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1. Vestibular sensitivity and locomotor behavior in early Paleocene mammals

   Bertrand, O. C. ; Shelley, S. L. ; Williamson, T. E. ; Wible, J. R. ; Chester, S. G. ; Holbrook, Luke T. ; Lyson, T. R. ; Meng, Jin ; Smith, Jin ; Smith, T. ; et al ( June 2023 , European Association of Vertebrate Palaeontologists)

   The end-Cretaceous extinction triggered the collapse of ecosystems and a drastic turnover of mammalian communities. During the Mesozoic, mammals were ecologically diverse, but less so than extant species. Modern ecological richness was established by the Eocene, but questions remain about the ecology of the first wave of mammals radiating after the extinction.Postcranial fossils are often used to determine locomotor behavior; however, the semicircular canals of theinner ear also represent a reliable proxy. These canals detect the angular acceleration of the head duringl ocomotion and transmit neuronal signals to the brain to allow stabilization of the eyes and head. Accordingly, vestibular sensitivity to rapid rotational head movements is higher in species with a larger canal radius of curvature and more orthogonal canals. We used high-resolution computed tomography scanning to obtain inner ear virtual endocasts for 30 specimens. We supplemented these with data from the literature to constructa database of 79 fossil from the Jurassic to the Eocene and 262 extant mammals. We compared data on canal morphology and another lifestyle proxy, the size of the petrosal lobules, which have a role in maintaining eyes’ movements and position. We find that Paleocene mammals exhibited a lower average and more constricted range of Agility Indices (AI), a new measure of canal radius size relative to body size, compared to Mesozoic, Eocene and extant taxa. Inthe early Paleocene, body mass and canal radius increased, but the former outpaced the latter leading to an AIdecline. Similarly, their petrosal lobules were relatively smaller on average compared to other temporal groups, which suggests less ability for fast movements. Additionally, Paleocene mammals had similar AIs to extant scansorial and terrestrial quadrupeds. In contrast, the lack of canal orthogonality change from the Mesozoic to the Paleocene indicates no trend toward lower vestibular sensitivity regardless of changes in body size. This result may reflect functional differences between canal orthogonality and radius size. Our results support previous work on tarsal morphology and locomotor behavior ancestral state reconstruction suggesting that ground dwelling mammals were more common than arboreal taxa during the Paleocene. Ultimately, this pattern may indicate that the collapse of forested environments immediately after extinction led to the preferential survivorship of more terrestrially adapted mammals. 

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2. LOCOMOTOR BEHAVIOR OF PALEOCENE MAMMALS: INSIGHTS FROM THE SEMICIRCULAR CANALS OF THE INNER EAR

   Bertrand, O.C. ; Shelley, S.L. ; Williamson, T.E. ; Wible, J.R. ; Chester, S. G. ; Holbrook, L. T. ; Lyson, T. R. ; Meng, J. ; Miller, ?I. M. ; Smith, T. ; et al ( June 2023 , European Association of Vertebrate Palaeontologists)

   Alba, D.M. ; Marigó, J. ; Nacarino-Meneses, C. ; Villa, A. (Ed.)

   The end-Cretaceous mass extinction triggered the collapse of ecosystems and a drastic turnover in mammalian communities leading to the demise of many ecologically specialized species. While Mesozoic mammals were ecomorphologically diverse, recognizable ecological richness was only truly established in the Eocene. Questions remain about the ecology of the first wave of mammals radiating after the extinction. Here, we use the semicircular canals of the inner ear as a proxy for locomotor behavior. Thirty new inner ear virtual endocasts were generated using high-resolution computed tomography scanning. This sample was supplemented by data from the literature to construct a dataset of 79 fossils spanning the Jurassic to the Eocene alongside 262 extant mammals. Vestibular sensitivity was measured using the radius of curvature against body mass and the residuals of this relationship were analyzed. The petrosal lobule size relative to body mass were compared with the inner ear data as they have a role in maintaining gaze stabilization during motion. Paleocene mammals exhibited smaller canal radius of curvature, compared to Mesozoic, Eocene, and extant taxa. In the early Paleocene, canal radius and associated petrosal lobules were relatively smaller on average compared to other temporal groups, suggesting less ability for fast movements. 

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3. Journal of Vertebrate Paleontology, Program and Abstracts

   Bertrand, O.C. ; Brusatte, S. L. ; Shelley, S. L. ; Wible, J. R. ; Williamson, T. E. ; Chester, S. G. ; Holbrook, L. T. ; Lyson, T. R. ; Smith, T. ; Meng, J. ( October 2020 , Society of Vertebrate paleontology)

   The end-Cretaceous mass extinction, 66 million years ago, profoundly reshaped the biodiversity of our planet. After likely originating in the Cretaceous, placental mammals (species giving live birth to well-developed young) survived the extinction and quickly diversified in the ensuing Paleocene. Compared to Mesozoic species, extant placentals have advanced neurosensory abilities, enabled by a proportionally large brain with an expanded neocortex. This brain construction was acquired by the Eocene, but its origins, and how its evolution relates to extinction survivorship and recovery, are unclear, because little is known about the neurosensory systems of Paleocene species. We used high-resolution computed tomography (CT) scanning to build digital brain models in 29 extinct placentals (including 23 from the Paleocene). We added these to data from the literature to construct a database of 98 taxa, from the Jurassic to the Eocene, which we assessed in a phylogenetic context. We find that the Phylogenetic Encephalization Quotient (PEQ), a measure of relative brain size, increased in the Cretaceous along branches leading to Placentalia, but then decreased in Paleocene clades (taeniodonts,phenacodontids, pantodonts, periptychids, and arctocyonids). Later, during the Eocene, the PEQ increased independently in all crown groups (e.g., euarchontoglirans and laurasiatherians). The Paleocene decline in PEQ was driven by body mass increasing much more rapidly after the extinction than brain volume. The neocortex remained small, relative to the rest of the brain, in Paleocene taxa and expanded independently in Eocene crown groups. The relative size of the olfactory bulbs, however, remained relatively stable over time, except for a major decrease in Euarchontoglires and some Eocene artiodactyls, while the petrosal lobules (associated with eye movement coordination) decreased in size in Laurasiatheria but increased in Euarchontoglires. Our results indicate that an enlarged, modern-style brain was not instrumental to the survival of placental mammal ancestors at the end-Cretaceous, nor to their radiation in the Paleocene. Instead, opening of new ecological niches post-extinction promoted the diversification of larger body sizes, while brain and neocortex sizes lagged behind. The independent increase in PEQ in Eocene crown groups is related to the expansion of the neocortex, possibly a response to ecological specialization as environments changed, long after the extinction. 

