It was 17 years ago this month, 1980 in Atlanta, Georgia, that the first short course on echinoderms was held. The conveners were Johnny Waters and Tom Broadhead. The Sunday short courses had been started only two years before, in 1978, with the mollusks being the first short course and arthropods the second. Neither of these first two short courses had accompanying printed notes. Thus, we echinodermers were the ones who began this tradition of short course notes that has continued to this day.

Those participants in that 1980 short course who are also on the program today are, in addition to Johnny Waters, a convener of both short courses, Dave Meyer, Jim Sprinkle, Bill Ausich, and Ron Parsley. Many new names and faces have appeared in the study of fossil echinoderms or those who study living echinoderms in such a manner that their work has bearing on the ancient record of these animals.

Comatulid crinoids, or featherstars, are the dominant group of living crinoids and occur in a wide range of modern marine environments from the intertidal zone to the abyss. This chapter outlines current understanding of comatulid classification, distribution, feeding ecology, diets, predation, and taphonomy. A detailed introduction to morphological features, terms and associated symbology is given together with a discussion of practical aspects of working with specimens and the difficulties associated with species identifications. An artificial key to the families of comatulids is included.

Stalked crinoids (sea lilies) are not extinct, but are restricted to depths below 100 m and comprise over 80 living species. Over the past 20 years, a wide range of new information on the biology of stalked crinoids has been acquired from deep-sea photography and submersible studies. Taphonomic studies show that stalked crinoids can disarticulate and undergo breakage and abrasion without significant transportation, although crinoids can survive the rigors of transportation without damage if decay has not weakened connective tissues. Because stalks can remain articulated longer than crowns, the occurrence of articulated fossil crinoid stalks should not be interpreted as proof of rapid burial. Stalked crinoids are passive suspension feeders forming filtration fans oriented normal to bottom currents of low velocity. Mutable collagenous tissues comprising the ligaments of stalk, cirri, and arms are significant in providing variable stiffness that enables stalked crinoids to maintain elevation of the stalk and the filtration fan feeding posture. Regenerated arms and stalks lacking crowns suggest that stalked crinoids are subject to predation by fish in deep water, although predation pressure is likely lower than in shallow water.

Recent advances in crinoid functional morphology have been concentrated predominantly in two areas, one involving the study of mutable collagenous tissues (MCTs), and the other dealing with the implications of crinoids' rheophilic nature. The presence of MCTs in crinoid cirri and stalks, and their importance to the function of these structures has been demonstrated in several studies. It has been recently reported that MCTs have contractile properties (Birenheide and Motokawa, 1996); if this is corroborated, it would help resolve many fundamental problems in crinoid functional morphology. Studies of the crinoids in their natural setting have confirmed their predominantly rheophilic nature. The application of the rheophilic paradigm and the principles of physics and engineering to crinoids has provided insights into several areas of crinoid functional morphology, including: (1) mechanics of particle interception—direct interception (and not sieving) as the dominant mode of particle capture by the tube feet (no mucus net); (2) the importance of feeding posture and orientation—(a) bilateral symmetry of the feeding posture (oral side downcurrent), (b) the problems associated with maintaining such a posture under bi- or multi-directional flow (swiveling of pinnules, arms, and /or entire filters); (3) the effect of filter resistivity on fluid interception—solid filters need high ambient current velocities; and (4) the influence of nearest neighbors on fluid interception—side-by-side neighbors might be favored over upstream–downstream neighbors.

Crinoids were a common component of Paleozoic benthic paleocommunities, yet they have been under-utilized in paleoecological analyses. Recent efforts to incorporate disarticulated ossicles into these analyses have greatly increased the robustness of paleoecological patterns noted for the Crinoidea. Analyses of crinoid functional morphology, particularly filtration dynamics, have provided testable hypotheses concerning the distribution of crinoids among benthic environments. These models predict that crinoids with dense-mesh filtration fans should be most common in high energy, shoreward paleoenvironments, whereas open-fan crinoids should be most common in low-energy, offshore paleoenvironments. Review of the Paleozoic fossil record appears to support these general predictions—from the Late Ordovician to the end of the Paleozoic, dense-fan crinoids are most abundant in nearshore paleoenvironments, whereas open-fan crinoids are most abundant offshore.

