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1. THE PHYLOGENY OF THE EARLY PALAEOCENE ORDER TAENIODONTA USING NEW SPECIMENS FROM NEW MEXICO, USA

   Kynigopoulou, Z.; Shelley, S.L.; Williamson, T. E.; Wible, J. R.; Brusatte, S. L. (June 2022, European Association of Vertebrate Paleontology Abstract Volume)

   After the Cretaceous-Palaeogene (K-Pg) mass extinction mammals thrived in the Cenozoic. However, the phylogenetic affinities of early Palaeogene ‘archaic’ mammals that lived immediately after the extinction remain unresolved. Taeniodonta is a group of puzzling ‘archaic’ mammals that appeared in the early Palaeocene of North America. They are arranged into two subgroups; the Conoryctidae and Stylinodontidae and are characterised by their extreme degree of dental wear, indicating an abrasive diet, which led to hypsodonty in the most derived species. Due, in part, to their worn teeth and their rarity in the fossil record, the position of taeniondonts in the mammalian phylogenetic tree remains unresolved. New fossils from San Juan basin, New Mexico, USA, including unworn teeth of four genera and postcranial elements of an early taeniodont, Conoryctes, shed light on their dental and postcranial anatomy. Both in the forelimb and hind limp of Conoryctes, there are anatomical adaptations towards fossoriality. Using these specimens, we scored taeniodonts and other Palaeogene mammals into a phylogenetic data matrix (620 characters, 135 taxa). We then conducted a phylogenetic analysis using parsimony. Our results show that Taeniodonta is a monophyletic group within Eutheria. We also found that Onychodectes is basal to the two subgroups previously proposed. Based on the new postcranial fossils and revised phylogeny, we concluded that digging behaviours were likely ancestral for taeniodonts. Therefore, a more fossorial mode of life may have been beneficial for their surviving and thriving in the wake of the K-Pg extinction. 

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2. Postcranial morphology of taeniodonts (Mammalia: Taeniodonta) indicates fossorial adaptations in the Palaeogene

   Kynigopoulou, Z.; Shelley, S. L.; Williamson, T. E.; Brusatte, S. L. (October 2019, Journal of Vertebrate Paleontology, Abstracts and Programs)

   After the Cretaceous-Paleogene (K-Pg) mass extinction mammals, which originated during the Mesozoic, managed to survive and thrive. However, the tempo and mode of evolution for eutherians (placentals and close relatives) after the extinction are still unclear. An ideal group to investigate the post KPg evolution of mammals is the taeniodonts, as they are among the few taxa to purportedly cross the boundary. They then underwent a radiation in the early Paleogene and are defined primarily by their unusual dentition which is suited to chew an abrasive and tough diet. Ten genera of taeniodonts are currently recognized and are commonly arranged into two families. The Conoryctidae is usually considered to have a more generalized body plan while Stylinodontidae possess relatively extreme digging adaptations and more highly derived dentitions with enlarged canines. We conducted a phylogenetic analysis by applying parsimony and Bayesian techniques to a dataset of characters gathered from extensive observation of new specimens. We found limited support for the conoryctid-stylinodontid division and the genera Conoryctes and Onychodectes are placed as key basal taxa outside the clade of the more robust derived taxa (Wortmania, Ectoganus, Psittacotherium, Stylinodon). We then assessed postcranial bones to determine functional modes for taeniodonts and to test changes across phylogeny. Qualitatively, most taeniodonts, including Onychodectes, possess indicators of digging, i.e., a well-developed deltopectoral crest and broad distal end of the humerus for increasing flexion, pronation and supination, a long olecranon process of the ulna and enlarged manual unguals. Then we conducted quantitative multivariate analyses (linear discriminant analysis), using 9 forelimb linear measurements and 29 tarsal ones, comparing taeniodonts to a suite of extant mammals with known locomotor mode and other Paleogene taxa. Our results suggest Onychodectes to be terrestrial/semifossorial and comparable with the numbat (Myrmecobius fasciatus). Ectoganus and Stylinodon are semi-fossorial and fall out near the gopher, Pappogeomys merriami and the aardvark (Orcyteropus afer). Therefore, our study indicates that digging behaviors are ancestral for taeniodonts, and suggest that burrowing may have been integral to their survival across the KPg boundary and their subsequent radiation. Grant Information: European Research Council Starting Grant (ERC StG 2017, 756226, PalM), National Science Foundation (EAR- 1325544, 1654952, DEB-1654952, 1654949) 