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4. Virtual endocranial and inner ear endocasts of the Paleocene ‘condylarth’ Chriacus : new insight into the neurosensory system and evolution of early placental mammals

   https://doi.org/10.1111/joa.13084  

   Bertrand, Ornella C. ; Shelley, Sarah L. ; Wible, John R. ; Williamson, Thomas E. ; Holbrook, Luke T. ; Chester, Stephen G. B. ; Butler, Ian B. ; Brusatte, Stephen L. ( October 2019 , Journal of Anatomy)

   The end‐Cretaceous mass extinction allowed placental mammals to diversify ecologically and taxonomically as they filled ecological niches once occupied by non‐avian dinosaurs and more basal mammals. Little is known, however, about how the neurosensory systems of mammals changed after the extinction, and what role these systems played in mammalian diversification. We here use high‐resolution computed tomography (CT) scanning to describe the endocranial and inner ear endocasts of two species,Chriacus pelvidensandChriacus baldwini, which belong to a cluster of ‘archaic’ placental mammals called ‘arctocyonid condylarths’ that thrived during theca. 10 million years after the extinction (the Paleocene Epoch), but whose relationships to extant placentals are poorly understood. The endocasts provide new insight into the paleobiology of the long‐mysterious ‘arctocyonids’, and suggest thatChriacuswas an animal with anencephalization quotient (EQ)range of 0.12–0.41, which probably relied more on its sense of smell than vision, because the olfactory bulbs are proportionally large but the neocortex and petrosal lobules are less developed. Agility scores, estimated from the dimensions of the semicircular canals of the inner ear, indicate thatChriacuswas slow to moderately agile, and its hearing capabilities, estimated from cochlear dimensions, suggest similarities with the extant aardvark.Chriacusshares many brain features with other Paleocene mammals, such as a small lissencephalic brain, large olfactory bulbs and small petrosal lobules, which are likely plesiomorphic for Placentalia. The inner ear ofChriacusalso shares derived characteristics of the elliptical and spherical recesses with extinct species that belong to Euungulata, the extant placental group that includes artiodactyls and perissodactyls. This lends key evidence to the hypothesized close relationship betweenChriacusand the extant ungulate groups, and demonstrates that neurosensory features can provide important insight into both the paleobiology and relationships of early placental mammals.

    

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5. Journal of Vertebrate Paleontology, Program and Abstracts

   Holpin, Sofia ; Shelley, Sarah L.' ; Williamson, Thomas E. ; Brusatte, Stephen ( October 2020 , Society of Vertebrate Paleontology)

   With the dawn of the Paleogene, the mammalian survivors of the Cretaceous–Paleogene mass extinction, 66 million years ago, found themselves in an emptied landscape. Within a million years of the bolide impact, placental mammals reached a diversity and abundance never seen during the Age of Dinosaurs. The North American ‘condylarths’ were amongst the first mammals to diversify during the early Paleogene and are often considered the ancestral ‘stock’ from which other euungulate groups evolved. Amongst these, Phenacodontidae are often regarded to lie at the base of the perissodactyl family tree, but their phylogenetic position, and that of other ‘condylarths’, remain contentious. Tetraclaenodon, a medium-sized herbivorous phenacodontid from the Torrejonian (~64 to ~62 Ma) of North America is generally recognized as the oldest member of Phenacodontidae, and thus is instrumental for untangling the evolutionary relationships of ‘condylarths’ and perissodactyls. Here we present new information on Tetraclaenodon based on a description of new and previously known fossil material from the San Juan Basin of New Mexico, U.S.A., which we studied using high-resolution computed tomography (CT) scanning. From CT scans of the cranium, we segmented the brain endocast, which is relatively small and smooth (lissencephalic), similar to that of other Paleocene mammals. The petrosal lobules, which are involved in eye movement coordination, are small. The semi-circular canals associated with balance, provide an agility score of 3 indicating that Tetraclaenodon was probably moderately agile, similar to the extant raccoon dog or the aardwolf. A multivariate analysis of tarsal measurements for a sample of Paleocene and extant mammals, which informs locomotor style, indicates that Tetraclaenodon was most suited to terrestrial locomotion. This is in line with anatomical and myological features of the limbs of Tetraclaenodon and other phenacodontids, early perissodactyls and extant mammals. These findings contradict previous studies that designated Tetraclaenodon as a scansorial mammal, capable of habitually climbing trees. Our results shed light on the locomotory adaptations of Tetraclaenodon in comparison to more cursorial phenacodontids and perissodactyls, such as Phenacodus and Hyrachyus. The earliest member of the perissodactyl stem lineage apparently lacked the more cursorial adaptations of their relatives in the late Paleocene and onwards. 

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