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1. 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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2. 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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3. New unworn dentitions illuminate the dental morphology and diversity of puzzling early eutherian mammals, Taeniodonta

   Kynigopoulou, Z.; Shelley, S. L.; Williamson, T. E.; Bertrand, O.; Butler, I. B.; Brusatte, S. L. (November 2021, Palaeontological Association Annual Meeting)

   Taeniodonta is a group of North America Palaeogene mammals that lived after the end-Cretaceous mass extinction. Taeniodonts show an extreme degree of dental wear, indicative of an abrasive diet, leading to hypsodonty in the most derived species. The rarity of fossils and their highly worn teeth makes their dental morphology difficult to study. We examined five new partial mandibles from the San Juan Basin, New Mexico, USA, most of which preserve unworn molars. One of the specimens preserves a deciduous ultimate premolar and using 3D micro-CT we were able to segment and study the unworn permanent tooth embedded in the jaw. We then conducted multivariate analyses on dental measurements to compare the new specimens to known teeth of early taeniodonts. We assigned the new specimens to at least three genera of Conoryctidae, a taeniodont subclade. Our results suggest that there is a broader dental diversity of the studied genera than previously thought. Morphological observations also suggest that progressive loss of cingulids and the addition of cuspids started early in the evolution of taeniodonts. These distinctive dental specializations strengthen the hypothesis that early Palaeocene mammals were able to rapidly adapt to fill the vacant ecological niches after the end-Cretaceous extinction 

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4. 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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5. Shifts in trophic architecture and ecospace stability determining survivorship and extinction at the end-Cretaceous

   Chiarenza, Alfio Alessandro; García-Girón, Jorge; Alahuhta, Janne; DeMar, D. G.; Jani, Heino; Mannion, P. D.; Williamson, T. E.; Wilson Mantilla, G. P.; Brusatte, S. L. (November 2022, Journal of Vertebrate Paleontology)

   An explanation for why some species, such as non-avian dinosaurs, became extinct, whereas others, including mammals, survived the Cretaceous/Paleogene (K/Pg) mass extinction, 66 million years ago (Ma) is still debated. What were the mechanisms behind community restructuring and the emergence of new ecological opportunities after the K/Pg event, selectively driving extinction and survivorship patterns? Using Markov networks, ecological niche partitioning and Earth System models, we reconstructed disruptions in continental food web dynamics, simulating long-term trajectories in ecospace occupancy through the latest Cretaceous (83.6–66.0 Ma) and early Paleogene (66.0–61.6 Ma). This method uses partial correlation networks to represent how different trophic groups interact in a food web and builds on empirical spatial co-variations to explore dependencies between trophic groups. Our analyses are based on a spatiotemporally and taxonomically standardized dataset, comprising more than 1,600 fossil occurrences representing more than 470 genera of fish, salamanders, frogs, albanerpetontids, lizards, snakes, champsosaurs, turtles, crocodylians, dinosaurs (including birds), and mammals across the best sampled region for this interval, the Western Interior of North America. We explicitly tested whether: 1) shifts in food web architecture underwent major restructuring before and after the K/Pg transition, including whether some trophic guilds were more prone to these shifts than others; and 2) any of these changes were associated with fluctuations in the realized niche space, helping to explain survivorship and extinction patterns at the boundary. We find a shift in latest Cretaceous dinosaur faunas, as medium-sized species counterbalanced a loss of large herbivores, but that dinosaur niches were otherwise resilient and static until the K/Pg boundary. Smaller terrestrial vertebrates, including mammals, followed a consistent trajectory of increasing trophic impact and relaxation of ecological niche limits that began in the Cretaceous and continued after the extinction. Patterns of mammalian ecological radiation and niche restructuring indicate that these taxa did not simply proliferate after the extinction; rather, their earlier ecological diversification might have helped them survive the K/Pg event, whereas the static niche of dinosaurs might have contributed to their demise. 

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