Using platyceratid gastropod behaviour to test functional morphology

During the Mississippian (Tournaisian), numerous crinoid genera of the subclass Camerata evolved exaggerated anal tubes, cylindrical extensions of the tegmen with the anus at the distal end. Additionally, camerates exhibit higher frequency of platyceratid gastropod infestation than any other crinoid clade leading some researchers to speculate that anal tubes evolved in response to platyceratid parasitism. To test the infestation avoidance role of anal tubes, platyceratid distribution was analyzed among 636 tubed and 675 tubeless crinoids from Mississippian strata in North America. Results demonstrate significantly higher infestation frequency in tubeless crinoids. Rather than attach to the anal vent, as is typical for platyceratids, the gastropods that infested tubed crinoids are always found at the tube base and acquired nutrients from their hosts via drilling. It is likely that infesting tubeless crinoids was a more cost effective trophic strategy than drilling tubed crinoids.     

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.     

Tube foot preservation in the Devonian crinoid Codiacrinus from the Lower Devonian Hunsrück Slate,Germany

Fossilized tube feet are described on Codiacrinus schultzei Follmann from the Lower Devonian Hunsrück Slate of Germany. This is the first definitive proof of tube feet on any fossil crinoid. Three lightly pyritized, flattened tube feet are preserved in a single interray of this cladid crinoid. The tube feet were at least 7 mm long. Their preservation is very similar to the tube feet reported previously from a Hunsrück ophiuroid, except that the Codiacrinus tube feet have small papillae, similar to living crinoids.     

Testing reduced evolutionary rates during the Late Palaeozoic Ice Age using the crinoid fossil record

In 2003, Stanley & Powell reported depressed rates of origination and extinction in marine invertebrates during the Late Palaeozoic Ice Age (LPIA). Using a database of crinoid genera, rates of origination, extinction and genus duration were calculated at the stage level from the Early Devonian to the Late Permian. This 165 m.y. time span includes non‐glacial intervals before and after the LPIA, which spanned the Serpukhovian to Sakmarian, providing background rates for comparison. Data generated on crinoid evolutionary rates during the Middle to Late Palaeozoic were analysed and compared to Stanley & Powell's data to determine whether crinoid evolutionary patterns support their findings or suggest an alternative hypothesis. Rates of origination and extinction in all crinoid clades were reduced during the LPIA compared to the combined background intervals before and after the LPIA. However, crinoid diversity was higher during the LPIA than the surrounding time intervals. The difference in diversity trends between crinoids and other marine invertebrates is due to the advanced cladids clade. Unstable, fluctuating environmental conditions during the LPIA may have created habitats suitable for opportunistic crinoid genera that reduced both the probability of origination and extinction. The increased diversity of the advanced cladids is likely due to their unique adaptation of muscular arm articulations, which allowed them to thrive in marine settings with increased siliciclastic influx brought on by the Alleghenian orogeny. Despite the advanced cladids’ departure from the expected diversity count, the results of analyses performed on the updated crinoid database provide independent confirmation of Stanley & Powell's original hypothesis of depressed evolutionary rates in marine invertebrates during the LPIA.     

Assessment of hidden diversity of crinoids and their symbionts in the Bay of Nhatrang,Vietnam

Crinoid associates represent an abundant and diverse, but poorly explored, component of the hidden biodiversity of coral-reef ecosystems. We studied data from 5 years of collecting in the Bay of Nhatrang (BN), Vietnam, to assess the diversity of crinoids and their symbionts, to compare it with other areas of the Indo-West Pacific, and to elucidate the extent to which the observed diversity of crinoids and their symbionts corresponds to their true diversity. In total, about 2,287 specimens of symbionts belonging to 70 species were found on 203 specimens of crinoids belonging to 33 species. Among the crinoids, the most numerous species were Himerometra robustipinna (36 specimens) and Cenometra bella (29 specimens), among the symbionts the polychaete Paradyte crinoidicola (c. 850 specimens) and the galatheid crustacean Allogalathea elegans (180 specimens). Species accumulation curves suggest that we have sampled most of the crinoid diversity in the BN, whereas the diversity of their symbionts remained undersampled. Estimated species richness of crinoids was higher than previously observed richness, and varied from 39 (estimated by bootstrap) to 46 (jackknife 2). Estimated species richness of symbionts was higher than observed richness, and varied from 71 (bootstrap) to 93 (jackknife 2). We suggest a slight increase in the number of crinoid species to result from more detailed studies of nocturnal species, and an increase in the number of symbiotic species when studies of nocturnal crinoid associates and sibling species among decapods are included. Our study revealed a rather rich crinoid fauna in the bay compared to other areas of the Indo-West Pacific, and the highest species richness of crinoid associates known from anywhere in the World Ocean.     

