Richards Spur Research Articles

Size and shape heterodonty in the early Permian synapsid Mesenosaurus efremovi.

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.

Visual methods for documenting the preservation of large-sized synapsids at Richards Spur

Large isolated skeletal elements, including those of sphenacodontid and ophiacodontid synapsids from the upland cave systems of the Richards Spur locality, Oklahoma, are described. Multiple forms of visual representation, including coquille and stipple drawings, are used to document and examine the isolated elements. A fragmentary anterior portion of a dentary has the sphenacodontid four-leaf clover plicidentine attachment within the tooth roots, but the teeth are all of uniform size, and the symphyseal area is relatively slender and does not curve upward, suggesting that the specimen may not belong to any known member of the clade. A larger humerus with only the distal end preserved and a complete astragalus have distinct characteristics which are attributable to the sphenacodontid Dimetrodon Cope, 1878. A second, smaller humerus was identified to belong to Ophiacodon Marsh, 1878, cf. O. navajovicus and represents the first record of an ophiacodontid at Richards Spur. Finally, two large sphenacodontid interclavicles were discovered, with one having unusual growths representing a pathological condition. Typically, large amniotes are quite rare in this early Permian upland ecosystem, but the discovery of the new material shows that large synapsids are present at Richards Spur.

A new trematopid from the lower Permian of Oklahoma and new insights into the genus Acheloma.

Modern-day terrestrial amphibians pale in comparison to their monstrous ancient relatives, the Late Carboniferous and Early Permian trematopid temnospondyls. With a skeleton that clearly indicated a terrestrial mode of life and armed with an impressive set of large, recurved marginal dentition and palatal fangs for holding their prey-this group of terrestrial temnospondyls roamed North America and Central Europe as a top predator. Lack of substantial informative fossil material has previously limited our understanding of trematopid diversity and ontogeny. Fortunately, this has improved in the last few decades with the help of exceptional localities like the Early Permian locality Richards Spur. While multiple species of dissorophid temnospondyl have been described from Richards Spur, only one trematopid species has been confidently recognized -Acheloma dunni. Here, we report on the presence of a new large, relatively mature trematopid skull from this famous locality, found encased within a limestone rich block composed mostly of skeletal remains from several other taxa. With the help of neutron computed tomography (nCT), a non-invasive method of analyzing internal and external morphologies, this specimen has revealed several features consistent with the genus, Acheloma, but distinct from Acheloma cumminsi and Acehloma dunni. The identification of these new features, in addition to the characters it shares with other Acheloma species, not only constitute it as a new species of this genus, but also challenges the notion of having synonymized Acheloma dunni with Acheloma cumminsi. In this study, we also unveil new anatomical characters that are potentially independent of ontogeny and could therefore help clarify some of the phylogenetic relationships of this fascinating group of Paleozoic terrestrial predators.

Paleozoic cave system preserves oldest-known evidence of amniote skin.

The richest and most diverse assemblage of early terrestrial tetrapods is preserved within the infilled cave system of Richards Spur, Oklahoma (289-286 Mya1). Some of the oldest-known terrestrial amniotes2,3 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.4 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.

New specimens of the early Permian apex predator Varanops brevirostris at Richards Spur, Oklahoma, with histological information about its growth pattern.

An articulated pelvic region and additional isolated material of Varanops brevirostris, which are indistinguishable from those of the generotype from the Cacops bonebed, demonstrate the presence of this large varanopid at the Richards Spur locality. The articulated specimen includes lumbar, sacral, and anterior caudal vertebrae, partial pelvis, femur, and proximal part of tibia, confirming the autapomorphies previously suggested for this species. These include the presence of distinct blade-like shapes of the neural spines in the sacral region, the presence of deeply excavated pubis, and the presence of a distinct transverse ridge on the ventral surface of the femur distal to the intertrochanteric fossa. It has also been found that the transverse ridges and grooves become larger during ontogeny since the juvenile specimen did not exhibit a well-developed ridge. Histological analysis of isolated limb bones and neutron computed tomography (nCT) of the articulated specimen indicate that the latter likely belonged to an adult individual. This is in contrast to the other varanopid at Richards Spur, the significantly smaller, more gracile predator Mesenosaurus efremovi, which also shows the presence of growth lines and the external fundamental system with an estimated minimum age of fourteen.