The partitioning of crinoid diversity through the Paleozoic shifted through time. Beta diversity was highest in the Ordovician, implying that the early diversification of crinoids was focused on partitioning the benthic landscape among taxa. Beta diversity was quite low by the late Paleozoic, however, local and within-habitat alpha diversity was much greater than during the Ordovician. This resulted in generally higher levels of eurytopy in the late Paleozoic compared to the Ordovician. Patterns of faunal disassembly associated with regional extinctions in North America during the Ordovician and Permian underscore the differences in the paleoecology of these crinoid faunas.

Issues of scale are becoming increasingly important to paleobiological interpretations of the fossil record. Nevertheless, a number of biological processes display scale-independent behaviors. The effects of predation on the distribution of dense populations of epifaunal, suspension-feeding ophiuroids are scale-independent, at scales ranging from the microecological to the macroevolutionary. On a microecological scale (meters to kilometers, hours to days), dense ophiuroid populations are limited in shallow-water environments by predatory fishes and crabs. On a larger, ecological scale (tens to hundreds of kilometers, decades to centuries), circumstantial evidence indicates that oceanographically driven, multidecadal cycles of predator abundance determine the abundance of ophiuroids throughout the western English Channel. On a macroevolutionary scale (millions to tens of millions of years, global spatial scale), dense, autochthonous assemblages of ophiuroids declined in conjunction with the Mesozoic diversification of modern shell-crushing predators: teleostean fishes, decapod crustaceans, and neoselachian sharks. The sporadic reappearance of dense ophiuroid populations in a late Eocene, shallow marine deposit in Antarctica suggests that predator-prey relationships were disrupted as temperatures declined in the region at that time. Scale-independence is a useful model for explaining and predicting patterns of distribution of dense ophiuroid populations in time and space.

Preservational style of fossil echinoid tests allows assessment of the likelihood of post-mortem transport out of the preferred sediment type of the living echinoid. Sedimentologic study of the matrix of untransported specimens allows a check on functional morphologic inferences of the species' preferred sediment types. Functional morphologic analysis allows inference of a species preferred sediment type because the petals, fasciolaes, tubercles, ambulacral pores, ambulacral shape, and test profile control the echinoid's ability to burrow, and the grain size of sediment into which it is capable of doing so. Past studies have achieved better than 90% accuracy in predicting the grain size of thin sections of rocks containing echinoids, simply by interpretation of their functional morphology. Most mistaken predictions are attributable to species living in sediments that are less difficult to burrow in (sands) than those to which they are adapted (muds). Other species may live in sediments in which they are not well adapted by assuming an epifaunal mode of life.

Relative proportions of species in an echinoid fauna preferring various sediment grain sizes, plotted for each of a number of localities, has proven useful in inferring generalized facies patterns within regions.

Plots of temporal changes in echinoid species diversity through time correspond well to changes in proportions of species inferred to have preferred various substrate conditions, suggesting an environmental and taphonomic component to simple diversity curves.

Taphonomic study of echinoderms provides useful information on sedimentary conditions before, during, and after burial. Taphonomic studies of Recent echinoderms indicate that much skeletal disarticulation occurs within a few days after death. However, experiments also indicate that within a short period after death echinoderm carcasses remain rather resistant to disarticulation, and thus may be transported a considerable distance by currents; following periods of a few hours of decay, more delicate portions of echinoderm skeletons are readily disarticulated. Some skeletal modules (e.g., crinoid pluricolumnals) may resist disarticulation for periods of months in quiet- and or cool-water environments. Anoxia promotes intact preservation by excluding scavenging metazoans. Echinoderm ossicles may undergo minor abrasion and/or corrosion if left exposed, and less dense stereom corrodes much more rapidly than dense plates, such as echinoid spines. However, heavily abraded ossicles may indicate prefossilization and reworking.

