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The mineralized skeletons of echinoderms are characterized by their complex, open-cell porous microstructure (also known as stereom), which exhibits vast variations in pore sizes, branch morphology, and three-dimensional (3D) organization patterns among different species. Quantitative description and analysis of these cellular structures in 3D are needed in order to understand their mechanical properties and underlying design strategies. In this paper series, we present a framework for analyzing such structures based on high-resolution 3D tomography data and utilize this framework to investigate the structural designs of stereom by using the spines from the sea urchin Heterocentrotus mamillatus as a model system. The first paper here reports the proposed cellular network analysis framework, which consists of five major steps: synchrotron-based tomography and hierarchical convolutional neural network-based reconstruction, machine learning-based segmentation, cellular network registration, feature extraction, and data representation and analysis. This framework enables the characterization of the porous stereom structures at the individual node and branch level (~10 µm), the local cellular level (~100 µm), and the global network level (~1 mm). We define and quantify multiple structural descriptors at each level, such as node connectivity, branch length and orientation, branch profile, ring structure, etc., which allows us to investigate the cellular network construction of H. mamillatus spines quantitatively. The methodology reported here could be tailored to analyze other natural or engineering open-cell porous materials for a comprehensive multiscale network representation and mechanical analysis. Statement of Significance The mechanical robustness of the biomineralized porous structures in sea urchin spines has long been recognized. However, quantitative cellular network representation and analysis of this class of natural cellular solids are still limited in the literature. This constrains our capability to fully understand the mechanical properties and design strategies in sea urchin spines and other similar echinoderms’ porous skeletal structures. Combining high-resolution tomography and computer vision-based analysis, this work presents a multiscale 3D network analysis framework, which allows for extraction, registration, and quantification of sea urchin spines’ complex porous structure from the individual branch and node level to the global network level. This 3D structural analysis is relevant to a diversity of research fields, such as biomineralization, skeletal biology, biomimetics, material science, etc.

Living crinoids are exclusively passive suspension feeders and benthic as adults. However, in the past they adapted to a broad range of ecological niches. For instance, the stratigraphically important middle Paleozoic scyphocrinoids are hypothesized to have been planktonic, employing their inferred gas-filled globular, chambered structure at the distal end of the stem, the so-called lobolith, as a buoyancy device with the crinoid calyx suspended below it. Here, we evaluate this hypothesis using evidence from skeletal micromorphology and theoretical biomechanical modeling. Lobolith walls are typically composed of ossicles, which are exclusively composed of constructional labyrinthic stereom. In plates from the distal side of the lobolith, this stereom extends into microperforate stereom layer, forming wavy ridges and spines. No microscale adaptations for preventing gas leaks and/or ingress of water (such as internal and external imperforate stereom layers) are known. Furthermore, theoretical calculations suggest that the scyphocrinoid tow-net mode of feeding would have resulted in small relative velocities between the towed filter and the ambient water, thus making it an ineffective passive filter feeder. We suggest that the lobolith of these crinoids acted as a modified holdfast rather than as a floating buoy. Its globular shape and distally positioned microspines served as adaptations for living in unconsolidated sediments, analogous to iceberg- and snowshoe-like strategies used by some mollusks and brachiopods. Like modern isocrinids, scyphocrinoids could have maintained an upright feeding posture by extending the distal portion of the stalk along the bottom. In this recumbent posture, the distal part of the stalk with the lobolith might have functioned as a drag anchor. As a consequence of the ~3-m-long stem, even with this posture, the benthic scyphocrinoids could have risen to the highest epifaunal tier in the Paleozoic.

Three incomplete thecae of the stylophoran Lagynocystis cf. pyramidalis (Barrande, 1887) are described from the Upper Ordovician of the Baltic Paleobasin for the first time. These specimens of Lagynocystis are the first records of this genus from the Ordovician outside peri-Gondwana, thus significantly expanding its paleogeographical distribution of this genus. Due to the preservation of the specimens the microstructure of the stereom is studied for the first time in this genus.

A hardground surface in the middle Silurian (Wenlock: Sheinwoodian) Massie Formation is well exposed at the New Point stone quarry in Napoleon, southeastern Indiana, USA, where it is densely encrusted by pelmatozoan echinoderm attachment structures. Among the most common attachment structures encrusting the hardground are terminal dendritic radix structures attributable to the monobathrid camerate crinoid Eucalyptocrinites and the hemicosmitid rhombiferan Caryocrinites. Radicles belonging to both of these genera are often modified by secretion of secondary stereom, resulting in swollen, irregular calcitic masses that extend onto the substrate and can engulf nearby bioclasts. Herein, we describe a specimen that resembles a portion of a swollen radicular attachment structure; however, close inspection reveals that is actually an interaction between radicles of Caryocrinites and Eucalyptocrinites, as indicated by the presence of both a trilobate lumen, characteristic of Caryocrinites, as well as a pentalobate lumen, characteristic of Eucalyptocrinites. Hence, what appears to be part of a single structure is actually parts of two interwoven structures belonging to entirely different taxa. Increased attention to echinoderm attachment structures consisting of overgrowths and secondarily mineralized radicles may result in recognition of additional specimens that represent analogous biotic interactions rather than simple holdfast occurrences.

Introduction: The sea star odontophore is the structure positioned between the paired oral ossicles, with which they articulate through proximal and distal processes. The internal anatomy structures may be used as taxonomic characters for a precise differentiation between species, so it is necessary to describe the structures variation throughout growth. Objective: To describe the odontophore shape and variation of Luidia superba A. H. Clark, 1917 from specimens of the Gulf of California deposited in the Echinoderm National Collection, ICML UNAM. Methods: A total of 735 specimens were examined to describe the external characters, from which 55 selected specimens, within a range of R = 14 mm and R = 210 mm, were dissected to extract the odontophores and analyzed with geometric morphometrics. Results: Scanning electronic microscopy (SEM) images of the odontophores showing the variations in shape throughout growth are presented. Differences in shape between size groups were confirmed with a Canonical Variables Analysis (P < 0.05). Conclusions: There are three main groups in this size ranges where specialization of the stereom can be observed through the ontogenetic series; the variation in shape of the odontophore shown here is a precedent for the use of internal anatomy as new taxonomic characters of identification. Resumen Introducción: El odontóforo es la estructura posicionada entre las placas orales pareadas con las que se articula a través de procesos proximales y distales. Las estructuras de la anatomía interna se pueden usar como caracteres taxonómicos para la diferenciación más precisa entre especies, por lo que es necesario describir la variación de las estructuras a lo largo del crecimiento. Objetivo: Describir la forma y la variación del odontóforo de Luidia superba A. H. Clark, 1917 de ejemplares del Golfo de California depositados en la Colección Nacional de Equinodermos, ICML UNAM. Métodos: Se revisaron un total de 735 ejemplares para describir los caracteres externos y de las cuales se seleccionaron 55 ejemplares, dentro de un intervalo de R = 14 mm a R = 210 mm, de los cuales se extrajeron los odontóforos y fueron analizados utilizando morfometría geométrica. Resultados: Se presentan imágenes de microscopía electrónica de barrido (MEB) de los odontóforos que muestran las variaciones de la forma durante el crecimiento. Se confirmaron diferencias significativas de la forma entre los grupos de tallas mediante un CVA (p< 0.05). Conclusiones: Se observa especialización del estereoma a lo largo de la serie ontogenética; la variación en la forma del odontóforo aquí mostrada es precedente para el uso de estructuras de la anatomía interna como nuevos caracteres de identificación.

