A broad sample of extant turtles possesses a series of paired bones in the neck that are situated between the cervical vertebrae. These paired bones were originally proposed to be cervical rib remnants, but have more recently been interpreted as vestiges of intercentra. Here, we document, for the first time, the neck development of a pleurodire turtle, Emydura subglobosa, and identify blastematous structures, which partially recapitulate the ribs and intercentra of the plesiomorphic tetrapod condition. We identify blastematous "bridges" between intercentra and the corresponding ribs, which we homologize with the vestiges visible in extant turtles and with the remnant parapophyseal articulation processes of the intercentra of some stem taxa. Only the unpaired, median part of the intercentrum of the atlas is retained in adult turtles, but intercentra are recapitulated along the entire vertebral column during development; they are embedded in the cervical myosepta and serve as attachment sites for neck musculature. We also identify two rib rudiments in the occipital region, which may indicate that at least two vertebrae are integrated into the cranium of turtles in particular, and of amniotes in general.
Abstract Plesiosaurians evolved four wing-like flippers that are morphologically similar to each other and were most likely used in underwater flight. Plesiosaurians have been the subject of a long research history as well as a long history of misidentifications and misinterpretations, especially transposition of parts of or entire fore- and hind flippers. We identified the transposed fore- and hind flippers in a mounted Cryptoclidus eurymerus specimen (GPIT-PV-30092) on display in the Paleontological Collection of Tübingen University. It is likely that the fore- and hind flippers were accidentally transposed when the skeleton was mounted, although, amongst plesiosaurians, the fore- and hind flippers of Cryptoclidus eurymerus are some of the least similar-looking ones. This occurred either during a remounting of the skeleton from a free-standing armature on the ground to a freely “flying” skeleton hanging from the ceiling, or after a research project conducted on the specimen in the 1970s. We summarize osteological characteristics that can be used to correctly identify fore- and hind flippers of this species, and for better future assessment of the plesiosaurian locomotory system.
Almost all extant testudinids are highly associated with terrestrial habitats and the few tortoises with high affinity to aquatic environments are found within the genus Manouria.Manouria belongs to a clade which forms a sister taxon to all remaining tortoises and is suitable as a model for studying evolutionary transitions within modern turtles.We analysed the feeding behaviour of Manouria emys and due to its phylogenetic position, we hypothesise that the species might have retained some ancestral features associated with an aquatic lifestyle.We tested whether M. emys is able to feed both in aquatic and terrestrial environments.In fact, M. emys repetitively tried to reach submerged food items in water, but always failed to grasp them -no suction feeding mechanism was applied.When feeding on land, M. emys showed another peculiar behaviour; it grasped food items by its jaws -a behaviour typical for aquatic or semiaquatic turtles -and not by the tongue as generally accepted as the typical feeding mode in all tortoises studied so far.In M. emys, the hyolingual complex remained retracted during all food uptake sequences, but the food transport was entirely lingual based.The kinematical profiles significantly differed from those described for other tortoises and from those proposed from the general models on the function of the feeding systems in lower tetrapods.We conclude that the feeding behaviour of M. emys might reflect a remnant of the primordial condition expected in the aquatic ancestor of the tortoises.
Almost all extant land turtles are highly associated with terrestrial habitats and the few tortoises with high affinity to aquatic environment are found within the genus Manouria . Manouria belongs to a clade which forms the sister taxon to all remaining tortoises and is suitable to be used as a model for studying evolutionary transitions from water to land within modern turtles. We analysed the feeding behaviour of M. emys and due to its phylogenetic position, we hypothesise that the species might have retained some ancestral characteristics associated to aquatic lifestyle. We tested whether M. emys is able to feed both in aquatic and terrestrial environments as mud turtles do. In fact, M. emys repetitively tried to reach submerged food items in water, but always failed to grasp them and no suction feeding mechanism was applied. When feeding on land, M. emys showed another peculiar behaviour; it grasped food items by its jaws – a behaviour typical for aquatic or semiaquatic turtles – and not by the tongue as in the typical feeding mode in all tortoises studied so far. In M. emys , the hyolingual complex remained retracted during all food uptake sequences, but the food transport was entirely lingual based. The kinematical profile significantly differed from those described for other tortoises and from those proposed from the general models on the function of the feeding systems in lower tetrapods. We conclude that the feeding behaviour of M. emys might reflect a remnant of the primordial condition expected in the aquatic ancestor of tortoises.
Embryogenesis is the first step in the ontogenetic life journey of any individual, and is thus a starting point for natural selection to cause evolutionary change. There are slight variations in the timing of embryonic development, known as heterochrony, which may eventually lead to major differences in adult anatomy. To test this hypothesis, the embryonic development of three closely related lizard species, Darevskia armeniaca, Lacerta agilis, and L. viridis, which are adapted to different habitats, was compared by analyzing discrete timing characters. Both intra- and interspecific variation was detected. The latter may be interpreted as embryonic pre-adaptions to later adult lifestyles, demonstrating that developmental penetrance manifests within a few million years. Traits with large intraspecific temporal variation, such as limb-related features, were susceptible to natural selection. In particular, the mountain-dwelling, climbing species D. armeniaca showed embryonic preadaptions by an early developing limb anlagen. This observation demonstrated interspecific variation, which was elusive in a previous comparative study based on purely metric data of developing limb lengths, and highlighted the importance of multiple data sources to draw robust conclusions about evolutionary change. Timing differences indicated unexplored ecological adaptations of the poorly understood lifestyle of these lizards. Thus, embryonic research provides a platform to explore superficially hidden evolutionary adaptations of all organisms on Earth.
Abstract Development in marsupials is specialized towards an extremely short gestation and highly altricial newborns. As a result, marsupial neonates display morphological adaptations at birth related to functional constraints. However, little is known about the variability of marsupial skull development and its relation to morphological diversity. We studied bony skull development in five marsupial species. The relative timing of the onset of ossification was compared to literature data and the ossification sequence of the marsupial ancestor was reconstructed using squared-change parsimony. The high range of variation in the onset of ossification meant that no patterns could be observed that differentiate species. This finding challenges traditional studies concentrating on the onset of ossification as a marker for phylogeny or as a functional proxy. Our study presents observations on the developmental timing of cranial bone-to-bone contacts and their evolutionary implications. Although certain bone contacts display high levels of variation, connections of early and late development are quite conserved and informative. Bones that surround the oral cavity are generally the first to connect and the bones of the occipital region are among the last. We conclude that bone contact is preferable over onset of ossification for studying cranial bone development.