Various groups of echinoderms (e.g., pelmatozoans, asterozoans, echinoids) have differing propensities for degradation and, therefore, produce different arrays of preserved fossil material primarily depending upon the relative rates of burial, bottom-water oxygenation, and turbulence. Echinoderms may be divided into three groups based upon the relative ease of skeletal disarticulation. Type 1 echinoderms include weakly articulated forms (e.g., asteroids and ophiuroids) that rapidly disintegrate into individual ossicles. Type 2 includes those echinoderms whose bodies contain portions in which are more tightly sutured, as well as portions in which the ossicles are somewhat more delicately bound (e.g., crinoids, regular echinoids). Such echinoderms display more varied taphonomic grades from fully intact to mixtures of isolated ossicles and articulated modules. Type 3 comprises those echinoderms (e.g., irregular echinoids) in which major portions of the skeleton are so resistant to disarticulation that they may be broken across sutures rather than coming apart at plate boundaries.

Comparative taphonomy of particular types of echinoderm skeletal remains leads to recognition of distinctive taphofacies that characterize particular depositional environments. Taphofacies include two types of characteristic modes of fossil preservation: event taphonomic signatures and background taphonomic signatures. Depending upon normal conditions of environmental energy and rates of sedimentation, the background condition of various types of echinoderms for a given facies may range from articulated, unabraded skeletal modules (in Types 2 and 3) to highly corroded and/or abraded ossicles. Conversely, the occurrence of fully intact fossil echinoderms provides unambiguous evidence of rapid and deep burial of benthic communities. Such well-preserved fossil assemblages can provide a wealth of information regarding the paleobiology of echinoderms, as well as the nature of the depositional events and burial histories.

This paper presents a preliminary classification and characterization of background and event aspects of echinoderm taphofacies for carbonate- (9 taphofacies, including reefs and hardgrounds) and siliciclastic-dominated (5 taphofacies) environments. In each case, we recognize a spectrum of echinoderm taphofacies that coincides with a gradient of environments, ranging from nearshore, high energy shoreface through proximal and distal storm-influenced shelf, to deeper ramp and dysoxic basinal settings. Most taphofacies also feature particular styles of obrution (smothered bottom) Lagerstätten. These range from scattered lenses of articulated fossils in some high energy sandstone and grainstone facies to bedding planes of articulated, pyrite coated specimens in dark shales. We classify and discuss the genesis of these types of Lagerstätten and list typical examples. Finally, we present a simple model that integrates the occurrence of various echinoderm taphofacies with concepts of cyclic and sequence stratigraphy.

The crystallographic orientations of echinoderm skeletal elements can supplement standard morphological comparisons in the exploration of echinoderm evolution. At a coarse scale, many echinoderms share a crystallographic pattern in which c axes radiate away from the axis of pentaradial symmetry. Within this common pattern, however, c axes of different taxa can differ dramatically in their degree of variability, angles of inclination, and relationships to the external morphology of skeletal elements. Crystallographic data reflect a variety of taxon-specific influences and therefore reveal different information in different taxa. In echinoids, orientations of c axes in coronal plates correlate well with high-level taxonomic groupings, while c axes of apical plates record modes of larval development. In blastoids, c axes of radial plates have a structural interpretation, with the c axis oriented parallel to the orientation of the surface of the radial plate during its initial growth stages. In crinoids, c axes do not correlate with taxonomic group, plate morphology, or developmental sequence, but instead correlate with relative positions of skeletal elements on the calyx. Although their full potential has yet to be explored, the varied crystallographic patterns in echinoderms have been used to clarify skeletal structure, characterize developmental anomalies, and infer homologies of skeletal plates both within specimens and between groups. A axes are less constrained in their orientations than c axes and offer less promise of revealing novel paleobiological information.