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3. Postcranial morphology of taeniodonts (Mammalia: Taeniodonta) indicates fossorial adaptations in the Palaeogene

   Kynigopoulou, Z.; Shelley, S. L.; Williamson, T. E.; Brusatte, S. L. (July 2019, Evolution and Paleoenvironment of Early Modern Vertebrates during the Paleogene International Symposium)

   Mammals originated during the Mesozoic and survived the Cretaceous-Palaeogene (K-Pg) mass extinction. Their evolution from small, opportunistic animals to more specialised animals with diverse locomotor behaviours following the extinction is still unclear. An ideal group to address this question is the Taeniodonta which are among the few eutherians that purportedly crossed the K-Pg boundary and diversified in the early Palaeogene. They are known thus far from the Palaeogene of North America and are characterised by their unique dentition adapted for an abrasive diet and their robust skeleton. There are 10 genera of taeniodonts classified into two families. The Conoryctidae are smaller with more generalised body plan, whereas the Stylinodontidae reached large body size (up to 100kg) and evolved crown hypsodonty. We focused our study on the postcranial functional morphology of the two taeniodont subgroups. We conducted linear discriminant analysis using 9 linear measurements of the humerus, comparing Onychodectes (a conoryctid) and Stylinodon, Ectoganus, Psittacotherium (stylinodonts) with extant mammals of known locomotion. We also used 29 linear tarsal measurements to evaluate the locomotor behaviour of Onychodectes and Conoryctes (conoryctids) and Ectoganus alongside a sample of extant mammals and other Palaeogene taxa. Our results show that Onychodectes, which is one of the most basal taeniodonts, might have been terrestrial/semi-fossorial, similar to the numbat. Postcranial features of Onychodectes show it possessed digging adaptations i.e. a long olecranon process of the ulna, enlarged manual unguals and a well-developed deltopectoral crest and broad distal end of the humerus. We find stylinodontid taeniodonts to be distinctly more fossorial, comparable to the striped skunk, gopher and the aardvark. Our study suggests that digging is an ancestral behaviour for taeniodonts implying the importance of burrowing for surviving the K-Pg extinction. 

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4. Size variation in populatinos of Tetraclaenodon (Mammalia, 'Condylarthra'), from the Torrejonian NALMA of the San Juan Basin, New Mexico, reveals new insights into their evolution and paleoenvironment

   Holpin, S.; Williamson, T. E.; Shelley, S. L.; Brusatte, S. L. (October 2019, Society of Vertebrate Paleontology, Abstracts and Programs)

