Crinoid holdfasts from the Silurian of Gotland

Crinoid holdfasts occur throughout the Silurian sequence of Gotland, with a marked concentration to reefs and immediate reef surroundings. Four morphological groups are recognized: (1) Discoid holdfasts. (2) Cirriferous holdfasts, represented by (a) Rhizocrinus -like holdfasts, (b) large radices with stout, often branching cirri, and (c) rhizoid holdfasts, formed by a complicated net of pseudocirri. (3) Creeping stems attached to the substrate by thin strands of stereome. (4) Coiling stems. Discoid attachments, cirriferous holdfasts (types b and c) and coiling stems show little preference to substrate and adapt morphologically to conditions at hand. Rhizoid holdfasts display the greatest variation and apparently occur within many different groups of crinoids. Rhizocrinusttke holdfasts have been found only in quiet-water deposits, while creeping stems were concentrated to more turbulent environments. Coiling stems were epizoic, attached to rugose corals, bryozoans, crinoid stems and similar supports. Growth, mode of life (attached or free), settling and fixation of larvae, and relation to substrate and other organisms ate discussed.     

A radicle solution: morphology and biomechanics of the Eucalyptocrinites ‘root’ system

Eucalyptocrinites is one the most common and familiar mid‐Palaeozoic crinoids and is the exemplar of a form with dendritic radicular holdfasts. American museums have hundreds of specimens of Eucalyptocrinites holdfasts from the Silurian Waldron Shale of Indiana and Kentucky, USA. The radix (‘root’) system of Eucalyptocrinites can be described as comprised of links (branches) that meet at nodes. Measured values include the x‐ and y‐coordinates of the nodes, the distances of the nodes from the stem, the angles between branches, branch length and branch width. Rose diagrams show clearly that the root systems are not isotropic but have preferred orientations. Branch angles are highly variable, but cluster around 60 degrees. Branch lengths and distal branch widths are relatively constant, but branch width increases variably towards the column. The branching pattern can be modelled as a self‐similar (‘fractal’) structure. Several specimens labelled as Eucalyptocrinites show a distinct fivefold symmetry without branching and very likely represent a different taxon. The Eucalyptocrinites radix system, along with the probably stiff dististele, most likely functioned as a rigid plate that resisted rotational forces due to currents acting on the crown. Upstream radicles experienced tension, whereas downstream roots were compressed. This force distribution may explain the observed anisotropies in radix morphology. The ‘roots’ of Eucalyptocrinites and other crinoids have long been compared with the root systems of plants. Although there are superficial similarities, there are fundamental differences.     

Phenotypic variability, deformations and post-traumatic reactions of the stem ofEncrinus cf.liliiformis Lamarck (Crinoidea) from the Middle Triassic of Qingyan, south-western China

Well-preserved stem remains ofEncrinus cf.liliiformis Lamarck 1801 from the Upper Anisian and lowest Ladinian (Middle Triassic) of Qingyan, Guizhou Province, south-western China, are described. The characteristic morphological features of the columnals, especially the sculptural characteristics of the articular facets, vary in the different parts of the stem. Numerous fine, short, peripheral crenulae and perilumina with a tendency to pentalobate shapes in the proximal stem region change to few long, coarse crenulae and circular perilumina in the distal stem part. The holdfasts exhibit various differentiations: the basic discoid type occurs on flat surfaces, encrusting holdfasts are characteristic of irregular substrates; juvenile holdfasts often show an irregular outline, adult ones have a more even, circular shape. Some special and unusual structures allow palaeoecological interpretations: Holdfasts and stems joined by callous growth probably result from the lack of suitable hard substrates for attachment on the soft seafloor; the larvae often had to settle very closely to each other on a relatively small substrate. After breakage of stem,Encrinus was able to continue living despite the traumatic loss of its basal fixation. Broken ends underwent skeletal regeneration. Some specimens strongly suggest that juvenile individuals sometimes even were able to re-attach secondarily, e.g. by coiling around stems of other crinoids and cementing to these by callous outgrowths.     

