Fenglu Han

Fenglu Han
China University of Geosciences · Department of Paleontology

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64
Publications
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Publications

Publications (64)
Article
Ceratopsian dinosaurs underwent great changes, including a shift of locomotion mode, enlarged horns and frills, and increased body size. These changes occur alongside the evolution of endocranial morphology and physiology such as the size and shape of the flocculus, hearing range, olfactory ratio, and the reptile encephalization quotient (REQ). How...
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The Upper Cretaceous Ganzhou Basin of Jiangxi Province, South China is among one of the richest egg fossil sites in the world and yields abundant exquisite egg clutches and embryonic eggs, especially in oviraptor-osaurs. Hadrosaurian, troodontid, and potential dromaeosaurid egg fossils were also reported, though they were rare. All these reported e...
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Nanhsiungchelys (Testudines: Nanhsiungchelyidae) has been established in the 1960s, but its plastral morphology is not well understood. Here, we report an anterior lobe of the plastron of Nanhsiungchelys from the Upper Cretaceous of Ganzhou Basin, China. The plastron of the new specimen bears a V-shaped notch anteriorly which is absent or unknown i...
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Our understanding of pre-Cretaceous dinosaur reproduction is hindered by a scarcity of evidence within the fossil record. Here we report three adult skeletons and five clutches of embryo-containing eggs of a new sauropodomorph from the Lower Jurassic of southwestern China, displaying several significant reproductive features that are either unknown...
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Background Tooth replacement patterns of early-diverging ornithischians, which are important for understanding the evolution of the highly specialized dental systems in hadrosaurid and ceratopsid dinosaurs, are poorly known. The early-diverging neornithischian Jeholosaurus , a small, bipedal herbivorous dinosaur from the Early Cretaceous Jehol Biot...
Article
Large quadrupedal sauropod dinosaurs of the group Titanosauria were globally distributed in the Late Cretaceous. Many titanosaurian species have been discovered in eastern Asia, but most of them are controversial and represented by poorly preserved remains. Here, we describe a new titanosaur, Gandititan cavocaudatus gen. et sp. nov., based on a par...
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Nanhsiungchelyidae are a group of large turtles that lived in Asia and North America during the Cretaceous. Here we report a new species of nanhsiungchelyid, Nanhsiungchelys yangi sp. nov., from the Upper Cretaceous of Nanxiong Basin, China. The specimen consists of a well-preserved skull and lower jaw, as well as the anterior parts of the carapace...
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Jurassic strata are widely distributed in the eastern part of Tibet Autonomous Region, and have yielded many dinosaur bones. However, none of these specimens has been studied extensively, and some remain unprepared. Here we provide a detailed description of some new sauropod material, including several cervical vertebrae and a nearly complete scapu...
Preprint
Nanhsiungchelyidae are a group of large turtles that lived in Asia and North America during the Cretaceous. Here we report a new species of nanhsiungchelyid, Nanhsiungchelys yangi sp. nov., from the Upper Cretaceous of Nanxiong Basin, China. This is the second valid species of Nanhsiungchelys, and the holotype consists of a well-preserved skull and...
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The dental system of ceratopsids is among the most specialized structure in Dinosauria by the presence of tooth batteries and high-angled wear surfaces. However, the origin of this unique dental system is poorly understood due to a lack of relative knowledge in early-diverging ceratopsians. Here we study the dental system of three earliest-divergin...
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Here we describe a newly discovered basal ichthyosauromorph from the Lower Triassic of South China, Baisesaurus robustus gen. et sp. nov. The only known specimen of this new species was collected from the Lower Triassic (Olenekian) Luolou Formation in the Zhebao region of Baise City, on the northwest margin of the Nanpanjiang Basin, and comprises a...
Preprint
Full-text available
The dental system of ceratopsids is among the most specialized structure in Dinosauria, and includes high angled wear surfaces, split tooth roots, and multiple teeth in each tooth family. However, the early evolution of this unique dental system is generally poorly understood due to a lack of knowledge of the dental morphology and development in ea...
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Turtle eggs containing embryos are exceedingly rare in the fossil record. Here, we provide the first description and taxonomic identification, to our knowledge, of a fossilized embryonic turtle preserved in an egg, a fossil recovered from the Upper Cretaceous Xiaguan Formation of Henan Province, China. The specimen is attributed to the Nanhsiungche...
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Lystrosaurus represents one of the most successful dicynodonts, a survivor of the end-Permian mass extinction that remained abundant in the Early Triassic, but many aspects of its paleobiology are still controversial. The bone histology of Lystrosaurus species from South Africa and India has provided important information on their growth strategy a...
Article
Bone histology has provided valuable information on the life history of dinosaurs, and the presence of growth lines provides useful information for age estimation, growth variation, and the reconstruction of paleobehavior. Here, we present new data recovered from five individuals of the non-iguanodontian ornithopod dinosaur Jeholosaurus shangyuanen...
