Tiering of benthic marine suspension-feeding communities on soft substrata has varied throughout the Phanerozoic. Epifaunal tiering was most developed during the middle and late Paleozoic and the Triassic to Jurassic, with large-scale reductions in tiering occurring during the Permian-Triassic extinctions and after the Jurassic. Infaunal tiering reached its highest level of organization after the Paleozoic.

The phylogenetic relationships between major groups of plesiomorphic pentaradial echinoderms, the Paleozoic crinoids, blastozoans, and edrioasteroids, are poorly understood because of a lack of widely recognized homologies. Here, we present newly recognized oral region homologies, based on the Universal Elemental Homology model for skeletal plates, in a wide range of fossil taxa. The oral region of echinoderms is mainly composed of the axial, or ambulacral, skeleton, which apparently evolved more slowly than the extraxial skeleton that forms the majority of the body. Recent phylogenetic hypotheses have focused on characters of the extraxial skeleton, which may have evolved too rapidly to preserve obvious homologies across all these groups. The axial skeleton conserved homologous suites of characters shared between various edrioasteroids and specific blastozoans, and between other blastozoans and crinoids. Although individual plates can be inferred as homologous, no directly overlapping suites of characters are shared between edrioasteroids and crinoids. Six different systems of mouth (peristome) plate organization (Peristomial Border Systems) are defined. These include four different systems based on the arrangement of the interradially-positioned oral plates and their peristomial cover plates, where PBS A1 occurs only in plesiomorphic edrioasteroids, PBS A2 occurs in plesiomorphic edrioasteroids and blastozoans, and PBS A3 and PBS A4 occur in blastozoans and crinoids. The other two systems have radially-positioned uniserial oral frame plates in construction of the mouth frame. PBS B1 has both orals and uniserial oral frame plates and occurs in edrioasterid and possibly edrioblastoid edrioasteroids, whereas PBS B2 has exclusively uniserial oral frame plates and is found in isorophid edrioasteroids and imbricate and gogiid blastozoans. These different types of mouth frame construction offer potential synapomorphies to aid in parsimony-based phylogenetics for exploring branching order among stem groups on the echinoderm tree of life.

Phylogenetic relationships among early crinoids are evaluated by maximizing parsimonious-informative characters that are unordered and unweighted. Primarily Tremadocian-Darriwilian (Early-Middle Ordovician) taxa are analysed. Stratigraphic congruence metrics support the best phylogenetic hypothesis derived using parsimony methods. This study confirms the traditionally recognized lineages of Palaeozoic crinoids and provides new information on the branching order of evolving lineages. Camerates are basal crinoids with progressively more tipward groups (from an Ordovician perspective) being protocrinoids, cladids (paraphyletic), hybocrinids and disparids. The Protocrinoida should be maintained, but the Aethocrinida should be placed within the Cladida. The results of this study identify phylogenetic structure amongst the major early crinoid lineages and delineate the relative positions of crinoid higher taxa along a tree. Each valid higher taxon discussed herein requires a comprehensive treatment to delimit within-lineage phylogenetic relationships.Key words: Echinodermata, Ordovician, phylogenetic methods, stratigraphic congruence.C R I N O I D S are regarded as the most primitive of the five living echinoderm classes (asteroids, crinoids, echinoids, holothurians and ophiuroids), but the phylogenetic position of crinoids amongst Palaeozoic echinoderms has been widely debated. The oldest known crinoids are from the early Tremadocian (Ordovician) (Guensburg and Sprinkle 2001, 2003, 2009 Guensburg 2010); and as part of the Great Ordovician Biodiversification Event (GOBE), they diversified to become the dominant stalked echinoderm clade by the early Late Ordovician (Sandbian). Crinoids became progressively more dominant amongst stalked echinoderms in most marine settings through the remainder of the Palaeozoic and were instrumental in establishment of the structure of Palaeozoic epifaunal suspension-feeding communities (Ausich and Bottjer 1982; Bottjer and Ausich 1987).However, a consensus has been difficult to reach concerning the phylogenetic position of crinoids amongst Ordovician clades and the early diversification of the Crinoidea. At the heart of this issue is a fundamental question of echinoderm phylogeny as a whole. Are stalked echinoderms a clade or a grade of echinoderm evolution? In this contribution, we adopt results from echinoderm universal elemental homology studies that have demonstrated that crinoids are nested within clades of other stalked echinoderms (Sumrall 2008(Sumrall , 2010 Sumrall and Waters 2012; Kammer et al. 2013; Sumrall 2014). With stalked echinoderms, including crinoids, recognized as a clade (the Pelmatozoa Leuckart, 1848), phylogenetic relationships of early crinoid evolution are examined.Various hypotheses for the origin and early evolution of crinoids have relied on the supposition that specific characters had special significance in identifying ancestordescendant relationships (e.g. ambulacral floor plates or lintels), that a priori interpretations of morphological tr...