Contractile tissues in the cirri of ancient crinoids: criteria for recognition

Stalked isocrinid and 'stalkless' comatulid crinoids are able to relocate by crawling on or swimming with their muscular arms. Reattachment is achieved using cirri containing contractile tissues which produce aboral flexure. The following cirral adaptations for active attachment were observed during a SEM study of two comasterid comatulids, Davidaster rubiginosa (Pourtalès) and D. discoidea (Carpenter): synarthrial articulations; fulcral ridges corresponding to short axes of ossicles; cirri flattened laterally; each cirrus with a claw at the tip and an aboral attachment pad; cirri serrated aborally and distally. Epizoans are only able to encrust single ossicles in actively motile cirri. These adaptations are associated with crinoids that have muscular arms which are used in relocation. Such a suite of characters is unknown in Palaeozoic crinoids; the ability to relocate only evolved in crinoids during the Mesozoic.□ Crinoids, cirri, comatulids, evoiution, functional morphology .     

Attachment of the adhesive holdfast organelle to the cellular stalk of Caulobacter crescentus

Caulobacters attach to surfaces in the environment via their holdfasts, attachment organelles located at the base of the flagellum in swarmer cells and later at the end of the cellular stalk in the stalked cells which develop from the swarmer cells. There seems to be little specificity with respect to the types of surfaces to which holdfasts adhere. A notable exception is that the holdfast of one cell does not adhere to the cell surface of another caulobacter, except by joining holdfasts, typically forming "rosettes" of stalked cells. Thus, the localized adhesion of the holdfasts to the cells is in some way a specialized attachment. We investigated this holdfast-cell attachment by developing an adhesion screening assay and analyzing several mutants of Caulobacter crescentus CB2A selected to be defective in adhesion. One class of mutants made a normal holdfast by all available criteria, yet the attachment to the cell was very weak, such that the holdfast was readily shed. Another class of mutants made no holdfast at all, but when mixed with a wild-type strain, a mutant of this class participated in rosette formation. The mutant could also attach to the discarded holdfast produced by a shedding mutant. In addition, when rosettes composed of holdfast-defective and wild-type cells were examined, an increase in the number of holdfast-defective cells was correlated with a decrease in the ability of the holdfast material at the center of the rosette to bind colloidal gold particles. Gold particles are one type of surface to which holdfasts adhere well, suggesting that the stalk end and the colloidal gold particles occupy the same sites on the holdfast substance. Taken together, the data support the interpretation that there is a specialized attachment site for the holdfast at the base of the flagellum which later becomes the end of the stalk, but not a specialized region of the holdfast for attachment to this site. Also, attachment to the cell is accomplished by bond formations that occur not only at the time of holdfast production. Thus, we propose that the attachment of the holdfast to the cell is a true adhesion process and that the stalk tip and base of the flagellum must have compositions distinctly different from that of the remainder of the caulobacter cell surface.     

Survival of crinoid stems following decapitation: evidence from the Ordovician and palaeobiological implications

It was recently discovered that the stems of extant crinoids may survive after detachment of the crown, presumably feeding by the absorption of nutrients through the ectoderm. Previously, only one analogous, albeit morphologically dissimilar, pattern of crownless survival has been recognized from the fossil record. Certain Upper Ordovician (Cincinnatian) crinoid pluricolumnals from Kentucky, Ohio and Indiana, derived from the disparids Cincinnaticrinus spp., have rounded terminations reminiscent of some modern bourgueticrinid overgrowths. Such specimens have hitherto been interpreted as distal terminations of mature individuals that have become detached from their attachment structures and taken to an eleutherozoic existence. However, it is considered more probable that they represent overgrowths of the column following predatory decapitation. If this new interpretation is correct, then post-decapitation survival of crinoid stems is now recognized for most of the history of the crinoids, 'lethal' predation on crinoid crowns occurred at least as early as the Late Ordovician and ancient crinoid populations can no longer be determined merely by counting crowns.     

Radial perforations in crinoid stems from the Silurian of Gotland

Radiating intracolumnal canals are a characteristic feature of large (diameter 10 mm or more) crinoid stems from the Silurian of Gotland. They are found in nodals as well as in internodals where the columnal height exceeds one millimetre. They were formed secondarily in the median and distal portions of crinoid stems with pseudocirriferous holdfasts. Intercolumnal canals are found in the distal parts of stems with true cirri regardless of the size of the stem. It is suggested that these canals played an important role in crinoid physiology. The crinoids are believed to have sustained a large proportion of their tissues through cutaneous digestion and uptake of dissolved substances from the surrounding sea water. The intra- and intercolumnal canals increased the surface of the axial canal in relation to volume. They provided a connection between the axial canal and the surrounding sea water, thus facilitating nutrient transport to the tissues.     

