The extraxial axial theory is used to investigate homology of ambulacral and oral plating because it predicts terminal branching and terminal addition of plates in the axial skeleton, although exceptions to the former may occur in some Paleozoic echinoderms. The variety of morphological designs and anomalous individuals also provide tests of plate homology. Homology of ambulacra is generally accepted, with the hydropore and/or single gonopore in Carpenter’s CD interray. In the 2-1-2 ambulacral pattern the unbranched ambulacrum is always in Carpenter’s A ray. All ambulacral morphology requires just three instructions: ‘grow,’ ‘branch,’ and ‘stop.’ The range of variation in echinoderms with fewer than five ambulacra implies that both the ‘branch’ and ‘stop’ instructions acted independently in all five rays. Numbers of ambulacra may or may not correlate with numbers of orals. Two basic patterns of ‘cystoid’ oral plating occur; with a single radial (circum-oral, CO) plate from each ambulacrum plus a sixth in the CD interray, and with all six interradial peri-oral (PO) plates, with two in the CD interambulacrum. Five ‘orals’ may involve loss of PO3 or PO6. Erect ambulacral structures are lost first in taphonomy and so poorly known. All ambulacral skeletal elements bear the same topological relationship to ambulacral soft tissues. Where branched ambulacra occur, the trunk or flooring plates are often modified first brachiolars or pinnulars. Both brachioles and pinnules may arise from facets developed on one or two flooring plates. Terminal addition of plates, spacing of brachioles/pinnules, and lack of musculature to open cover plates all suggest that ‘cystoids’ had extensions of the water vascular system in their ambulacra.

Universal elemental homology (UEH) underpins recent understanding of peristomial and ambulacral elemental homology of pentaradiate echinoderms by providing a uniform set of terminology to construct phylogenetic characters. Variation in the expression of these elements provides evidence for phylogenetic relationships. Two nonhomologous sets of plates border the peristome and are associated with two nonhomologous sets of floor plates forming the ambulacral food groove. Some edrioasteroid-grade and eocrinoid-grade echinoderms have ambulacral systems formed from oral frame plates and adradial floor plates, whereas most blastozoans and crinoids bear oral plates and abradial floor plates. These plates are expressed in a variety of ways among echinoderms, but nearly all can be reconciled with the underlying model. Arguments against UEH are methodologically flawed and confuse many terms and interpretations.

The loss of information resulting from taphonomic degradation could represent a significant bias in the study of morphological diversity. This potential bias is even more concerning given the uneven effect of taphonomy across taxonomic groups, depositional facies, and stratigraphic successions and in response to secular changes through the Phanerozoic. The effect of taphonomic degradation is examined using character-based morphological data sets describing disparity in Paleozoic crinoids and blastozoans. Characters were sequentially excluded from the analyses following progressive taphonomic loss to determine how morphologic metrics, such as the relative distribution of taxa in morphospace and partial disparity, changed with increasing taphonomic alteration. Blastozoans showed very little change in these metrics with decreasing preservational quality, which is a result of characters that create distance in morphospace being recognizable in isolated plates. The opposite result is present in crinoids as the characters that are important in structuring the morphospace require intact modules (i.e., the calyx) to accurately assess. Temporal and stratigraphic trends produced encouraging results in that patterns could be largely recovered even with exaggerated taphonomic biases. However, certain parts of a stratigraphic sequence should be avoided and morphological outliers could potentially play a larger role through time, though both of these biases can be easily identified and avoided. The methods presented in this study provide a way to assess potential taphonomic biases in character-based studies of morphological diversity.

Fossilized tests of 1,053 Echinocyamus stellatus (Capeder, 1906) from the Miocene Globigerina Limestone Formation exposed on the northern coast of Gozo (Maltese Islands) were analyzed for predation traces. Specimens mixed by time-averaging processes can be clearly separated into two distinct samples according to their preservation as phosphatized or nonphosphatized individuals. Overall, 11.1% of the tests reveal holes that are referred to the ichnospecies Oichnus simplex (Bromley, 1981). Because of the hole morphology and diameter, the holes are interpreted as predatory drill holes, most likely produced by cassid gastropods. Redeposited phosphatized echinoids derived from an earlier period of reduced sedimentation rates show drilling frequencies of 20.5%. Younger, autochthonous, nonphosphatized echinoids show drilling frequencies of 8.1%. In both samples, predators predominantly targeted the aboral side of the echinoid test, particularly on the petalodium.

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.

