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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.

Deuterostomes are a morphologically disparate clade, encompassing the chordates (including vertebrates), the hemichordates (the vermiform enteropneusts and the colonial tube-dwelling pterobranchs) and the echinoderms (including starfish). Although deuterostomes are considered monophyletic, the interrelationships between the three clades remain highly contentious. Here we report, Yanjiahella biscarpa, a bilaterally symmetrical, solitary metazoan from the early Cambrian (Fortunian) of China with a characteristic echinoderm-like plated theca, a muscular stalk reminiscent of the hemichordates and a pair of feeding appendages. Our phylogenetic analysis indicates that Y. biscarpa is a stemechinoderm and not only is this species the oldest and most basal echinoderm, but it also predates all known hemichordates, and is among the earliest deuterostomes. This taxon confirms that echinoderms acquired plating before pentaradial symmetry and that their history is rooted in bilateral forms. Yanjiahella biscarpa shares morphological similarities with both enteropneusts and echinoderms, indicating that the enteropneust body plan is ancestral within hemichordates.

The Cambrian edrioasteroid “Totiglobus” spencensisWen et al., 2019 is redescribed on the basis of a new and exquisitely preserved specimen from the Cambrian Wuliuan of the Spence Shale Member, Langston Formation (Utah). This new occurrence is associated with soft-body preservation of several invertebrate groups and other shelly fossils. The description of “T.” spencensis was originally based on a single poorly preserved specimen. As a result, some of its features, such as curvature of the ambulacra and morphology of floor and cover plates, were misinterpreted or unavailable. The new material allows a new placement in Sprinkleoglobus spencensis n. comb. (Wen et al., 2019) on the basis of the general shape of the theca, attachment disc, and biserial flooring plates with podial pores and multiple cover plate series. The attachment disc of the new specimen lies in contact with a trilobite librigena, supporting evidence that this taxon is one of the earliest known examples of attachment to hard, mineralized substrata among Edrioasteroidea. Edrioasteroids are a group of extinct echinoderms that originated in the Cambrian during the early emergence of animal body plans. Fossils of this group look like modern starfish with five main rays (ambulacra) that converge in a central mouth. But this resemblance is only superficial. Edrioasteroids lived permanently attached to the sea floor or to hard substrates. Herein we redescribe the poorly known edrioasteroid Sprinkleoglobus spencensis n. comb. (Wen et al., 2019) on the basis of a new exquisitely preserved specimen from the Spence Shale Member, Langston Formation of Utah. This new specimen allows interpretation of its anatomy and will help us understand early echinoderm evolution.

Restudy of Proexenocrinus inyoensisStrimple and McGinnis, 1972, shows that this earliest-known rhodocrinitid diplobathrid camerate crinoid (late Floian, Early Ordovician) expresses the only known record of ambulacral floor plates within pinnules. These pinnule floor plates are remarkably conserved plesiomorphic expressions, with anatomy similar to floor plates of some of the earliest pentaradiate echinoderms, although on a smaller scale. Proexenocrinus floor plates provide direct skeletal evidence that the general resemblance of blastozoan (eocrinoid, diploporan, rhombiferan) brachioles and crinoid pinnules is the product of homoplasy. Proexenocrinus posterior cup morphology is interpreted to include an anitaxis, a distinctive posterior interray morphology.

New specimens of Cyclocystoides scammaphoris Smith and Paul, 1982, are here reported from the Upper Ordovician Platteville Formation of northern Illinois, Plattin and Decorah groups of east-central Missouri, and Lebanon Limestone of central Tennessee. These fossils reveal skeletal details that provide insight into the anatomy of cyclocystoids. Of particular significance is a network of channels that likely originate near the center of the central disk and extend along the oral side of the radial plates, bifurcating distally two or three times before entering the radial facets on the proximal surface of each marginal ossicle. From here, the network enters a series of facet canals that extend upward through each marginal ossicle, exiting in a linear row of pores. The canals are very similar in size and distribution to the nerve canals in living echinoderms. The axes of the canals, which number up to 500 in some specimens, and those of the radial ducts project proximally away from the oral surface at an elevation angle of about 25°, apparently forming a network that could have converged within the upper part of the body cavity. This origin and function are made clear by the connection between the channel on each radial plate and the radial facet canal pores within each marginal ossicle.