An intriguing new diapsid reptile with evidence of mandibulo-dental pathology from the early Permian of Oklahoma revealed by neutron tomography.

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.

Revised Description of the Early Permian Recumbirostran “Microsaur” Nannaroter mckinziei Based on New Fossil Material and Computed Tomographic Data

The early Permian Richards Spur locality of Oklahoma has produced abundant material of numerous terrestrial fossil tetrapods, including various “microsaurs,” several of which are considered to belong to the clade Recumbirostra. We present a new partial skull of the recumbirostran “microsaur”Nannaroter mckinziei; through computed tomography (CT) analysis of both this new specimen and the holotype, we provide an updated description of the taxon. This new description provides novel information regarding several regions that could not be examined previously due to either being absent in the holotype or difficult to access. This includes missing and obscured aspects of the skull roof, braincase, lower jaw, and the palatal region. Furthermore, the new information obtained from this description was used to update phylogenetic character codings ofNannaroter, and a revised phylogenetic analysis was conducted. The results of this updated analysis are congruent with those of other recent phylogenetic analyses of recumbirostran “microsaurs.” This new information adds to the ever-growing body of early tetrapod CT data, which has been, and will continue to be, important in revealing details regarding early tetrapod anatomy, interrelationships, paleoecology, and evolution.

Multiple Tooth-Rowed Parareptile From the Early Permian of Oklahoma

The Dolese Limestone Quarry near Richards Spur, Oklahoma includes an elaborate system of caves which have been infilled with early Permian fossil rich sediments. In operation for more than a century, the quarry yielded vast numbers of disarticulated skeletal elements of the most diverse assemblage of fully terrestrial tetrapods from the Paleozoic. Excavations carried out in this century are distinct in producing large numbers of articulated and semiarticulated skeletons, including numerous new taxa. Dolese is therefore unique among early Permian localities in being home to a diverse assemblage of small parareptiles, including two species ofDelorhynchus. Here we describe a new species of acleistorhinid, characterized by the presence of multiple tooth rows on the dentary, that can be identified with confidence as a third new species ofDelorhynchus. The multiple tooth rowed condition is deemed not to be a pathological condition, and appears to have formed in the same manner as in the captorhinid eureptileCaptorhinus agutithrough uneven growth of the tooth-bearing element.

The First Age of Reptiles? Comparing Reptile and Synapsid Diversity, and the Influence of Lagerstätten, During the Carboniferous and Early Permian

Terrestrial ecosystems during the Pennsylvanian (late Carboniferous) and Cisuralian (early Permian) are usually described in the literature as being dominated by synapsids, the mammal-line amniotes. The pelycosaurs (a paraphyletic grouping of synapsid families) have been considered more speciose, abundant, and ecologically diverse than contemporary reptile-line amniotes. However, this dominance has never been subjected to quantitative testing accounting for sampling bias. Moreover, in recent years the amniote phylogeny has undergone numerous revisions, with suggestions that varanopids and recumbirostran microsaurs fall within reptiles, and that diadectomorphs may be pelycosaurian-grade synapsids. An examination of local species richness (alpha diversity) of synapsids and reptiles during the Pennsylvanian and Cisuralian at different spatial scales shows that these taxonomic revisions have substantial impacts on relative diversity patterns of synapsids and reptiles. Synapsids are only found to be consistently more diverse through the early Permian when using the “traditional” taxonomy. The recent taxonomic updates produce diversity estimates where reptile diversity is consistent with, or in some cases higher than that of synapsids. Moreover, biases in preservation may affect patterns. Where preservation favors smaller vertebrates, e.g., Richards Spur, South Grandfield, reptiles overwhelmingly dominate. If smaller vertebrates are expected to make up the bulk of amniote diversity, as they do in the present day, such lagerstätten may be more representative of true diversity patterns. Therefore, the dominance of pelycosaurs during this interval should be reconsidered, and this interval may be considered the First Age of Reptiles.

