatosaurus izanagii and Kamuysaurus japonicus from the time equivalent formations shows the contemporaneous existence of a derived and a basal hadrosaurid in Asia for the first time (Fig. 1G). This contemporaneous occurrence may be due to a different paleoenvironment preserved where Japan was more coastal in nature and richer in vegetation with plants such as plane trees (Platanus sp.), elm (Ulmus sp.), and cycads (Zamiophyllum sp.) than the fluvial environments preserved in the mainland Asian continent. An alternative scenario is that these dinosaurs did not co-occur and instead there was some level of provinciality within the coastal environments of this part of Asia where Kamuysaurus japonicus occupied a northern area and Yamatosaurus izanagii was restricted to a more southern area.

Phylogeny. Phylogenetic analysis was conducted on the data matrix modified from Takasaki, et al. ⁶¹, using TNT ver 1.5 ⁶². The immature Edmontosaurus OTUs were removed from the original data matrix since their phylogenetic relationships are out of the scope of this study. Nanyangosaurus zhugeii was also excluded from the analysis because of its poor preservation. Protohadros byrdi is included in the data matrix by scoring the phylogenetic characters based on the published literature ⁶³. Lophorhothon atopus was re-scored based on the recently published paper by Gates and Lamb ⁶⁴. The resultant data matrix consists of 71 OTUs and 354 characters (Supplementary Data S1). A phylogenetic analysis was conducted using TNT ver. 1.5 ⁶², with the Ouranosaurus nigeriensis set as the outgroup. All of the characters were equally weighted and unordered. The maximum number of trees was set to 99,999 in memory. A traditional search with 10,000 replicates of Wagner trees using random additional sequences, followed by the TBR branch swapping that held 10 trees per replicate was performed. Supports for the clade were evaluated by bootstrap resampling using standard absolute frequencies (1,000 replicates) and Bremer decay indices calculations.

Evolutionary rates. Branch evolutionary rates of the phylogenetic characters related to pectoral girdle + forelimb were examined using R package Claddis version 0.6.5 ⁵⁴ in R v.4.0.3 ⁶⁵, based on the current morphological data matrix and the phylogenetic hypothesis. OTUs with character completeness less than 50% are pruned prior to the analyses since missing data leads to extreme results ⁵⁴. Removal of the taxa with low character completeness also provides much better resolution to the phylogenetic tree. The remaining polytomies are randomly resolved prior to the branch partition analyses then given branch lengths of 10^− 6 of the total tree lengths. One-hundred time-calibrated phylogenetic trees which incorporate the ranges of the time of first and last appearances are constructed based on the equal method ⁶⁶ using the R package paleotree ⁶⁷. The analysis was repeated twice, with and without Yamatosaurus izanagii. The trees were calibrated based on the first and the last appearances of each OTUs (Supplementary Data S2). The analysis was repeated twice, with and without Yamatosaurus izanagii. The analyses are conducted using the R script provided as the Supplementary Data S3.

Ancestral range reconstruction. Ancestral habitat ranges were reconstructed using R package BioGeoBEARS ⁶⁸ in R v.4.0.3 ⁶⁵ using the R script in the Supplementary Data S4. Although recent biogeographic works generally conduct analyses over multiple models ^56,69, Ree and Sanmartín ⁷⁰ recently pointed out conceptual problems in “+J” models. Since our interest is on habitat range reconstruction rather than model selection, this study conservatively performed only the DEC analysis following the suggestions by Ree and Sanmartín ⁷⁰. A phylogenetic hypothesis based on the majority consensus tree was used. The remaining polytomies were resolved then given branch lengths of 10^− 6 of the total tree lengths. To incorporate the ambiguities in time ranges of the OTUs, 100 time-calibrated phylogenetic trees were produced using the R package paleotree ⁶⁷, based on the “equal” method ⁶⁶. The marginal probabilities at each node are averaged over the 100 time-calibrated trees. Three analyses with the “starting”, “harsh”, and “relaxed” dispersal multiplier matrices were conducted following Poropat, et al. ⁷¹. In the “starting” and the “relaxed” matrices, the dispersal multiplier values between the following continents are set to 0.5 and 1, respectively: Asia and Laurasia, Asia and Europe, Laramidia and Appalachia, Laramidia and South America, Appalachia and Europe, and Europe and Africa (Supplementary Data S5 – S7). The maximum range size allowed for any species to occupy was set 3. The dates and the continents of each OTUs are compiled from previous literature (Supplementary Data S2, S8).