Hydrospires are internal structures in blastoids that primarily served a respiratory function. Historically, hydrospires have been modeled as passive-flow respiratory structures with a vertical orientation. This project constructed virtual 3D models of blastoids from legacy acetate peel collections at the Naturalis Museum in the Netherlands. Computational fluid dynamic (CFD) simulations of the blastoid models reconstructed in living position indicated that hydrospires likely were oriented horizontally when the blastoid was in feeding mode in current velocities>0.5 cm/s to 10 cm/s. In this range of current velocities, passive water flow through the hydrospires did not produce conditions optimized for efficient gas exchange. However, optimal water flow through the hydrospires could be achieved if the excurrent velocity of water exiting the hydrospire through the spiracle was approximately one-half the velocity of ambient environmental currents. Maintaining such a ratio in the dynamic current systems in which blastoids lived suggests that cilia-driven active water flow through the hydrospires is a better model for optimizing respiratory effectiveness.

The primitive blastozoan Felbabkacystis luckae n. gen. n. sp. is described from the Drumian Jince Formation, Barrandian area (Czech Republic) from eleven fairly well-preserved specimens. Its unique body plan organization is composed of a relatively long, stalk-like imbricate structure directly connected to the aboral imbricate cup of the test and of an adoral vaulted tessellate test supporting the ambulacral and brachiolar systems. Its bipartite test, called prototheca, highlights the evolution of the body wall among blastozoans. Felbabkacystis n. gen. shows the combination of plesiomorphic (imbricate stalk-like appendage) and derived features (highly domed peristome, elongate epispires). The new genus is interpreted as a transitional form between calyx-bearing and theca-bearing blastozoans, and is attributed to the new family Felbabkacystidae. The lithology, the associated fauna, and the possession of a long stalk suggest that Felbabkacystis was probably a low-level suspension feeder living in relatively deep settings.

The genus Rhopalocystis (Eocrinoidea, Blastozoa) is characterized by both a short stratigraphic range (Fezouata Shale, middle Tremadocian to middle Floian, Lower Ordovician) and a reduced geographic extension (Agdz-Zagora area, central Anti-Atlas, Morocco). Since the original description of its type species (R. destombesi Ubaghs, 1963), three successive revisions of the genus Rhopalocystis have led to the erection of nine additional species. The morphological disparity within this genus is here critically reassessed on the basis of both historical material and new recently collected samples. The detailed examination of all specimens, coupled with morphometric and cladistic analyses, points toward a relatively strong support for five morphotypes. A systematic revision of Rhopalocystis is thus suggested, with only five valid taxa: R. destombesi, R. fraga Chauvel, 1971, R. grandis Chauvel, 1971, R. havliceki Chauvel, 1978, and R. zagoraensis Chauvel, 1971. The five others are considered as junior synonyms (R. dehirensis Chauvel and Régnault, 1986, R. lehmani Chauvel and Régnault, 1986, R. sp. A, R. sp. B, and R. sp. C).

A diverse crinoid fauna is described from the Upper Ordovician (Katian) Fombuena Formation from the eastern Iberian Chains of Spain. New crinoids include the diplobathrid camerates Fombuenacrinus nodulus n. gen. n. sp., Goyacrinus gutierrezi n. gen. n. sp., Dalicrinus hammanni n. gen. n. sp., and Ambonacrinus decorus n. gen. n. sp.; the monobathrid camerate Eopatelliocrinus hispaniensis n. sp.; and the cladid Picassocrinus villasi n. gen. n. sp. A new occurrence of Heviacrinus melendezi Gil Cid, Domínguez Alonso, and Silván Pobes, 1996 is also documented from the Castillejo Formation (Darriwilian, Middle Ordovician) from the eastern Iberian Chains of Spain. The Fombuena Formation comprises a Gondwanan crinoid assemblage from a high paleolatitude and has the highest crinoid diversity of any currently known Katian Gondwanan fauna. This assemblage is compared to other Katian age faunas around the globe, and its paleobiogeographic implications are discussed.

Moroccodiscus smithi represents a new cyclocystoid genus and species based on moldic specimens from the Middle Ordovician Taddrist Formation (Darriwilian) of SE Morocco. This represents the earliest articulated member of the Cyclocystoidea and is the first complete cyclocystoid described from the Ordovician of Gondwana, as well as the first cyclocystoid ever recorded from Africa. The anatomy and morphology of this new species were studied using a combination of conventional paleontological methods and nondestructive X-ray computed tomography. Because Moroccodiscus differs from other cyclocystoids, in particular by lacking cupules attached to the marginal ossicles, it is assigned to the new family Moroccodiscidae. This new taxon illustrates the relatively poorly known early diversification of these enigmatic extinct echinoderms and sheds light on the mode of life of cyclocystoids, including injuries to plate circlets during early ontogeny and folding of these disk-like specimens at the time of death. The overall thecal shape was very similar in cyclocystoids and many domal edrioasteroids, probably because they were both sessile or attached, benthic, suspension feeders. However, many oral surface, ambulacral, and marginal ring features had become very different, indicating that these two groups had either converged because of similar life modes or were only distantly related sister groups.

