Iron-coated Komodo dragon teeth and the complex dental enamel of carnivorous reptiles.

Komodo dragons (Varanus komodoensis) are the largest extant predatory lizards and their ziphodont (serrated, curved and blade-shaped) teeth make them valuable analogues for studying tooth structure, function and comparing with extinct ziphodont taxa, such as theropod dinosaurs. Like other ziphodont reptiles, V. komodoensis teeth possess only a thin coating of enamel that is nevertheless able to cope with the demands of their puncture-pull feeding. Using advanced chemical and structural imaging, we reveal that V. komodoensis teeth possess a unique adaptation for maintaining their cutting edges: orange, iron-enriched coatings on their tooth serrations and tips. Comparisons with other extant varanids and crocodylians revealed that iron sequestration is probably widespread in reptile enamels but it is most striking in V. komodoensis and closely related ziphodont species, suggesting a crucial role in supporting serrated teeth. Unfortunately, fossilization confounds our ability to consistently detect similar iron coatings in fossil teeth, including those of ziphodont dinosaurs. However, unlike V. komodoensis, some theropods possessed specialized enamel along their tooth serrations, resembling the wavy enamel found in herbivorous hadrosaurid dinosaurs. These discoveries illustrate unexpected and disparate specializations for maintaining ziphodont teeth in predatory reptiles.

New soft tissue data of pterosaur tail vane reveals sophisticated, dynamic tensioning usage and expands its evolutionary origins.

Pterosaurs were the first vertebrates to achieve powered flight. Early pterosaurs had long stiff tails with a mobile base that could shift their center of mass, potentially benefiting flight control. These tails ended in a tall, thin soft tissue vane that would compromise aerodynamic control and efficiency if it fluttered during flight like a flag in the wind. Maintaining stiffness in the vane would have been crucial in early pterosaur flight, but how this was achieved has been unclear, especially since vanes were lost in later pterosaurs and are absent in birds and bats. Here we use Laser-Stimulated Fluorescence imaging to reveal a cross-linking lattice within the tail vanes of early pterosaurs. The lattice supported a sophisticated dynamic tensioning system used to maintain vane stiffness, allowing the whole tail to augment flight control and the vane to function as a display structure.

Exceptional preservation and foot structure reveal ecological transitions and lifestyles of early theropod flyers.

Morphology of keratinised toe pads and foot scales, hinging of foot joints and claw shape and size all inform the grasping ability, cursoriality and feeding mode of living birds. Presented here is morphological evidence from the fossil feet of early theropod flyers. Foot soft tissues and joint articulations are qualitatively assessed using laser-stimulated fluorescence. Pedal claw shape and size are quantitatively analysed using traditional morphometrics. We interpret these foot data among existing evidence to better understand the evolutionary ecology of early theropod flyers. Jurassic flyers like Anchiornis and Archaeopteryx show adaptations suggestive of relatively ground-dwelling lifestyles. Early Cretaceous flyers then diversify into more aerial lifestyles, including generalists like Confuciusornis and specialists like the climbing Fortunguavis. Some early birds, like the Late Jurassic Berlin Archaeopteryx and Early Cretaceous Sapeornis, show complex ecologies seemingly unique among sampled modern birds. As a non-bird flyer, finding affinities of Microraptor to a more specialised raptorial lifestyle is unexpected. Its hawk-like characteristics are rare among known theropod flyers of the time suggesting that some non-bird flyers perform specialised roles filled by birds today. We demonstrate diverse ecological profiles among early theropod flyers, changing as flight developed, and some non-bird flyers have more complex ecological roles.

Preserved soft anatomy confirms shoulder-powered upstroke of early theropod flyers, reveals enhanced early pygostylian upstroke, and explains early sternum loss.