Acknowledgments

The authors deeply appreciate Mr. Shingo Kishimoto for donating the Yamatosaurus type specimen for scientific contribution. We are grateful to Kazumi Wada (Museum of Nature and Human Activities, Hyogo, Japan) for his outstanding preparation of the specimen (MNHAH D1-033516) and Haruo Saegusa and Yutaka Furutani (Museum of Nature and Human Activities, Hyogo, Japan) for providing us the information on the fossil locality. We also thank Tadahiro Ikeda (Museum of Nature and Human Activities, Hyogo, Japan), Brandon Stanley (Royal Tyrrell Museum of Palaeontology, Canada), Kevin Seymour (Royal Ontario Museum, Canada), Carl Mehling (American Museum of Natural History, USA), Margaret Currie (Canadian Museum of Nature, Canada), Xing Xu (Institute of Vertebrate Paleontology and Paleoanthropology, China), and Amanda Millhouse (Smithsonian National Museum of Natural History, USA) for providing access specimens. We thank Olivia Fiorillo for additional helpful comments. This work was supported by JSPS KAKENHI Grant Number JP19K04052 to Y.K.

Additional Information

Author contributions Y.K. designed and directed the project. All wrote the description of Yamatosaurus, performed statistical analyses, worked on comparisons, and contributed to discussions and shared role of editing the manuscript.

Competing interests

The author(s) declare no competing interests.

