Explore the vast range of titles available.

Knowledge of phylogenetic relationships among species is fundamental to understanding basic patterns in evolution and underpins nearly all research programs in biology and paleontology. However, most methods of phylogenetic inference typically used by paleontologists do not accommodate the idiosyncrasies of fossil data and therefore do not take full advantage of the information provided by the fossil record. The advent of Bayesian ‘tip-dating’ approaches to phylogeny estimation is especially promising for paleosystematists because time-stamped comparative data can be combined with probabilistic models tailored to accommodate the study of fossil taxa. Under a Bayesian framework, the recently developed fossilized birth–death (FBD) process provides a more realistic tree prior model for paleontological data that accounts for macroevolutionary dynamics, preservation, and sampling when inferring phylogenetic trees containing fossils. In addition, the FBD tree prior allows for the possibility of sampling ancestral morphotaxa. Although paleontologists are increasingly embracing probabilistic phylogenetic methods, these recent developments have not previously been applied to the deep-time invertebrate fossil record. Here, I examine phylogenetic relationships among Ordovician through Devonian crinoids using a Bayesian tip-dating approach. Results support several clades recognized in previous analyses sampling only Ordovician taxa, but also reveal instances where phylogenetic affinities are more complex and extensive revisions are necessary, particularly among the Cladida. The name Porocrinoidea is proposed for a well-supported clade of Ordovician ‘cyathocrine’ cladids and hybocrinids. The Eucladida is proposed as a clade name for the sister group of the Flexibilia herein comprised of cladids variously considered ‘cyathocrines,’ ‘dendrocrines,’ and/or ‘poteriocrines’ by other authors.

A major goal of biological classification is to provide a system that conveys phylogenetic relationships while facilitating lucid communication among researchers. Phylogenetic taxonomy is a useful framework for defining clades and delineating their taxonomic content according to well-supported phylogenetic hypotheses. The Crinoidea (Echinodermata) is one of the five major clades of living echinoderms and has a rich fossil record spanning nearly a half billion years. Using principles of phylogenetic taxonomy and recent phylogenetic analyses, we provide the first phylogeny-based definition for the Clade Crinoidea and its constituent subclades. A series of stem- and node-based definitions are provided for all major taxa traditionally recognized within the Crinoidea, including the Camerata, Disparida, Hybocrinida, Cladida, Flexibilia, and Articulata. Following recommendations proposed in recent revisions, we recognize several new clades, including the Eucamerata Cole 2017, Porocrinoidea Wright 2017, and Eucladida Wright 2017. In addition, recent phylogenetic analyses support the resurrection of two names previously abandoned in the crinoid taxonomic literature: the Pentacrinoidea Jaekel, 1918 and Inadunata Wachsmuth and Springer, 1885. Last, a phylogenetic perspective is used to inform a comprehensive revision of the traditional rank-based classification. Although an attempt was made to minimize changes to the rank-based system, numerous changes were necessary in some cases to achieve monophyly. These phylogeny-based classifications provide a useful template for paleontologists, biologists, and non-experts alike to better explore evolutionary patterns and processes with fossil and living crinoids.

Geometric morphometrics facilitates the quantification and visualization of variation in shape and proportion through the comparison of homologous features. Eublastoidea, a Paleozoic echinoderm clade with a conservative body plan, is an ideal group for morphometric analysis, because their plate junctions are homologous and identifiable on all species. Eublastoids have previously been grouped taxonomically by generalized shape types (e.g., globose). These shapes are often used in taxonomic descriptions and as characters in phylogenetic analyses. The underlying homology of these broad shape types has never been explored. Herein we apply the first comprehensive use of three-dimensional geometric morphometrics (3D GMM) on fossil echinoderms to investigate taxonomic assignments, temporal distribution, and whether the varying proportions of skeletal elements that produce the gross thecal morphology are distinguishable. Taxonomic assignments specifically at the ordinal and family levels show varying amounts of overlap in morphospace, suggesting that many assignments may not be reevaluated. Our results suggest that none of the generalized shape types are distinct in morphospace and, therefore, likely do not capture the homologous changes in taxa. The plate circlet ratios showed trends specifically relating to the deltoid plate circlet, which has the most variability. We reanalyzed previous work and subsetted our data to be more comparable and found that there are key differences between methodologies and landmarks that will require future evaluation. Applying modern technological methods to previously explored questions allows for an updated understanding of this important fossil clade and provides a framework for others to assess fossil clades in a similar manner.

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.