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The gross dental morphology of the holotype of the theropod Sinraptor dongi from the Jurassic Shishugou Formation of China is comprehensively described. We highlight a combination of dental features that appear to be restricted to Sinraptor: i) crowns with denticulated mesial and distal carinae extending from the root, and an irregular surface text...
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Titanosauriform sauropod dinosaurs were once considered rare in the Upper Cretaceous of Asia, but a number of titanosauriforms from this stratigraphic interval have been discovered in China in recent years. In fact, all adequately known Cretaceous Asian sauropods are titanosauriforms, but only a few have been well studied, lending significance to a...
Article
Graphical Abstract Highlights d Two new alvarezsaurian dinosaurs are described from Northwest China d They are intermediate between Late Jurassic and Late Cretaceous alvarezsaurians d They showcase the evolution of highly specialized alvarezsaurian forelimb d Specialized alvarezsaurian forelimb morphology evolved slowly, in a mosaic fashion In Brie...
Article
Ceratopsia includes some of the best-known ornithischian dinosaurs. Many species are erected based on cranial elements alone, and the postcranial skeletons are either missing or undescribed in many taxa. Here we provide the first detailed postcranial description of Yinlong downsi based on the holotype and eight other well-preserved skeletons. Yinlo...
Raw Data
This is the word version of character lists of Ornithischians for Han et al (2017). This file includes more than 200 figures and is easier for you to edit. Enjoy
Article
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The partial skeleton of a leptoceratopsid dinosaur, Ischioceratops zhuchengensis gen. et sp. nov., was excavated from the bone-beds of the Upper Cretaceous Wangshi Group of Zhucheng, Shandong Province, China. This fossil represents the second leptoceratopsid dinosaur specimen recovered from the Kugou locality, a highly productive site in Zhucheng....
Data
Character list for analysis of ornithischian phylogenetic relationships. Table A. Codings for S1 phylogenetic analysis, in TNT format. (DOC)
Data
Character list for analysis of ceratopsian phylogenetic relationships. Table A. Codings for S3 phylogenetic analysis, in TNT format (DOC)
Article
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Ceratopsia is one of the best studied herbivorous ornithischian clades, but the early evolution of Ceratopsia, including the placement of Psittacosaurus, is still controversial and unclear. Here, we report a second basal ceratopsian, Hualianceratops wucaiwanensis gen. et sp. nov., from the Upper Jurassic (Oxfordian) Shishugou Formation of the Jungg...
Article
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Yinlong downsi, from the Upper Jurassic Shishugou Formation, Xinjiang, northwestern China, is the oldest known ceratopsian dinosaur. Here we provide a detailed description of the skull and mandible based on the holotype, three partial skulls, and disarticulated materials from several other specimens. Yinlong can be diagnosed by six autapomorphies:...
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The homology of the 'semilunate' carpal, an important structure linking non-avian and avian dinosaurs, has been controversial. Here we describe the morphology of some theropod wrists, demonstrating that the 'semilunate' carpal is not formed by the same carpal elements in all theropods possessing this feature and that the involvement of the lateralm...
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A new ankylosaurid, Chuanqilong chaoyangensis gen. et sp. nov., is described here based on a nearly complete skeleton from the Lower Cretaceous Jiufotang Formation of Baishizui Village, Lingyuan City, Liaoning Province, China. Chuanqilong chaoyangensis can be diagnosed on the basis of two autapomorphies (glenoid fossa for quadrate at same level as...
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Background Southwestern Henan Province in central China contains many down-faulted basins, including the Xixia Basin where the Upper Cretaceous continental sediments are well exposed. The Majiacun Formation is a major dinosaur-bearing stratigraphic unit that occurs in this basin. Methodology/Principal Findings A new basal hadrosauroid dinosaur, Zh...
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Jeholosaurus shangyuanensis is a small ornithischian dinosaur from the Lower Cretaceous Yixian Formation of the Lujiatun locality, Liaoning Province, China. Here, we provide the first detailed description of its postcranial skeleton based on the holotype and four other well-preserved skeletons, and compare it with material of other primitive cerapo...
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The alvarezsauroid Linhenykus monodactylus from the Upper Cretaceous of Inner Mongolia, China is the first known monodactyl non-avian dinosaur, providing important information on the complex patterns of manual evolution seen in alvarezsauroids. Here we provide a detailed description of the osteology of this taxon. Linhenykus shows a number of featu...
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Comment on “Embryological evidence identifies wing digits in birds as digits 1, 2, and 3.”
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The Troodontidae represents one of the most bird-like theropod groups and plays an important role in our understanding of avian origins. Although troodontids have been known for over 150 years, few known derived troodontid specimens preserve significant portions of both the forelimb and the hindlimb. Here, we report a new troodontid taxon, Linheven...
