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.
Apektocrinus ubaghsinew genus and species is a monospecific taxon assigned to the new family Apektocrinidae based on additional preparation of a single previously studied specimen.Apektocrinusis among the oldest known crinoids (Early Tremadoc, Early Ordovician). Although expressing crinoid apomorphies, it is interpreted as retaining plesiomorphies in its arms reflecting early edrioasteroid rather than blastozoan (eocrinoid) ancestry. Apomorphies represent basal crinoid and cladid (crownward) levels of phylogeny.Restudy fortifies previous reports of the presence of a basal echinoderm plesiomorphy; floor plates above brachials in the arms ofApektocrinus, as well as in other approximately contemporary crinoids.Apektocrinusfurnishes the first record of podial basins in crinoid arms. Arms and calyx ofApektocrinusmerge gradually, facilitated by continuations of interbrachials (extraxial body plates) extending onto the arms and separating floor plates from brachials. These arm interbrachials, which diminish and pinch out distally as floor plates nestle into the brachial (adoral) groove, have not been recognized as such in crinoids.
Although echinoderm debris is locally common, articulated specimens are rare in Late Cambrian rocks from the Great Basin and Rocky Mountains of the western United States and are mostly associated with hardgrounds. The fauna, including cornute stylophorans, trachelocrinid eocrinoids, solute homoiosteleans, and rare edrioasteroids, includes several members of the archaic Cambrian Evolutionary Fauna, which had already passed its maximum diversity for echinoderms. In addition to the low diversity, articulated specimen abundance is very low, averaging only about one-tenth that found in overlying Lower Ordovician units. The transition between the Cambrian and Paleozoic Evolutionary Faunas for echinoderms in North America apparently occurred rapidly very close to the Cambrian-Ordovician boundary, because no unequivocal examples of the Paleozoic fauna (such as crinoids, glyptocystitid rhombiferans, asteroids, or echinoids) were found in the Late Cambrian sections.New taxa include several cothurnocystid stylophorans assigned to Acuticarpus delticus, new genus and species, Acuticarpus? republicensis, new species, and Archaeocothurnus goshutensis, new genus and species; Scotiaecystis? species, a poorly preserved cornute stylophoran with lamellipores; Minervaecystis? species, a fragmentary solute homoiostelean based on several steles; Tatonkacystis codyensis, new genus and species, a well-preserved trachelocrinid eocrinoid with five unbranched arms bearing numerous brachioles; an unnamed, poorly preserved, epispire-bearing eocrinoid; an unnamed, poorly preserved, globular eocrinoid? lacking epispires; and an unnamed, heavily weathered, edrioasterid edrioasteroid. Nearly all holdfasts found in these Upper Cambrian units are single-piece blastozoan types, probably belonging to trachelocrinid and other eocrinoids. Distinctive columnals and thecal plates of several additional undescribed eocrinoids and other echinoderms were locally abundant and are also described.
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