Terminology and functional morphology of attachment structures in pelmatozoan echinoderms

Brett, Carlton E. 1981 12 IS: Terminology and functional morphology of attachment structures in pelmatozoan echinoderms. Lethaia . Vol. 14, pp. 343–370. Oslo. 1SSN 0024–1164.
Fixation on the seafloor is a fundamental, though little studied, aspect of the biology and distributional ecology of pelmatozoan echinoderms. Morphology of pelmatozoan attachment structures (holdfasts) and their component appendages is summarized herein and a detailed terminology proposed. Three major modes of fixation are differentiated in living and fossil stemmed echinoderms - primary, composite and secondary attachments - in addition to the direct thecal attachment in stemless forms. Specific holdfast morphologies, which characterize certain major taxa (families and orders) of pelmatozoans, may limit the available life-modes in these echinoderms. Significantly, numerous lineages of echinoderms have independently developed secondary attachment modes associated with the evolution of flexible tapered stem ends and/or true cirri. Cirri are rare in early Paleozoic pelmatozoans but appeared in most major groups by the late Paleozoic. * Pelmatozoa. attachment structures, holdfasts, terminology, functional morphology .     

Presumed postlarval pentacrinoids from the Lower Devonian Hunsrück Slate,Germany

Several tiny crinoids with crowns as small as 1 mm, or less, in width are newly recognized from the Hunsrück Slate of southwestern Germany. The presence of erect arms above an amorphous calyx in some specimens can be inferred. Based on comparison with the size and gross morphology of developmental stages in living crinoids, these tiny Hunsrück crinoids are judged to be at an early postlarval stage that is analogous to the pentacrinoid stage just after development from the stalked, but armless, smaller cystidean larval stage found in both living comatulids and isocrinids. Some of these tiny crinoids have a stalk up to 4 mm long attached to a now pyritized former substrate. Their clustered occurrence suggests gregarious settlement of larvae. Taxonomic identification of these presumed pentacrinoids is not possible, even to the sub‐class level, although they are preserved with larger juveniles of the cladids Propoteriocrinus and Lasiocrinus. These larger juveniles exhibit 3‐D pyritized calcite plates, whereas the probable pentacrinoids appear to be preserved as flattened, micro‐crystalline pyritized dermal tissues that enclosed lightly calcified, porous ossicles. The pentacrinoids were likely buried within weeks or months of hatching, based on developmental stages in similar‐sized living crinoids. These tiny crinoids, presumably pentacrinoids, are a further example of the extraordinarily detailed preservation of delicate tissues in pyrite from the Hunsrück Slate. They are most likely the pentacrinoid stage from one or more of the crinoid taxa (30 genera) present in the Hunsrück Slate. Assuming these are not microcrinoids, they are the first report of pentacrinoids from the fossil record and document that a Palaeozoic sister group to modern crinoids had similar developmental stages.     

Infestation of Middle Devonian (Givetian) camerate crinoids by platyceratid gastropods and its implications for the nature of their biotic interaction

One of the classic examples of biotic interactions preserved in the fossil record is that between crinoids and infesting platyceratid gastropods. This relationship, spanning an interval from the Middle Ordovician to the end of the Permian, is recognized by the firm attachment and positioning of platyceratids over the anal vent of their hosts. Several hypotheses have been proposed to explain this interaction; the most widely accepted is that the gastropods were coprophagous commensals, feeding on crinoid excrement without any significant detriment to their hosts. The purpose of this investigation was to test this hypothesis. Two species of Middle Devonian camerate (Monobathrida, Compsocrinina) crinoids, Gennaeocrinus variabilis Kesling & Smith 1962 and Corocrinus calypso (Hall 1862), were used in this investigation. The data consisted of 426 individuals of G. variabilis collected near Rockport, Michigan, 30 of which were infested, and 188 individuals of C. calypso collected near Arkona, Ontario, Canada, of which 25 were infested. Length and volume were measured for each crinoid to determine whether a significant difference existed in the size of infested versus uninfested individuals. The results indicated that for both species of crinoids individuals infested by snails were significantly smaller than uninfested individuals (p < 0.05). We explored a variety of scenarios to explain this pattern and conclude that they falsify the null hypothesis that the crinoid-gastropod relationship was strictly commensal. The smaller size of the infested crinoids is interpreted as a consequence of nutrient-stealing by the parasitic gastropods, a strategy that finds analogs in modern seas. Moreover, the absence of platyceratids on the largest crinoids suggests that large size may have inferred immunity from lasting infestation.     