Anatomy of the first flying feathered dinosaurs, modern birds and crocodylians, proposes an ancestral flight system divided between shoulder and chest muscles, before the upstroke muscles migrated beneath the body. This ancestral flight system featured the dorsally positioned deltoids and supracoracoideus controlling the upstroke and the chest-bound pectoralis controlling the downstroke. Preserved soft anatomy is needed to contextualize the origin of the modern flight system, but this has remained elusive. Here we reveal the soft anatomy of the earliest theropod flyers preserved as residual skin chemistry covering the body and delimiting its margins. These data provide preserved soft anatomy that independently validate the ancestral theropod flight system. The heavily constructed shoulder and more weakly constructed chest in the early pygostylian Confuciusornis indicated by a preserved body profile, proposes the first upstroke-enhanced flight stroke. Slender ventral body profiles in the early-diverging birds Archaeopteryx and Anchiornis suggest habitual use of the pectoralis could not maintain the sternum through bone functional adaptations. Increased wing-assisted terrestrial locomotion potentially accelerated sternum loss through higher breathing requirements. Lower expected downstroke requirements in the early thermal soarer Sapeornis could have driven sternum loss through bone functional adaption, possibly encouraged by the higher breathing demands of a Confuciusornis-like upstroke. Both factors are supported by a slender ventral body profile. These data validate the ancestral shoulder/chest flight system and provide insights into novel upstroke-enhanced flight strokes and early sternum loss, filling important gaps in our understanding of the appearance of modern flight.

The exquisitely preserved integument of Psittacosaurus and the scaly skin of ceratopsian dinosaurs.

The Frankfurt specimen of the early-branching ceratopsian dinosaur Psittacosaurus is remarkable for the exquisite preservation of squamous (scaly) skin and other soft tissues that cover almost its entire body. New observations under Laser-Stimulated Fluorescence (LSF) reveal the complexity of the squamous skin of Psittacosaurus, including several unique features and details of newly detected and previously-described integumentary structures. Variations in the scaly skin are found to be strongly regionalized in Psittacosaurus. For example, feature scales consist of truncated cone-shaped scales on the shoulder, but form a longitudinal row of quadrangular scales on the tail. Re-examined through LSF, the cloaca of Psittacosaurus has a longitudinal opening, or vent; a condition that it shares only with crocodylians. This implies that the cloaca may have had crocodylian-like internal anatomy, including a single, ventrally-positioned copulatory organ. Combined with these new integumentary data, a comprehensive review of integument in ceratopsian dinosaurs reveals that scalation was generally conservative in ceratopsians and typically consisted of large subcircular-to-polygonal feature scales surrounded by a network of smaller non-overlapping polygonal basement scales. This study highlights the importance of combining exceptional specimens with modern imaging techniques, which are helping to redefine the perceived complexity of squamation in ceratopsians and other dinosaurs.

Oldest preserved umbilical scar reveals dinosaurs had 'belly buttons'.

BACKGROUND: In egg-laying amniotes, the developing embryo is tethered to a number of the extraembryonic membranes including the yolk sac and allantois that deliver oxygen and nutrients and remove metabolic waste products throughout embryonic development. Prior to, or soon after hatching, these membranes detach from the animal leaving a temporary or permanent umbilical scar (umbilicus) equivalent to the navel or 'belly button' in some placental mammals, including humans. Although ubiquitous in modern mammals and reptiles (including birds), at least early in their ontogeny, the umbilicus has not been identified in any pre-Cenozoic amniote. RESULTS: We report the oldest preserved umbilicus in a fossil amniote from a ~130-million-year-old early-branching ceratopsian dinosaur, Psittacosaurus. Under laser-stimulated fluorescence (LSF), the umbilicus is revealed as an elongate midline structure delimited by a row of paired scales on the abdomen. The relatively late ontogenetic stage (close to sexual maturity) estimated for the individual indicates that the umbilicus was probably retained throughout life. CONCLUSIONS: Unlike most extant reptiles and birds that lose this scar within days to weeks after hatching, the umbilicus of Psittacosaurus persisted at least until sexual maturity, similar to some lizards and crocodylians with which it shares the closest morphological resemblance. This discovery is the oldest record of an amniote umbilicus and the first in a non-avian dinosaur. However, given the variability of this structure in extant reptilian analogues, a persistent umbilical scar may not have been present in all non-avian dinosaurs.

Quadrupedal water launch capability demonstrated in small Late Jurassic pterosaurs.