Permian millipedes from the Fort Sill fissures of southwestern Oklahoma, with comments on allied taxa and millipedes preserved in karstic environments

AbstractPermian millipedes are rare, especially so considering the relative abundance of millipedes in Carboniferous rocks. We report an early Permian millipede fauna containing three new genera and species of millipedes (Oklahomasoma richardsspurensenew genus new species,Karstiulus fortsillensisnew genus new species, andDolesea subtilanew genus new species) found in fossil-producing pockets of the Fort Sill fissures exposed in the Dolese Quarry near Richards Spur, southwest Oklahoma, USA. These are the first new genera of invertebrates to be described from this site, one of the most prolific fossil-vertebrate sites in the world. We also comment on taxa with morphological similarities and note previously described occurrences of Permian millipedes as well as occurrences of fossil myriapods (millipedes and centipedes) in karst deposits (caves and fissure fills) in Europe, Africa, Asia, North America, and the Caribbean. In contrast with the forms found at Richards Spur, most of these previous accounts of millipedes found in caves and fissure fills are of Pleistocene forms that are closely allied to modern taxa. The taxa from Richards Spur bear some similarities to Pennsylvanian forms. Karst (cave and fissure) faunas should be ranked with concretion faunas, cannel coals, and amber faunas as a major source of fossil myriapods.UUID:http://zoobank.org/5d58e1fb-4e5b-4597-9cd9-5cc9e2096b4d

Computed tomographic analysis of the cranium of the early Permian recumbirostran ‘microsaur’ Euryodus dalyae reveals new details of the braincase and mandible

AbstractRecumbirostran ‘microsaurs’ are a clade of Palaeozoic tetrapods that possess numerous morphological adaptations for fossorial ecologies. Re‐study of many ‘microsaurs’ using tomographic methods has provided substantial new data on poorly known anatomy that informs their debated phylogenetic position. Recent studies have identified suites of features among recumbirostrans that place the group within crown Amniota, contrary to hypothesized positions on the amniote stem or the lissamphibian stem. Herein we describe the cranial anatomy of the early Permian gymnarthrid Euryodus dalyae through tomographic analysis of the holotype from South Grandfield, Oklahoma and new specimens from karst deposits near Richards Spur. The braincase of E. dalyae is composed of well‐ossified pleurosphenoids, orbitosphenoids that brace against the skull roof, and unpaired median ossifications. The otic capsules are well‐ossified, and the occiput is unconsolidated. Analysis of the mandibles, typically obscured in articulated specimens, reveals a second tooth row on the dentary, a feature previously unknown in ‘microsaurs’ that is reminiscent of the condition of the co‐occurring captorhinid Captorhinus aguti. The Richards Spur specimens share many of these features, including the second tooth row, but the neurocranium of the scanned specimen (OMNH 53519) differs from that of the holotype of E. dalyae (e.g. absence of unpaired median ossifications), and these specimens are referred to Euryodus sp. These data add to the growing neurocranial dataset of ‘microsaurs’, which is essential for iterative reevaluation of early tetrapod phylogeny. This discovery of multiple tooth rows in ‘microsaurs’ provides further support for the hypothesized close relationships between ‘microsaurs’ and reptiles.

Postcranial anatomy and histology of Seymouria, and the terrestriality of seymouriamorphs.

