What makes an ophiuroid? A morphological study of the problematic Ordovician stelleroid Stenaster and the palaeobiology of the earliest asteroids and ophiuroids

Extant asteroids and ophiuroids [EchinodermataJ are distinguished by differences in arm support, water vascular system structures and in details of arm and jaw structure. However, some lower Palaeozoic taxa show combinations of both asteroid-like and ophiuroid-like characters and their morphology and functional biology is poorly understood. This paper redescribes one such taxon, the middle-upper Ordovician stellate echinoderm Stenaster and clarifies its phylogenetic status. Characters in common with extant and Ordovician ophiuroids, include arm support due primarily to ambulacral ossicles, presence of extensive longitudinal arm musculature, a mobile jaw and an internalised radial water vessel with internalised podial pores. In addition, Stenaster lacks several characters which are conventionally considered to be asteroid-like, for example an axillary, madreporitc, marginal ossicles and a true ambulacral groove. However, in overall shape Stenaster is remarkably asteroid-like, showing short, broad-based arms shared podial basins and a small disc. A cladistic analysis of early asteroids, ophiuroids and somasteroid taxa consistently places Stenaster within the ophiuroids and suggests secondary convergence to asteroids. In functional terms, Stenaster is interpreted as an ophiuroid which has secondarily adopted a semi-infaunal, deposit-feeding mode of life, analogous to that of some extant paxillosid asteroids.     

Towards a Better Understanding of the Origins of Microlens Arrays in Mesozoic Ophiuroids and Asteroids

Echinoderms are characterized by a calcite endoskeleton with a unique microstructure, which is optimized for multiple functions. For instance, some light-sensitive ophiuroids (Ophiuroidea) and asteroids (Asteroidea) possess skeletal plates with multi-lens arrays that are thought to act as photosensory organs. The origins of these lens-like microstructures have long been unclear. It was recently proposed that the complex photosensory systems in certain modern ophiuroids and asteroids could be traced back to at least the Late Cretaceous (ca. 79 Ma). Here, we document similar structures in ophiuroids and asteroids from the Early Cretaceous of Poland (ca. 136 Ma) that are approximately 57 million years older than the oldest asterozoans with lens-like microstructures described thus far. We use scanning electron microscopy, synchrotron tomography, and electron backscatter diffraction combined with focused ion beam microscopy to describe the morphology and crystallography of these structures in exceptional detail. The results indicate that, similar to Recent light-sensitive ophiuroids, putative microlenses in Cretaceous ophiuroids and asteroids exhibit a shape and crystal orientation that would have minimized spherical aberration and birefringence. We suggest that these lens-like microstructures evolved by secondary deposition of calcite on pre-existing porous tubercles that were already present in ancestral Jurassic forms.     

Resolving phylogenetic signal from noise when divergence is rapid: a new look at the old problem of echinoderm class relationships

Resolving evolutionary relationships in groups that underwent fast radiation in deep time is a problem for molecular phylogeny, as the scant phylogenetic signal that characterises short internal branches is generally swamped by more recent substitutions. We implement an approach, that maps how the support for rival phylogenies changes when analysing subsets of sites with either faster and more heterogeneous rates or slower and more homogeneous rates, to address a long-standing problem in deuterostome phylogeny - the interrelationships of the eleutherozoan echinoderm classes. We show that miRNA genes are phylogenetically uninformative as to the relationships of asteroids, echinoids and ophiuroids, consistent with a rapid radiation of these groups as suggested by their fossil record. Using three nuclear rRNAs and seven nuclear housekeeping genes, we map the support for the three possible phylogenetic arrangements of asteroids, ophiuroids and echinoids when moving between subsets of the data with very similar or very different rates of evolution. Only one of the three possible topologies (asteroids (ophiuroids + echinoids)) strengthens when the most rate-homogeneous subset of data are analysed. The other two possible pairings become stronger in a less reliable data subset, which includes the fastest and thus homoplasy-rich data in our alignment. Thus, while superficial analysis of our concatenated alignment identifies asteroids and ophiuroids as sister taxa, more thorough analyses suggest that ophiuroids may be more closely related to echinoids. Divergence of these echinoderm groups, using a relaxed molecular clock, is estimated to have occurred within ∼5 million years. Our results illustrate that the analytic approach of phylogenetic signal dissection can be a powerful tool to investigate rapid radiations in deep geologic time.     

