The study of morphological characteristics and growth information in fish scales is a crucial component of modern fishery biological research, while it has been less studied in fossil materials. This ...paper presents a detailed morphological description and growth analysis of a fossil ctenoid scale obtained from the Upper Cretaceous Campanian lacustrine deposits in northeastern China. The morphological features of this fossil scale are well-preserved and consistent with the structures found in ctenoid scales of extant fish species and display prominent ring ornamentation radiating outward from the central focus, with grooves intersecting the rings. A comparative analysis of the morphological characteristics between the fossil ctenoid scale and those well-studied extant fish Mugilidae allows us to explore the applicability of modern fishery biological research methods to the field of fossil scales. The scale length, scale width, the vertical distance from the focus to the apex of the scale, and the total number of radii have been measured. The age of the fish that possessed this ctenoid scale has been estimated by carefully counting the annuli, suggesting an age equal to or more than seven years. The distribution of growth rings on the scale potentially reflects the warm paleoclimatic condition and fish-friendly paleoenvironment prevalent during that period. This paper, moreover, serves as a notable application of fishery biological methods in the examination of fossil materials.
'Mycterosaurus' smithae, from the Cisuralian (early Permian) of Colorado, was first described in 1965 as a second species of the genus Mycterosaurus. While the type species of this genus, M. ...longiceps, has been shown by multiple cladistic analyses to belong to the basal synapsid family Varanopidae, 'M.' smithae has been largely ignored since its original description. Additional preparation and synchrotron scanning has revealed new significant information that supports the assignment of this species to a new genus: Vaughnictis gen. nov. Vaughnictis lacks many of the characteristics of mycterosaurines and varanopids in general: it lacks the slender femur, the linguo-labially compressed and strongly recurved teeth, and the lateral boss on the postorbital characteristic of this family. Instead, it possesses coronoid teeth, a large supratemporal, and a large pineal foramen positioned midway along the length of the parietal, features that support its assignment to Eothyrididae. Moreover, the postcranium shares many characters with the eothyridid Oedaleops. An expanded version of a recently published phylogenetic analysis of pelycosaurian-grade synapsids positions Vaughnictis as the sister taxon of Eothyris within the clade Eothyrididae. The addition of data on the postcranium of eothyridids and the inclusion of the recently-described basal caseid Eocasea confirms the recently-disputed position of caseasaurs as the most basal synapsids. As the parsimony analysis produced low support values and a lack of resolution due to missing data, additional analyses were undertaken using Bayesian and Implied Weights methods, which produced better resolution and relationships with higher support values. While the results are similar, alternative positions for the enigmatic Moscovian age (Carboniferous) synapsid Echinerpeton are suggested by Bayesian analysis; the parsimony analysis found it to be an ophiacodontid, while the Bayesian and Implied Weights analysis found it to be the sister to the Sphenacomorpha.
The initial stages of diapsid evolution, the clade that includes extant reptiles and the majority of extinct reptilian taxa, is surprisingly poorly known. Notwithstanding the hypothesis that ...varanopids are diapsids rather than synapsids, there are only four araeoscelidians and one neodiapsid present in the late Carboniferous and early Permian. Here we describe the fragmentary remains of a very unusual new amniote from the famous cave deposits near Richards Spur, Oklahoma, that we recognize as a diapsid reptile, readily distinguishable from all other early amniotes by the unique dentition and lower jaw anatomy. The teeth have an unusual reeding pattern on the crown (long parallel ridges with rounded surfaces), with some teeth posteriorly tilted and strongly recurved, while a ventral protuberance forms the anterior terminus of the dentary. Overall, the lower jaw is unusually slender with a flattened ventral surface formed by the dentary and splenial anteriorly and the angular in the mid-region. The presence of a very slender triradiate jugal revealed through computed tomography confirms the existence of a large lower temporal fenestra, while the medial edge of the maxilla and the anatomy of the palatine confirm the presence of a large suborbital fenestra. Computed tomography of this new taxon reveals maxillary innervation that is characteristically reptile, not synapsid. Although no other definitively identifiable skull roof elements exist, the suborbital fenestra borders preserved on the palatine and maxilla supports the hypothesis that this is a diapsid reptile. Interestingly, the right dentary shows evidence of pathology, a rarely reported occurrence in Paleozoic amniotes, with several empty tooth sockets filled by bone. This small predator with delicate subthecodont implanted dentition provides strong evidence that diapsid reptiles were already diversifying rapidly in the early Permian, but likely were relatively rare members of terrestrial vertebrate assemblages.
