Christian F. Kammerer

The early evolution of synapsids, and the influence of sampling on their fossil record

Abstract Synapsids dominated the terrestrial realm between the late Pennsylvanian and the Triassic. Their early evolution includes some of the first amniotes to evolve large size, herbivory, and macro-predators. However, little research has focused on the changes in diversity occurring during this early phase in their evolutionary history, with more effort concentrating on later events such the Permo-Triassic extinction. Here we assess synapsid diversity, at both the species and genus levels, between the Carboniferous (Moscovian) and the Middle Permian (Capitanian). A raw, taxic diversity (richness) estimate is generated, and we use two separate methods to correct for sampling biases in this curve. To remove the effect of anthropogenic sampling bias, we apply a recently published modification of the residual diversity method, and then generate a supertree, using matrix representation with parsimony to infer ghost lineages and obtain a phylogenetic diversity estimate. The general diversity pattern reflects the initial diversification of synapsids in the late Pennsylvanian and early Cisuralian, which was followed by an extinction event during the Sakmarian. Diversity recovered during the Artinskian and Kungurian, coinciding with the radiation of Caseidae, although other families begin to decline. A second extinction event occurred across the Kungurian/Roadian boundary, in which Edaphosauridae and Ophiacodontidae died out although Caseidae and Therapsida diversified. The sampling-corrected curves reveal further extinction during the Roadian, although therapsids were again unaffected. Pelycosaurian-grade synapsids survived during the Wordian and Capitanian, but were a minor part of an otherwise therapsid-dominated fauna. Evidence of significant anthropogenic sampling bias calls into question previous diversity studies that have not employed sampling correction.

On the Validity and Phylogenetic Position of Eubrachiosaurus browni, a Kannemeyeriiform Dicynodont (Anomodontia) from Triassic North America

The large dicynodont Eubrachiosaurus browni from the Upper Triassic Popo Agie Formation of Wyoming is redescribed. Eubrachiosaurus is a valid taxon that differs from Placerias hesternus, with which it was previously synonymized, by greater anteroposterior expansion of the scapula dorsally and a very large, nearly rectangular humeral ectepicondyle with a broad supinator process. Inclusion of Eubrachiosaurus in a revised phylogenetic analysis of anomodont therapsids indicates that it is a stahleckeriid closely related to the South American genera Ischigualastia and Jachaleria. The recognition of Eubrachiosaurus as a distinct lineage of North American dicynodonts, combined with other recent discoveries in the eastern USA and Europe, alters our perception of Late Triassic dicynodont diversity in the northern hemisphere. Rather than being isolated relicts in previously therapsid-dominated regions, Late Triassic stahleckeriid dicynodonts were continuing to disperse and diversify, even in areas like western North America that were otherwise uninhabited by coeval therapsids (i.e., cynodonts).

The African cynodont Aleodon (Cynodontia, Probainognathia) in the Triassic of southern Brazil and its biostratigraphic significance

In this contribution we report the first occurrence of the enigmatic African probainognathian genus Aleodon in the Middle-early Late Triassic of several localities from the state of Rio Grande do Sul in southern Brazil. Aleodon is unusual among early probainognathians in having transversely-expanded postcanine teeth, similar to those of gomphodont cynognathians. This genus was previously known from the Manda Beds of Tanzania and the upper Omingonde Formation of Namibia. The Brazilian record of this genus is based upon multiple specimens representing different ontogenetic stages, including three that were previously referred to the sectorial-toothed probainognathian Chiniquodon theotonicus. We propose a new species of Aleodon (A. cromptoni sp. nov.) based on the specimens from Brazil. Additionally, we tentatively refer one specimen from the upper Omingonde Formation of Namibia to this new taxon, strengthening biostratigraphic correlations between these strata. Inclusion of A. cromptoni in a phylogenetic analysis of eucynodonts recovers it as the sister-taxon of A. brachyrhamphus within the family Chiniquodontidae. The discovery of numerous specimens of Aleodon among the supposedly monospecific Chiniquodon samples of Brazil raises concerns about chiniquodontid alpha taxonomy, particularly given the extremely broad geographic distribution of Chiniquodon. The discovery of Brazilian Aleodon and new records of the traversodontid Luangwa supports the hypothesis that at least two subzones can be recognized in the Dinodontosaurus Assemblage Zone.