Examination of nearly 500 specimens of Mitrocystites mitra, and Mitrocystella incipiens from the Lower Ordovician of Bohemia housed in the Barrande and Hanus collections of the National Museum, Prague, Czech Republic and new material from the Horǎk Museum, Rokycany, has yielded a wealth of new morphological data. Peripores and lateripores in M.mitra are surrounded by fine-mesh stereom clearly indicating they are sutural pores in origin. Internal tubular structures associated with these pores are probably gonadal in function; not eyes (pores) and nervous system as viewed in Jefferies’ ‘concept Calcichordata’. Both M.mitra and M. incipiens have a unique fully plated cloaca associate with the digit-like anal valvular plates. Mitrocystella incipiens, in some specimens, retains vestigial lateripores and the newly discovered aulacophore is longer than the theca. Within the Mitrocystitidae we can observe the possession and loss of basic echinoderm structures i.e. external hydropore and sutural pores. In addition, stereom construction of the plates, sutural pore systems, and well developed anal valves are thecal characters shared with the Blastozoa.

It has been argued that skeletal Mg/Ca ratio in echinoderms is mostly governed by Mg2+ and Ca2+ concentrations in the ambient seawater. Accordingly, well-preserved fossil echinoderms were used to reconstruct Phanerozoic seawater Mg2+ /Ca2+ ratio. However, Mg/Ca ratio in echinoderm skeleton can be affected by a number of environmental and physiological factors, the effects of which are still poorly understood. Notably, experimental data supporting the applicability of echinoderms in paleoenvironmental reconstructions remain limited. Here, we investigated the effect of ambient Mg2+ /Ca2+ seawater ratio and diet on skeletal Mg/Ca ratio and growth rate in two echinoid species (Psammechinus miliaris and Prionocidaris baculosa). Sea urchins were tagged with manganese and then cultured in different Mg2+ /Ca2+ conditions to simulate fluctuations in the Mg2+ /Ca2+ seawater ratios in the Phanerozoic. Simultaneously, they were fed on a diet containing different amounts of magnesium. Our results show that the skeletal Mg/Ca ratio in both species varied not only between ossicle types but also between different types of stereom within a single ossicle. Importantly, the skeletal Mg/Ca ratio in both species decreased proportionally with decreasing seawater Mg2+ /Ca2+ ratio. However, sea urchins feeding on Mg-enriched diet produced a skeleton with a higher Mg/Ca ratio. We also found that although incubation in lower ambient Mg2+ /Ca2+ ratio did not affect echinoid respiration rates, it led to a decrease or inhibition of their growth. Overall, these results demonstrate that although skeletal Mg/Ca ratios in echinoderms can be largely determined by seawater chemistry, the type of diet may also influence skeletal geochemistry, which imposes constraints on the application of fossil echinoderms as a reliable proxy. The accuracy of paleoseawater Mg2+ /Ca2+ calculations is further limited by the fact that Mg partition coefficients vary significantly at different scales (between species, specimens feeding on different types of food, different ossicle types, and stereom types within a single ossicle).

Simple Summary Arbacia lixula, Paracentrotus lividus and Sphaerechinus granularis play a key role in many sublittoral biocommunities of the Mediterranean Sea. However, their skeletons seem to differ, both morphologically and in chemical composition. Thus, the skeletal elements display different properties, which are affected not only by the environment, but also by the vital effect of each species. We studied the microstructural morphology and crystalline phase of the test and spines, while also examining the effect of time on their elemental composition. Results showed morphologic differences among the three species both in spines and tests. They also seem to respond differently to possible time-related changes. The mineralogical composition of P. lividus appears to be quite different compare to the other two species. The results of the present study may contribute to a better understanding of the skeletal properties of these species, this being especially useful in predicting the effects of ocean acidification. More specifically, since the skeleton plays a key role for the survival of sea urchins in general, a potential change in any skeletal structure, either morphologically or chemically, may affect these organisms directly while also affecting their ecosystem indirectly. Abstract In the Mediterranean Sea, the species Arbacia lixula, Paracentrotus lividus and Sphaerechinus granularis often coexist, occupying different subareas of the same habitat. The mechanical and chemical properties of their calcitic skeletons are affected both by their microstructural morphology and chemical composition. The present study describes the main morphologic features and the possible temporal differences in elemental composition of the test and spines of the three species, while also determining the molar ratio of each element of their crystalline phase. Scanning electron microscopy showed major differences in the ultrastructure of the spines, while minor differences in the test were also noticed. More specifically, the spines of all three sea urchins possess wedges, however A. lixula exhibits bridges connecting each wedge, while barbs are observed in the wedges of S. granularis. The spines of P. lividus are devoid of both microstructures. Secondary tubercles are absent in the test of A. lixula, while the tests and spines of all three species are characterized by different superficial stereom. Energy dispersive x-ray spectroscopy detected that Ca, Mg, S, Na and Cl were present in all specimen. Mg and Mg/Ca showed significant differences between species both in test and spines with S. granularis having the highest concentration. The spines of P. lividus exhibited lowest values between all species. Differences between spines and test were observed in all elements for P. lividus except S. A. lixula exhibited different concentrations between test and spines for Ca, Mg and Mg/Ca, whereas S. granularis for Mg, Cl and Mg/Ca. Finally, temporal differences for Ca were observed in the test of P. lividus and the spines of S. granularis, for Mg in test of S. granularis, for S in the spines of A. lixula and the test and spine of S. granularis, for Na in the test of P. lividus and A. lixula and for Cl and Mg/Ca in the test P. lividus. Powder X-ray diffractometry determined that, out of all three species, the spines of P. lividus contained the least Mg, while the test of the same species exhibited higher Mg concentration compared to A. lixula and S. granularis. The current study, although not labeling the specimens attempts to estimate potential time-related elemental differences among other results. These may occur due to changes in abiotic factors, probably water temperature, salinity and/or pH. Divergence in food preference and food availability may also play a key role in possible temporal differences the skeletons of these species

Spines of the slate pencil sea urchin Heterocentrotus mamillatus Linnaeus, 1758, are in focus of biomimetic research as they feature a “graceful” failure behaviour under uniaxial compression dissipating energy and resisting high loads even after high strain. This study elucidates and quantifies the organization of calcitic trabeculae and pores in large primary spines of the slate pencil urchin H. mamillatus by image analysis from scanning electron microscopy, X-ray micro-computed tomography (µCT) and gravimetry. This study delivers a detailed distribution of porosities within the whole spine and shows that parts of the spines have a much higher porosity then hitherto thought. The central part (medulla) of the high-magnesium calcitic stereom of H. mamillatus spines has a porosity range of 75% to nearly 90%. From this innermost structure, more than 200 radially aligned, but often sinuous trabeculae extend to the spine rim. The structure of this complicated meshwork (radiating layer) is best seen in basal cross sections and was confirmed by µCT scans. The radiating layer has a porosity range from 40–70% and is irregularly separated by the dense growth layers (15–35% porosity). Growth layers were classified in proximal and distal growth layers with numbers ranging within a single animal between 3–14 and 2–7, respectively. These growth layers are characteristic for H. mamillatus spines and play a major role in their remarkable mechanical properties. The porosity of the spine increases from base to tip. Biological and mechanical implications of the variations are discussed.