Escalating predation on crinoids in the Devonian: negative community-level evidence

Previous morphological studies suggest that predation on stalked crinoids increased in the Devonian. However, there was no concomitant decline in the occurrence of dense, shallow-water stalked crinoid assemblages in North America from the Ordovician and Silurian to the Mississippian. The evolution of crinoid defensive adaptations may have kept pace with rising durophagy, forestalling the expected decline of crinoid communities. It is also possible that the demise of reefs after the Frasnian-Famennian mass extinction indirectly decreased predation pressure on crinoids by removing shelter for predatory fish. A third possibility is that stalked crinoid abundance was affected neither by new predators nor by the decline of reefs. □ Crinoidea, dense crinoid assemblages, Echinodermata, North America, Paleozoic, predation.     

Before the extinction – Permian platyceratid gastropods attached to platycrinitid crinoids and an abnormal four-rayed Platycrinites s.s. wachsmuthi (Wanner) from West Timor

The occurrence of Paleozoic gastropods attached to echinoderms has been recognized for nearly 170 years. Specimens have been illustrated from each of the geologic periods from the Ordovician to Permian. We illustrate two occurrences of Permian platycrinitid camerate crinoids from West Timor, Platycrinites s.s. wachsmuthi (Wanner) and Neoplatycrinus dilatatus Wanner, that have the platyceratid attached on the tegmen on the former and along the radial summit on the latter. Both platyceratids barely cover the anal opening and suppress the development of the arms of at least one ray. The D ray arms are suppressed on Platycrinites s.s. wachsmuthi, whereas the development of the C ray arms and perhaps the most adjacent arms of the D and B rays are suppressed on Neoplatycrinus dilatatus. An additional specimen of Neoplatycrinus major retains the impressed conch outline of a formerly attached platyceratid on the tegmen. This is the first report of a platyceratid/Neoplatycrinus association. The West Timor occurrences are among the youngest known of the platyceratid/platycrinitid association before the End-Permian extinction of the camerates. In addition, an abnormal four-rayed theca of Platycrinites s.s. wachsmuthi is described.     

Phylogenetic implications of the Protocrinoida: Blastozoans are not ancestral to crinoids

A traditional, widely cited hypothesis for over a century posits the origin of the crinoids from blastozoans. The blastozoan hypothesis is contradicted by the discovery of a new crinoid order, the Protocrinoida. Protocrinoids exhibit many traits that are consistent with a basal crinoid phylogenetic position, but inconsistent with a blastozoan ancestry. Protocrinoids are among the oldest crinoids and are therefore stratigraphically correctly placed. The blastozoan hypothesis in contrast, relies on putative homologies between blastozoans and crinoids taken from taxonomically and stratigraphically disparate representatives of both groups; these disparities indicate homoplasy rather than propinquity of descent. Data supporting these ideas are reviewed here. These findings reinforce insightful observations made by Georges Ubaghs decades ago with less data.     

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.     

Host-specific pit-forming epizoans on Silurian crinoids

Brett, Carlton E. 197807 15: Host-specific pit-forming epizoans on Silurian crinoids. Lethaia , Vol. 11. pp. 217–232. Oslo. ISSN 0024–1164.
Circular-parabolic pits occur commonly on the endoskeletal remains of certain Paleozoic crinoids. Detailed study of several hundred specimens, representing about 30 pelmatozoan species from the Upper Silurian Rochester Shale of New York and Ontario, reveals that such pits occur exclusively in seven species of crinoids. Furthermore, there are consistent differences in the morphology and orientation of holes occurring on the different crinoid species. This suggests that distinct epizoan species settled selectively on given hosts. The relationship between the hole-producing epizoans and crinoid hosts is inferred to have been a form of dependent commensalism. Preliminary surveys of other Paleozoic crinoid assemblages reveal similar host-selectivity by pit-producing epizoans. Crinoidepizoan pairs apparently co-evolved through considerable spans of geologic time as related genera and species of different ages, from Silurian to Pennsylvanian, exhibit similar pits.     