Evolutionary history of regeneration in crinoids (Echinodermata)

The fossil record indicates that crinoids have exhibited remarkable regenerative abilities since their origin in the Ordovician, abilities that they likely inherited from stem-group echinoderms. Regeneration in extant and fossil crinoids is recognized by abrupt differences in the size of abutting plates, aberrant branching patterns, and discontinuities in carbon isotopes. While recovery is common, not all lost body parts can be regenerated; filling plates and overgrowths are evidence of non-regenerative healing. Considering them as a whole, Paleozoic crinoids exhibit the same range of regenerative and non-regenerative healing as Recent crinoids. For example, Paleozoic and extant crinoids show evidence of crown regeneration and stalk regrowth, which can occur only if the entoneural nerve center (chambered organ) remains intact. One group of Paleozoic crinoids, the camerates, may be an exception in that they probably could not regenerate their complex calyx-plating arrangements, including arm facets, but their calyxes could be healed with reparative plates. With that exception, and despite evidence for increases in predation pressure, there is no compelling evidence that crinoids have changed though time in their ability to recover from wounds. Finally, although crinoid appendages may be lost as a consequence of severe abiotic stress and through ontogenetic development, spatiotemporal changes in the intensity and frequency of biotic interactions, especially direct attacks, are the most likely explanation for observed patterns of regeneration and autotomy in crinoids.     

Variations in holdfast attachment mechanics with developmental stage, substratum-type, season, and wave-exposure for the intertidal kelp species Hedophyllum sessile (C. Agardh) Setchell

Biomechanical models that describe physical and biological interactions on wave-exposed shores typically assume that a species' attachment properties are similar between seasons and sites. We tested this assumption using Hedophyllum sessile to investigate how macroalgal biomechanical attachment properties vary with developmental stage, substratum-type, season, and wave-exposure. Hedophyllum sessile is an intertidal kelp species that is able to survive in wave-exposed areas in the Northeast Pacific. For both juveniles and adults, holdfast attachment force and strength were measured at a wave-exposed and wave-protected site in Barkley Sound, British Columbia, Canada. Substratum and wave-exposure effects on attachment properties were tested in juvenile populations. Adult populations were sampled prior to (in July 1996) and after (in November 1996) a series of storms. Site and seasonal wave-exposure effects on attachment properties were tested in these adult populations. Comparisons to known attachment properties of other temperate macroalgal species were also made. Causes for these patterns are discussed but were not isolated in these studies. Juveniles' attachment properties differed on different substrata types and between wave-exposures, with the highest attachment forces and the most attached juveniles in articulated coralline algal turfs. Adult attachment is firm ( approximately 100 N), but relatively weak ( approximately 0.07 MNm(-2)). Adult attachment did not vary with site wave-exposure, but there was a shift within each site to more resistant holdfasts after a series of early winter storms. Seasonal increases in storm swells correlated to more thallus tattering and selected against large, loose holdfasts. The data presented here suggest that results from holdfast attachment field studies in one season cannot be extrapolated to another due to a complex set of dynamics. This is the first documentation of seasonal patterns in macroalgal attachment properties.     

NEW CRINOIDS (ECHINODERMATA) FROM THE LLANDOVERY (LOWER SILURIAN) OF THE BRITISH ISLES

Abstract:  British Llandovery crinoids remain poorly known. Three species are documented herein that were originally described, but not published, by W. H. C. Ramsbottom. Clematocrinus ramsbottomi Fearnhead sp. nov. (Tortworth Inlier, Gloucestershire; Telychian) has a heteromorphic column, N434243414342434, radices directed away from the crown, and ten long, uniserial arms with a pustular aboral sculpture and long, slender pinnules. Clematocrinus spp. are widely distributed in the Silurian of England. Ptychocrinus mullochillensis Fearnhead and Donovan sp. nov. (Girvan district, Strathclyde; Rhuddanian) is the second report of this Upper Ordovician–Lower Silurian genus from outside North America. Although incompletely known, this species is distinguished by its hidden infrabasals, 20 arms and sunken interbrachial plates with a strongly stellate sculpture. Petalocrinus bifidus (Bather MS) Donovan and Fearnhead sp. nov. (Woolhope Inlier, Herefordshire; Telychian) is locally common enough to give its name to a mappable lithostratigraphic unit, the Petalocrinus Limestone. Unlike other Petalocrinus species known from the Silurian of China, northern Europe and North America, the fused arms of P. bifidus are divided in two by a proximal adoral ridge and an associated distal notch. Unlike extant crinoids, Petalocrinus probably lived as a rheophobe; the fused arms may have acted to deter predators.     