Twelve specimens of EumorphocystisBranson and Peck, 1940 provide the basis for new findings and a more informed assessment of whether this blastozoan (a group including eocrinoids, blastoids, diploporites, rhombiferans) constitutes the sister taxon to crinoids, as has been recently proposed. Both Eumorphocystis and earliest-known crinoid feeding appendages express longitudinal canals, a demonstrable trait exclusive to these taxa. However, the specimen series studied here shows that Eumorphocystis canals constrict proximally and travel within ambulacrals above the thecal cavity. This relationship is congruent with a documented blastozoan pattern but very unlike earliest crinoid topology. Earliest crinoid arm cavities lie fully beneath floor plates; these expand and merge directly with the main thecal coelomic cavity at thecal shoulders. Other associated anatomical features echo this contrasting comparison. Feeding appendages of Eumorphocystis lack two-tiered cover plates, podial basins/pores, and lateral arm plating, all features of earliest crinoid ‘true arms.’ Eumorphocystis feeding appendages are buttressed by solid block-like plates added during ontogeny at a generative zone below floor plates, a pattern with no known parallel among crinoids. Eumorphocystis feeding appendages express brachioles, erect extensions of floor plates, also unknown among crinoids. These several distinctions point to nonhomology of most feeding appendage anatomy, including longitudinal canals, removing Eumorphocystis and other blastozoans from exclusive relationship with crinoids. Eumorphocystis further differs from crinoids in that thecal plates express diplopores, respiratory structures not present among crinoids, but ubiquitous among certain groups of blastozoans. Phylogenetic analysis places Eumorphocystis as a crownward blastozoan, far removed from crinoids.

Intermediate morphologies of a new fossil crinoid shed light on the pathway by which crinoids acquired their distinctive arms. Apomorphies originating deep in echinoderm history among early nonblastozoan pentaradiate echinoderms distinguish Tremadocian (earliest Ordovician) crinoid arms from later taxa. The brachial series is separated from the ambulacra, part of the axial skeleton, by lateral plate fields. Cover plates are arrayed in two tiers, and floor plates expressed podial basins and pores. Later during the Early Ordovician, floor plates contacted and nestled into brachials, then were unexpressed as stereom elements entirely and cover plates were reduced to a single tier. Incorporation of these events into a parsimony analysis supports crinoid origin deep in echinoderm history separate from blastozoans (eocrinoids, ‘cystoids'). Arm morphology is exceptionally well-preserved in the late Tremadocian to early Floian Athenacrinus broweri new genus new species. Character analysis supports a hypothesis that this taxon originated early within in the disparid clade. Athenacrinus n. gen. (in Athenacrinidae new family) is the earliest-known crinoid to express what is commonly referred to as ‘compound' or ‘biradial’ morphology. This terminology is misleading in that no evidence for implied fusion or fission of radials exists, rather it is suggested that this condition arose through disproportionate growth.

For many years the earliest record of the class Crinoidea was a single late Tremadocian genus. In the past decade, five crinoid genera were described from the early and middle Tremadocian, near the base of the Ordovician. Together these six genera represent a diverse assemblage with all but one expressing existing subclass apomorphies. Two of the recently described genera were initially assigned to their own order (plesion) Protocrinoida but not to a subclass. Here they are placed in the camerates based on apomorphies of the tegmen complex. Protocrinoids exhibit plesiomorphies unlike typical camerates. Two genera group with cladids, one expressing dendrocrinine apomorphies and the other cyathocrinine. One genus is placed within disparids, with iocrinid apomorphies. Based on its ancient age and trait mosaic, the protocrinoid Titanocrinus is designated as outgroup in a phylogenetic analysis using all other Early Ordovician and select Middle Ordovician taxa as an ingroup. Character compilation and phylogenetic analysis posit early class-level plesiomorphies inherited from an unknown ancestry but lost during subsequent crinoid evolution. Class-level apomorphies also emerge, some of which were subsequently lost and others retained. Results are generally robust and consistent with earlier subdivisions of the class, but supporting lower rank reorganizations. Strong support for the camerate branch low in the crinoid tree mirrors findings of earlier workers. Cladids branch from a series of intermediate nodes and disparids nest highest. Branching of disparids from cladids could be homoplastic.