Electron microscopic study of the cortical reaction of an ophiuroid echinoderm.

The egg coats of an ophiuroid echinoderm (Ophiopholis aculeata) are described by electron microscopy before and after fertilization. The unfertilized egg is closely invested by a vitelline coat about 40 A thick, and the peripheral cytoplasm is crowded with cortical granules five or six deep. During the cortical reaction, which rapidly follows insemination, exocytosis of cortical granules takes place. Some of the cortical granule material is evidently added to the vitelline coat to form a composite structure, the fertilization envelope, which is made up of a 400 A thick middle layer separating inner and outer dense layers, each about 50 A thick. The elevation of the fertilization envelope from the egg surface creates a perivitelline space in which the hyaline layer soon forms. The hyaline layer is about 2 micron thick, finely granular, and apparently derived from cortical granule material. The extracellular layers of the early developmental stages of ophiuroids and echinoids are quite similar in comparison to those of asteroids; this finding helps support Hyman's argument that the ophiuroids are more closely related to the echinoids than to the asteroids.     

Comparative Morphology of Echinoderm Sperm and Possible Phylogenetic Implications

The sperm morphology of at least 7 crinoids, 11 ophiuroids,23 asteroids, 23 holothuroids, and 37 echinoids is known. Forall the known crinoids, ophiuroids, and asteroids, the spermare spherical, while those of echinoids are conical. Holothuroidsperm are basically spherical, with two exceptions: cylindricalin Cucumaria lubrica and tabloid in Cucumaria pseudocurata.The possible implications of sperm morphology in the mode offertilization and in echinoderm phylogeny are discussed.     

Larval budding, metamorphosis, and the evolution of life-history patterns in echinoderms

Abstract. The oral surface and mouth of juvenile asteroids and echinoids with indirect development forms on the lower left side of the larval body, thus establishing a new axis of body symmetry. In contrast, the juvenile mouth of ophiuroids and holothuroids develops from the larval one, and the larval and adult body axes roughly coincide. Explaining how two such disparate modes of development arose in evolution has been a perennial problem for echinoderm biologists, but recent observations on larval budding in asteroids may provide an answer. The juvenile mouth of asteroids forms near the base of the left posterolateral lobe. The posterolateral lobes are also the principal site of bud formation in asteroid larvae that propagate asexually, and buds form mouths. By accelerating the development of oral and ectodermal structures belonging to the bud, and combining these with internal organs derived from the parent larva, a composite individual could be constructed with the same orientation and positioning as the juvenile rudiment in asteroids. Whether this also explains the position of the juvenile rudiment in echinoids is a more complex question, depending in part on whether asexual propagation is derived, and restricted to asteroids and ophiuroids, or is more primitive and hence widespread among stem echinoderms.     

Echinoderms from Marino Ballena National Park, Pacific, Costa Rica

A total of 25 species of echinoderms (four asteroids, six ophiuroids, five echinoids and ten holothurians) were recorded at Marino Ballena National Park, using 25 m2 quadrants, parallel to the coast, at seven sites. The ophiuroids were the most abundant group with 581 individuals and the asteroids the less abundant (48 individuals). Echinoderms densities were low, with the exception of the ophiuroids. Diversity, density and the number of groups were higher where sedimentation was lower. We suggest that sedimentation is having a negative effect on the diversity of echinoderms and on the development of the coral reefs in this park.     

Preservation of tube feet in an ophiuroid (Echinodermata) from the Lower Devonian Hunsrück Slate of Germany and a redescription of<Emphasis Type="Italic">Bundenbachia beneckei</Emphasis> and<Emphasis Type="Italic">Palaeophiomyxa grandis</Emphasis>