Uintacrinoids (Uintacrinoidea) are among the most bizarre of all the crinoids. These stratigraphically important Late Cretaceous crinoids are characterized by an atypical large stalkless globular cup without cirriferous centrodorsal and extremely long arms. Owing to this unusual morphology, the mode of life of these crinoids has confused palaeontologists for more than a century. While uintacrinoids have been commonly regarded as pelagic forms, either swimming actively or floating passively by means of a putative buoyancy mechanism, some recent authors have favoured a benthic mode of life. However, evidence supporting a benthic lifestyle hypothesis is still limited. Here we describe the stereom microstructure of Marsupites and Uintacrinus. In Marsupites, thecal plates are typically comprised of coarse to medium, clearly structural, galleried stereom, which alternates with zones of fine and deeply penetrating galleried stereom oriented perpendicular to the plate margins. The latter stereom type corresponds to the external ribbed ornament of the theca, and is indicative of collagenous sutural fibers for binding adjacent plates. In Uintacrinus, thecal plates are almost entirely composed of fine galleried stereom diagnostic for collagenous sutural fibers. In both taxa, a perforate stereom layer, that is known to increase plate strengthening and resistance to abrasion in some Recent echinoderms, is developed most externally, and also produces a characteristic interior growth banding pattern. No skeletal adaptations for maintaining buoyancy, such as skeletal lightening and hollow architecture, that are known to occur in nektonic and pseudoplanktonic crinoids, are observed. Instead, the stereom organization in uintacrinoids closely resembles those observed in Recent benthic crinoids. Our findings thus strongly support a benthic mode of life of uintacrinoids. Their widespread distribution within a short time interval can be best explained by having a very long-term planktonic larval stage as previously hypothesized by some authors.

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 FloianAthenacrinus browerinew genus new species. Character analysis supports a hypothesis that this taxon originated early within in the disparid clade.Athenacrinusn. 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.UUID:http://zoobank.org/b383e039-3298-4472-a7e3-e81684f87cfe

10 pages; International audience; The extinct echinoderm clade Stylophora consists of some of the strangest known deuterostomes. Stylophorans are known from complete, fully articulated skeletal remains from the middle Cambrian to the Pennsylvanian, but remain difficult to interpret. Their bizarre morphology, with a single appendage extending from a main body, has spawned vigorous debate over the phylogenetic significance of stylophorans, which were long considered modified but bona fide echinoderms with a feeding appendage. More recent interpretation of this appendage as a posterior “tail-like” structure has literally turned the animal back to front, leading to consideration of stylophorans as ancestral chordates, or as hemichordate-like, early echinoderms. Until now, the data feeding the debate have been restricted to evaluations of skeletal anatomy. Here, we apply novel elemental mapping technologies to describe, for the first time, soft tissue traces in stylophorans in conjunction with skeletal molds. The single stylophoran appendage contains a longitudinal canal with perpendicular, elongate extensions projecting beyond hinged biserial plates. This pattern of soft tissues compares most favorably with the hydrocoel, including a water vascular canal and tube feet found in all typical echinoderms. Presence of both calcite stereom and now, an apparent water vascular system, supports echinoderm and not hemichordate-like affinities.

The skeleton of spines and tests of the species of sea urchins Strongylocentrotus intermedius, Mesocentrotus nudus, Scaphechinus mirabilis, and Echinocardium cordatum from the Sea of Japan is composed of a spongy stereom, consisting of calcite with a high content of magnesium. It was found that the tests and spines of the skeletons of sea urchins are composed of calcium–organic composite materials inlaid with other metals: Mg, Fe, Zn, and Rb. In the four species of sea urchins studied, the strength and other mechanical properties of the tests and spines differ and depend on the chemical composition and structural organization of their components. It was shown that the content of volatile substances correlates with their fragility or elasticity. It is revealed that the chemical composition of the tests of two species of the spherical sea urchins S. intermedius and M. nudus indicates significant differences between these two species of sea urchins.

One of the most profound environmental changes thought to be reflected in chemical composition of numerous geological archives is Mg/Ca ratio of the seawater, which has varied dramatically throughout the Phanerozoic. Echinoderms that today typically form high magnesium calcite skeletons are increasingly being utilized as a proxy for interpreting secular changes in seawater chemistry. However, accurate characterization of the diagenetic changes of their metastable high magnesium calcite skeletons is a prerequisite for assessing their original, major-element geochemical composition. Here we expand the existing models of diagenesis of echinoderm skeleton by integration of various analytical methods that up to now rarely have been used to assess the diagenetic changes of fossil echinoderms. We validated the preservation of a suite of differently preserved echinoderm ossicles, mostly crinoids, ranging in age from the Cambrian through Recent. In 13 of 99 fossil echinoderm ossicles we found well-preserved porous microstructure (stereom), non-luminescent behaviour or blotchy dark color in cathodoluminescence, and distinct nanostructural features (layered and nanocomposite structure). Moreover, in representatives of such preserved samples, distribution of sulphates associated with organic matter is identical to those in Recent echinoderms. Only such ossicles, despite of local micrometer-scale diagenetic changes, were herein considered well-preserved, retaining their original major-element skeletal composition. By contrast, majority of samples show transformation to the stable low magnesium calcite that leads to obliteration of the primary geochemical and micro/nanostructural features and is accompanied with increase in cathodoluminescence emission intensity. Using only well-preserved fossil echinoderm samples, we found purely random variation in Mg/Ca in echinoderm skeletons through the observed time series; any periodicities in echinoderm skeletal Mg/Ca ratio which might be related to the secular transitions in calcite and aragonite seas were not confirmed. These findings suggest that, in contrast to some groups of organisms with relatively weak control over their biomineralization (such as algae, sponges, and bryozoans), in which polymorph mineralogies consistently changed according to the seawater type, application of fossil echinoderms, in particular crinoids, to seawater Mg/Ca reconstructions is unreliable. These data emphasize a key-role of physiological factors (the so-called vital effects) in echinoderm biomineralization.

Exceptionally well-preserved ossicles of xenomorphic stalked crinoids (Echinodermata) were found into the Late Ypresian clay of the Tuilerie de Gan (Pyrenees-Atlantiques, southwestern France). Three kinds of columnal articulation correspond to three different genera, including the new genus Eocenocrinus. These columnals are attributed to Paraconocrinus romanensis, Democrinus londinensis and Eocenocrinus hessi n. gen., n. sp. Associated benthic fauna as well as columnal characters related to water depth confirm an increasing depth to an epibathyal environment as previously suggested by mollusk and foraminifer associations. Paraconocrinus romanensis associated with Democrinus londinensis mark the maximum water depth, which did not exceed 300 m. Distal columnal series with proximal roots of P. romanensis and D. londinensis were collected with their ossicles connected, indicating that these species lived on a muddy bottom. E. hessi n. gen., n. sp. is suspected to have been attached on hard substrate. Without diagenetic crystallizing, the crinoid stereom and its micro-biocorrosion appear as in extant specimens. We attribute to Eocenocrinus n. gen. large columnals previously described from northeastern Italy and recognize two species: E. bayani n. gen., n. sp. (Middle Lutetian) and E. didymus (Priabonian). E. hessi n. gen., n. sp. may represent the oldest known species in the family Phrynocrinidae.

Recent studies have shown that in some light-sensitive species of an brittlestar (Asteroidea, Echinodermata), the upper surface of the dorsal arm plate bears arrays of hemispherical microstructures which in combination with underlying neural bundles and intraskeletal chromatophores probably function as a compound eye. These calcitic lenses possess superior properties such as light weight, mechanical strength, and very low aberration and birefringence; they display a unique focusing effect, signal enhancement, intensity adjustment, angular selectivity, and photochromic activity. The discovery of these unique optical structures revealed that brittlestar visual system is more sophisticated than initially thought and has inspired active interest toward designing of biomimetic highly tunable optical elements for a wide variety of cutting-edge technological applications. Up to this moment, analogous spherical calcitic lenses have been only reported in a few species of modern brittlestars and starfish. Similar calcitic microlenses have been also observed in the Late Cretaceous fossil echinoderms. Here, we report the structural evidence for the presence of calcitic microlenses in an extant species of starfish Archaster typicus. The close resemblance in microstructure and location between the transparent regions of compact stereom described above and microlenses in the photosensitive brittlestar Ophiocoma wendtii suggests that these regions may be involved with the photoreceptor system in A. typicus.