Colonization of substrates: Vendian sedentary benthos

Studies of fossils sampled from a single fossiliferous layer (monotopic series) and a summary of previous data on taphonomy and morphology of Vendian macroorganisms suggest that most strategies of attachment of sedentary benthos were formed in the Vendian. These included: (1) free living with possible organic gluing; (2) fixation on the substrate using sucker-like structures and various incrustations; (3) fixation of the basal parts by shallow submergence in the sediment; (4) anchoring into the sediment with discoidal and rhizoid-like holdfasts; and (5) partial submergence and infaunal lifestyle. The most widespread fixation in the Vendian was the attachment by discoid organs, which can be interpreted as symbiotrophic structures. Symbiotrophy in Vendian organisms submerged in the anoxic sediment can apparently be supported by indirect evidence: structural diversity of basal organs, their complex morphology, large area of contact with substrate, unlimited isometric growth, predominant fossilization of attachment discs; inconsistent systematization of organisms.     

Late Cambrian hard substrate communities from Montana/Wyoming: the oldest known hardground encrusters

Hardground surfaces from the Late Cambrian Snowy Range Formation in Montana/Wyoming are the oldest known non-reefal hard substrates exhibiting encrusting fossils. These surfaces range in age from Early Franconian to early Trempealeauan. Hardgrounds were developed on slightly hummocky to planar, truncated surfaces of glauconite-rich, carbonate, flat pebble conglomerates, which were deposited during episodes of storm scouring in shallow subtidal environments of the Montana/Wyoming shelf. Snowy Range hardgrounds are encrusted by a low diversity assemblage of fossils dominated by simple discoidal holdfasts of pelmatozoans, probably crinoids, and including small conical spongiomorph algae? and probable stromatolites. Macroborings (e.g. Trypanites) are notably absent from all hardground surfaces, although sharp-walled, vertical, cylindrical holes (borings?) occur in micrite clasts imbedded in certain flat pebble conglomerates. No evidence of faunal succession or microecologic partitioning of irregular surfaces was observed on these Cambrian hardgrounds.     

Characterization of the Adhesive Holdfast of Marine and Freshwater Caulobacters

Caulobacters are prosthecate (stalked) bacteria that elaborate an attachment organelle called a holdfast at the tip of the cellular stalk. We examined the binding of lectins to the holdfasts of 16 marine Caulobacter strains and 10 freshwater species or strains by using a panel of fluorescein-conjugated lectins and fluorescence microscopy. The holdfasts of all the marine isolates bound to only wheat germ agglutinin (WGA) and other lectins that bind N-acetylglucosamine (GlcNac) residues. The freshwater caulobacters showed more variability in holdfast composition. Some bound only to WGA and comparable lectins as the marine strains did. Others bound additional or other lectins, and some did not bind to the lectins tested. The binding of WGA appeared to involve the regions of the holdfast involved with adhesion; a holdfast bound to WGA was significantly less adhesive to glass. Competition experiments with WGA-binding holdfasts and oligomers of GlcNac demonstrated that trimers of GlcNac (the preferred substrate for WGA binding) were more effective than dimers or monomers in preventing WGA binding to holdfasts, suggesting that stretches of contiguous GlcNac residues occur in the WGA-binding holdfasts. In addition, differences between freshwater and marine holdfasts in the strength of WGA binding were noted. The effect of a number of proteolytic and glycolytic enzymes on holdfast integrity was examined; the proteases had no effect for all caulobacters. None of the glycolytic enzymes had an effect on marine caulobacter holdfasts, but chitinase and lysozyme (both attack oligomers of GlcNac) disrupted the holdfasts of those freshwater caulobacters that bound WGA. Despite some similarity to chitin, holdfasts did not bind Calcofluor and no measurable effects on holdfast production were detectable after cell growth in the presence of diflubenzuron or polyoxin D, inhibitors of chitin synthesis in other systems. Finally, the holdfasts of all caulobacters bound to colloidal gold particles, without regard to the coating used to stabilize the gold particles. This binding was stronger or more specific than WGA binding; treatment with colloidal gold particles prevented WGA binding, but the reverse was not the case.     

