Nanhsiungchelyidae is a group of large Pan-Trionychia that lived in North America and Asia during the Cretaceous. Despite a long research history, their taxonomy and internal nodes are still controversial. Here, we erect a new taxon, Xianyuechelys yingliangi gen. et sp. nov., from the Upper Cretaceous Lianhe Formation of Ganzhou Basin, China. Xianyuechelys yingliangi is attributed to Nanhsiungchelyidae based on its carapace being covered by a sculpturing of large pits. However, the new species also possesses an unique combination of characteristics among nanhsiungchelyids such as a wide, crescentic nuchal, absence of a pair of anterolateral processes, and the fourth and fifth vertebral scutes that contact at a point. The holotype is a juvenile or sub-adult based on the incomplete ossification of carapace, but its morphological characteristics can mostly be regarded as taxonomically informative rather than ontogenetic in origin. Phylogenetic analysis suggests Xianyuechelys yingliangi forms a monophyletic group with Nanhsiungchelys spp. and Anomalochelys angulata, supported by two unambiguous synapomorphies: wide vertebral scutes and wide neurals. The phylogenetic analysis also suggests Basilemys, Jiangxichelys and 'Zangerlia' form a monophyletic group. Our new finding increases the known diversity of nanhsiungchelyids in southern China.

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 partial skeleton recovered from the lower Upper Cretaceous of Ganxian County, Ganzhou City, southern China, and comprising six articulated cervical vertebrae, two partial dorsal vertebrae, and a complete sacrum preserved in articulation with the first 17 caudal vertebrae and part of the right pelvis. Gandititan can be diagnosed on the basis of the following autapomorphies: long and narrow fossae present on the dorsal and ventral parts of the lateral surfaces of the cervical centra, sacral neural spines forming a dorsal platform with wave-shaped lateral margins, anteriormost six caudal vertebrae with bifurcated neural spines, the presence of prominent triangular flanges on the transverse processes of the anteriormost caudal vertebrae, a pair of slit-like foramina present on the ventral surface of each anterior caudal centrum, lateral surfaces of neural arches strongly excavated, additional spinoprezygapophyseal laminae present, and a prominent lamina extending horizontally between the bases of the pre- and postzygapophyses in some anterior caudal vertebrae. An expanded phylogenetic analysis places Gandititan as the sister taxon to Abdarainurus, within a clade of non-lithostrotian titanosaurs that also includes the Chinese titanosaurs Dongyangosaurus, Baotianmansaurus and Huabeisaurus, as well as the Argentine titanosaur Andesaurus. Such results imply the possible existence of a previously unrecognized group of titanosaurs in eastern Asia, and potential dispersal of titanosaurs between Asia and South America during the mid-Cretaceous.

Two nanhsiungchelyid carapaces from the Upper Cretaceous of Ganzhou Basin, Jiangxi Province, Southern China, are reported and assigned to Nanhsiungchelys sp. The new material reveals an unusual carapace morphology that was either not or poorly preserved in previously known Nanhsiungchelys specimens, including differentiated neurals, alternating costals, significantly modified scutes with vertebrals 4–5 contacting each other at a point, reduced pleurals 2–4 and greatly expanded lateral and posterior marginals. The discovery extends the geographical distribution of Nanhsiungchelys to the Ganzhou Basin.

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 in other Nanhsiungchelyidae. Besides, we also further describe the plastron of the holotype of Nanhsiungchelys yangi. These two specimens reveal that the plastron of Nanhsiungchelys spp. is partly hollow, and may have additional ossifications of the epiplastra. All of these suggest an unusual plastral morphology and evolution of Nanhsiungchelys.

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, Nanhsi-ungchelys 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 and plastron. The diagnostic features of Nanhsiungchelys include a large entire carapace length (& SIM;55.5 cm), a network of sculptures consisting of pits and ridges on the surface of the skull and shell, shallow cheek emargination and temporal emargination, deep nuchal emargination, and a pair of anterolateral processes on the carapace. However, Nanhsiungchelys yangi differs from the other species of Nanhsiungchelys mainly in having a triangular-shaped snout (in dorsal view) and wide anterolateral processes on the carapace. Additionally, some other characteristics (e.g., the premaxilla is higher than wide, the maxilla is unseen in dorsal views, a small portion of the maxilla extends posterior and ventral of the orbit, and the parietal is bigger than the frontal) are strong evidence to distinguish Nanhsiungchelys yangi from Nanhsiungchelys wuchingensis. A phylogenetic analysis of nanhsiungchelyids places Nanhsiungchelys yangi and Nanhsiungchelys wuchingensis as sister taxa. Nanhsiungchelys yangi and some other nanhsiungchelyids bear distinct anterolateral processes on the carapace, which have not been reported in any extant turtles and may have played a role in protecting the head. The Nanxiong Basin was extremely hot during the Late Cretaceous, and so we suggest that nanhsiungchelyids might have immersed themselves in mud or water to avoid the heat, similar to some extant tortoises. If they were capable of swimming, our computer simulations of fluid flow suggest the anterolateral processes could have reduced drag during locomotion.

提起白垩纪,你可能会想到在陆地奔跑的恐龙或者在天空翱翔的翼龙.实际上,在一亿年前的亚洲和北美洲,还生活着一类体型硕大的龟鳖.这些龟鳖被称为南雄龟,是和恐龙同行的生物.