Pterosaurs thrived in and around water for 160 + million years but their take-off from water is poorly understood. A purportedly low floating position and forward centre of gravity barred pterosaurs from a bird-like bipedal running launch. Quadrupedal water launch similar to extant water-feeding birds and bats has been proposed for the largest pterosaurs, such as Anhanguera and Quetzalcoatlus. However, quadrupedal water launch has never been demonstrated in smaller pterosaurs, including those living around the Tethys Sea in the Late Jurassic Solnhofen Lagoon. Using Laser-Stimulated Fluorescence, we singled out aurorazhdarchid specimen MB.R.3531 that alone preserved specific soft tissues among more than a dozen well-preserved Solnhofen pterosaur specimens. These soft tissues pertain to primary propulsive contact surfaces needed for quadrupedal water launch (pedal webbing and soft tissues from an articulated forelimb) that permit robust calculations of its dynamic feasibility without the need to make assumptions about contact areas. A first-principles-based dynamics model of MB.R.3531 reveals that quadrupedal water launch was theoretically feasible and that webbed feet significantly impacted launch performance. Three key factors limiting water launch performance in all pterosaurs are identified, providing a foundation for understanding water launch evolution: available propulsive contact area, forelimb extension range and forelimb extension power about the shoulder.

Newly detected data from Haestasaurus and review of sauropod skin morphology suggests Early Jurassic origin of skin papillae.

Discovered in 1852, the scaly skin belonging to Haestasaurus becklesii was the first to be described in any non-avian dinosaur. Accordingly, it has played a crucial role in the reconstruction of sauropod integument and dinosaurs more broadly. Here, we reassess this historic specimen using Laser-Stimulated Fluorescence (LSF), revealing extensive, previously unknown regions of skin that augment prior interpretations of its integumentary morphology and taphonomy. Under white light, polygonal-subrounded, convex scales are visible on one side of the block ('side A'), but LSF reveals extensive smaller and more flattened scales, which are diagenetically fragmented, on the reverse block surface ('side B'). Contrary to the prior interpretation that the visible scales are the epidermal undersides, the presence of convex, intrascale papilliform textures on side A suggests that the external skin surface is exposed. We define intrascale papillae and provide a review of sauropod skin morphology, which clarifies that intrascale papillae are unique to and widespread across stem Neosauropoda, and likely have an evolutionary origin in the Early Jurassic. Intrascale papillae may ultimately have been integral to the evolution of gigantism in this charismatic clade.

Morphology and distribution of scales, dermal ossifications, and other non-feather integumentary structures in non-avialan theropod dinosaurs.

Modern birds are typified by the presence of feathers, complex evolutionary innovations that were already widespread in the group of theropod dinosaurs (Maniraptoriformes) that include crown Aves. Squamous or scaly reptilian-like skin is, however, considered the plesiomorphic condition for theropods and dinosaurs more broadly. Here, we review the morphology and distribution of non-feathered integumentary structures in non-avialan theropods, covering squamous skin and naked skin as well as dermal ossifications. The integumentary record of non-averostran theropods is limited to tracks, which ubiquitously show a covering of tiny reticulate scales on the plantar surface of the pes. This is consistent also with younger averostran body fossils, which confirm an arthral arrangement of the digital pads. Among averostrans, squamous skin is confirmed in Ceratosauria (Carnotaurus), Allosauroidea (Allosaurus, Concavenator, Lourinhanosaurus), Compsognathidae (Juravenator), and Tyrannosauroidea (Santanaraptor, Albertosaurus, Daspletosaurus, Gorgosaurus, Tarbosaurus, Tyrannosaurus), whereas dermal ossifications consisting of sagittate and mosaic osteoderms are restricted to Ceratosaurus. Naked, non-scale bearing skin is found in the contentious tetanuran Sciurumimus, ornithomimosaurians (Ornithomimus) and possibly tyrannosauroids (Santanaraptor), and also on the patagia of scansoriopterygids (Ambopteryx, Yi). Scales are surprisingly conservative among non-avialan theropods compared to some dinosaurian groups (e.g. hadrosaurids); however, the limited preservation of tegument on most specimens hinders further interrogation. Scale patterns vary among and/or within body regions in Carnotaurus, Concavenator and Juravenator, and include polarised, snake-like ventral scales on the tail of the latter two genera. Unusual but more uniformly distributed patterning also occurs in Tyrannosaurus, whereas feature scales are present only in Albertosaurus and Carnotaurus. Few theropods currently show compelling evidence for the co-occurrence of scales and feathers (e.g. Juravenator, Sinornithosaurus), although reticulate scales were probably retained on the mani and pedes of many theropods with a heavy plumage. Feathers and filamentous structures appear to have replaced widespread scaly integuments in maniraptorans. Theropod skin, and that of dinosaurs more broadly, remains a virtually untapped area of study and the appropriation of commonly used techniques in other palaeontological fields to the study of skin holds great promise for future insights into the biology, taphonomy and relationships of these extinct animals.