   After the end-Cretaceous mass extinction, approximately 75% of life on land and in the sea disappeared. The mammals of the early Cenozoic rapidly diversified and dispersed, rising to numerical and ecological dominance beyond their Mesozoic norms. Among those initial groups that ushered in the Age of Mammals, Paleocene and Eocene ‘condylarths’ are thought to include the ancestors of modern odd-toed ungulates (horses, tapirs, rhinos). Tetraclaenodon is the oldest genus of the ‘condylarth’ group Phenacodontidae and one of the most abundant fossils from the San Juan Basin (SJB) of New Mexico. Tetraclaenodon was a medium sized (mean body mass ca. 10kg), terrestrial mammal which was lightly built and had an omnivorous to herbivorous bunodont dentition. Here we use multivariate and statistical analyses to investigate body mass and dental variation in 110 teeth of Tetraclaenodon spanning the Torrejonian (Paleocene) interval of the SJB. The specimens were grouped into six time bins by their biostratigraphical reference, from Tj1 (~63.8 Ma) through Tj6 (~62.7 Ma). Measurements of the length, mesial and distal width of the lower first molars (m1) were subject to principal component analysis (PCA), and m1 area was used to predict body mass using a regression equation. The PCA morphospace ordinates specimens along a PC1 axis that accounts for 90.05% of total variance and is significantly correlated with body size. A PERMANOVA test finds a significant difference in morphospace occupation (non-overlap) between clusters of specimens from Tj1-3 and Tj4-6, but there are no significant differences between the individual time bins within each cluster. This trend is also seen in the body size estimates: Mann-Whitney tests recover significant differences between the two clusters. These results suggest that Torrejonian populations of Tetraclaenodon were relatively constant in size throughout Tj1-3, but between Tj3 and Tj4 underwent an increase in body mass and subsequently stabilized (at this resolution) for the remainder of the Torrejonian. A similar trend is seen in contemporary populations of the periptychid ‘condylarth’ Periptychus, suggesting that there were selective environmental pressures acting on these herbivorous species. These body size differences may reflect the emergence of a new, larger Tetraclaenodon species in Tj4, or may be associated to an environmental change, perhaps relating to climate or vegetation. In either case, this illustrates dynamic evolution of mammals during the few million years after the extinction. Grant Information: European Research Council Starting Grant (ERC StG 2017, 756226, PalM), National Science Foundation (EAR- 1654952) 

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5. Testing the phylogeny of Periptychidae and "archaic" Paleocene mammals uner different optimality criteria

   Puschel, H. P.; Shelley, S. L.; Williamson, T. E.; Wible, J. R.; Brusatte, S. L. (October 2019, Journal of Vertebrate Paleontology, Abstracts and Program)

   It is now well established that the end-Cretaceous mass extinction had enormous repercussions for mammalian evolution. Following the extinction, during the Paleocene, mammals started to radiate, occupying new and diverse ecological niches. However, the phylogenetic relationships between the socalled “archaic” mammals of this time, and their position within Placentalia, remain contentious. The Periptychidae are a clade of distinctive “archaic” ungulates, composed of ~17 genera of small to large bodied, highly bunodont, terrestrial herbivores that were among the first placental mammals to appear after the end-Cretaceous mass extinction. Although the Periptychidae has been historically considered a distinctive “condylarth” subgroup, their higherlevel relationships have been rarely tested. Here, we present an inclusive cladistic analysis to determine and test the phylogenetic affinities of Periptychidae and other key Paleocene groups within Placentalia under different cladistic optimality criteria. We scored 140 taxa for 503 dental, cranial and postcranial characters, incorporating new morphological and taxonomic data. The data were then subject to parsimony and Bayesian tree of morphological evolution, running 5000000 generations with samples every 200 generations and discarding 25% of the samples as burn-in. Stationarity was achieved and a 50 percent majority rule consensus tree from the sampled trees was obtained. The parsimony analysis recovered 48 most parsimonious trees. The two consensus trees derived from the different analyses are largely congruent and recover a monophyletic Periptychidae, although the parsimony consensus tree is better resolved. These results are consistent with simulation studies showing that parsimony tends to be more precise (more nodes reconstructed) than Bayesian analyses, although less accurate. The main topological differences between the results relate to the position of poorly known Puercan (earliest Paleocene) species. Our results affirm the monophyly of Periptychidae and its nesting within a group of “condylarths” positioned at the base of Laurasiatheria and closely related to Artiodactyla. Within Periptychidae we found support for the three major subfamilial divisions in both analyses. These results highlight the importance of using different optimality criteria when resolving a phylogeny and provide a new insight into how placental mammals were evolving after the end-Cretaceous extinction. Grant Information: CONICYT PFCHA/DOCTORADO BECAS CHILE/2018, European Research Council Starting Grant (ERC StG 2017, 756226, PalM), National Science Foundation (NSF EAR 1654952, DEB 1654949) 

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