Seymouria is the best known of the seymouriamorphs, a group of Permo-Carboniferous reptiliomorphs with both terrestrial and aquatic taxa. The majority of research on Seymouria has focused on cranial anatomy, with few detailed descriptions or illustrations of the postcrania. We utilized neutron computed tomography (nCT) and histological sampling to provide updated, detailed figures that clarify details of the postcranial anatomy and to assess the development and histology of Seymouria through specimens from the early Permian Richards Spur locality. The correlation of morphological and histological data indicate rapid metamorphosis in this terrestrially capable stem amniote, with the youngest specimen being postmetamorphic despite being distinctly younger than premetamorphic individuals of Discosauriscus, the only other seymouriamorph to have been histologically sampled. The microanatomical data (e.g., semi-open medullary cavity) also substantiate the hypothesis that Seymouria was terrestrial based on interpretation of external features, although the persistence of a modestly developed medullary spongiosa in comparison to either Discosauriscus or to other co-occurring terrestrial tetrapods suggests additional nuances that require further exploration. In the absence of clearly recognizable postmetamorphic stages in several seymouriamorph taxa, it is difficult to determine the evolutionary trajectory of terrestriality within the clade. Our analysis provides the first histological characterization of the life history of Seymouria and highlights the need for further study of seymouriamorph ontogeny.

Bone histology of varanopids (Synapsida) from Richards Spur, Oklahoma, sheds light on growth patterns and lifestyle in early terrestrial colonizers.

Varanopids were a group of small to medium-sized synapsids whose fossil record spans the Carboniferous through middle Permian. Although their phylogenetic relationships have received some interest in recent years, little is known about other aspects of their palaeobiology, including their skeletal growth, allometry and habitat preference. Here, we describe varanopid long bone histology based on a sample of well-preserved femora from the lower Permian Richards Spur fissure fill locality, Comanche County, Oklahoma, USA. The sample includes five femora from at least two varanopid taxa-Mycterosaurus and the large varanodontine Varanops brevirostris-and four additional mycterosaurine femora not diagnosed to genus. Prior work on femoral bone compactness provided a baseline to make lifestyle inferences and evaluate whether varanopids were ancestrally terrestrial. Moreover, the large availability of specimens spanning different sizes made possible an assessment of size-related ontogenetic histovariability. All specimens revealed moderately dense cortical bone tissues composed of sparsely vascularized parallel-fibred and lamellar bone with radially arranged rows of longitudinal canals (mostly simple), and many preserved regularly spaced growth marks (annuli and lines of arrested growth) as in modern varanids. We show that bone histology has the potential to explain how ballast was shed and the skeleton lightened for terrestrial mobility in ancestral synapsids and their basal amniote kin, as well as how adjustments in postnatal growth influenced the evolution of larger body sizes in the terrestrial frontier. This article is part of the theme issue 'Vertebrate palaeophysiology'.

New information on the early Permian lanthanosuchoid Feeserpeton oklahomensis based on computed tomography.

The cave deposits of the Lower Permian Richards Spur locality in Oklahoma, USA, have produced an incredible number of terrestrial tetrapod taxa, many of which are currently only known from this locality. One of the many recent taxa to be described from the locality was the small lanthanosuchoid parareptile Feeserpeton oklahomensis. Represented by a well-preserved, near complete skull, F. oklahomensis would have been a small predatory reptile, likely preying upon arthropods, and contributes to the extensive tetrapod fauna that was present at Richards Spur. New computed tomography data of the holotype and only specimen has allowed us to visualize and describe previously obscured and inaccessible anatomy of this taxon. These areas include the mandibular ramus, the palate, the sphenethmoid, the epipterygoids, and the braincase. Furthermore, this new anatomical information allowed formerly unknown character codings to be updated, thus we also performed new phylogenetic analyses that incorporated this new information. The results of these updated phylogenetic analyses are very similar to those of past studies, with F. oklahomensis being found as the sister taxon to all other lanthanosuchoids.

A new varanopid synapsid from the early Permian of Oklahoma and the evolutionary stasis in this clade.