Background: Fluorochrome staining is among the most widely used techniques to study growth dynamics of echinoderms. However, it fails to detect fine-scale increments because produced marks are commonly diffusely distributed within the skeleton. In this paper we investigated the potential of trace element (manganese) labeling and subsequent cathodoluminescence (CL) imaging in fine-scale growth studies of echinoderms. Results: Three species of sea urchins (Paracentrotus lividus, Echinometra sp. and Prionocidaris baculosa) were incubated for different periods of time in seawater enriched in different Mn2+ concentrations (1mg/L; 3mg/L; 61.6mg/L). Labeling with low Mn2+ concentrations (at 1mg/L and 3mg/L) had no effect on behavior, growth and survival of sea urchins in contrast to the high Mn2+ dosage (at 61.6mg/L) that resulted in lack of skeleton growth. Under CL, manganese produced clearly visible luminescent growth fronts in these specimens (observed in sectioned skeletal parts), which allowed for a determination of the average extension rates and provided direct insights into the morphogenesis of different types of ossicles. The three species tend to follow the same patterns of growth. Spine growth starts with the formation of microspines which are simultaneously becoming reinforced by addition of thickening layers. Spine septa develop via deposition of porous stereom that is rapidly (within less than 2days) filled by secondary calcite. Development of the inner cortex in cidaroids begins with the formation of microspines which grow at ~3.5μm/day. Later on, deposition of the outer polycrystalline cortex with spinules and protuberances proceeds at ~12μm/day. The growth of tooth can be rapid (up to ~1.8mm/day) and starts with the formation of primary plates (pp) in plumula. Later on, during the further growth of pp in aboral and lateral directions, secondary extensions develop inside (in chronological order: lamellae, needles, secondary plate, prisms and carinar processes), which are increasingly being solidified towards the incisal end. Interradial growth in the ambital interambulacral test plates exceeds meridional growth and inner thickening. Conclusions: Mn2+ labeling coupled with CL imaging is a promising, low-cost and easily applicable method to study growth dynamics of echinoderms at the micro-length scale. The method allowed us to evaluate and refine models of echinoid skeleton morphogenesis., SCOPUS: ar.j, info:eu-repo/semantics/published

A new species of Apiocrinites is described from the Matmor Formation (Middle Jurassic, upper Callovian) of Hamakhtesh Hagadol, southern Israel. Apiocrinites feldmani n. sp. is a small species associated with the larger A. negevensis in a calcareous sponge and coral patch reef community. During life the columns of A. feldmani were commonly and preferentially infested with a soft-bodied parasite that grew with the crinoid and became embedded in its skeleton. These parasites embedded at the articulation between columnals, forcing the columnals to grow around them and producing with time a conical pit surrounded by swollen stereom. If the parasite died while the crinoid was still growing, the conical pit was roofed over by continued growth of columnals, resulting in a swelling with no external opening. Because the crinoids invested energy in forming extra skeleton around these parasites and because the crinoid stems were consequently deformed and likely lost flexibility, we consider these parasites to have caused significant harm. Curiously, these parasites apparently did not infect the larger and more common contemporaneous A. negevensis that lived in the same community.

The material for the present paper is based on 50 kg of processed sediment from the Early Kimmeridgian of Geisingen. The most frequent crinoids comprise cups and brachials of the roveacrinid Saccocoma quenstedti while Crassicoma feifeli is represented by a few cups only; these remains are not described. Cups and primibrachials of Eugeniacrinites cariophilites are the commonest cyrtocrinids, while cups of Pilocrinus moussoni are rather rare and not further treated, as are columnals of the two species. The ontogenetic series of cups and axillary primibrachials of E. cariophilites starts from ossicles of about 0.5 mm in height; and growth is allometric in both types. Postlarval cups resemble phyllocrinids; the distinctive shape with lanceolate interradial extensions and wide articular facets appears at a later stage. The spearhead-shaped axillary primibrachials are synapomorphic. The youngest of these ossicles with a muscular facet proximally were able to form a flat roof over the radial cavity. First primibrachials with synostosial distal facet appeared at a slightly later stage with a width of 1 mm; growth is isometric. A significant number of cups are mutilated by bite marks. The predatory attacks were sublethal in most cases as shown by secondary stereom growth and regenerated facets. First primibrachials are not deformed, but deformations occur in a small number of second primibrachials. The results suggest that the predators, presumably fishes, targeted the soft parts in the cup. Enigmatic brachials with a large wing to one side and a facet to an arm are thought to belong to E. cariophilites and presumably attached to deformed cups; the wings may have been an antipredatory adaptation providing some shelter to the cup. The various antipredator morphologies in cyrtocrinid history are discussed. The presently available growth series of Eugeniacrinites invites discussion of early cyrtocrinid phylogeny. Eugeniacrinites seems to have been derived from Phyllocrinidae; and Sclerocrinidae with Cyrtocrinus as the first representative in the Middle Jurassic is thought to have originated from Early Jurassic Tetracrinidae such as Bilecicrinus.

Cidaroid sea urchins are the sister clade to all other extant echinoids and have numerous unique features, including unusual primary spines. These lack an epidermis when mature, exposing their high-magnesium calcite skeleton to seawater and allowing the settlement of numerous epibionts. Cidaroid spines are made of an inner core of classical monocrystalline skeleton and an outer layer of polycrystalline magnesium calcite. Interestingly, cidaroids survived the Permian-Triassic crisis, which was characterized by severe acidification of the ocean. Currently, numerous members of this group inhabit the deep ocean, below the saturation horizon for their magnesium calcite skeleton. This suggests that members of this taxon may have characteristics that may allow them to resist ongoing ocean acidification linked to global change. We compared the effect of acidified seawater (pH 7.2, 7.6, or 8.2) on mature spines with a fully developed cortex to that on young, growing spines, in which only the stereom core was developed. The cortex of mature spines was much more resistant to etching than the stereom of young spines. We then examined the properties of the cortex that might be responsible for its resistance compared to the underlying stereomic layers, namely morphology, intramineral organic material, magnesium concentration, intrinsic solubility of the mineral, and density. Our results indicate that the acidification resistance of the cortex is probably due to its lower magnesium concentration and higher density, the latter reducing the amount of surface area in contact with acidified seawater. The biofilm and epibionts covering the cortex of mature spines may also reduce its exposure to seawater. © 2014, The American Microscopical Society, Inc.

Growth dynamics of the primary spine of the cidaroid sea urchin Phyllacanthus imperialis was assessed for the first time using pulsed (26) Mg-labeling and NanoSIMS isotopic imaging. The sea urchin was incubated twice (for 48 h) in artificial seawater with elevated level of (26) Mg. After each labeling event, the sea urchin was returned for 72 h to seawater with natural isotopic abundance of (26) Mg. NanoSIMS ion microprobe was subsequently used to visualize the labeled regions of the spine with submicrometer lateral resolution. The growth of the new skeleton was restricted to the distalmost and peripheral portions of the spine. Skeletogenesis involved mostly the deposition of continuous thickening layers and lateral growth involving bridges between previously formed trabeculae. The timescale of formation of individual thickening layers (ca. 1 µm in width) on the stereom trabeculae was on the order of 1 day. Longitudinal growth occurred mainly at the periphery in the form of small portions of the thickening deposits or more massive microspines that appeared to branch and fuse with those above and below. These microspines were found to grow at about 10 µm/day. These results reveal that the skeletal growth of a juvenile cidaroid spine is complex and highly heterogeneous, with different extension rates depending on the stage of the stereom development and/or direction of the growth fronts. The growth pattern observed here at the submicrometer scale provides direct evidence supporting the earlier suggestions that the lamellar structure of echinoderm stereom is formed by periodic deposition of continuous mineral layers.