Evidence from the earliest-known crinoids (Tremadocian, Early Ordovician), called protocrinoids, is used to hypothesize initial steps by which elements of the calyx evolved. Protocrinoid calyces are composed of extraxial primary and surrounding secondary plates (both of which have epispires along their sutures) that are unlike those of more crownward fossil and extant crinoids in which equivalent calycinal plating is strongly organized. These reductions inspired several schemes by which to name the plates in these calyces. However, the primary-secondary systems seen in protocrinoids first appeared among Cambrian stem radial echinoderms, with primaries representing centers around which secondaries were sequentially added during ontogeny. Therefore, the protocrinoid calyx represents an intermediate condition between earliest echinoderms and crownward crinoids. Position and ontogeny indicate certain primaries remained as loss of secondaries occurred, resulting in abutting of primaries into the conjoined alternating circlets characteristic of crinoids. This transformative event included suppression of secondary plating and modification or, more commonly, elimination of respiratory structures. These data indicate subradial calyx plate terminology does not correspond with most common usage, but rather, supports an alternative redefinition of these traditional expressions. Extension and adoral growth of fixed rays during calyx ontogeny preceded conjoined primaries in earliest crinoids. Restriction with modification or elimination of calyx respiratory structures also accompanied this modification. Phylogenetic analyses strongly support crinoid origination from early pentaradiate echinoderms, separate from blastozoans. Accordingly, all Tremadocian crinoids express a distinctive aggregate of plesiomorphic and apomorphic commonalities; all branch early within the crinoid clade, separate from traditional subclass-level clades. Nevertheless, each taxon within this assemblage expresses at least one diagnostic apomorphy of camerate, cladid, or disparid clades.

Biological variation fuels evolutionary change. Across longer timescales, however, polymorphisms at both the genomic and phenotypic levels often persist longer than would be expected under standard population genetic models such as positive selection or genetic drift. Explaining the maintenance of this variation within populations across long time spans via balancing selection has been a major triumph of theoretical population genetics and ecology. Although persistent polymorphisms can often be traced in fossil lineages over long periods through the rock record, paleobiology has had little to say about either the long-term maintenance of phenotypic variation or its macroevolutionary consequences. I explore the dynamics that occur when persistent polymorphisms maintained over long lineage durations are filtered into descendant lineages during periods of demographic upheaval that occur at speciation. I evaluate these patterns in two lineages: Ectocion, a genus of Eocene mammals, and botryocrinids, a Mississippian cladid crinoid family. Following origination, descendants are less variable than their ancestors. The patterns by which ancestral variation is sorted cannot be distinguished from drift. Maintained and accumulated polymorphisms in highly variable ancestral lineages such as Barycrinus rhombiferus Owen and Shumard, 1852 may fuel radiations as character states are sorted into multiple descendant lineages. Interrogating the conditions under which trans-specific polymorphism is either maintained or lost during periods of demographic and ecological upheaval can explain how population-level processes contribute to the emergent macroevolutionary dynamics that shape the history of life as preserved in the fossil record.

Among invertebrates and both in modern and ancient marine environments, certain echinoderms have been and are some of the most active and widespread bioturbators and bioeroders. Bioturbation and/or bioerosion of regular and irregular echinoids, starfish, brittle stars, sea cucumbers and crinoids are known from modern settings, and some of the resulting traces have their counterparts in the fossil record. By contrast, surficial trails or trackways produced by other modern echinoderms, e.g., sand dollars, exhibit a lower preservation rate and have not yet been identified in the fossil record. In addition, the unique features of the echinoderm skeleton (e.g., composition, rapid growth, multi-element architecture, etc.) may promote the production of related traces produced by the reutilization of echinoderm ossicles (e.g., burrow lining), predation (e.g., borings), or parasitism (e.g., swellings or cysts). Finally, the skeletal robustness of some echinoids may promote their post mortum use as benthic islands for the settlement of hard-substrate dwellers.

