Xingsheng Jin

The First Dictyoolithus Egg Clutches from the Lishui Basin, Zhejiang Province, China

ABSTRACT Three clutches and eight additional eggs are described from the Upper Cretaceous Chichengshan Formation in the Lishui Basin of Zhejiang Province, China. The three clutches contain 6–18 partial or complete spherical eggs, 14–18 cm in diameter. The eggshell thickness and reticulate pore system are identical to Dictyoolithus hongpoensis Zhao, 1994. However, the eggshell lacks superimposed shell units previously reported in this oospecies. The difference in microstructure most likely results from the reliance on thin section analysis in the original study; furthermore, superimposed shell units are inadequately documented in all Dictyoolithus oospecies due to the absence of scanning electron microscopy (SEM) or low (×45) magnification of SEM imaging. Cladistic analysis and the presence of the following characters support a theropod affinity of Dictyoolithus hongpoensis: relatively narrow spacing of nucleation sites, mammillary cones with blocky calcite cleavage, prismatic columns, and parallel laminae within the shell units. We reevaluate previous assignment of Dictyoolithus eggs to the Dinosauroid-spherulitic Basic Type and advocate discontinuation of Basic Type and Morphotype in fossil egg classification.

The first in situ turtle clutch from the Cretaceous Tiantai Basin, Zhejiang Province, China

Abstract The first in situ turtle egg clutch reported from China comes from the Lower Cretaceous (Albian) Liangtoutang Formation in the Tiantai Basin, Zhejiang Province. This clutch originally contained a minimum of 27 eggs, but four eggs separated from the clutch during excavation. The spherical eggs vary from 34 to 52 mm in diameter. The eggshell is 0.7–1.0 mm thick and has straight, narrow shell units with parallel margins and a height-to-width ratio of 2.5–3:1. Two eggs from this clutch previously were used to establish the new oogenus and oospecies Tiantaioolithus jiangi Fang et al., 2003, within the oofamily Testudoolithidae Fang et al., 2003. Based on our examination of all available eggs from this clutch, we propose the following: (1) Testudoolithidae Fang et al., 2003, is a junior homonym for Testudoolithidae Hirsch, 1996; (2) Tiantaioolithus Fang et al., 2003, is a junior subjective synonym of Testudoolithus Hirsch, 1996; and (3) the eggs pertain to a distinct oospecies, namely Testudoolithus jiangi (new combination). A previously unreported, isolated egg from the Upper Cretaceous (stage unknown) Chichengshan Formation in the Tiantai Basin is also referred to Testudoolithus jiangi based on its similar size, shell thickness, and shell unit height-to-width ratio; this specimen thus extends the fossil record of this oospecies into the Late Cretaceous. Taphonomic assessment of the egg block suggests that the egg clutch was buried in the substrate in a manner similar to modern turtles. The large, spherical eggs and large number of eggs in the clutch indicate the eggs were laid by a turtle taxon of large body size. The thick eggshell and sparse pores penetrating the shell indicate adaptation for a relatively arid, terrestrial environment.

Eggs and clutches of the Spheroolithidae from the Cretaceous Tiantai basin, Zhejiang Province, China

Numerous discoveries in the Tiantai basin of Zhejiang Province, China, enrich our understanding of the parataxonomy, paleobiology and taphonomic histories of fossil eggs from a diverse array of Cretaceous oofamilies. We describe the most abundant of these egg types catalogued in the Zhejiang Museum of Natural History, Spheroolithus cf. zhangtoucaoensis (oofamily Spheroolithidae). Scanning electron microscopy, here utilised for the first time on Spheroolithus eggs from Tiantai, and petrographic microscopy reveal 0.81–1.37-mm thick eggshell composed of a single structural layer of calcite with slightly flaring shell units, irregular pores, horizontal accretion lines and a sweeping extinction pattern under crossed polars. This contradicts previous reports of the presence of two structural layers in Tiantai Spheroolithus. Clutches consist of 2–13 eggs arranged in an irregular, single-layered pattern. As these eggs are among the oldest Asian examples of Spheroolithus currently known, they may shed light on the early evolution of this oogenus. This study establishes the definitive presence of the Spheroolithidae in the Tiantai basin, contra recent reports, and provides the framework for ongoing examination of egg diversity and taphonomy in the Tiantai basin.