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 and plastron. The diagnostic features of Nanhsiungchelys include a large entire carapace length (∼55.5 cm), a network of sculptures consisting of pits and ridges on the surface of the skull and shell, shallow cheek emargination and temporal emargination, deep nuchal emargination, and a pair of anterolateral processes on the carapace. However, Nanhsiungchelys yangi differs from the other species of Nanhsiungchelys mainly in having a triangular-shaped snout (in dorsal view) and wide anterolateral processes on the carapace. Additionally, some other characteristics (e.g., the premaxilla is higher than wide, the maxilla is unseen in dorsal views, a small portion of the maxilla extends posterior and ventral of the orbit, and the parietal is bigger than the frontal) are strong evidence to distinguish Nanhsiungchelys yangi from Nanhsiungchelys wuchingensis. A phylogenetic analysis of nanhsiungchelyids places Nanhsiungchelys yangi and Nanhsiungchelys wuchingensis as sister taxa. Nanhsiungchelys yangi and some other nanhsiungchelyids bear distinct anterolateral processes on the carapace, which have not been reported in any extant turtles and may have played a role in protecting the head. The Nanxiong Basin was extremely hot during the Late Cretaceous, and so we suggest that nanhsiungchelyids might have immersed themselves in mud or water to avoid the heat, similar to some extant tortoises. If they were capable of swimming, our computer simulations of fluid flow suggest the anterolateral processes could have reduced drag during locomotion.

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 and plastron. The diagnostic features of Nanhsiungchelys include a large entire carapace length (∼55.5 cm), a network of sculptures consisting of pits and ridges on the surface of the skull and shell, shallow cheek emargination and temporal emargination, deep nuchal emargination, and a pair of anterolateral processes on the carapace. However, Nanhsiungchelys yangi differs from the other species of Nanhsiungchelys mainly in having a triangular-shaped snout (in dorsal view) and wide anterolateral processes on the carapace. Additionally, some other characteristics ( e.g. , the premaxilla is higher than wide, the maxilla is unseen in dorsal views, a small portion of the maxilla extends posterior and ventral of the orbit, and the parietal is bigger than the frontal) are strong evidence to distinguish Nanhsiungchelys yangi from Nanhsiungchelys wuchingensis . A phylogenetic analysis of nanhsiungchelyids places Nanhsiungchelys yangi and Nanhsiungchelys wuchingensis as sister taxa. Nanhsiungchelys yangi and some other nanhsiungchelyids bear distinct anterolateral processes on the carapace, which have not been reported in any extant turtles and may have played a role in protecting the head. The Nanxiong Basin was extremely hot during the Late Cretaceous, and so we suggest that nanhsiungchelyids might have immersed themselves in mud or water to avoid the heat, similar to some extant tortoises. If they were capable of swimming, our computer simulations of fluid flow suggest the anterolateral processes could have reduced drag during locomotion.

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 lower jaw, as well as the anterior parts of the carapace and plastron. The diagnostic features of Nanhsiungchelys include a huge estimated body size (~55.5 cm), a special network of sculptures on the surface of the skull and shell, weak cheek emargination and temporal emargination, deep nuchal emargination, and a pair of anterolateral processes on the carapace. However, Nanhsiungchelys yangi differs from the other species of Nanhsiungchelys in having a triangular-shaped snout and wide anterolateral processes. A phylogenetic analysis of nanhsiungchelyids places Nanhsiungchelys yangi and Nanhsiungchelys wuchingensis as sister taxa. Some nanhsiungchelyids bear special anterolateral processes on the carapace, which are unknown in extant turtles. Here we test the function of these processes in Nanhsiungchelys yangi using computational fluid dynamics, and the results suggest these processes could enhance locomotory performance by remarkably reducing drag force when the animal was swimming through water. ### Competing Interest Statement The authors have declared no competing interest.

AbstractNanhsiungchelyidae 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 yangisp. nov., from the Upper Cretaceous of Nanxiong Basin, China. This is the second valid species ofNanhsiungchelys, and the holotype consists of a well-preserved skull and lower jaw, as well as the anterior parts of the carapace and plastron. The diagnostic features ofNanhsiungchelysinclude a huge estimated body size (~55.5 cm), a special network of sculptures on the surface of the skull and shell, weak cheek emargination and temporal emargination, deep nuchal emargination, and a pair of anterolateral processes on the carapace. However,Nanhsiungchelys yangidiffers from the other species ofNanhsiungchelysin having a triangular-shaped snout and wide anterolateral processes. A phylogenetic analysis of nanhsiungchelyids placesNanhsiungchelys yangiandNanhsiungchelys wuchingensisas sister taxa. Some nanhsiungchelyids bear special anterolateral processes on the carapace, which are unknown in extant turtles. Here we test the function of these processes inNanhsiungchelys yangiusing computational fluid dynamics, and the results suggest these processes could enhance locomotory performance by remarkably reducing drag force when the animal was swimming through water.

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 Nanhsiungchelyidae (Pan-Trionychia), an extinct group of large terrestrial turtles (possibly the species Yuchelys nanyangensis). The egg is rigid, spherical, and is one of the largest and thickest shelled Mesozoic turtle eggs known. Importantly, this specimen allowed identification of other nanhsiungchelyid egg clutches and comparison to those of Adocidae, as Nanhsiungchelyidae and Adocidae form the basal extinct clade Adocusia of the Pan-Trionychia (includes living soft-shelled turtles). Despite the differences in habitat adaptations, nanhsiungchelyids (terrestrial) and adocids (aquatic) shared several reproductive traits, including relatively thick eggshells, medium size clutches and relatively large eggs, which may be primitive for trionychoids (including Adocusia and Carrettochelyidae). The unusually thick calcareous eggshell of nanhsiungchelyids compared to those of all other turtles (including adocids) may be related to a nesting style adaptation to an extremely harsh environment.