Monstrocrinus is a most unusual crinoid from the Devonian of Germany. It has long, simple to complex spines attached to most calyx plates. A stem was not associated with the initial specimens on which this genus was named. Thus, it was assumed that Monstrocrinus was a stemless crinoid that sat directly on the sea floor and, when alive, rolled around on its spines. For a variety of reasons, this seems unlikely, but the rolling crinoid lifestyle interpretation has persisted in the literature. More thorough cleaning of historical specimens and, more importantly, a new specimen with an attached column clearly refute the classical lifestyle interpretation and demonstrate that Monstrocrinus was a typical crinoid attached to a long column that was presumably anchored to the sea floor. In addition to revising the lifestyle of this crinoid, the varieties of spine types are redefined with names that more accurately reflect their morphology. Monstrocrinus occurs in Europe and South America, and one previously described species is placed in synonymy so that the genus is now comprised of three species. The diplobathrid camerate crinoid genus Monstrocrinus is morphologically reinterpreted on the basis of new finds from the upper Emsian (Lower Devonian) to lower Eifelian (Middle Devonian) of Germany (Rhenish Slate Mountains, North Rhine-Westphalia, and Rhineland-Palatinate). The most complete, new specimen has a long segment of column projecting outward from the basal concavity, which confirms that the Monstrocrinus crown was attached to a column as an adult. Thus, a free-living life mode of a passively rolling or drifting crinoid is refuted and Monstrocrinus is reinterpreted as an attached, stalked echinoderm. This finding is supported by re-examination of the historical type material. A column attachment with a central pentalobate axial canal was recognized for the first time in the calyx of the holotype of the type species Monstrocrinus securifer. The holotype of Monstrocrinus granosus is far more complete than previously thought. It is a partially disarticulated crown embedded together with an associated longer part of the column. The extraordinary spines on the calyx plates of Monstrocrinus can be divided into eight morphological types and into two superordinate categories: “Category A” is a physical part of the calyx plate, whereas “Category B” is an attachment onto the calyx plate. Taxonomically, M. aliformis from the upper Emsian of Spain is treated herein as a subjective junior synonym of M. securifer.

Jurassic cyrtocrinids from Spain are first documented here as representing relatively diverse assemblages from the western Tethys. The species Ascidicrinus pentagonus, Eugeniacrinites cariophilites, Gammarocrinites compressus, Pilocrinus moussoni, Sclerocrinus cf. S. strambergensis, and Tetracrinus moniliformis are described from Oxfordian marl levels belonging to the Yátova Formation, around Tosos (Zaragoza, NE Spain). Although based on partially disarticulated material, these fossils preserve cups, stem columnals, attachment structures, and brachial plates. Based on the sedimentology and associated invertebrate fauna, cyrtocrinids from this area lived below storm wave action but eventually were affected by storms, as opposed to their modern counterparts that occupy deep water environments. Some specimens preserve traces of interactions with other organisms, such as predation marks or epibiontic colonization during life and post-mortem. Attachment structures suggest cyrtocrinids mostly attached on sponges. We note that diversity of cyrtocrinids changed in step with the abundance of sponge reefs in the Jurassic, suggesting that both groups probably had an important link in that period related with similar ecological requirements.

Crinoids, the group known today as the sea lilies, were a major constituent of ocean environments from the late Carboniferous (323–299 million years ago). However, crinoid fossil-forming potential is poor, and they typically fell apart quickly after death. This limits our ability to study much about their life histories, including how they would have grown. Through the discovery of an area of exceptional fossil preservation in the Barnsdall Formation of Oklahoma, we have a rare chance to learn about the growth of one of these species of crinoids, Erisocrinus typus. Here we perform a growth analysis of a well-preserved series of fossils and discuss the patterns that it showed from its juvenile stage to adulthood. Crinoids were major constituents of late Carboniferous (Pennsylvanian) marine ecosystems, but their rapid disarticulation rates after death result in few well-preserved specimens, limiting the study of their growth. This is amplified for cladids, who had among the highest disarticulation rates of all Paleozoic crinoids due to the relatively loose suturing of the calyx plates. However, Erisocrinus typusMeek and Worthen, 1865 has been found in unusually large numbers, most preserved as cups but some as nearly complete crowns, in the Barnsdall Formation in Oklahoma. The Barnsdall Formation, a Koncentrat Lagerstätte, is composed predominantly of fine- to medium-grained sandstone, overlain by mudstone and shale; severe compaction of the fossils in the mudstone and shale layer in this formation allowed for exceptional preservation of the plates. Herein, we summarize a growth study based on 10 crowns of E. typus, showcasing a well-defined growth series of this species from the Barnsdall Formation, including fossils from juvenile stages of development, which are rarely preserved. We used high-resolution photographs imported into ImageJ and recorded measurements of the cup and arms for all nondistorted or disarticulated plates. Results show that the plates of the cup grew anisometrically with both positive and negative allometry. The primibrachial plates of E. typus grew with positive allometry. The brachial plates started as uniserial (i.e., cuneiform) as juveniles but shifted to be biserial. Erisocrinus typus broadly shares similar growth trajectories with other cladids. These growth patterns provide insight into feeding strategies and can aid in understanding crinoid evolutionary paleoecological trends.