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Archaeopteryx is widely accepted as being the most basal bird, and accordingly it is regarded as central to understanding avialan origins; however, recent discoveries of derived maniraptorans have weakened the avialan status of Archaeopteryx. Here we report a new Archaeopteryx-like theropod from China. This find further demonstrates that many featu...
Article
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Sixteen isolated theropod teeth were discovered in two areas in the upper Middle–lower Upper Jurassic Shishugou Formation of the Xinjiang Uyghur Autonomous Region, northwest China. This sample can be divided on the basis of qualitative features and simple quantitative metrics into seven tooth morphotypes, each of which probably represents a distinc...
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Digital reduction is a striking evolutionary phenomenon that is clearly exemplified in theropod dinosaurs by the functionally didactyl manus of tyrannosaurids, the flight-adapted manus of birds (Aves), and the tridactyl but digit II-dominated manus of alvarezsauroids. The enlargement of manual digit II in alvarezsauroids and the concurrent reductio...
Article
A new alvarezsaurid theropod, Xixianykus zhangi gen. et sp. nov., is described based on a partial postcranial skeleton collected from the Upper Cretaceous Majiacun Formation of Xixia County, Henan Province. The new taxon can be diagnosed by the following autapomorphies: sacral rib-transverse process complexes and zygapophyses fused to form separate...
Article
A new alvarezsaurid theropod, Xixianykus zhangi gen. et sp. nov., is described based on a partial postcranial skeleton collected from the Upper Cretaceous Majiacun Formation of Xixia County, Henan Province. The new taxon can be diagnosed by the following autapomorphies: sacral rib-transverse process complexes and zygapophyses fused to form separate...
Article
Big Finger Alvarezsauroidea are an enigmatic group of theropods presumed to be closely related to birds, though most specimens are younger than Archaeopteryx . Choiniere et al. (p. 571 ; see news story by Stone) now describe a more complete early specimen, dating to about 160 million years ago, which supports the conclusion that Alvarezsauroidea ar...
Article
Theropods have traditionally been assumed to have lost manual digits from the lateral side inward, which differs from the bilateral reduction pattern seen in other tetrapod groups. This unusual reduction pattern is clearly present in basal theropods, and has also been inferred in non-avian tetanurans based on identification of their three digits as...
Article
A detailed description of the skull and mandible of the Chinese cerapodan ornithischian dinosaur Jeholosaurus shangyuanensis (Lower Cretaceous, Yixian Formation) is presented for the first time and this information is used to reassess its phylogenetic position. Jeholosaurus can be distinguished from all other cerapodans on the basis of one autapomo...
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Recent fossil discoveries have substantially reduced the morphological gap between non-avian and avian dinosaurs, yet avians including Archaeopteryx differ from non-avian theropods in their limb proportions. In particular, avians have proportionally longer and more robust forelimbs that are capable of supporting a large aerodynamic surface. Here we...
Data
FIGURE 1. Holotype skull and mandibles of Jeholosaurus shangyuanensis (IVPP V 12529) in A, dorsal, B, ventral and C, left lateral views. D, close up on left maxillary tooth row. Scale bars = 20 mm (A – C) and 10 mm (D).
Data
FIGURE 5. Referred skull and mandible of Jeholosaurus shangyuanensis (IVPP V 15716) in right lateral view. A, whole specimen and B, close up on jugal to illustrate its nodular ornamentation. Scale bars = 20 mm (A) and 10 mm (B).
Data
FIGURE 8. Referred specimens of Jeholosaurus shangyuanensis. A – D, IVPP V 15718. E – F, IVPP V 15719. A, F, right lateral view, B, dorsal view, C, E, ventral view and D, caudal view. Scale bars = 20 mm.
Data
FIGURE 2. Outline drawings of the holotype skull of Jeholosaurus shangyuanensis (IVPP V 12529) in A, dorsal and B, ventral views. Abbreviations: an, angular; aof, antorbital fossa; bo, basioccipital; bs, basisphenoid; cr, cervical rib; d, dentary; ect, ectopterygoid; en, endocast; epi, epipterygoid; ex, exoccipital / opisthotic; f, frontal; j, juga...
Data
FIGURE 3. Referred skull and mandible of Jeholosaurus shangyuanensis (IVPP V 12530) in right lateral view. A, whole skull and B, close up on snout to illustrate the line of foramina that perforates the nasal (individual foramina are marked with an arrow). Scale bars = 20 mm (A) and 5 mm (B).
Data
FIGURE 6. Outline drawing of the referred skull and mandible of Jeholosaurus shangyuanensis (IVPP V 15716) in right lateral view. Abbreviations: an, angular; aof, antorbital fossa; aofo, antorbital fenestra; d, dentary; epi, epipterygoid; j, jugal; l, lachrymal; mx, maxilla; mxt, maxillary teeth; n, nasal; nf, nasal foramina; orn, ornamentation; pb...
Data
FIGURE 7. Referred skull and mandible of Jeholosaurus shangyuanensis (IVPP V 15717) A, dorsal, B, ventral and C

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