Predator-prey interactions are thought by many researchers to define both modern ecosystems and past macroevolutionary events. In modern ecosystems, experimental removal or addition of taxa is often used to determine trophic relationships and predator identity. Both characteristics are notoriously difficult to infer in the fossil record, where evidence of predation is usually limited to damage from failed attacks, individual stomach contents, onesided escalation, or modern analogs. As a result, the role of predation in macroevolution is often dismissed in favor of competition and abiotic factors. Here we show that the end-Devonian Hangenberg event (359 Mya) was a natural experiment in which vertebrate predators were both removed and added to an otherwise stable prey fauna, revealing specific and persistent trophic interactions. Despite apparently favorable environmental conditions, crinoids diversified only after removal of their vertebrate consumers, exhibiting predatory release on a geological time scale. In contrast, later Mississippian (359-318 Mya) camerate crinoids declined precipitously in the face of increasing predation pressure from new durophagous fishes. Camerate failure is linked to the retention of obsolete defenses or "legacy adaptations" that prevented coevolutionary escalation. Our results suggest that major crinoid evolutionary phenomena, including rapid diversification, faunal turnover, and species selection, might be linked to vertebrate predation. Thus, interactions observed in small ecosystems, such as Lotka-Volterra cycles and trophic cascades, could operate at geologic time scales and higher taxonomic ranks. Both trophic knock-on effects and retention of obsolete traits might be common in the aftermath of predator extinction.Carboniferous | durophagy | biodiversity | paleontology | macroecology

Tiering is the vertical distribution of organisms within the benthic boundary layer. Primary tierers are suspension-feeding organisms with a body or burrow that intersects the seafloor. Secondary tierers are suspension-feeders that maintain positions above or below the sediment-water interface as either epizoans on primary tierers and plants or by living in the burrows of primary tierers. Different primary tierers from soft substrata, nonreef, shallow subtidal shelf and epicontinental sea settings have had different tiering histories, resulting largely from contrasting constructional and phylogenetic constraints. Primary colonial tierers generally occupied lower epifaunal tiers during the Paleozoic and Mesozoic, but since the Cretaceous they have been dominant in the highest tier (+ 20 to +50 cm). Primary echinoderm tierers have been almost exclusively epifaunal, and from the Paleozoic through the Jurassic they were present throughout the epifaunal tiered structure. Although primary bivalve tierers have been both epifaunal and infaunal, they have occupied only lower epifaunal tiers, whereas they have adapted to all levels of the infaunal tiering structure, particularly from the late Paleozoic through the Recent. Brachiopods have lived primarily in tiers directly above or below the water-sediment interface and have not contributed significantly to tiering complexity.Of the numerous physical and biotic processes and constraints that affect shallow marine benthos, a few have contributed more significantly to changes in tiering patterns. Trends for increasing body size could have accounted for most of the development of tiering complexity up to +50 cm and down to –12 cm. Development of tiering above +50 cm could have been due to processes which would have yielded greater feeding capability, such as competitive interactions for a place from which to feed or adaptations to velocity gradients in the hydrodynamic boundary layer. The most significant process for development of infauanl tiering below –12 cm appears to have been as an adaptive response for predator avoidance.Unlike infaunal tiering, which never declined after it developed, epifaunal tiering has undergone a general reduction twice. Reduction in epifaunal tiering at the end of the Paleozoic appears to have been the result of the mass extinction at this time, whereas long-term biotic processes seem to have been more important for the tiering decline at the end of the Mesozoic. Tiering structure through the Phanerozoic was thus produced through interactions of a number of physical and biotic factors, tempered by constructional and phylogenetic constraints of each primary tierer group.

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A stepwise approach is employed to determine the phylogeny of the initial crinoid radiation during the Arenig and Llanvirn series of the Ordovician. Parsimony-based character analysis is completed first on Arenig crinoids and then for Arenig and Llanvirn crinoids combined.The topology from well-resolved trees of this early crinoid radiation indicates that the Crinoidea should be subdivided into six subclasses. A new subclass and new order, Aethocrinea and Aethocrinida, respectively, are proposed for crinoids with four circlets of plates in the aboral cup: lintels, infrabasals, basals, and radials. This aboral cup construction is best displayed byAethocrinus, one of the oldest known crinoids (?Tremadoc-Arenig). However, this primitive aboral cup construction is also present in two Llanvirn crinoids,PerittocrinusandTetracionocrinus.The Aethocrinea was a small, short-lived radiation of crinoids with this design that was different from those of other crinoids. Two families in the Aethocrinida are the Aethocrinidae, which includesAethocrinus, and the Perittocrinidae, which includesPerittocrinusandTetracionocrinus.In addition to the Aethocrinea, the following crinoid subclasses are recognized based on character analyses of these earliest crinoids: Cladida, Camerata, Disparida, Flexibilia, and Articulata.

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