Echinoderms underwent a major two-part radiation that produced all of the major groups found in the fossil record between the Early Cambrian and the Middle Ordovician. A small initial radiation in the Early and Middle Cambrian produced about nine classes containing low-diversity members of the Cambrian Evolutionary Fauna. These were characterized by primitive morphology, simple ambulacral feeding structures, and the early development of a multiplated stalk or stem for attachment to skeletal fragments on a soft substrate. Several groups became extinct at the end of the Middle Cambrian, leaving the Late Cambrian as a gap of very low diversity in the fossil record of echinoderms with only four classes preserved and very few occurrences of complete specimens, mostly associated with early hardgrounds. The survivors from this interval re-expanded in the Early Ordovician and were joined by many newly evolved groups to produce a much larger radiation of more advanced, diverse, and successful echinoderms representing the Paleozoic Evolutionary Fauna on both hard and soft substrates. At least 17 classes were present by the Middle Ordovician, the all-time high point for echinoderm class diversity, and nearly all of the major ways-of-life (except for deep infaunal burrowing) had been developed. With the rise to dominance of crinoids, many less successful or archaic groups did not survive the Middle Ordovician, and echinoderm class diversity dropped further because of the mass extinction at the end of the Ordovician. This weeding-out process of other less-successful echinoderm groups continued throughout the rest of the Paleozoic, and only five classes of echinoderms have survived to the Recent from this early Paleozoic radiation.

Stylophora and Homoiostelea are the largest classes of the subphylum Homalozoa. They have also been placed in the Calcichordata but that position is herein rejected. Stylophorans are divided into two orders the Cornuta and Ankyroida: cornutes have asymmetrical thecae, aulacophores with stylocones and cover plates over the food groove that open widely; ankyroids have essentially bilaterally symmetrical thecae, aulacophores with styloids and in most the cover plates do not open widely. Epispires, cothurnopores, and lamellipores in cornutes are respiratory structures not atypical of early echinoderms and are only superficially similar to chordate gill slits. The superior and inferior faces of cornute and ankyroid thecae and the aulacophores are homologous. There is no evidence that ‘mitrates’ (most ankyroids) are inverted or their aulacophores(calcichordate tail) have been lost and re-evolved.

Homoiosteles are superficially similar to stylophorans: the column or stele resembles the aulacophore and the theca in younger genera develope distinct marginal and somatic plate patterns. The earliest homoiosteles are attached by a holdfast, at least in juvenile stages, and this fixation may have imprinted some morphological features on steles of vagile genera. Earliest homoiosteles share significant characters with coeval species of the eocrinoid Gogia and it serves as outgroup.

Cladograms for Stylophora and Homoiostelea were generated by NONA, a phylogenetic program for personal computers.

It is now possible to reconstruct what happened when the chordates and echinoderms first split from each other. This involves a three-way comparison among: 1) the solute Coleicarpus, which is probably a stem-group dexiothete; 2) the Cincta, which seem to be the least crownward known echinoderms; and 3) the solute Castericystis, which is a stem-group chordate, probably the least crownward known. Counter-torsion of the tail, by which the effects of dexiothetism were nullified in the tail, took place in two phases, firstly in the fore tail and later in the hind tail. Echinoderms and chordates are descended from ancestors that were attached to, or lay on, the sea floor and were therefore much more liable to attack from above than beneath. This probably explains why the main nerve trunk in chordates is dorsal, rather than being ventral as in protostomes.