The Holocystites Fauna is an enigmatic group of North American diploporitans that presents a rare window into unusual middle Silurian echinoderm communities. Multiple systematic revisions have subdivided holocystitids on the basis of presumed differences in oral area plating and respiratory structures. However, these differences were based on a fundamental misunderstanding of the homologous elements of the oral area and the taphonomic process; taphonomic disarticulation of the oral area formed the basis for the erection of Pentacystis and Osgoodicystis as separate genera, and Osgoodicystis is interpreted as the junior synonym of Pentacystis. Holocystitids show a conservative peristomial bordering plate pattern that is shared among all described genera. The peristome is bordered by seven interradially positioned oral plates as is typical for oral plate–bearing blastozoans. A second open circlet of facetal plates lies distal to the oral plates; five of these facetal plates bear facets for feeding appendages (lost on the A ambulacrum in some taxa), while two lateral facets (present in all taxa except Pustulocystis) do not. Holocystitid taxa show minor modifications to this basic peristomial bordering plate pattern. As thecal morphologies are highly variable within populations, taxonomic revision of holocystitids is based on modifications of the plating of the oral area.

The Permian is regarded as one of the most crucial intervals during echinoid evolution because crown group echinoids are first widely known from the Permian. New faunas provide important information regarding the diversity of echinoids during this significant interval as well as the morphological characterization of the earliest crown group and latest stem group echinoids. A new fauna from the Capitanian Lamar Member of the Bell Canyon Formation in the Guadalupe Mountains of West Texas comprises at least three new taxa, including Eotiaris guadalupensis Thompson n. sp. an indeterminate archaeocidarid, and Pronechinus? sp. All specimens represented are silicified and known from disarticulated or semiarticulated interambulacral and ambulacral plates and spines. This assemblage is one of the most diverse echinoid assemblages known from the Permian and, as such, informs the paleoecological setting in which the earliest crown group echinoids lived. This new fauna indicates that crown group echinoids occupied the same environments as stem group echinoids of the Archaeocidaridae and Proterocidaridae. Furthermore, the echinoids described herein begin to elucidate the order of character transitions that likely took place between stem group and crown group echinoids. At least one of the morphological innovations once thought to be characteristic of early crown group echinoids, crenulate tubercles, was in fact widespread in a number of stem group taxa from the Permian as well. Crenulate tubercles are reported from two taxa, and putative cidaroid style U-shaped teeth are present in the fauna. The presence of crenulate tubercles in the archaeocidarid indicates that crenulate tubercles were present in stem group echinoids, and thus the evolution of this character likely preceded the evolution of many of the synapomorphies that define the echinoid crown group.

Three new ophiuran species, Enakomusium whymanae n. sp., Aspidophiura? seren n. sp., and Ophiotitanos smithi n. sp., and an unnamed specimen assignable to the genus Dermocoma are described from the Callovian to Oxfordian Oxford Clay Formation of Great Britain. These determinations are based on new finds and a critical reassessment of historic specimens. The Oxford Clay ophiuroids represent two loose assemblages, one from the middle Callovian Peterborough Member and the other from the lower Oxfordian Weymouth Member. Both assemblages accord well with coeval midshelf mud bottom ophiuroid communities in terms of taxonomic composition and relative abundance of taxa. The British Oxford Clay ophiuroids are particularly significant as they are one of the rare instances where multiple species are represented, almost exclusively, by exceptionally preserved articulated skeletons. This provides an important window into the understanding of mid-Upper Jurassic ophiuroid paleobiology.

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.

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 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.

The external expression of hydrospires in blastoids has provided a basis for major and minor group classification in the clade for over a century. Unfortunately, the complete anatomy of the hydrospires has never been comprehensively studied. This study examined and described the internal hydrospires of six spiraculate species by digitally extracting hydrospire data from a legacy data set of serial acetate peels. Although only six models have been currently generated, hydrospire morphology is variable both within and between previously escribed spiraculate families. Hydrospires were found to possess novel characters that were incorporated into a phylogenetic analysis of the six digitally modeled species and several related species. The addition of internal morphology into the phylogenetic analysis provides further resolution between groupings of blastoids.