Varanopids are a basal clade of small- to medium-sized non-therapsid synapsids, whose range extends from the late Pennsylvanian to the late middle Permian, and are found in North America, Russia, Europe and South Africa. The greatest varanopid diversity is observed at the fossiliferous cave deposits near Richards Spur, Oklahoma, well known for the preservation of a complex early Permian upland community. Two previously described varanopids, Mycterosaurus and Varanops, are known only from fragmentary disarticulated material at Richards Spur. A third putative varanopid, Basicranodon fortsillensis, represented by a partial parasphenoid, has been synonymized with Mycterosaurus longiceps. This study reports on a new varanopid taxon, represented by substantially more complete material, including three nearly complete skulls. Such comprehensive cranial material allows for a detailed study of the taxon and its relationship to other varanopids. This new varanopid bears great morphological similarity to Mesenosaurus romeri from the middle Permian Mezen River Basin of northern Russia. Phylogenetic analysis recovers a sister relationship between this taxon and Me. romeri. This relationship, in conjunction with a detailed morphological comparison, supports the placement of this taxon within Mesenosaurus, as a new species, Me. efremovi. These results reveal an unexpected extension of the geographical and temporal range of Mesenosaurus, contributing to our understanding of varanopid dispersal. The extended persistence of this basal clade of predatory synapsids, underscored by the apparent evolutionary stasis of this genus, is unusual among Palaeozoic tetrapods. This phenomenon implies an exceptionally high degree of extended ecological resilience across major faunal and environmental transitions.

Neurocranial anatomy of Seymouria from Richards Spur, Oklahoma

ABSTRACTSeymouriamorphs are a group of Permo-Carboniferous tetrapods with both terrestrial and aquatic members. Since their initial discovery, they have been proposed as phylogenetic intermediates on the amniote stem between temnospondyl amphibians and crown amniotes and are thus frequently used in phylogenetic analyses of both early amniotes and, more broadly, Paleozoic tetrapods. We utilized neutron computed tomography (nCT) to study the morphology of the first cranial material of Seymouria from the early Permian karst deposits found at the Dolese Brothers Limestone Quarry near Richards Spur, Oklahoma. The presence of a mixture of purported autapomorphies of both of the better-known species of Seymouria, S. baylorensis and S. sanjuanensis, prevents referral to either species; the potential for ontogenetic variation in many of these characters highlights the need to reevaluate the taxonomy of these species. Our description of the tomographic data focuses on the braincase and otic capsule and helps to clarify details of previously poorly known neurocranial anatomy, such as an orbitosphenoid that expands dorsally to abut the overlying frontals. The prootic and the opisthotic are complex structures that preserve clear impressions of all three semicircular canals and that frame the laterally extensive otic tube and possibly the perilymphatic duct. Our analysis provides additional details regarding a previously described but poorly known ossification of the synotic tectum and highlights the ongoing uncertainty regarding the homology of such ossifications across Paleozoic tetrapods.

Permian metabolic bone disease revealed by microCT: Paget's disease-like pathology in vertebrae of an early amniote.

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.

A New Captorhinid From the Permian Cave System Near Richards Spur, Oklahoma, and the Taxic Diversity of Captorhinus at This Locality