Experimental ocean acidification leads to a shift in resource allocation and to an increased [HCO3 −] within the perivisceral coelomic fluid (PCF) in the Baltic green sea urchin Strongylocentrotus droebachiensis. We investigated putative mechanisms of this pH compensation reaction by evaluating epithelial barrier function and the magnitude of skeleton (stereom) dissolution. In addition, we measured ossicle growth and skeletal stability. Ussing chamber measurements revealed that the intestine formed a barrier for HCO3 − and was selective for cation diffusion. In contrast, the peritoneal epithelium was leaky and only formed a barrier for macromolecules. The ossicles of 6 week high CO2-acclimatised sea urchins revealed minor carbonate dissolution, reduced growth but unchanged stability. On the other hand, spines dissolved more severely and were more fragile following acclimatisation to high CO2. Our results indicate that epithelia lining the PCF space contribute to its acid–base regulation. The intestine prevents HCO3 − diffusion and thus buffer leakage. In contrast, the leaky peritoneal epithelium allows buffer generation via carbonate dissolution from the surrounding skeletal ossicles. Long-term extracellular acid–base balance must be mediated by active processes, as sea urchins can maintain relatively high extracellular [HCO3 −]. The intestinal epithelia are good candidate tissues for this active net import of HCO3 − into the PCF. Spines appear to be more vulnerable to ocean acidification which might significantly impact resistance to predation pressure and thus influence fitness of this keystone species.

The external and internal morphologies of cidaroid and camarodont sea urchin primary spines are investigated giving an overview of the internal microstructure and structural properties. The investigated species comprise the cidaroids Eucidaris metularia, Phyllacanthus imperialis, Plococidaris verticillata and Prionocidaris baculosa as well as the camarodont Heterocentrotus mammillatus (Echinodermata: Class Echinoidea), and morphological descriptions are based on scanning electron microscopy and micro-computed tomography. Stereom types and densities are differentiated using pore and trabecular diameter measurements. Structural analysis was performed using three point bending tests resulting in the calculation of force, deflection and stress, strain relationships. All studied species possess primary spines with a medulla consisting of laminar stereom regardless of the age and position of the spine on the tests. Differences in stereom morphology occur in the radiating layer and the surface of the spines. Material densities and stereom types differ with respect to growth lines when present and the radiating layer. The primary spines also show large differences in their outer morphologies ranging from smooth, striated to tuberculate. H. mammillatus spines are shown to bear more stress resistance than those of the cidaroids. Differences in spine morphologies and reaction to stress are interpreted with respect to functional morphological response, to ambient environmental parameters and their strategies between and within evolutionary stages.

Middle Miocene tests of Clypeaster from L’Arrabassada (Tarragona, north-east Spain) show evidence of intense endoskeletozoan colonization, preserved as borings and associated carbonate secretions that allow gastrochaenid bivalves to be identified as the colonizers. Two modes of occurrence have been recognized for these bivalve dwelling cavities; ‘intrastereom clavate borings’ which are restricted to the echinoid stereom, and ‘semi-endoskeletal dwellings’, which penetrate across the test wall and extend as carbonate crypts into the sediment fill of the internal test cavity. Their size, density and position rule out a syn-vivo relationship with the echinoids and demonstrate that colonization was post mortem. Because of the endurance of clypeasteroid skeleton and the pronounced bell-shaped morphology of Clypeaster, the tests of these echinoids provided the most suitable substrates for hard-bottom colonizers on an otherwise sandy seafloor. The scenario described from Tarragona can be extended to other Neogene and Quaternary strata elsewhere; there is ample evidence for the long-term utilization of tests of Clypeaster by gastrochaenid bivalves in shoreface palaeoenvironments.

Pleurocystitid rhombiferans are among the most unusual echinoderms whose mode of life has been long debated. These echinoderms are usually interpreted as vagile epibenthic echinoderms, moving over the sea bottom by means of a flexible stem. Although their life habits and posture are reasonably well understood, the mechanisms that control the movement of stem are highly controversial. Specifically, it is unknown whether the stem flexibility was under the control of muscles or ligamentary mutable collagenous tissues (MCTs). Here, we reconstruct palaeoanatomy of the two Ordovician pleurocystitid rhombiferans (Pleurocystites and Amecystis) based on stereom microstructure.Weshow that the articular facets of columnals in pleurocystitid rhombiferans are composed of fine labyrinthic stereom. Comparison with modern echinoderms suggests that this type of stereom was associated with muscles implying that their stem was a muscular locomotory organ supporting an active mode of life, Department of Biogeology, Institute of Paleobiology, Polish Academy of Sciences, Polonia, Unidad de Zaragoza, Instituto Geológico y Minero de España, España, National Museum of Natural History, Smithsonian Institution, Estados Unidos

The stereom microstructure of columnal latera in various articulate crinoids is examined to ascertain its potential application in phylogenetic studies. Most show either a poorly ordered labyrinthic stereom or a more regularly ordered perforate stereom with lumina arranged in an en echelon pattern. Labyrinthic stereom is taxonomically widespread and is interpreted here as the plesiomorphic state. Perforate stereom, typically with an en echelon arrangement of elliptical lumina, appears confined to the Isocrinina and is considered to be apomorphic for that group. The absence of any development of perforate stereom in the bourgueticrinid Democrinus suggests that the bourgueticrinids have affinities with the comatulids rather than with the isocrinids. A second type of perforate stereom, with sinuous or straight lines of circular to elongate lumina, has been found only in Pentacrinites and may be autapomorphic for that group. The shape and spacing of stereom lumina on the columnal latera of different genera of isocrinids displays minor variation that may have some value for assessing phylogenetic affinity at genus and family level in taxa known only from columnals.

The early Middle Cambrian Monastery Creek Phosphorite (Beetle Creek Formation, Queensland, Australia) contains an assemblage of disarticulated echinoderm ossicles that are exquisitely preserved. Amongst this material we recognize pelmatozoan brachials, radials, basals and holomeric columnals. Although we cannot reconstruct the complete animal with precision, these elements represent the oldest known pelmatozoan with crinoid-like appendages. Key elements include isotomously to heterotomously branched uniserial appendage plates with a tripartite adoral food groove, a longitudinal central canal interpreted as housing entoneural nerve, and differentiated articulation facets. There are also epispire-bearing radials bearing one to four arm insertion-facets, each one pierced by a central neural canal. These canals run internal towards the oral area beneath the external food groove. Co-occuring material includes single truncated cone-shaped basals and holomeric columnals, both with a similar articulation pattern, and irregular, epispire-bearing thecal plates. This mosaic of crinozoan (uniserial isotomous to heterotomous arms with neural canal), blastozoan (epispire-bearing thecal plates, appendage leading to oral thecal food groove without direct connection with body cavity) and apomorphic characters (circumoral instead of basal entoneural plexus) is unexpected and demonstrates that crinoid-like pelmatozoans with uniserial, branched arms appeared significantly earlier than the Tremadocian, when the first articulated crinoid skeletons are found. It also raises questions about the polyphyletic appearance of feeding appendages among pelmatozoan echinoderms.