Laser-stimulated fluorescence reveals unseen details in fossils from the Upper Jurassic Solnhofen Limestones.

Laser-stimulated fluorescence (LSF) has seen increased use in palaeontological investigations in recent years. The method uses the high flux of laser light of visible wavelengths to reveal details sometimes missed by traditional long-wave ultraviolet (UV) methods using a lamp. In this study, we compare the results of LSF with UV-A-generated fluorescence on a range of fossils from the Upper Jurassic Solnhofen Limestone Konservat-Lagerstätte of Bavaria, Germany. The methodology follows previous protocols of LSF with modifications made to enhance laser beam intensity, namely keeping the laser at a constant distance from the specimen, using a camera track. Our experiments show that along with making surface details more vivid than UV-A or revealing them for the first time, LSF has the additional value of revealing shallow subsurface specimen detail. Fossil decapods from the Solnhofen Limestone reveal full body outlines, even under the matrix, along with details of segmentation within the appendages such as limbs and antennae. The results indicate that LSF can be used on invertebrate fossils along with vertebrates and may often surpass the information provided by traditional UV methods.

Pterosaurs evolved a muscular wing-body junction providing multifaceted flight performance benefits: Advanced aerodynamic smoothing, sophisticated wing root control, and wing force generation.

Pterosaurs were the first vertebrate flyers and lived for over 160 million years. However, aspects of their flight anatomy and flight performance remain unclear. Using laser-stimulated fluorescence, we observed direct soft tissue evidence of a wing root fairing in a pterosaur, a feature that smooths out the wing-body junction, reducing associated drag, as in modern aircraft and flying animals. Unlike bats and birds, the pterosaur wing root fairing was unique in being primarily made of muscle rather than fur or feathers. As a muscular feature, pterosaurs appear to have used their fairing to access further flight performance benefits through sophisticated control of their wing root and contributions to wing elevation and/or anterior wing motion during the flight stroke. This study underscores the value of using new instrumentation to fill knowledge gaps in pterosaur flight anatomy and evolution.

Response to Serrano and Chiappe.

In the recent study in Current Biology by Pei and colleagues1, we used two proxies - wing loading and specific lift - to reconstruct powered flight potential across the vaned feathered fossil pennaraptorans. The results recovered multiple origins of powered flight. We respectfully disagree with the criticism raised by Serrano and Chiappe2 that wing loading and specific lift, used in sequence, fail to discriminate between powered flight and gliding. We will explain this in reference to our original conservative approach.

Burmese Amber Compared Using Micro-Attenuated Total Reflection Infrared Spectroscopy and Ultraviolet Imaging.

Attenuated Total Reflection (ATR) spectroscopy coupled with a microscope allows for the analysis of specimens without any preparation, spatially correlated with the morphology of the specimen. These characteristics make micro-ATR systems very useful for studying gemstones and in particular amber samples. Indeed, in this report, the micro-ATR technique was used to characterize three Burmite samples, as case studies. Particularly, focusing the ATR crystal spot on the amber surface where a difference in the Ultraviolet (UV) reaction was previously detected, thereby relative differences in the IR spectrum could be analyzed. Here we present a methodology combining the UV imaging technique with the micro-Attenuated total reflection Fourier transform infrared spectroscopy (micro-ATR/FT-IR) analyses, in order to correlate the fluorescence information with the molecular vibrational modes of amber and their relative spatial distribution. Finally, this work is intended as an initial methodology study and part of the amber characterization project, focused on characterizing the Burmese amber collection of the Peretti Museum Foundation from several disciplines (i.e., palaeoentomology, vibrational spectroscopy, neutron tomography, etc.).