Abstract
The oldest known complex terrestrial vertebrate community included hypercarnivorous varanopids, a successful clade of amniotes with wide geographic and temporal distributions. Little is ...known about their dentition and feeding behaviour, but with the unprecedented number of specimens of the varanopid
Mesenosaurus
from cave deposits in Oklahoma, we show that it exhibited serrations on the tooth crowns, and exceptionally rapid rates of development and reduced longevity relative to other terrestrial amniotes. In contrast, the coeval large apex predator
Dimetrodon
greatly increased dental longevity by increasing thickness and massiveness, whereas herbivores greatly reduced tooth replacement rates and increased dental longevity. Insectivores and omnivores represented the primitive condition and maintained modest replacement rates and longevity. The varied patterns of dental development among these early terrestrial amniotes reveal a hidden aspect of dental complexity in the emerging diverse amniote community, very soon after their initial appearance in the fossil record.
The exquisite preservation of maxillary and mandibular fragments of Seymouria has allowed us to examine for the first time in detail the dental anatomy and patterns of development in this stem ...amniote. The results obtained through histological examination show that Seymouria has pleurodont implantation with ankylosis of the tooth to the labial side of the jawbone. The dentary and maxillary teeth exhibit similar dental characteristics, such as the attachment bone (alveolar bone) and cementum rising above the jawbone on the base of the tooth, and smooth carinae extending lingually toward the tooth apex. Additionally, the clear presence of plicidentine, infolding of dentine into the pulp cavity, was found within the tooth root extending into the tooth crown. Lastly, the tooth replacement pattern is alternating, illustrating that Seymouria retains the classic primitive condition for tetrapods, a pattern that is continued in amniotes. Our results provide an important basis for comparison with other stem amniotes and with amniotes.
Using histological examination to show that the stem amniote, Seymouria, exhibits pleurodont implantation with the clear presence of plicidentine within the tooth root extending into the crown. Additionally, the tooth attachment tissues were identified and the alternating tooth replacement pattern, illustrating that Seymouria retains the classic primitive condition for tetrapods. Our results provide an important basis for comparison with other stem amniotes and with amniotes.
Tooth implantation provides important phylogenetic and functional information about the dentitions of amniotes. Traditionally, only mammals and crocodilians have been considered truly thecodont, ...because their tooth roots are coated in layers of cementum for anchorage of the periodontal ligament, which is in turn attached to the bone lining the alveolus, the alveolar bone. The histological properties and developmental origins of these three periodontal tissues have been studied extensively in mammals and crocodilians, but the identities of the periodontal tissues in other amniotes remain poorly studied. Early work on dental histology of basal amniotes concluded that most possess a simplified tooth attachment in which the tooth root is ankylosed to a pedestal composed of "bone of attachment", which is in turn fused to the jaw. More recent studies have concluded that stereotypically thecodont tissues are also present in non-mammalian, non-crocodilian amniotes, but these studies were limited to crown groups or secondarily aquatic reptiles. As the sister group to Amniota, and the first tetrapods to exhibit dental occlusion, diadectids are the ideal candidates for studies of dental evolution among terrestrial vertebrates because they can be used to test hypotheses of development and homology in deep time. Our study of Permo-Carboniferous diadectid tetrapod teeth and dental tissues reveal the presence of two types of cementum, periodontal ligament, and alveolar bone, and therefore the earliest record of true thecodonty in a tetrapod. These discoveries in a stem amniote allow us to hypothesize that the ability to produce the tissues that characterize thecodonty in mammals and crocodilians is very ancient and plesiomorphic for Amniota. Consequently, all other forms of tooth implantation in crown amniotes are derived arrangements of one or more of these periodontal tissues and not simply ankylosis of teeth to the jaw by plesiomorphically retaining "bone of attachment", as previously suggested.
Bone remodeling is an essential physiological process in growth and healing. In modern systems deviations from normal bone physiology in the form of pathologies aid in the understanding of normal ...bone metabolism. Here we use external morphology and X-ray microtomography to diagnose and describe a metabolic bone disease in an amniote from the early Permian. The specimen consists of two fused tail vertebrae of a small varanopid from early Permian (289 million years old) cave deposits near Richards Spur, Oklahoma, USA. Inspection of the outer morphology reveals that the fusion encompasses the vertebral centra, zygopophyses and haemal arches, with the fusion zones distinctly swollen on the left side of the specimen. With visualization of its internal structure by microCT, this specimen is diagnosed as a complex metabolic bone disease. The radiological imaging suggests a pathologically high bone turnover rate, as shown by abnormal bone formation in some areas and increased bone resorption in others. This supports that the varanopid suffered from a metabolic bone disease similar to Paget's disease of bone as seen in humans today, which is linked to both genetic and viral factors. This finding extends the occurrence of Paget-like disease to the early Permian, and-provided a viral component was present-would also be by far the oldest evidence of viral infection in the fossil record.