Vecchia, F. M. D. Telmatosaurus and the other hadrosaurids of the Cretaceous European Archipelago. An overview. Nat. Nascosta. 32, 1–55 (2006).
Horner, J. R., Weishampel, D. B., Forster, C. A. & Hadrosauridae in The Dinosauria: Second Edition (eds. Weishampel, D. B., Dodson, P., & Osmólska, H.) 438–463(University of California Press, 2004).
Weishampel, D. B., Norman, D. & Grigorescu, D. Telmatosaurus transsylvanicus from the Late Cretaceous of Romania. Palaeontology. 36, 361–385 (1993).
You, H. L. & Li, D. Q. A new basal hadrosauriform dinosaur (Ornithischia: Iguanodontia) from the Early Cretaceous of northwestern China. Can. J. Earth Sci. 46, 949–957 https://doi.org/10.1139/e09-067 (2009).
Godefroit, P., Shulin, H., Tingxiang, Y. & Lauters, P. New hadrosaurid dinosaurs from the uppermost Cretaceous of northeastern China. Acta Palaeontol. Pol. 53, 47–74 https://doi.org/10.4202/app.2008.0103 (2008).
Prieto-Márquez, A. Global phylogeny of Hadrosauridae (Dinosauria: Ornithopoda) using parsimony and Bayesian methods. Zoological Journal of the Linnean Society. 159, 435–502 https://doi.org/10.1111/j.1096-3642.2009.00617.x (2010).
Xing, H. et al. A new basal hadrosauroid dinosaur (Dinosauria: Ornithopoda) with transitional features from the Late Cretaceous of Henan Province, China. PLoS ONE. 9, e98821 https://doi.org/10.1371/journal.pone.0098821 (2014).
Cullen, T. M., Fanti, F., Capobianco, C., Ryan, M. J. & Evans, D. C. A vertebrate microsite from a marine-terrestrial transition in the Foremost Formation (Campanian) of Alberta, Canada, and the use of faunal assemblage data as a paleoenvironmental indicator. Palaeogeogr., Palaeoclimatol., Palaeoecol. 444, 101–114 https://doi.org/10.1016/j.palaeo.2015.12.015 (2016).
Nakajima, J. et al. Dinosaur tracks at the Nemegt locality: Paleobiological and paleoenvironmental implications. Palaeogeogr., Palaeoclimatol., Palaeoecol. 494, 147–159 https://doi.org/10.1016/j.palaeo.2017.10.026 (2018).
Weishampel, D. B. et al. Dinosaur distribution in The Dinosauria: Second Edition (eds. Weishampel, D. B., Dodson, P., & Osmólska, H.) 515–606 (University of California Press, 2004).
Prieto-Márquez, A. Global historical biogeography of hadrosaurid dinosaurs. Zoological Journal of the Linnean Society. 159, 503–525 https://doi.org/10.1111/j.1096-3642.2010.00642.x (2010).
Takasaki, R., Fiorillo, A. R., Kobayashi, Y., Tykoski, R. S. & McCarthy, P. J. The first definite lambeosaurine bone from the Liscomb Bonebed of the Upper Cretaceous Prince Creek Formation, Alaska, United States. Sci Rep. 9, 5384 https://doi.org/10.1038/s41598-019-41325-8 (2019).
Erickson, G. M. et al. Complex dental structure and wear biomechanics in hadrosaurid dinosaurs. Science. 338, 98–101 https://doi.org/10.1126/science.1224495 (2012).
Weishampel, D. B. Evolution of jaw mehcanisms in ornithopod dinosaurs. Adv Anat Embryol Cell Biol. 87, https://doi.org/10.1007/978-3-642-69533-9_1 (1984).
Cuthbertson, R. S., Tirabasso, A., Rybczynski, N. & Holmes, R. B. Kinetic limitations of intracranial joints in Brachylophosaurus canadensis and Edmontosaurus regalis (Dinosauria: Hadrosauridae), and their implications for the chewing mechanics of hadrosaurids. Anat Rec (Hoboken). 295, 968–979 https://doi.org/10.1002/ar.22458 (2012).
Weishampel, D. B. The nasal cavity of lambeosaurine hadrosaurids (Reptilia: Ornithischia): comparative anatomy and homologies. Journal of Paleolimnology. 55, 1046–1057 (1981).
Evans, D. C. Nasal cavity homologies and cranial crest function in lambeosaurine dinosaurs. Paleobiology. 32, 109–125 https://doi.org/10.1666/04027.1 (2006).
Bell, P. R., Fanti, F., Currie, P. J. & Arbour, V. M. A mummified duck-billed dinosaur with a soft-tissue cock's comb. Curr. Biol. 24, 70–75 https://doi.org/10.1016/j.cub.2013.11.008 (2014).