Filtration models, guilds, and biofacies: Crinoid paleoecology of the Stanton Formation (Upper Pennsylvanian), midcontinent, North America

Filtration models make several predictions concerning the distribution of crinoids among benthic habitats: (1) generally, higher energy shoreward settings should contain crinoid assemblages dominated by taxa with dense mesh filtration fans; (2) generally, lower energy offshore settings should be dominated by crinoids with open mesh filtration fans; (3) diversity should be highest in nearshore settings, which generally contain low energy microhabitats within high energy settings where both dense fan and open fan forms can co-occur, whereas offshore assemblages should be dominated by open fan taxa; thus, open fan forms should be more eurytopic than dense fan forms. Also, previous empirical models for crinoid distribution patterns note that nearshore assemblages are dominated by large forms whereas offshore assemblages are dominated by small forms. Filtration fan morphology and body size interact significantly to determine how and where crinoids feed and thus are used here as criteria for distinguishing late Paleozoic crinoid guilds. The distribution patterns of crinoid taxa and guilds among the benthic paleoenvironments of the Upper Pennsylvanian (Missourian) Stanton Formation were used to test the predictions of the filtration models.

The results of quantitative analyses show that large, dense fan crinoids are significantly more abundant in shoreward assemblages than smaller, open fan crinoids, which are more abundant in offshore assemblages; these results are consistent with the aforementioned models. However, diversity patterns are not consistent with the filtration models, indicating that mid-depth and aerobic offshore assemblages are more taxonomically and trophically diverse than nearshore and dysaerobic offshore assemblages, ostensibly due to the overlapping ranges of characteristically nearshore and offshore taxa. Analysis of guild distribution patterns also indicates some contradictions to predicted patterns of stenotopy and eurytopy, indicating additional complexities to late Paleozoic crinoid paleoecology.

Five recurrent crinoid assemblages, or biofacies, have been identified from the Stanton Formation of midcontinent North America. The guild structure and diversity of these biofacies are characterized here as: (1) a moderate diversity nearshore biofacies dominated by large calyx, dense fan guilds; (2) a high diversity midshelf biofacies characterized by equitable contributions of many taxa and guilds; (3) a moderate diversity offshore biofacies dominated by non-pinnulate and small calyx, open fan pinnulate guilds; (4) a low diversity offshore dysaerobic biofacies dominated by small calyx pinnulate guilds; (5) a depauperate offshore biofacies dominated by a unique stalkless species, Paragassizocrinus tarri. Except for Biofacies 5, the number of guilds appears to increase offshore; however, the equitability of taxa among and within the guilds generally decreases offshore. This pattern indicates that the heterogeneity of resources, or the recognition by taxa of different classes of resources, as reflected by the variety of functional types, increased from nearshore to offshore, but that the abundance or accessibility of these resources, and/or the relaxation of competition within guilds, increased from offshore to nearshore.     

The complete mitochondrial genomes of the sea lily Gymnocrinus richeri and the feather star Phanogenia gracilis: signature nucleotide bias and unique nad4L gene rearrangement within crinoids

Complete DNA sequences have been determined for the mitochondrial genomes of the crinoids Phanogenia gracilis (15892 bp) and Gymnocrinus richeri (15966 bp). The mitochondrial genetic map of the stalkless feather star P. gracilis is identical to that of the comatulid feather star Florometra serratissima (Scouras, A., Smith, M.J., 2001. Mol. Biol. Evol. 18, 61-73). The mitochondrial gene order of the stalked crinoid G. richeri differs from that of F. serratissima and P. gracilis by the transposition of the nad4L protein gene. The G. richeri nad4L mitochondrial map position is unique among metazoa and is likely a derived feature in this stalked crinoid. Nucleotide compositional analyses of protein genes encoded on the major sense strand confirm earlier conclusions regarding a crinoid-distinctive T over C bias. All three crinoids exhibit high T levels in third codon positions, whereas other echinoderm classes favor A or C in the third codon position. The nucleotide bias is reflected in the relative synonymous codon usage patterns of crinoids versus other echinoderms. We suggest that the nucleotide bias of crinoids, in comparison to other echinoderms, indicates that a physical inversion of the origin of replication has occurred in the crinoid lineage. Evolutionary rate tests support the use of the cytochrome b (cob) gene in molecular phylogenetic analyses of echinoderms. A consensus echinoderm tree was generated based on cytochrome b nucleotide alignments that placed the asteroids as a sister group to a clade containing the ophiuroids and the (echinoids+holothuroids) with the crinoids basal to the rest of the echinoderm classes: [Crinoid,(Asteroid,(Ophiuroid,(Echinoid,Holothuroid)))].     

Diversity dynamics of post‐Palaeozoic crinoids – in quest of the factors affecting crinoid macroevolution

Following the end‐Permian biotic crisis which led to the near extinction of crinoids, this echinoderm class rebounded rapidly during the Mesozoic, resulting in forms with important morphological and behavioural novelties. However, quantitative patterns of crinoid diversity during the Mesozoic remain largely unexplored. Here, we report results of analyses of the evolutionary dynamics of post‐Palaeozoic crinoid genera spanning a time interval between 250 and 70 Myr. We show that crinoids reached their Mesozoic peak of genus‐level richness during the Late Jurassic. We also document a major reorganization of different ecological crinoid groups in the Mesozoic. More specifically, the diversity of sessile forms generally increased towards the mid‐Mesozoic but decreased significantly starting in the Cretaceous, whereas the number of motile crinoid genera increased linearly during the Mesozoic. The possible role of biotic and abiotic factors in crinoid evolution is discussed.     