Giant kelp vegetative propagation: Adventitious holdfast elements rejuvenate senescent individuals of the Macrocystis pyrifera “integrifolia” ecomorph

Recent findings on holdfast development in the giant kelp highlighted its key importance for Macrocystis vegetative propagation. We report here for the first time the development of adventitious holdfasts from Macrocystis stipes. Swellings emerge spontaneously from different areas of the stipes, especially in senescent or creeping individuals. After being manually fastened to solid substrata, these swellings elongated into haptera, which became strongly attached after 1 month. Within 4 months, new thalli increased in size and vitality, and developed reproductive fronds. Our results suggest the usage of these structures for auxiliary attachment techniques. These could act as a backup, when primary holdfasts are weak, and thus improve the survival rate of the giant kelp in natural beds.     

Piscinoodinium, a fish-ectoparasitic dinoflagellate, is a member of the class dinophyceae, subclass gymnodiniphycidae: convergent evolution with Amyloodinium

All dinoflagellates that infest the skin and gills of fish have traditionally been placed within the class Blastodiniphyceae. Their relatedness was primarily based upon a similar mode of attachment to the host, i.e., attachment disc with holdfasts. Results of recent molecular genetic analyses have transferred these parasites, including Amyloodinium, to the class Dinophyceae, subclass Peridiniphycidae. In our study, a small subunit rDNA gene from a parasitic dinoflagellate that has features diagnostic for species in the genus Piscinoodinium, i.e., typical trophont with attachment disc having rhizocysts, infesting the skin of freshwater tropical fish, places this organism within the dinophycean subclass Gymnodiniphycidae. This suggests a close relationship of Piscinoodinium spp. to dinoflagellates that include symbionts, e.g., species of Symbiodinium, and free-living algae, e.g., Gymnodinium spp. These molecular and morphological data suggest that evolution of this mode of fish ectoparasitism occurred independently in 2 distantly related groups of dinoflagellates, and they further suggest that the taxonomic status of parasites grouped as members of Piscinoodinium requires major revision.     

A re‐interpretation of the ambulacral system of Eumorphocystis (Blastozoa,Echinodermata) and its bearing on the evolution of early crinoids

Recent debates over the evolutionary relationships of early echinoderms have relied heavily on morphological evidence from the feeding ambulacral system. Eumorphocystis, a Late Ordovician diploporitan, has been a focus in these debates because it bears ambulacral features that show strong morphological similarity to early crinoid arms. Undescribed and well‐preserved specimens of Eumorphocystis from the Bromide Formation (Oklahoma, USA) provide new data illustrating that composite arms supported by a radial plate that bear a triserial arrangement of axial and extraxial components encasing a coelomic extension can also be found in blastozoans. Previous reports have considered these arm structures to be restricted to crinoids; these combined features have not been previously observed in blastozoan echinoderms. Phylogenetic analyses suggest that Eumorphocystis and crinoids are sister taxa and that shared derived features of these taxa are homologous. The evidence from the arms of Eumorphocystis suggests that crinoid arms were derived from a specialized blastozoan ambulacral system that lost feeding brachioles and strongly suggests that crinoids are nested within blastozoans.     

A Lower Devonian hexacrinitid crinoid (Camerata,Monobathrida) from south-west England

The most species-rich and widespread crinoid clade in the type area of the Devonian of south-west England is the monobathrid camerate family Hexacrinitidae Wachsmuth and Springer. These crinoids occur either as thecae (Middle Devonian) or pluricolumnals and columnals (Lower to Upper Devonian). The first new, nominal species of hexacrinitid, probably Oehlerticrinus Le Menn, to be described from this region since the nineteenth century is Oehlerticrinus peachi sp. nov. from the Lower Devonian Looe Basin of southern Cornwall. This specimen is mouldic and somewhat flattened, retaining the proxistele and arms. Diagnostic features include the heteromorphic proxistele with circlets of long, unbranched radices directed towards the crown; the high, box-like (=skyphosiform) theca with a flattened base; a thecal plate sculpture of tubercles and ridges arranged in triangles; and the pinnulate arms. Any uncertainty in identifying this species at the generic level rests with the arms being pinnulate, not ramulose as is common in Hexacrinites Austin and Austin, indicating that O. peachi is closer to Oehlerticrinus Le Menn.