Kobayashi, Y. et al. A new hadrosaurine (Dinosauria: Hadrosauridae) from the marine deposits of the Late Cretaceous Hakobuchi Formation, Yezo Group, Japan. Sci Rep. 9, 12389 https://doi.org/10.1038/s41598-019-48607-1 (2019).
Kubota, K. A list of Mesozoic dinosaur fossils from Japan. Hum Nat. 28, 97–115 (2017).
Suzuki, D., Saegusa, H. & Furutani, H. Newly found hadrosaurid fossil co-producing broadleaf fossils from Sumoto, west central Japan. Journal of Vertebrate Paleontology. 25, 120 (2005).
Morozumi, Y. Late Cretaceous (Campanian and Maastrichtian) ammonites from Awaji Island, Southwest Japan. Bull. Osaka Mus. Nat. Hist. 39, 1–58 (1985).
Iwaki, T. & Maeda, H. Mudstone facies and the fossil molluscan assemblages of the Upper Cretaceous Izumi Group in the southeastern Awaji Island, southwest Japan. Research Reports of Kochi University. Natural Science 38, 187–201, pls. 181–186(1989).
Karasawa, H., Kishimoto, S., Ohara, M. & Ando, Y. Late Cretaceous Decapoda from the Izumi Group of Japan, with descriptions of two new genera and one new species of Axiidea and one new family of Brachyura. Bull Mizunami Fossil Mus. 45, 43–85 21 pls (2019).
Hirayama, R., Kishimoto, S., Saegusa, H. & Ikeda, T. Skulls of fossil marine turtles from the Late Cretaceous Izumi Group in Sumoto, Hyogo Prefecture, western Japan. Abstracts with Programs, Regular Meeting, the Palaeontological Society of Japan. 163, 21–21 (2014).
Sato, T., Konishi, T., Hirayama, R. & Caldwell, M. W. A review of the Upper Cretaceous marine reptiles from Japan. Cretac. Res. 37, 319–340 https://doi.org/10.1016/j.cretres.2012.03.009 (2012).
Tanaka, T., Kobayashi, Y., Ikuno, K., Ikeda, T. & Saegusa, H. A marine Hesperornithiform (Avialae: Ornithuromorpha) from the Maastrichtian of Japan: Implications for the paleoecological diversity of the earliest diving birds in the end of the Cretaceous. Cretac. Res. 104492, https://doi.org/10.1016/j.cretres.2020.104492 (2020).
Kodama, K. Magnetostratigraphy of the Izumi Group along the Median Tectonic Line in Shikoku and Awaji Islands, Southwest Japan. J. Geol. Soc. Jpn. 96, 265–278 (1990).
Gradstein, F., Ogg, J. G., Schmitz, M. D. & Ogg, G. M. The Geologic Time Scale 2012(Elsevier, 2012).
Shigeta, Y., Tsutsumi, Y. & Misaki, A. U–Pb age of the Didymoceras awajiense Zone (upper Campanian, Cretaceous) in the Aridagawa area, Wakayama, southwestern Japan. Bull. Natl. Mus. Nat. Sci. Ser. C, Geol. & Paleontol. 43, 11–18 (2017).
Shigeta, Y., Izukura, M. & Tsutsumi, Y. An early Maastrichtian (latest Cretaceous) ammonoid fauna from the Soya Hill area, Hokkaido, northern Japan. Bull. Hobetsu Mus. 32, 7–41 (2017).
Owen, R. Report on British fossil reptiles, Part 2. Report of the British Association for the Advancement of Science. 11, 60–204 (1842).
Seeley, H. G. On the classification of the fossil animals commonly named Dinosauria. Proceedings of the Royal Society of London 43, 165–171, doi:10.1098/rspl.1887.0117 (1887).
Marsh, O. C. Principal characters of American Jurassic dinosaurs, part IV. American Journal of Science. 21, 417–423 https://doi.org/10.2475/ajs.s3-21.122.167 (1881).
Cope, E. D. Synopsis of the extinct Batrachia, Reptilia and Aves of North America. Trans. Am. Philos. Soc. 14, 1–252 (1870).
Freedman Fowler, E. A. & Horner, J. R. A new brachylophosaurin hadrosaur (Dinosauria: Ornithischia) with an intermediate nasal crest from the Campanian Judith River Formation of northcentral Montana. PLoS ONE. 10, e0141304 https://doi.org/10.1371/journal.pone.0141304 (2015).
Godefroit, P., Bolotsky, Y. L. & Bolotsky, I. Y. Osteology and relationships of Olorotitan arharensis, a hollow-crested hadrosaurid dinosaur from the latest Cretaceous of Far Eastern Russia. Acta Palaeontol. Pol. 57, 527–560 https://doi.org/10.4202/app.2011.0051 (2012).