Calcite isocrinid ossicles from the Middle Jurassic (Bathonian) clays in Gnaszyn (central Poland) show perfectly pre− served micro− and nanostructural details typical of diagenetically unaltered echinoderm skeleton. Stereom pores are filled with ferroan calcite cements that sealed off the skeleton from diagenetic fluids and prevented structural and geochemical alteration. In contrast with high−Mg calcite skeleton of modern, tropical echinoderms, the fossil crinoid ossicles from Gnaszyn contain only 5.0–5.3 mole% of MgCO3. This low Mg content can be a result of either a low temperature environ− ment (ca. 10C) and/or low Mg/Ca seawater ratio. Both conditions have been proposed for the Middle Jurassic marine en− vironment. Occurrence of Mg−enriched central region of stereom bars of Jurassic columnal ossicle of Chariocrinus an− dreae is consistent with the concept of magnesium ions involvement in earliest growth phases of calcium carbonate biominerals. Key wor ds: Echinodermata, Crinoidea, calcite, nanostructure, geochemistry, AFM, NanoSIMS, Jurassic.

Previously undescribed material in the Natural History Museum Basel provides the basis for a reappraisal of the enigmatic Middle Jurassic to Early Cretaceous genus Cyclocrinus D’ORBINGY known only from mostly single columnals and attachment discs. Only one of the previously described Jurassic species, C. rugosus, is recognized despite wide variation in morphology of columnals ranging from Bajocian to Oxfordian in age. A small percentage of columnals are axillary, show facets for side branches or are arched, which has led to the assumption that Cyclocrinus columnals represent radicular cirrals and should be assigned to the Order Bourgueticrinida. Both hypotheses are rejected. The prevalence of single columnals is explained by the restriction of loose, galleried (α) stereom to an area near the facets and around the axial canal; through-going ligament fibres responsible for the preservation of pluricolumnals were thus largely absent from much of the stereom. Cyclocrinus variolarius (SEELEY) from the Albian of England hardly differs from the Jurassic C. rugosus. The ordinal position of Cyclocrinus and of the family Cyclocrinidae SIEVERTS-DORECK is left open though the form has some resemblance to the Early Jurassic millericrinid genus Amaltheocrinus KLIKUSHIN.

Based on rich material (381 specimens examined) from two Cambrian echinoderm faunas, the early Cambrian Balang fauna and middle Cambrian Kaili fauna in Guizhou Province, South China, the taphonomy of gogiid echinoderms is described in detail, and the preservation of stereomic microstructure and organic remains of Cambrian gogiid echinoderms is reported here for the first time. Taphonomic considerations include entombment patterns, decay sequences, individual-specific diagenetic histories, unusual burial postures, selective disarticulation patterns, and post-mortem elongation. In particular, five categories of gogiid entombment patterns are proposed to describe the multi-directional orientations recorded at the burial time of articulated gogiids. Gogiid-bearing slabs of Guizhou material primarily (70%) display the type 2 entombment pattern (articulated gogiids preserved with fan-shaped brachioles); thus, most Guizhou gogiids were buried with brachioles preserved in feeding posture during obrution events. Balang gogiid faunas contain the oldest evidence of palaeoecological interactions among echinoderms and other indigenous taxa. In addition to pre-burial and post-burial decay, other potential causes for unusual disarticulation patterns exhibited by the gogiids from the lower Cambrian Balang Formation include pre-burial bio-disturbance and post-burial bioturbation based on ichnogenera, including Rusophycus and Planolites. Chemical analyses reveal that carbon, calcium, manganese and iron are the major elements responsible for the variety of colours exhibited by Guizhou gogiids. Three-dimensional stereomic microstructure (mean stereom pore size = 8.4–8.7 μm; average trabecular thickness = 4.5–4.6μm) occurs on the external surfaces of thecal plates in two gogiid species. Stereom preservation in calcite suggests that the dissolution of calcareous echinoderm plates, yielding characteristic mouldic preservation, is sub-Recent (after lithificaiton and exposure of gogiid-bearing, marine sedimentary successions on or near the land surface).

Paleogenomics propels the meaning of genomic studies back through hundreds of millions of years of deep time. Now that the genome of the echinoid Strongylocentrotus purpuratus is sequenced, the operation of its genes can be interpreted in light of the well-understood echinoderm fossil record. Characters that first appear in Early Cambrian forms are still characteristic of echinoderms today. Key genes for one of these characters, the biomineralized tissue stereom, can be identified in the S. purpuratus genome and are likely to be the same genes that were involved with stereom formation in the earliest echinoderms some 520 million years ago.

During the geologic history of crinoids, many groups have modified their distal stems to function as anchors in soft sediments, but more so in the Paleozoic than in later times. Major strategies were passively implanting anchors, cirral root systems, coils, and stemless reclining. In Paleozoic roots, the stereom usually consists of crystallographically individualized ossicles, but articulations may be hidden by a tough cuticle that protected the living tissue of immersed parts and stiffened them with cortical stereom. Post-Paleozoic representatives produced similar anchoring structures by different fabricational pathways, including non-articulated root systems.

This paper reports results of a novel approach, X-ray microCT, for quantifying stereom structures applied to ossicles of the sea urchin Lytechinus variegatus. MicroCT, a high resolution variant of medical CT (computed tomography), allows noninvasive mapping of microstructure in 3-D with spatial resolution approaching that of optical microscopy. An intact pyramid (two demipyramids, tooth epiphyses, and one tooth) was reconstructed with 17 microm isotropic voxels (volume elements); two individual demipyramids and a pair of epiphyses were studied with 9-13 microm isotropic voxels. The cross-sectional maps of a linear attenuation coefficient produced by the reconstruction algorithm showed that the structure of the ossicles was quite heterogeneous on the scale of tens to hundreds of micrometers. Variations in magnesium content and in minor elemental constitutents could not account for the observed heterogeneities. Spatial resolution was insufficient to resolve the individual elements of the stereom, but the observed values of the linear attenuation coefficient (for the 26 keV effective X-ray energy, a maximum of 7.4 cm(-1) and a minimum of approximately 2 cm(-1) away from obvious voids) could be interpreted in terms of fractions of voxels occupied by mineral (high magnesium calcite). The average volume fraction of mineral determined for a transverse slice of the demipyramid near where it joins an epiphysis was 0.46; for a slice 3.3 mm adoral it was 0.70. Local volume fractions of mineral approached 1, and, away from resolvable voids, considerable portions of the demipyramids had volume fractions of calcite at or below approximately 0.33. MicroCT imaging of a demipyramid before and after infiltration with a high absorptivity fluid (sodium polytungstate) confirmed the determination of the volume fractions of minerals.

Detailed examination of isolated thecal plates belonging to three discocystinid edrioasteroids, Spiraclavus nacoensis Sumrall, Hypsiclavus kinsleyi Sumrall, and Giganticlavus bennisoni Sumrall and Bowsher, reveals striking similarity in morphology among these species. Stereom observed in the ambulacral floor plates indicates that ligamentous connective tissue opened the ambulacral cover plates and muscle tissue closed them. The ambulacral floor plates are interpreted as rigid supports for the oral surface with the interambulacral areas acting as flexible integuments of plates. The aboral surface is interpreted as flexible and highly contractile. All discocystinid thecal openings are consistent in morphology with adaptations for thecal pressurization. Extension and contraction of the theca was accomplished by pumping water in and out of an inflatable sac associated with the periproctial opening. The pedunculate zone is interpreted as passively expanding and contracting by relaxing of mutable collagenous tissue and stiffening when the theca was in the desired position. All of these features illustrate that discocystinid edrioasteroids have highly–evolved morphology and function.