Archaeopteryx feather sheaths reveal sequential center-out flight-related molting strategy.

Modern flying birds molt to replace old and worn feathers that inhibit flight performance, but its origins are unclear. We address this by presenting and evaluating a ~150 million year old record of molting in a feathered dinosaur from the early bird Archaeopteryx. Laser-Stimulated Fluorescence revealed feather sheaths that are otherwise invisible under white light. These are separated by one feather and are not in numerical sequential order and are mirrored in both wings. This indicates that a sequential center-out molting strategy was already present at the origins of flight, which is used in living falcons to preserve maximum flight performance. This strategy would have been a welcome advantage for early theropod flyers that had poor flight capabilities. This discovery provides important insights into how birds refined their early flight capabilities before the appearance of the keeled sternum, pygostyle and triosseal canal.

Aerodynamics Show Membrane-Winged Theropods Were a Poor Gliding Dead-end.

The bizarre scansoriopterygid theropods Yi and Ambopteryx had skin stretched between elongate fingers that form a potential membranous wing. This wing is thought to have been used in aerial locomotion, but this has never been tested. Using laser-stimulated fluorescence imaging, we re-evaluate their anatomy and perform aerodynamic calculations covering flight potential, other wing-based behaviors, and gliding capabilities. We find that Yi and Ambopteryx were likely arboreal, highly unlikely to have any form of powered flight, and had significant deficiencies in flapping-based locomotion and limited gliding abilities. Our results show that Scansoriopterygidae are not models for the early evolution of bird flight, and their structurally distinct wings differed greatly from contemporaneous paravians, supporting multiple independent origins of flight. We propose that Scansoriopterygidae represents a unique but failed flight architecture of non-avialan theropods and that the evolutionary race to capture vertebrate aerial morphospace in the Middle to Late Jurassic was dynamic and complex.

Reanalysis of putative ovarian follicles suggests that Early Cretaceous birds were feeding not breeding.

We address the identity of putative ovarian follicles in Early Cretaceous bird fossils from the Jehol Biota (China), whose identification has previously been challenged. For the first time, we present a link to the botanical fossil record, showing that the "follicles" of some enantiornithine fossils resemble plant propagules from the Jehol Biota, which belong to Carpolithes multiseminalis. The botanical affinities of this "form-taxon" are currently unresolved, but we note that C. multiseminalis propagules resemble propagules associated with cone-like organs described as Strobilites taxusoides, which in turn are possibly associated with sterile foliage allocated to Liaoningcladus. Laser-Stimulated Fluorescence imaging furthermore reveals different intensities of fluorescence of "follicles" associated with a skeleton of the confuciusornithid Eoconfuciusornis zhengi, with a non-fluorescent circular micro-pattern indicating carbonaceous (or originally carbonaceous) matter. This is inconsistent with the interpretation of these structures as ovarian follicles. We therefore reaffirm that the "follicles" represent ingested food items, and even though the exact nature of the Eoconfuciusornis stomach contents remains elusive, at least some enantiornithines ingested plant propagules.

Disassociated rhamphotheca of fossil bird Confuciusornis informs early beak reconstruction, stress regime, and developmental patterns.

Soft tissue preservation in fossil birds provides a rare window into their anatomy, function, and development. Here, we present an exceptionally-preserved specimen of Confuciusornis which, through Laser-Stimulated Fluorescence imaging, is identified as preserving a disassociated rhamphotheca. Reconstruction of the in vivo position of the rhamphotheca validates the association of the rhamphotheca with two previous confuciusornithid specimens while calling that of a third specimen into question. The ease of dissociation is discussed and proposed with a fourth specimen alongside finite element analysis as evidence for preferential soft-food feeding. However, this proposition remains tentative until there is a better understanding of the functional role of beak attachment in living birds. Differences in post-rostral extent and possibly rhamphotheca curvature between confuciusornithids and modern birds hint at developmental differences between the two. Together, this information provides a wealth of new information regarding the nature of the beak outside crown Aves.