A New Dicynodont (Therapsida: Anomodontia) from the Permian of Southern Brazil and Its Implications for Bidentalian Origins

Dicynodonts were a highly successful group of herbivorous therapsids that inhabited terrestrial ecosystems from the Middle Permian through the end of the Triassic periods. Permian dicynodonts are extremely abundant in African deposits, but are comparatively poorly known from the other regions of Gondwana. Here we describe a new South American dicynodont, Rastodon procurvidens gen. et sp. nov., from the Boqueirão farm site of the Rio do Rasto Formation, Paraná Basin, Guadalupian/Lopingian of Brazil. Diagnostic features of R. procurvidens include uniquely anteriorly-curved maxillary tusks, well-developed ridges extending from the crista oesophagea anteriorly along the pterygoid rami, strong posterior angulation of the posterior pterygoid rami, and a bulbous, well-developed retroarticular process of the articular. Phylogenetic analysis indicates that R. procurvidens is the earliest and most basal member of Bidentalia, a cosmopolitan clade that includes Permian and Triassic dicynodonts whose dentition is usually reduced to a pair of maxillary tusks.

Redescription of the geikiid Pelanomodon (Therapsida, Dicynodontia), with a reconsideration of ‘Propelanomodon’

ABSTRACT Cryptodont dicynodonts are some of the most abundant therapsid taxa in the upper Permian fossil record. Despite extensive taxonomic study of these families, the species diversity of geikiid cryptodonts remains problematic, particularly for the set of edentulous taxa in the genera Pelanomodon, Propelanomodon, and Geikia. Here, all known specimens of tuskless geikiid dicynodonts from the upper Permian Karoo Basin of South Africa are reevaluated. The genus Pelanomodon is redescribed based on a series of skulls representing a range of sizes. All tuskless geikiids from the Karoo Basin are considered to represent a single species (Pelanomodon moschops) that exhibited cranial dimorphism as adults. The nominal species Pe. moschops and Pe. rubidgei differ only in degree of facial boss development and are most parsimoniously interpreted as sexual dimorphs. Propelanomodon is considered to represent the juvenile morphotype of Pelanomodon moschops, as indicated by the geographic and stratigraphic overlap of the two taxa, the lack of any small skulls of the Pelanomodon morphotype in well-sampled localities, and the presence of specimens of intermediate size and morphology. ‘Propelanomodon’ specimens are distinctive among juvenile dicynodonts in having a significantly narrower intertemporal region than adults. Analysis of intertemporal width relative to total skull size in Pelanomodon and the abundant cryptodont taxa Aulacephalodon, Oudenodon, and Tropidostoma suggests that Pelanomodon had a unique growth trajectory. Based on available stratigraphic data, Pelanomodon was restricted to the uppermost Permian Daptocephalus Assemblage Zone of the South African Beaufort Group and was a victim of the end-Permian mass extinction. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Novel Hind Limb Morphology in a Kannemeyeriiform Dicynodont from the Manda Beds (Songea Group, Ruhuhu Basin) of Tanzania

ABSTRACT A partial hind limb (femur and tibia) from the Triassic Lifua Member of the Manda Beds (Tanzania) is recognized as a new morphotype of kannemeyeriiform dicynodont. The femur of this specimen is the largest known dicynodont postcranial element from the Manda Beds and indicates an animal nearing the size of the Late Triassic Stahleckeria potens from Brazil and Namibia. This specimen also resembles both Stahleckeria and the related Argentine stahleckeriid Ischigualastia in having an unusually elongate, straight femoral shaft and a massive and bulbous femoral head, but it differs in its compact distal end of the femur and the relative gracility of both its femur and tibia. The recognition of this specimen as a novel form of Manda dicynodont indicates that as many as six distinct kannemeyeriiforms are present in the mid-to-upper Lifua Member fauna, equaling or exceeding the species richness of previously known Laurasian faunas and substantially exceeding the richness of coeval Gondwanan faunas.