Selina Cole

Relatively few Hirnantian (Late Ordovician) crinoids are known, and none has been previously described from the palaeocontinent of Baltica. This has impaired our ability to understand the patterns of extinction and biogeographic dispersal surrounding the Late Ordovician mass extinction, which triggered a major turnover in crinoid faunas. Here, we describe Tallinnicrinus toomae gen. et sp. nov., an anthracocrinid diplobathrid from the Hirnantian of northern Estonia. Tallinnicrinus is the youngest member of the Anthracocrinidae and the first representative of the family to occur in Baltica. Morphologically , Tallinnicrinus is unusual in that the radial and basal plates are in a single circlet of 10 plates, similar to the anthracocrinid Rheocrinus Haugh, 1979 from the Katian of Laurentia. Phylogenetic analysis further confirms a close relationship between Tallinnicrinus and Laurentian anthracocrinids, suggesting biogeographic dispersal of the lineage from Laurentia to Baltica during the late Katian or early Hirnantian. The occurrence of this new taxon establishes that the family Anthracocrinidae survived the first pulse of the Late Ordovician mass extinction. However, the lineage remained a 'dead clade walking' because it failed to diversify in the wake of the end-Katian extinction and ultimately went extinct itself by the end of the Ordovician.

Identifying correlates of extinction risk is important for understanding the underlying mechanisms driving differential rates of extinction and variability in the temporal durations of taxa. Increasingly, it is recognized that the effects of multiple, potentially interacting variables and phylogenetic relationships should be incorporated when studying extinction selectivity to account for covariation of traits and shared evolutionary history. Here, I explore a variety of biological and ecological controls on genus longevity in the global fossil record of diplobathrid crinoids by analyzing the combined effects of species richness, habitat preference, body size, filtration fan density, and food size selectivity. I employ a suite of taxic and phylogenetic approaches to (1) quantitatively compare and rank the relative effects of multiple factors on taxonomic longevity and (2) determine how phylogenetic comparative approaches alter interpretations of extinction selectivity.

I find controls on diplobathrid genus duration are hierarchically structured, where species richness is the primary predictor of duration, habitat is the secondary predictor, and combinations of ecological and biological traits are tertiary controls. Ecology plays an important but complex role in the generation of crin-oid macroevolutionary patterns. Notably, tolerance of environmental heterogeneity promotes increased genus duration across diplobathrid crinoids, and the effects of traits related to feeding ecology vary depending on habitat lithology. Finally, I find accounting for phylogeny does not consistently decrease the significance of correlations between traits and genus duration, as is commonly expected. Instead, the strength of relationships between traits and duration may increase, decrease, or remain statistically similar, and both the magnitude and direction of these shifts are generally unpredictable. However, traits with strong correlations and/or moderately large effect sizes (Cohen's f 2 > 0.15) under taxic approaches tend to remain qualitatively unchanged under phylogenetic approaches.

Order Diplobathrida is a major clade of camerate crinoids spanning the Ordovician–Mississippian, yet phylogenetic relationships have only been inferred for Ordovician taxa. This has hampered efforts to construct a comprehensive tree of life for crinoids and develop a classification scheme that adequately reflects diplobathrid evolutionary history. Here, I apply maximum parsimony and Bayesian phylogenetic approaches to the fossil record of diplobathrids to infer the largest tree of fossil crinoids to date, with over 100 genera included. Recovered trees provide a framework for evaluating the current classification of diplobathrids. Notably, previous suborder divisions are not supported, and superfamily divisions will require significant modification. Although numerous revisions are required for families, most can be retained through reassignment of genera. In addition, recovered trees were used to produce phylogeny-based estimates of diplobathrid lineage diversity. By accounting for ghost lineages, phylogeny-based richness estimates offer greater insight into diversification and extinction dynamics than traditional taxonomy-based approaches alone and provide a detailed summary of the ~150 million-year evolutionary history of Diplobathrida. This study constitutes a major step toward producing a phylogeny of the Crinoidea and documenting crinoid diversity dynamics. In addition, it will serve as a framework for subsequent phylogeny-based investigations of macroevolutionary questions.