Prieto-Márquez, A., Erickson, G. M. & Ebersole, J. A. Anatomy and osteohistology of the basal hadrosaurid dinosaur Eotrachodon from the uppermost Santonian (Cretaceous) of southern Appalachia. PeerJ. 4, e1872 https://doi.org/10.7717/peerj.1872 (2016).
Xing, H., Mallon, J. C. & Currie, M. L. Supplementary cranial description of the types of Edmontosaurus regalis (Ornithischia: Hadrosauridae), with comments on the phylogenetics and biogeography of Hadrosaurinae. PLoS ONE. 12, e0175253 https://doi.org/10.1371/journal.pone.0175253 (2017).
Tsogtbaatar, K., Weishampel, D. B., Evans, D. C. & Watabe, M. A new hadrosauroid (Plesiohadros djadokhtaensis) from the late Cretaceous Djadokhtan fauna of southern Mongolia in Hadrosaurs (eds. Eberth, D. A. & Evans, D. C.) 108–135(Indiana University Press, 2015).
Jinyou, M. O., Zhongru, Z., Wei, W. & Xing, X. U. The first hadrosaurid dinosaur from southern China. Acta Geologica Sinica - English Edition. 81, 550–554 https://doi.org/10.1111/j.1755-6724.2007.tb00978.x (2007).
Prieto-Márquez, A., Fondevilla, V., Sellés, A. G., Wagner, J. R. & Galobart, Ã. Adynomosaurus arcanus, a new lambeosaurine dinosaur from the Late Cretaceous Ibero-Armorican Island of the European archipelago. Cretac. Res. 96, 19–37 https://doi.org/10.1016/j.cretres.2018.12.002 (2019).
Holliday, C. M. New insights into dinosaur jaw muscle anatomy. Anat Rec (Hoboken). 292, 1246–1265 https://doi.org/10.1002/ar.20982 (2009).
Godefroit, P., Dong, Z., Bultynck, P., Li, H. & Feng, L. New Bactrosaurus (Dinosauria: Hadrosauroidea) material from Iren Dabasu (Inner Mongolia, PR China). Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, Sciences de la Terra 68, 3–70(1998).
Prieto-Márquez, A. Cranial and appendicular ontogeny of Bactrosaurus johnsoni, a hadrosauroid dinosaur from the Late Cretaceous of northern China. Palaeontology. 54, 773–792 https://doi.org/10.1111/j.1475-4983.2011.01053.x (2011).
Evans, D. C. Cranial anatomy and systematics of Hypacrosaurus altispinus, and a comparative analysis of skull growth in lambeosaurine hadrosaurids (Dinosauria: Ornithischia). Zoological Journal of the Linnean Society. 159, 398–434 https://doi.org/10.1111/j.1096-3642.2009.00611.x (2010).
LeBlanc, A. R., Reisz, R. R., Evans, D. C. & Bailleul, A. M. Ontogeny reveals function and evolution of the hadrosaurid dinosaur dental battery. BMC Evol Biol. 16, 152 https://doi.org/10.1186/s12862-016-0721-1 (2016).
Horner, J. R. & Currie, P. J. Embryonic and neonatal morphology and ontogeny of a new species of Hypacrosaurus (Ornithischia, Lambeosauridae) from Montana and Alberta in Dinosaur Eggs and Babies (eds. Carpenter, K., Hirrsch, K. F., & Horner, J. R.) 312–336(Cambridge University Press, 1994).
Brochu, C. A. Closure of neurocentral sutures during crocodilian ontogeny: implications for maturity assessment in fossil archosaurs. Journal of Vertebrate Paleontology. 16, 49–62 https://doi.org/10.1080/02724634.1996.10011283 (1996).
Takasaki, R., Chiba, K., Kobayashi, Y., Currie, P. J. & Fiorillo, A. R. Reanalysis of the phylogenetic status of Nipponosaurus sachalinensis (Ornithopoda: Dinosauria) from the Late Cretaceous of southern Sakhalin. Hist. Biol. 30, 694–711 https://doi.org/10.1080/08912963.2017.1317766 (2017).
Prieto-Márquez, A., Weishampel, D. B. & Horner, J. R. The dinosaur Hadrosaurus foulkii, from the Campanian of the east coast of North America, with a reevaluation of the genus.Acta Palaeontologica Polonica51 (2006).
Bramble, K., LeBlanc, A. R. H., Lamoureux, D. O., Wosik, M. & Currie, P. J. Histological evidence for a dynamic dental battery in hadrosaurid dinosaurs. Sci Rep. 7, 15787 https://doi.org/10.1038/s41598-017-16056-3 (2017).
Dilkes, D. W. Appendicular myology of the hadrosaurian dinosaur Maiasaura peeblesorum from the Late Cretaceous (Campanian) of Montana. Transactions of the Royal Society of Edinburgh: Earth Sciences. 90, 87–125 https://doi.org/10.1017/s0263593300007185 (2000).