Echinoderm skeletons are described within a hierarchical framework ranging from complete organisms to the ultrastructural level. They consist of numerous elements which can be isolated, connected by soft tissue or locked together in rigid structures. The top level considers skeletons as a whole with all associated elements and the basic symmetry of the echinoderms. The next level deals with the structural analysis and modeling of echinoids with respect to the growth parameters and stress resistance of the corona. The flexibility and movement has also been studied for the stalks and arms of recent and fossil crinoids. The next level deals with the elaborate morphology and types of symmetry found in single skeletal elements. The numerous types of stereom architectures found within the elements of all echinoderms are highly correlated to specific functions. A high number of recent studies concern the last hierarchical level on ultrastructure and biomineralization. Lightweight aspects of the skeleton are especially present at the level of conjoined plates, single elements and the stereom.

The microstructure of many echinoid species has long fascinated scientists because of its high porosity and outstanding mechanical properties. We have used X-ray microtomography to examine the test of Echinocardium cordatum (heart urchin), a burrowing cousin of the more commonly known sea urchins. Three dimensional imaging demonstrates that the bulk of the test is composed of only two distinct, highly porous, fenestrated regions (stereom), in which the thickness of the struts is constant. Different degrees of porosity are achieved by varying the spacing of the struts. Drawing an analogy to vertebrate trabecular bone, where for example, human bone has a connectivity density of ≈1/mm 3 , we measure up to 150,000 strut connections per mm 3 . Simulations of mechanical loading using finite element calculations indicate that the test performs at very close to the optimum expected for foams, highlighting the functional link between structure and mechanical properties.

New information pertaining to the biostratinomy of Uintacrinus assemblages has been derived from re-examination of museum collections and analysis of in situ material. Individuals are preserved as thin lenses in dense aggregations with articulated calyces and arms only on the lower surface, in contrast to disarticulated material on the upper surface. Calyces may be imbricated within a lens and are mostly compressed laterally. Specimens also may be preserved oral side up or down. Some specimens displaying the oral side retain soft-part preservation of the tegmen, anal tube, and ambulacra. A thin, jet black organic lamination is visible beneath calyx plates. SEM analysis reveals the presence of possible microbial spherules and filaments on this lamination. Dense aggregations reveal a number of new preservational features, including marginal indentations and lacunae that suggest cohesive behavior of the layer prior to burial. SEM also reveals well-preserved stereom in articulated crinoids on the lower surface, in contrast to the upper surface in which the stereom is apparently infilled with calcium carbonate. Some specimens retain a black, organic lamination within the crinoidal layer itself that may represent remnants of a microbial mat. It is proposed that a microbial mat of necrolytic origin provided cohesion and that microbial sealing during decay may help to explain other instances of similar crinoid preservation, including both benthic and pelagic forms, in which articulation is confined to lower surfaces.

The stereom of living Echinus esculentus skeletons is composed of Mg calcite (interambulacral plates 6 mole % MgCO3; spines 4 mole % MgCO3) and is compared with the stereom of Pennsylvanian echinoderms from the Holder Formation, Dry Canyon. Some Dry Canyon echinoderms had their pore system (55% volume) filled by magnesian ferroan calcite cement before the stereom transformed (type 1) into small ( 20 µm diameter in ossicles that underwent type 2 transformation indicates ionic transfer over tens of microns. The margins of some (type 2) ossicles set in a clay matrix are Fe-rich and contain barite crystals; the Fe2+ and Ba2+ were transferred from the surrounding clay hundreds of microns into these ossicles. Type 1 transformation was catalyzed by internal water from the stereom; the presence of a ferroan calcite crystal casket confined alteration to the skeletal Mg calcite. Type 2 transformation involved both stereom and pore-filling Mg calcite cement. The unstable dolomite crystals formed by both transformation processes were protected from further change by the crystal caskets in which they are encased. The transformation of Dry Canyon echinoderms was probably triggered, or accelerated, by rising temperature caused by burial. The bulk composition of stereom that has undergone type 1 transformation can be used as a proxy for the original, major-element chemistry of echinoderm skeletons. Minor amounts of Mg calcite cement that occur in Dry Canon (type 1) ossicles indicate the chemistry of porewater close to the sediment-water interface. Fossil echinoderms are preserved in many ways, and it is suggested that their textural variations can be used to map the distribution of diagenetic reactions. The stereom is destroyed in most fossil echinoderms, and even in those where the stereom shape is preserved its internal structure is changed. Echinoderm Mg calcite does not transform with perfect textural preservation.

Interambularcral plates of echinoid Heterocentrotus trigonarius, composed of Mg calcite 1 (≈14 mol% MgCO3), were heated in three timed series of experiments at 300°C. Dried plate fragments and fragments with added water were heated separately in pressurized bombs. X-ray powder diffractometry, unit cell dimensions, and phase compositions are used to monitor reaction progress. After 10 h heating in the bombs dolomite (43.5 mol% MgCO3) and Mg calcite appear (4–7 mol% MgCO3); by 20 h all Mg calcite 1 is consumed, and at 120 h dolomite composition has evolved to ≈47 mol% MgCO3 and calcite to ≈2 mol% MgCO3. Whole plates heated at 300°C in an open muffle furnace develop dolomite (≈42 mol% MgCO3) and Mg calcite 2 (≈6 mol% MgCO3) after 10 h and remain compositionally invariant throughout subsequent heating to 620 h. Limited skeletal water catalyzes the early reaction but escapes from the open furnace and consequently reaction ceases after ≈10 h. The experimentally produced dolomite has relative Mg-Ca ordering of 75% to 79%. The stabilization of echinoid Mg calcite by heating at 300°C to a mixture of dolomite and calcite occurs through a dissolution/precipitation reaction. The alteration fabric produced within the stereom consists of irregularly shaped, branched dolomite crystals > 5 μm homoaxially set in a calcite 2 (bomb) or Mg calcite 2 (furnace) matrix. Round and tubular pores 1 to 5 μm are randomly distributed throughout this fabric. The stereom pore system remains intact during furnace heating but is destroyed during heating in bombs. The texture of experimentally stabilized echinoid skeletons is different from that of fossil echinoderms that are composed of microrhomic dolomite homoaxially set in a single calcite crystal.

Echmatocrinus from the Middle Cambrian Burgess Shale of British Columbia was originally described as the earliest crinoid(?) known from the fossil record. Recently, Conway Morris and Ausich & Babcock have questioned whether Echmatocrinus is in fact an echinoderm, comparing it instead to cnidarians with a polyp-like body and pinnate tentacles, and other authors are beginning to use this reinterpretation. We studied the well-preserved holotype of Echmatocrinus brachiatus, two paratypes, and 18 new specimens recovered from different levels in the Burgess Shale sequence at three localities. All are preserved as pyrite films in dark shale with relatively little relief, suggesting a lightly skeletized body. Complete specimens have a long, slightly tapering, large-plated attachment stalk, a conical cup or calyx with numerous small to medium-sized irregular plates, and 7–10 short arms with heavier plating and (in the holotype) soft appendages alternating from opposite sides of several arms. Several morphologic features indicate that Echmatocrinus is an echinoderm and has crinoid affinities: (1) Sutured plates, shown by darker depressed sutures, slightly raised plate centers, and oriented plate ornament, cover all major parts of the body; (2) reticulate surface ornament in the pyrite film on the plates of all specimens matches the ornament in the Burgess Shale edrioasteroid Walcottidiscus, an undoubted echinoderm, but not the pyritized surfaces of other metazoans in the fauna; (3) this distinctive ornament may represent the surface expression of microporous stereom; (4) possible ligament or muscle pads are present between the arm ossicles to fold and unfurl the more heavily plated arms. Within the echinoderms, only crinoids commonly have a calyx attached by a stalk or stem to the substrate and bear erect, moveable, uniserial arms for feeding. Although Echmatocrinus shows some resemblance to octocorals in overall body shape as an attached suspension feeder, almost all the details are different, indicating that Echmatocrinus is most likely unrelated to this group. All complete specimens of Echmatocrinus are attached to hard substrates, either another fossil or skeletal debris. The new specimens indicate that Echmatocrinus was twice as common (about 0.02%) in the Burgess Shale fauna as previously recorded and represents one of the earliest attached, medium-level, skeletized, suspension feeders or microcarnivores in the fossil record.

