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Tanystropheus longobardicus is one of the most remarkable and iconic Triassic reptiles. Mainly known from the Middle Triassic conservation Lagerstätte of Monte San Giorgio on the Swiss-Italian border, it is characterized by an extraordinarily long and stiffened neck that is almost three times the length of the trunk, despite being composed of only 13 hyper-elongate cervical vertebrae [1-8]. Its paleobiology remains contentious, with both aquatic and terrestrial lifestyles having been proposed [1, 9-12]. Among the Tanystropheus specimens, a small morphotype bearing tricuspid teeth and a large morphotype bearing single-cusped teeth can be recognized, historically considered as juveniles and adults of the same species [4]. Using high-resolution synchrotron radiation microtomography (SRμCT), we three-dimensionally reconstruct a virtually complete but disarticulated skull of the large morphotype, including its endocast and inner ear, to reveal its morphology for the first time. The skull is specialized toward hunting in an aquatic environment, indicated by the placement of the nares on the top of the snout and a "fish-trap"-type dentition. The SRμCT data and limb bone paleohistology reveal that the large morphotype represents a separate species (Tanystropheus hydroides sp. nov.). Skeletochronology of the small morphotype specimens indicates that they are skeletally mature despite their small size, thus representing adult individuals of Tanystropheus longobardicus. The co-occurrence of these two species of disparate size ranges and dentitions provides strong evidence for niche partitioning, highlighting the surprising versatility of the Tanystropheus bauplan and the complexity of Middle Triassic nearshore ecosystems., Competing Interests: Declaration of Interests The authors declare no competing interests., (Copyright © 2020 Elsevier Inc. All rights reserved.)

The turtle shell is a complex structure that currently serves a largely protective function in this iconically slow-moving group [1]. Developmental [2, 3] and fossil [4-7] data indicate that one of the first steps toward the shelled body plan was broadening of the ribs (approximately 50 my before the completed shell [5]). Broadened ribs alone provide little protection [8] and confer significant locomotory [9, 10] and respiratory [9, 11] costs. They increase thoracic rigidity [8], which decreases speed of locomotion due to shortened stride length [10], and they inhibit effective costal ventilation [9, 11]. New fossil material of the oldest hypothesized stem turtle, Eunotosaurus africanus [12] (260 mya) [13, 14] from the Karoo Basin of South Africa, indicates the initiation of rib broadening was an adaptive response to fossoriality. Similar to extant fossorial taxa [8], the broad ribs of Eunotosaurus provide an intrinsically stable base on which to operate a powerful forelimb digging mechanism. Numerous fossorial correlates [15-17] are expressed throughout Eunotosaurus' skeleton. Most of these features are widely distributed along the turtle stem and into the crown clade, indicating the common ancestor of Eunotosaurus and modern turtles possessed a body plan significantly influenced by digging. The adaptations related to fossoriality likely facilitated movement of stem turtles into aquatic environments early in the groups' evolutionary history, and this ecology may have played an important role in stem turtles surviving the Permian/Triassic extinction event., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

The origin of the turtle shell has perplexed biologists for more than two centuries. It was not until Odontochelys semitestacea was discovered, however, that the fossil and developmental data could be synthesized into a model of shell assembly that makes predictions for the as-yet unestablished history of the turtle stem group. We build on this model by integrating novel data for Eunotosaurus africanus-a Late Guadalupian (∼260 mya) Permian reptile inferred to be an early stem turtle. Eunotosaurus expresses a number of relevant characters, including a reduced number of elongate trunk vertebrae (nine), nine pairs of T-shaped ribs, inferred loss of intercostal muscles, reorganization of respiratory muscles to the ventral side of the ribs, (sub)dermal outgrowth of bone from the developing perichondral collar of the ribs, and paired gastralia that lack both lateral and median elements. These features conform to the predicted sequence of character acquisition and provide further support that E. africanus, O. semitestacea, and Proganochelys quenstedti represent successive divergences from the turtle stem lineage. The initial transformations of the model thus occurred by the Middle Permian, which is congruent with molecular-based divergence estimates for the lineage, and remain viable whether turtles originated inside or outside crown Diapsida., (Copyright © 2013 Elsevier Ltd. All rights reserved.)