An avian egg from the Lower Cretaceous (Albian) Liangtoutang Formation of Zhejiang Province, China

ABSTRACT Mesozoic avian eggs are rare, especially from the mid-Cretaceous basins of Zhejiang Province, China. Here we report an avian egg from the Lower Cretaceous (Albian) Liangtoutang Formation. The specimen (JYM F0033) measures 50 mm × 32 mm and the 166-µm-thick eggshell consists of three structural layers of calcite. The mammillary layer (ML) and overlying continuous layer (CL) each measure approximately 46 µm, whereas the outermost, external layer (EL) measures 74 µm. Ratios of these layers are ML:CL:EL = 1:1:1.6. An external layer that exceeds the thickness of the continuous layer represents an autapomorphy of this new egg type, which we herein establish as Pachycorioolithus jinyunensis oogen. et oosp. nov. within Pachycorioolithidae oofam. nov. Documentation of eggshell features and their first occurrence in the fossil record provides phylogenetically important information that may potentially aid in clarifying the evolution of avian reproduction and biology.

A psittacosaurid-like basal neoceratopsian from the Upper Cretaceous of central China and its implications for basal ceratopsian evolution.

Psittacosauridae (parrot-beaked dinosaurs) represents the first major radiation of ceratopsians (horned dinosaurs). However, psittacosaurids are divergent from the general morphology found in other ceratopsians, and this has resulted in their uncertain systematic position among ceratopsians. Here we describe a new basal neoceratopsian dinosaur, Mosaiceratops azumai gen. et sp. nov. based on a partial semi-articulated skeleton recovered from the Upper Cretaceous Xiaguan Formation of Neixiang County, Henan Province, China. Although our phylogenetic analysis supports this taxon as the most basal neoceratopsian, Mosaiceratops exhibits many features previously considered unique to the Psittacosauridae among the basal Ceratopsia. These include a relatively highly positioned external naris, a proportionally large premaxilla, the nasal extending ventral to the external naris, slender postorbital and temporal bars, a large notch between the basal tubera, and the edentulous premaxilla. Thus, the discovery of Mosaiceratops reduces the morphological disparity between the Psittacosauridae and other basal ceratopsians. Character optimization suggests that basal neoceratopsians have re-evolved premaxillary teeth; a major reversal previously unknown in any dinosaur clade. The new specimen also highlights the mosaic nature of evolution among early ceratopsians and supports the phylogenetic hypothesis that the Psittacosauridae is a relatively derived clade, rather than the most basal group of the Ceratopsia.

An early juvenile specimen of Bolong yixianensis (Ornithopoda: Iguanodontia) from the Lower Cretaceous of Ningcheng County, Nei Mongol, China

We describe an early juvenile specimen (ZMNH M8812) of Bolong yixianensis from the Yixian Formation (Lower Cretaceous) of Ningcheng County, Nei Mongol, China. The specimen consists of an almost complete skeleton preserved two-dimensionally on a slab. The short and deep skull proportions and unfused neurocentral sutures in most preserved vertebrae suggest that the ZMNH M8812 is a juvenile individual. Osteohistological study confirms a very early developmental stage. The study reveals the ontogenetic changes of Bolong for the first time. The specimen revealed one additional autapomorphy for Bolong yixianensis: the lingual face of the maxillary crown is bounded by thickened mesial and distal margins and bisected by a prominent median principal ridge. The study revealed the following ontogenetic trends of Bolong: increased tooth rows in both maxilla and dentary, increased robustness of the jugal and scapula, the radius and ulna become more robust and shorter relative to the hindlimb and the metatarsals become proportionally shorter. ZMNH M8812 represents the first juvenile non-hadrosaurid iguanodontian specimen described from the Lower Cretaceous of eastern Asia.

Lay, brood, repeat: nest reuse and site fidelity in ecologic time for two Cretaceous troodontid dinosaurs

ABSTRACT Whereas ‘biological site fidelity’ refers to the regular reuse of a favored locale (e.g., breeding ground or nest) by an individual animal, ‘paleontological site fidelity’ typically refers to repeated use of a nesting locality by a herd or species over geologic time scales. Two new Cretaceous specimens from the Two Medicine Formation of Montana, U.S.A., and the Liantoutang Formation of Zhejiang, China, each preserve two closely superimposed clutches of the egg form Prismatoolithus. These eggs belong to the Troodontidae, small theropod dinosaurs sharing a close ancestry with Aves. In both specimens, eggs of a lower clutch are truncated at a level below what would normally preserve in an undisturbed hatched clutch. These traces differ from past examples of dinosaur site fidelity in (1) the close or cross-cutting relationship of the clutches, (2) the precise overlay of clutch atop clutch, and (3) the implication of nest reuse and, thus, site fidelity on an ecologic rather than geologic scale and at approximately the individual rather than species level. Given the likely extended occupation of troodontid nesting sites by attending adults, factors such as nesting success and territoriality may, in addition to favorable substrates, have influenced the behaviors recorded by these specimens. The arrangements of eggs as clutches within the geologic record represent trace fossils. Thus, they record past in situ behavior, providing important insight into dinosaur nesting. In addition, they can serve as independent indicators of substrate conditions and sedimentary history, potentially refining our understanding of paleoenvironments.