Morphological data can be used effectively in phylogenetic analyses to determine relationships among echinoderm clades. These data in the form of characters are simply hypotheses that any observed morphological state among taxa results from a single character state transformation and is therefore, homologous. All such character states must be scored as potentially homologous unless the hypothesis of homology can be rejected by the tests of similarity, conjunction (a priori), or character congruence (a posteriori). Fossils are not always more incomplete than extant forms and incompleteness originates from non-preservation and long phylogenetic branches. The greatest strength of fossil data lies in its ability to effectively shorten long phylogenetic branches by occurring on the tree nearer to the nodes than extant terminal taxa and thus circumventing positively misleading results encountered in parsimony analysis under long branch conditions.

New, competing ideas on crinoid plate circlet homologies and the desire for a phylogenetically-based classification have led to a reexamination of the Crinoidea. The most primitive crinoids had four plate circlets, from bottom to top: lintels, infrabasals, basals, and radials. Dicyclic camerates, cladids, flexibles, and dicyclic articulates are composed of infrabasals, basals, and radials. Monocyclic camerates and monocyclic articulates are composed of basals and radials. These all follow traditional ideas on homologies. In contrast, disparids with “compound radials” are composed of lintels, infrabasals, and radials, rather than “basals,” “inferradials,” and “superradials;” and disparids without “compound radials” have lintels and infrabasals rather than “basals” and “radials.”

Parsimony-based phylogenetic interpretation of Ordovician crinoids with rhombiferans as the outgroup shows that crinoids are divisible into six clades, two paraphyletic and four monophyletic, including four-circlet crinoids, cladids, disparids, camerates, flexibles, and articulates.

The impressive array of disparity within the Echinodermata can be explained by the interplay of components (particularly skeletal elements) making up two major body wall regions: axial and extraxial. Axial skeleton comprises paired plate columns of the ambulacra, formed according to the Ocular Plate Rule (OPR) and in association with the water vascular system. Extraxial skeleton (subdivided into two subtypes: perforate and imperforate) is not formed according to the OPR, and new elements can be added anywhere and at any time within extraxial body wall. Recent work on early development of echinoderms reveals that axial skeleton is formed as an integral part of the rudiment, but that extraxial skeleton is derived from the non-rudiment part of the larval body. In addition to displaying such fundamental embryological and ontogenetic differences, the body wall regions have distinctive distributions and topologies that can be used to formulate criteria for their identification in any echinoderm regardless of how esoteric their morphology might be. Like the system of homologies that has long been established for vertebrates, the model of axial and extraxial skeletal types can be used to explore relationships among Recent and fossil taxa alike. Application of the model also leads to reassessment of previously published morphological characters and phylogenies.

Single data sets, whether derived from morphological or molecular evidence, provide one-off estimates of the correct phylogeny. Their reliability can only be gauged by statistical approaches such as bootstrapping or clade decay, but these test only whether there are sufficient characters in the data matrix to justify the groupings identified. They do not test whether the characters themselves are reliable. Consequently, confidence in the correctness of phylogenetic interpretations comes primarily from discovering the same (or statistically indistinguishable) patterns from independent data sets.

Congruence studies are most advanced for echinoids, where four independent data sets (two morphological and two molecular) provide strong corroboration for a single phylogenetic scheme. Analysis of all four data sets combined generates a highly robust hypothesis of relationships. The situation is very different for asteroids. Two analyses based on morphological data have reached very different conclusions. Three independent molecular data sets also have been compiled, but none has a statistically reliable signal concerning higher taxon relationships. Even combining all three molecular data sets fails to generate a statistically robust solution, implying that the major lines of asteroids diverged rapidly from one another. For ophiuroids, both morphological and molecular data generate topologies that for the most part lack statistical robustness. There is currently no cladistic analysis of holothurian relationships based on morphological data, and only a few taxa have been sequenced. The molecular data is, however, congruent and does permit an initial assessment of relationships. Nothing definite can be deduced about crinoid relationships since even fewer molecular sequences are known and morphological analysis remains sketchy.

Class-level relationships derived from two morphological and two molecular data sets also show considerable congruence, though a single definitive solution has yet to emerge.