Fossilized organic remains are important sources of information because they provide a unique form of biological and evolutionary information, and have the long-term potential for genomic ...explorations. Here we report evidence of protein preservation in a terrestrial vertebrate found inside the vascular canals of a rib of a 195-million-year-old sauropodomorph dinosaur, where blood vessels and nerves would normally have been present in the living organism. The in situ synchrotron radiation-based Fourier transform infrared (SR-FTIR) spectra exhibit the characteristic infrared absorption bands for amide A and B, amide I, II and III of collagen. Aggregated haematite particles (α-Fe
O
) about 6∼8 μm in diameter are also identified inside the vascular canals using confocal Raman microscopy, where the organic remains were preserved. We propose that these particles likely had a crucial role in the preservation of the proteins, and may be remnants partially contributed from haemoglobin and other iron-rich proteins from the original blood.
Paleozoic synapsids represent the first chapter in the evolution of this large clade that includes mammals. These fascinating terrestrial vertebrates were the first amniotes to successfully adapt to ...a wide range of feeding strategies, reflected by their varied dental morphologies. Evolution of the marginal dentition on the mammalian side of amniotes is characterized by strong, size and shape heterodonty, with the late Permian therapsids showing heterodonty with the presence of incisiform, caniniform, and multicuspid molariform dentition. Rarity of available specimens has previously prevented detailed studies of dental anatomy and evolution in the initial chapter of synapsid evolution, when synapsids were able to evolve dentition for insectivory, herbivory, and carnivory. Numerous teeth, jaw elements, and skulls of the hypercarnivorous varanopid Mesenosaurus efremovi have been recently discovered in the cave systems near Richards Spur, Oklahoma, permitting the first detailed investigation of the dental anatomy of a Paleozoic tetrapod using multiple approaches, including morphometric and histological analyses. As a distant stem mammal, Mesenosaurus is the first member of this large and successful clade to exhibit a type of dental heterodonty that combines size and morphological (shape) variation of the tooth crowns. Here we present the first evidence of functional differentiation in the dentition of this early synapsid, with three distinct dental regions having diverse morphologies and functions. The quality and quantity of preserved materials has allowed us to identify the orientation and curvature of the carinae (cutting edges), and the variation and distribution of the ziphodonty (serrations) along the carinae. The shape‐related heterodonty seen in this taxon may have contributed to this taxon's ability to be a successful mid‐sized predator in the taxonomically diverse community of early Permian carnivores, but may have also extended the ecological resilience of this clade of mid‐sized predators across major faunal and environmental transitions.
Here we present the first evidence of functional differentiation in the dentition of an early synapsid, Mesenosaurus efremovi. Mesenosaurus is the first member of this large and successful clade to exhibit a type of dental heterodonty that combines size and morphological (shape) variation of the tooth crowns, including variation and distribution of the ziphodonty (serrations) along the crowns cutting edges.
The richest and most diverse assemblage of early terrestrial tetrapods is preserved within the infilled cave system of Richards Spur, Oklahoma (289-286 Mya
). Some of the oldest-known terrestrial ...amniotes
are exquisitely preserved here because of early impregnation and encasement of organic material by oil-seep hydrocarbons within rapidly deposited clay-rich cave sediments under toxic anoxic conditions.
This phenomenon has also afforded the preservation of exceedingly rare integumentary soft tissues, reported here, providing critical first evidence into the anatomical changes marking the transition from the aquatic and semiaquatic lifestyles of anamniotes to the fully terrestrial lifestyles of early amniotes. This is the first record of a skin-cast fossil (3D carbonization of the skin proper) from the Paleozoic Era and the earliest known occurrence of epidermal integumentary structures. We also report on several compression fossils (carbonized skin impressions), all demonstrating similar external morphologies to extant crocodiles. A variety of previously unknown ossifications, as well as what are likely palpebral ossifications of the deeper dermis layer of the skin, are also documented. These fossils also serve as invaluable references for paleontological reconstructions. Chromatographic analysis of extractable hydrocarbons from bone and cave samples indicates that the source rock is the Devonian age Woodford Shale. Hydrocarbons derived from ancient marine organisms interacting with geologically younger terrestrial vertebrates have therefore resulted in the oldest-known preservation of amniote skin proper.
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