Most major crinoid clades first appear in the fossil record during the Ordovician, but the evolutionary origins of many higher taxa remain unresolved. Here, the first camerate crinoids from the Katian (Upper Ordovician) of Estonia with complete calyces are described. Phylogenetic analyses are conducted to determine the affinities of two new genera and to assess the evolutionary origins of the families Opsiocrinidae (Diplobathrida) and Glyptocrinidae (Monobathrida). Sakucrinus krossi gen. et sp. nov. is identified as the earliest known member of Opsiocrinidae and demonstrates that: (1) the family originated much earlier than was previously assumed; (2) features previously considered plesiomorphic for the family are instead derived; and (3) the family is likely closely related to the Dimerocrinitidae. Family Pachycephalocrinidae fam. nov. is established to accept the monocyclic glyptocrinid Pachycephalocrinus jaanussoni gen. et sp. nov. Pachycephalocrinus jaanussoni displays morphological features that are unique among monobathrid crinoids, including a large periproct in the calyx side. Inclusion of Pachycephalocrinus in the first phylogenetic analysis focusing on Ordovician monobathrid crinoids elucidates evolutionary relationships among Ordovician monobathrids and calls into question the validity of superorder Compsocrinina. The new genera described here provide important insight into the timing of origination and morphological diversity of higher taxa during the early evolutionary history of camerate crinoids.

The subclass Camerata (Crinoidea, Echinodermata) is a major group of Paleozoic crinoids that represents an early divergence in the evolutionary history and morphologic diversification of class Crinoidea, yet phylogenetic relationships among early camerates remain unresolved. This study conducted a series of quantitative phylogenetic analyses using parsimony methods to infer relationships of all well-preserved Ordovician camerate genera (52 taxa), establish the branching sequence of early camerates, and test the monophyly of traditionally recognized higher taxa, including orders Monobathrida and Diplobathrida. The first phylogenetic analysis identified a suitable outroup for rooting the Ordovician camerate tree and assessed affinities of the atypical dicyclic family Reteocrinidae. The second analysis inferred the phylogeny of all well-preserved Ordovician camerate genera. Inferred phylogenies confirm: (1) the Tremadocian genera Cnemecrinus and Eknomocrinus are sister to the Camerata; (2) as historically defined, orders Monobathrida and Diplobathrida do not represent monophyletic groups; (3) with minimal revision, Monobathrida and Diplobathrida can be re-diagnosed to represent monophyletic clades; (4) family Reteocrinidae is more closely related to camerates than to other crinoid groups currently recognized at the subclass level; and (5) several genera in subclass Camerata represent stem taxa that cannot be classified as either true monobathrids or true diplobathrids. The clade containing Monobathrida and Diplobathrida, as recognized herein, is termed Eucamerata to distinguish its constituent taxa from more basally positioned taxa, termed stem eucamerates. The results of this study provide a phylogenetic framework for revising camerate classification, elucidating patterns of morphologic evolution, and informing outgroup selection for future phylogenetic analyses of post-Ordovician camerates.

A period of reduced reefal abundance and size occurred around the time of the Silurian–Devonian boundary, between the middle Silurian and Middle Devonian
reef building peaks. In southeastern Laurentia, stromatoporoid and coral reefs became especially rare during this time, and very few buildups have been reported from the
region. This paper reports on the sedimentological, faunal, and paleoenvironmental characteristics of a newly discovered latest Silurian stromatoporoid-dominated buildup in the Keyser Limestone of the Virginia Appalachians. The buildup is classified as a parabiostrome, because over 95 % of the stromatoporoids in the buildup have been disturbed. Based on qualitative and statistical comparisons of facies characteristics, the buildup most likely formed from a stromatoporoid-tabulate buildup that was repeatedly damaged by high-energy events. Multivariate analyses of point
count data reveal a decrease in stromatoporoids and an increase in tabulate corals and matrix moving vertically upward through the buildup, indicating changes in faunal composition as the buildup grew. The results of this study provide unique insight into reef development in one of the oldest Silurian reefal buildups from the central Appalachian Basin during a time interval when reefal buildups were poorly represented.