Lloyd, G. T. Estimating morphological diversity and tempo with discrete character-taxon matrices: implementation, challenges, progress, and future directions. Biological Journal of the Linnean Society. 118, 131–151 https://doi.org/10.1111/bij.12746 (2016).
Maidment, S. & Barrett, P. Osteological correlates for quadrupedality in ornithischian dinosaurs. Acta Palaeontol. Pol. https://doi.org/10.4202/app.2012.0065 (2012).
Longrich, N. R., Suberbiola, X. P., Pyron, R. A. & Jalil, N. E. The first duckbill dinosaur (Hadrosauridae: Lambeosaurinae) from Africa and the role of oceanic dispersal in dinosaur biogeography. Cretac. Res. 104678, https://doi.org/10.1016/j.cretres.2020.104678 (2020).
Fiorillo, A. R. Dinosaurs of Alaska: implications for the Cretaceous origin of Beringia. Geological Society of America Special Papers. 442, 313–326 https://doi.org/10.1130/2008.442(15 (2008).
Fiorillo, A. R. Alaska Dinosaurs: An ancient arctic world(CRC Press, 2018).
Vecchia, F. M. D. Tethyshadros insularis, a new hadrosauroid dinosaur (Ornithischia) from the Upper Cretaceous of Italy. Journal of Vertebrate Paleontology. 29, 1100–1116 https://doi.org/10.1671/039.029.0428 (2009).
Weishampel, D. B. & Jianu, C. M. Transylvanian dinosaurs(Johns Hopkins University Press, 2011).
Takasaki, R., Fiorillo, A. R., Tykoski, R. S. & Kobayashi, Y. Re-examination of the cranial osteology of the Arctic Alaskan hadrosaurine with implications for its taxonomic status. PLoS ONE. 15, e0232410 https://doi.org/10.1371/journal.pone.0232410 (2020).
Goloboff, P. A. & Catalano, S. A. TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics. 32, 221–238 https://doi.org/10.1111/cla.12160 (2016).
Head, J. J. A new species of basal hadrosaurid (Dinosauria, Ornithischia) from the Cenomanian of Texas. Journal of Vertebrate Paleontology. 18, 718–738 https://doi.org/10.1080/02724634.1998.10011101 (1998).
Gates, T. A. & Lamb, J. Redescription of Lophorhothon atopus (Ornithopoda: Dinosauria) from the Late Cretaceous of Alabama based on new material. Can. J. Earth Sci. https://doi.org/10.1139/cjes-2020-0173 (2021).
R Core Team. R: A language and environment for statistical computing (R Foundation for Statistical Computing, Vienna, Austria, 2020) https://www.R-project.org/.
Brusatte, S. L., Benton, M. J., Ruta, M. & Lloyd, G. T. Superiority, competition, and opportunism in the evolutionary radiation of dinosaurs. Science. 321, 1485–1488 https://doi.org/10.1126/science.1161833 (2008).
Bapst, D. W. paleotree: an R package for paleontological and phylogenetic analyses of evolution. Methods in Ecology and Evolution. 3, 803–807 https://doi.org/10.1111/j.2041-210X.2012.00223.x (2012).
Matzke, N. J. Probabilistic historical biogeography: new models for founder-event speciation, imperfect detection, and fossils allow improved accuracy and model-testing. Frontiers of Biogeography. 5, https://doi.org/10.21425/f5fbg19694 (2013).
Xu, X. et al. A new Middle Jurassic diplodocoid suggests an earlier dispersal and diversification of sauropod dinosaurs. Nat Commun. 9, 2700 https://doi.org/10.1038/s41467-018-05128-1 (2018).
Ree, R. H. & Sanmartín, I. Conceptual and statistical problems with the DEC + J model of founder-event speciation and its comparison with DEC via model selection. Journal of Biogeography. 45, 741–749 https://doi.org/10.1111/jbi.13173 (2018).
Poropat, S. F. et al. New Australian sauropods shed light on Cretaceous dinosaur palaeobiogeography. Sci Rep. 6, 34467 https://doi.org/10.1038/srep34467 (2016).

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A New Basal Hadrosaurid (Dinosauria: Ornithischia) From the Latest Cretaceous Kita-ama Formation in Japan: the Rise of Hadrosaurs

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