Muscular articulations in modern stalked crinoids are only present in the arms. Although it has been suggested that certain coiled-stemmed fossil taxa may have been functionally adapted to utilize muscles, evidence supporting this interpretation is lacking. Here, we use cathodoluminescence and SEM to reveal the skeletal microstructure of the enigmatic coiled-stemmed taxon Ammonicrinus (Flexibilia). Based on the well-established link between skeletal microstructure and the nature of infilling soft tissues in modern echinoderms, we reconstructed the palaeoanatomy of the Middle Devonian ammonicrinids. We show that their median columnals with elongated lateral columnal enclosure extensions (LCEE) have stereom microstructure unexpectedly resembling that in the crinoid muscular arm plates. In particular, large ligamentary facets, that are present on each side of a transverse ridge, are mainly comprised of fine galleried stereom that is indicative of the mutable collagenous tissues. In contrast, fine labyrinthic stereom, commonly associated with muscles, is situated in the periphery on each side of the surface of elongated LCEE. Our findings thus strongly suggest that the muscles may have also been present in the stem of ammonicrinids. These results reassess the previous hypotheses about evolution of muscles in crinoids and provide new insights into the mode of life of Ammonicrinus.

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 tooth stereom microstructure of 43 species of ophiuroids, from 9 families, was examined using light and scanning electron microscopy. Two basic types of teeth, uniform and compound, are present in ophiuroids. From a mechanical standpoint, compound teeth may provide a tougher grinding surface for ingestion of prey. Tooth types seem to be conserved at the family level, with only two exceptions, and are considered in light of known feeding strategies. Macrophagous feeders generally possess uniform teeth whereas microphagous feeders possess compound teeth. Ophiocomina nigra and Ophiopteris papillosa (Ophiocomidae) possess tooth types that are not in accord with other species in this family. Perhaps genera within this family are in need of reclassification.

Echinoderms possess a skeleton with a unique and distinctive meshlike microstructure called stereom that is underpinned by a specific family of genes. Stereom is thus considered the major echinoderm synapomorphy and is recognized already in some Cambrian echinoderm clades. However, data on the skeletal microstructures of early echinoderms are still sparse and come only from isolated ossicles of limited taxonomic value in which the primary calcium carbonate has been replaced or thinly coated by phosphates, silica or iron oxides. Here, we applied cathodoluminescence (CL) to reveal stereom microstructures of the diagenetically altered calcitic skeletons of some Cambrian echinoderms (in particular Protocinctus, Stromatocystites and Dibrachicystidae). CL not only provides insights into the diagenesis of their skeletons but also reveals primary microstructural details that are not visible under transmitted light or SEM. Different stereom types (resembling labyrinthic, fascicular, galleried, microperforate and imperforate microfabrics) comparable to those observed in extant echinoderms have been recognized for the first time in these Cambrian taxa. Our results show that the stereom microstructures widely occur in various Cambrian echinoderm clades which suggest that they likely evolved the same genetically controlled biomineralization mechanisms as those observed in modern echinoderms. These results underline that the CL technique can be a powerful tool in the detection of the microstructural organization in even severely recrystallized echinoderm specimens. Given the close association between the skeletal microstructure and the investing soft tissues, the method presented here opens new possibilites for revealing skeletal growth and soft tissue palaeoanatomy of fossil echinoderms.

The sea urchin (Echinodermata: Echinoidea) masticatory apparatus, or Aristotle\'s lantern, is a complex structure composed of numerous hard and soft components. The lantern is powered by various paired and unpaired muscle groups. We describe how one set of these muscles, the lantern protractor muscles, has evolved a specialized morphology. This morphology is characterized by the formation of adaxially-facing lobes perpendicular to the main orientation of the muscle, giving the protractor a frilled aspect in horizontal section. Histological and ultrastructural analyses show that the microstructure of frilled muscles is largely identical to that of conventional, flat muscles. Measurements of muscle dimensions in equally-sized specimens demonstrate that the frilled muscle design, in comparison to that of the flat muscle type, considerably increases muscle volume as well as the muscle\'s surface directed towards the interradial cavity, a compartment of the peripharyngeal coelom. Scanning electron microscopical observations reveal that the insertions of frilled and flat protractor muscles result in characteristic muscle scars on the stereom, reflecting the shapes of individual muscles. Our comparative study of 49 derived “regular” echinoid species using magnetic resonance imaging (MRI) shows that frilled protractor muscles are found only in taxa belonging to the families Toxopneustidae, Echinometridae, and Strongylocentrotidae. The onset of lobe formation during ontogenesis varies between species of these three families. Because frilled protractor muscles are best observed in situ, the application of a non-invasive imaging technique was crucial for the unequivocal identification of this morphological character on a large scale. Although it is currently possible only to speculate on the functional advantages which the frilled muscle morphology might confer, our study forms the anatomical and evolutionary framework for future analyses of this unusual muscle design among sea urchins.

The Aristotle's lantern, or masticatory apparatus, of regular sea-urchins is a complex musculo-skeletal system which is thought to have contributed significantly to the evolutionary success of these animals. This paper gives an account of the antomical relationships and functional morphology of both skeletal and soft tissue components in the lantern and related structures of the sea-urchin Stylocidaris affinis (Cidaridae), and compares these features with their equivalent in the previously described lantern of the sea-urchin Paracentrotus lividus (Echinidae, Camarodonta). There are major differences in the skeletons of these lanterns which involve mostly the arrangement and morphology of elements participating in movement, i.e. joints and articular surfaces, and which highlight the generally heavier and less mobile nature of the lantern in the Cidaridae. There are remarkably few differences, however, in the microstructure of the skeletal stereom. Significant dissimilarities were found in the anatomical arrangement of muscles and ligamentous structures and in their macro- and microstructure. The implications of these morphological features for the functioning of the lantern of the Cidaridae are discussed in the context of an integrated model of lantern biomechanics.

When echinoids feed on sponges, silicate spicules of the sponge were found to enter their body either by penetration through the wall of the food canal into coelomic cavities or by penetration into skeletal plates and spines. The spicules, which have penetrated into the coelom, obviously evoke a kind of protective answer. They were found to be entangled by clusters of cell remnants, the so-called brown bodies. The brown bodies contain melanin and gather at special sites of the echinoid body; these are the Stewart Organs, the gills and the inner side of the ambulacral plates. Sometimes the silicate spicule becomes surrounded by a calcareous sheath. The length of the sponge spicules makes their removal impossible, so that they are stored. The spicules penetrating into the plates are partly incorporated into the stereom. The four species examined in this study were Asthenosoma ijimai, Araeosoma owstoni, Diadema setosum (collected in Sagami Bay, Japan in 1991) and Hapalosoma gemmiferum (collected in Suruga Bay, Japan in 1991).