Evaluating deformation in Spheroolithus dinosaur eggs from Zhejiang, China

Lack of stratigraphic context for dinosaur eggs inhibits understanding of dinosaur reproductive biology and the taphonomic processes of egg preservation. Past taphonomic work suggests two features, compression ridges (sharp edge of broken eggshell around egg circumference) and deformation asymmetry (proportion of crushed to rounded sides of the egg), as geopetal structures. We examined these features across a large sample of Spheroolithus eggs from the Cretaceous of Zhejiang, China, to test their utility. On 103 isolated eggs, we determined asymmetry ratios (crushed side egg height divided by rounded side egg height) and observed an average asymmetry ratio of 0.71. Additional observations of in situ eggs demonstrate the stratigraphic downside as more rounded and less fractured, the stratigraphic upside as flatter with heavier fracturing and compression ridges as parallel to original bedding plane. Burial-caused fractures on the upper side of the egg allowed sediment to partially fill, subsequently supporting the bottom portion. Examining these features within 16 clutches allowed differentiation of biotic versus taphonomically altered arrangements. Three common clutch arrangements include planar (minimal egg overlap), offset (extreme overlap) and agglomerate (randomly arranged, closely packed). Analysis of egg strike and dip across clutches favours planar clutches as the principal configuration for Spheroolithus clutches.

A new juvenile specimen of Yunnanosaurus robustus (Dinosauria: Sauropodomorpha) from Early to Middle Jurassic of Chuxiong Autonomous Prefecture, Yunnan Province, China

An almost complete skeleton with partial cranial material (ZMNH-M8739) is recovered from the Early or Middle Jurassic of southwest China. ZMNH-M8739 is identified as a juvenile individual of basal sauropodomorph dinosaur, Yunnanosaurus robustus Young, 1951. The revised diagnoses are as follows: absence of anteroposterior expansion on the medial end of astragalus and dorsoventrally compressed medium shaft of the metatarsal IV. Unfused neural arch and finely grooved long bone surface texture indicate that this individual is in the immature growth stage. ZMNH-M8739 possesses the tooth–tooth wear facet on its mesial maxillary and dentary teeth. However, the distal maxillary teeth have coarse serrations. Such a characteristic dentition could represent a unique feeding mechanism of this animal. Finally, ZMNH-M8739 constitutes a monophyletic group with Y. robustus (holotype), and Y. huangi is nesting this clade in the phylogenetic tree of the present analysis. Comparison of juvenile and adult specimen reveals distinctive growth changes of Y. robustus. This clade is positioned in an unnamed clade at a sister taxon of Sauropoda. Finally, some members of the so-called prosauropod dinosaurs constitute a monophyletic group in the present result.

The most basal ankylosaurine dinosaur from the Albian–Cenomanian of China, with implications for the evolution of the tail club

The tail club knob is a highly specialized structure thought to characterize a subgroup of the ankylosaurine ankylosaurians, and the oldest documented tail club knob in the fossil record occurred in the Campanian ankylosaurine Pinacosaurus. Here we report a new ankylosaurid Jinyunpelta sinensis, gen. et sp. nov., from the Albian–Cenomanian Liangtoutang Formation, Jinyun County, Zhejiang, China. This is the first definitive and the best preserved ankylosaurid dinosaur ever found in southern China. Jinyunpelta possesses unique cranial features differs from other ankylosaurs including two paranasal apertures level with and posterior to the external naris, a triangular fossa on the anterodorsal edge of the maxilla, an antorbital fossa in the junction between the maxilla, lacrimal and jugal, and an anterior process of the prearticular that lies ventral to the splenial. Our phylogenetic analysis suggests Jinyunpelta as the most basal ankylosaurine dinosaur. Jinyunpelta has a tail club with interlocking caudal vertebrae and a well-developed tail club knob, it represents the oldest and the most basal ankylosaurian known to have a well-developed tail club knob. The new discovery thus demonstrates that a large and highly modified tail club evolved at the base of the ankylosaurine ankylosaurs at least about 100 million years ago.