The early Permian cave system in the Dolese Brothers Limestone Quarry near Richards Spur, Oklahoma represents a unique depositional environment that has been interpreted as preserving an upland biota. The quarry and the region around it represent Paleozoic cave systems that underwent periods of flooding not unlike present-day conditions that are commonly associated with monsoonal episodes. The Richards Spur locality is particularly rich in captorhinid eureptiles which represent one of the earliest reptilian clades to have evolved a specialized dentition. Although the multiple-tooth rowed Captorhinus aguti is the most abundant captorhinid at Richards Spur, at least one other species has been described (Captorhinus magnus) and assigned to the same genus, but five other captorhinid taxa have also been found. We describe a new member of the genus Captorhinus (Captorhinus kieranensis) and explore details of the dental anatomy against the two other members of the genus at Richards Spur, C. aguti and C. magnus, as well as with a member of the genus not presently known from Richards Spur (C. laticeps). Findings suggest that the nature of the ogival dentition described previously as a synapomorphy uniting C. aguti with C. magnus is not supported and we propose a more informative method for differentiating among dental characters within the clade. The discovery of a new species of Captorhinus provides additional evidence for captorhinid taxic diversity at Richards Spur and is supportive of niche partitioning, which is likely associated with reducing intra-specific competition within the clade. Finally, we argue that the captorhinid fossils at Richards Spur likely includes one additional, currently undescribed multiple-tooth rowed form, that renders the current practice of assigning disarticulated cranial remains, specifically dental fragments, to the species C. aguti problematic.

A small caseid synapsid, Arisierpeton simplex gen. et sp. nov., from the early Permian of Oklahoma, with a discussion of synapsid diversity at the classic Richards Spur locality.

The fossil record of caseids, a clade of faunivorous to large herbivorous Permian synapsids, is unusual in having a poorly documented history. Although Kungurian caseids are common in the well-known continental deposits of North America, and the fossil record of the group extends into the middle Permian (Guadalupian), with the presence of the large caseid Ennatosaurus in the Mezen Basin faunal assemblage, only two other occurrences are known in older Permian age sediments. One is an undescribed caseid from the Bromacker Quarry in Germany, and the second is Oromycter from the lower Permian of Richards Spur, Oklahoma. The former is known from several articulated skeletons, but the latter is known only from a handful of skeletal elements, including elements of the snout and lower jaw, some phalanges, and a few vertebrae. Here the fragmentary tooth bearing elements and dorsal vertebrae of another small caseid from Richards Spur are described, with a discussion of its significance in the context of caseid evolution, and the continuously expanding faunal list and taxic diversity at this locality.

A Juvenile Specimen of the Trematopid Acheloma From Richards Spur, Oklahoma and Challenges of Trematopid Ontogeny

Trematopids are a clade of terrestrial dissorophoid temnospondyls documented primarily from terrestrial Permo-Carboniferous environments in North America and Europe. Here we describe the complete skull and articulated mandibles of a diminutive trematopid specimen (OMNH 79318) from the Early Permian karst deposits near Richards Spur, Oklahoma. Based on aspects of the neurocranium (e.g., unossified sphenethmoid, prootics, epipterygoids), the specimen represents one of the best examples of a markedly immature trematopid, an important data point for understanding the early ontogeny of trematopids. Specifically, it provides evidence that variation in otic notch structure can be ontogenetically influenced, not only among eucacopine dissorophids but also among trematopids. We provisionally refer the specimen to cf. Acheloma based on the presence of a denticulate vomerine ridge and other qualitative features. However, we emphasize that the taxonomic referral is complicated by several factors that more broadly confound trematopid taxonomy. This includes a low sample size (n = 1) of many taxa and marked size, and presumed ontogenetic, disparity between the known size range of different taxa. Complementary reexamination of both Acheloma cumminsi and Acheloma dunni as part of this study also reveals that the former possesses lateral exposures of palate bones, the presence / absence of which was the only formal character that previously differentiated the two species, although other qualitative features (e.g., size of the internarial fontanelle) may differentiate these two species. With respect to OMNH 79318, the taxonomic referral is tentative because the specimen also shares many qualitative attributes with Phonerpeton pricei, a trematopid represented only by small-bodied, probably immature individuals. However, many of these shared features are likely to be influenced by ontogeny or size. The subsequent challenges that we encountered in our taxonomic referral suggest that ontogeny may be confounding taxonomy in both diagnoses and phylogenetic analyses of trematopids and emphasize the need for careful study of how this affects our understanding of trematopid intrarelationships.