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 palatal 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

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 kierani) 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 (Captorhinus 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. In addition, 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. Finally, we offer a method for a comprehensive examination of the dental characteristics, which can then be applied to explore taxic diversity at Richards Spur and examine one of the earliest examples of niche specialization. As a consequence of this research, additional insight into exploring biological interactions between Paleozoic taxa can be examined, with an opportunity to shed light on what might have driven these evolutionary processes

New material of the ‘microsaur’ Llistrofus from the cave deposits of Richards Spur, Oklahoma and the paleoecology of the Hapsidopareiidae

The Hapsidopareiidae is a group of “microsaurs” characterized by a substantial reduction of several elements in the cheek region that results in a prominent, enlarged temporal emargination. The clade comprises two markedly similar taxa from the early Permian of Oklahoma, Hapsidopareion lepton and Llistrofus pricei, which have been suggested to be synonymous by past workers. Llistrofus was previously known solely from the holotype found near Richards Spur, which consists of a dorsoventrally compressed skull in which the internal structures are difficult to characterize. Here, we present data from two new specimens of Llistrofus. This includes data collected through the use of neutron tomography, which revealed important new details of the palate and the neurocranium. Important questions within “Microsauria” related to the evolutionary transformations that likely occurred as part of the acquisition of the highly modified recumbirostran morphology for a fossorial ecology justify detailed reexamination of less well-studied taxa, such as Llistrofus. Although this study eliminates all but one of the previous features that differentiated Llistrofus and Hapsidopareion, the new data and redescription identify new features that justify the maintained separation of the two hapsidopareiids. Llistrofus possesses some of the adaptations for a fossorial lifestyle that have been identified in recumbirostrans but with a lesser degree of modification (e.g., reduced neurocranial ossification and mandibular modification). Incorporating the new data for Llistrofus into an existing phylogenetic matrix maintains the Hapsidopareiidae’s (Llistrofus + Hapsidopareion) position as the sister group to Recumbirostra. Given its phylogenetic position, we contextualize Llistrofus within the broader “microsaur” framework. Specifically, we propose that Llistrofus may have been fossorial but was probably incapable of active burrowing in the fashion of recumbirostrans, which had more consolidated and reinforced skulls. Llistrofus may represent an earlier stage in the step-wise acquisition of the derived recumbirostran morphology and paleoecology, furthering our understanding of the evolutionary history of “microsaurs.

A Phylogeny and Timescale for the Evolution of Pseudocheiridae (Marsupialia: Diprotodontia) in Australia and New Guinea

Pseudocheiridae (Marsupialia: Diprotodontia) is a family of endemic Australasian arboreal folivores, more commonly known as ringtail possums. Seventeen extant species are grouped into six genera (Pseudocheirus, Pseudochirulus, Hemibelideus, Petauroides, Pseudochirops, Petropseudes). Pseudochirops and Pseudochirulus are the only genera with representatives on New Guinea and surrounding western islands. Here, we examine phylogenetic relationships among 13 of the 17 extant pseudocheirid species based on protein-coding portions of the ApoB, BRCA1, ENAM, IRBP, Rag1, and vWF genes. Maximum parsimony, maximum likelihood, and Bayesian methods were used to estimate phylogenetic relationships. Two different relaxed molecular clock methods were used to estimate divergence times. Bayesian and maximum parsimony methods were used to reconstruct ancestral character states for geographic provenance and maximum elevation occupied. We find robust support for the monophyly of Pseudocheirinae (Pseudochirulus + Pseudocheirus), Hemibelidinae (Hemibelideus + Petauroides), and Pseudochiropsinae (Pseudochirops + Petropseudes), respectively, and for an association of Pseudocheirinae and Hemibelidinae to the exclusion of Pseudochiropsinae. Within Pseudochiropsinae, Petropseudes grouped more closely with the New Guinean Pseudochirops spp. than with the Australian Pseudochirops archeri, rendering Pseudochirops paraphyletic. New Guinean species belonging to Pseudochirops are monophyletic, as are New Guinean species belonging to Pseudochirulus. Molecular dates and ancestral reconstructions of geographic provenance combine to suggest that the ancestors of extant New Guinean Pseudochirops spp. and Pseudochirulus spp. dispersed from Australia to New Guinea ∼12.1–6.5 Ma (Pseudochirops) and ∼6.0–2.4 Ma (Pseudochirulus). Ancestral state reconstructions support the hypothesis that occupation of high elevations (>3000 m) is a derived feature that evolved on the terminal branch leading to Pseudochirops cupreus, and either evolved in the ancestor of Pseudochirulus forbesi, Pseudochirulus mayeri, and Pseudochirulus caroli, with subsequent loss in P. caroli, or evolved independently in P. mayeri and P. forbesi. Divergence times within the New Guinean Pseudochirops clade are generally coincident with the uplift of the central cordillera and other highlands. Diversification within New Guinean Pseudochirulus occurred in the Plio-Pleistocene after the establishment of the Central Range and other highlands