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.     

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.     

Early phylogeny of crinoids within the pelmatozoan clade

Phylogenetic relationships among early crinoids are evaluated by maximizing parsimonious‐informative characters that are unordered and unweighted. Primarily Tremadocian–Darriwilian (Early–Middle Ordovician) taxa are analysed. Stratigraphic congruence metrics support the best phylogenetic hypothesis derived using parsimony methods. This study confirms the traditionally recognized lineages of Palaeozoic crinoids and provides new information on the branching order of evolving lineages. Camerates are basal crinoids with progressively more tipward groups (from an Ordovician perspective) being protocrinoids, cladids (paraphyletic), hybocrinids and disparids. The Protocrinoida should be maintained, but the Aethocrinida should be placed within the Cladida. The results of this study identify phylogenetic structure amongst the major early crinoid lineages and delineate the relative positions of crinoid higher taxa along a tree. Each valid higher taxon discussed herein requires a comprehensive treatment to delimit within‐lineage phylogenetic relationships.     

Taphonomy of Lower Carboniferous crinoids from the Hook Head Formation, Ireland

Ausich, W.I. & Sevastopulo, G.D. 1994 10 15: Taphonomy of Lower Carboniferous crinoids from the Hook Head Formation, Ireland.
Lower Carboniferous strata at Hook Head, County Wexford, Ireland, record tempestite deposition on a mixed carbonate and siliciclastic shelf-ramp. Crinoidal remains occur in all facies, which allows for a comparative taphonomic analysis of crinoid preservation. Preservation of crinoid crowns was controlled by a balance between mean storm intensity and frequency. The best preservation occurs at moderate depths. Taphonomic resistance among Hook Head crinoids is interpreted as follows (from most to least resistant to disarticulation): monobathrid camerates -cladids - diplobathrid camerates — flexibles (with Platycrinites and Dichocrinus disarticulation more like cladids). Crinoidea, taphonomy, preservation, Lower Carboniferous, Ireland .     

Functional morphologies of the columns of Upper Ordovician Xenocrinus and Dendrocrinus

Two common Upper Ordovician crinoids, Xenocrinus and Dendrocrinus , had distinctive columns that showed marked contrasts of both form and function. Xenocrinus baeri (Meek) had a tetragonal facet geometry but functioned in the normal manner for a symplectially articulated column. Column flexure was by bending, the four directions of curvature being limited by the tetragonal arrangement of crenulae. In contrast, columns of Dendrocrinus casei Meek, with a characteristic pentastellate symmetry, were able to twist, a functional adaptation not previously reported from crinoids; this was controlled by the unusual geometry of the crenularium. This suggests that some or all crinoid columns may be subject to twisting stresses, perhaps associated with changes in the current direction, that crenulae may resist passively. Functional morphology, column, crinoids, X enocrinus , D endrocrinus     

Host‐specific acrothoracid barnacles on Middle Devonian platyceratid gastropods

Borings, attributed to acrothoracic barnacles, occur on the platyceratid gastropod Naticonema lineatum (Conrad) from the Middle Devonian Hamilton Group of western New York and rarely in specimens as old as the Early Devonian. These latter are the oldest known acrothoracid borings are in the fossil record. The borings are consistently developed as laterally compressed, inequilateral pouches exclusively on these gastropods, commonly as dense infestations.

Naticonema shells yielding borings typically occur associated with partially articulated qrinoid remains, and they are sometimes found attached to crinoids in a manner similar to coprophagous Platyceras. In addition to barnacle borings, Naticonema shells often bear thin encrustations of bryozoans which are usually perforated by these borings but sometimes also overgrow them. Barnacles bored live hosts; gastropods prevented shell penetration by producing cyst‐like secondary secretions of calcite beneath acrothoracid boreholes.

The relative antiquity of these borings and their association with coprophagous platyceratids makes their discovery particularly significant in revealing aspects of the early ecology of barnacles. Attachment to the host commensal gastropods was one of the first successful life modes of these crustaceans prior to their later diversification to other habitats. Mississippian and Pennsylvanian occurrences of similarly bored gastropods demonstrate continuity of the barnacle‐gastropod‐crinoid ecological association from the Middle to Late Paleozoic.