The preservation of non-mineralized tissues in the fossil record is extremely rare. The Lower Devonian Hunsrück Slate of Germany has long been known for the preservation of non-mineralized tissues in pyrite but whether or not these remnants represent true soft tissues has been questioned. This is especially true for struetures visible only on radiographs that are too delicate for excavation by traditional methods. Here we report the discovery of well-preserved pyritized tube feet in six fully prepared specimens of the protasterid brittle starBundenbachia beneckei from the Hunsrück Slate. This discovery represents the first report of fossilized ophiuroid tube feet in the fossil record. The successful excavation of the delicate tube feet was made possible by improved airbrasive techniques developed by German fossil collectors. The relatively large size of the fossil tube feet inBunden-bachia beneckei is consistent with earlier inferences on size based on the presence of large podial basins. Protasterid ophiuroids lack the specialized arm musculature and articulations that provide increased flexibility and strength to the arms of modern ophiuroids with typically reduced tube feet. How-ever, tube foot form and perhaps function inBundenbachia might have been similar to those of living asteroids in which large tube feet are used primarily for locomotion and food-manipulation thus compensating for a lack of specialized arm musculature and articulation. Hence, feeding and life mode of protasterid ophiuroids was not necessarily limited to sedentary, infaunal microphagy as traditionally suggested. Two Hunsrück protasterid ophiuroids,Bundenbachia benecki andPalaeophiomyxa grandis are redescribed and compared.      

Asteroids,ophiuroids and holothurians from the southeastern Weddell Sea (Southern Ocean)

Until the early 1980s, the composition and distribution of the asteroid (starfish), ophiuroid (brittle star) and holothurian (sea cucumber) bottom fauna of the southeastern Weddell Sea was virtually unknown. This southernmost part of the Atlantic sector of the Southern Ocean is a typical high-latitude Antarctic region located in the circumpolar permanent pack-ice zone. It became accessible for large-scale scientific surveys only through the availability of modern ice-breaking research vessels, such as the German RV “Polarstern”. Here, we describe a dataset of the faunal composition and abundance of starfish, brittle star and sea cucumber assemblages in this area, based on collections from trawl catches carried out during three “Polarstern” cruises in 1983, 1984 and 1985. The set comprises a total of 4,509 records of abundances of 35 asteroid species (with a total of 2,089 specimens) and 38 ophiuroid species (with a total of 18,484 specimens) from 34 stations, as well as of 66 holothurian species (with a total of 20,918 specimens) from 59 stations including zero-abundances (absences). A synthesizing zoogeographical community analysis confirms the presence of three distinct assemblages of asteroids, ophiuroids, and holothurians with highest species richness on the eastern shelf. Overall, starfishes, brittle stars and sea cucumbers were present at all sites investigated in the study area but composition and abundance of asterozoan (asteroids and ophiuroids together) and holothurian fauna varied considerably. A synthesizing zoogeographical community analysis confirms the presence of three distinct assemblages of asteroids, ophiuroids, and holothurians with highest species richness on the eastern shelf. In the case of asterozoans, water depth and latitude seemed to be the most important drivers of assemblage distribution and composition. One of the holothurian assemblages was part of the rich macrozoobenthic community dominated by a diverse and abundant epifauna, mainly sponges and gorgonians. Another one was mainly composed of vagrant deposit-feeding species inhabiting a predominantly non-colonised substratum. In addition, a mixed holothurian assemblage was identified.     

A new ciliary feeding structure in an ophiuroid echinoderm

The genital shields which form the walls of the bursal slits in Ophiura texturata are covered by a precisely orientated arrangement of ciliated ridges and non-ciliated grooves. An electron microscopic examination has revealed many mucous cells associated with this structure and a catecholamine-containing nerve plexus underlying it. An examination of the currents produced by this ciliated structure suggest that it is associated with suspension feeding and preliminary results indicate that the secretion of mucus is under neurol control. Specialized structures of this type have not been previously described in ophiuroids and are only present in the members of certaln families. The interest in these structures is not just in relation to feeding mechanisms in ophiuroids but they also provide a useful specialized preparation for the study of some aspects of the function of the subepidermal nerves in echinoderms.     

Sea urchins, sea stars and brittle stars from Terra Nova Bay (Ross Sea, Antarctica)

Although echinoderms constitute some of the most conspicuous taxa of the Antarctic benthic communities, the echinoderm fauna of Terra Nova has not been described yet. The present study provides the first species list of echinoids, ophiuroids and asteroids from Terra Nova Bay (30–500 m depth) and describes the depth distribution of these species. Preliminary observations of the summer reproductive condition of some of the species are also included.     

Toxicity of shallow-water Antarctic echinoderms

Summary The toxicity of 23 species of shallow-water antarctic echinoderms from McMurdo Sound was investigated using Gambusia affinis (Vertebrata: Pisces) as a test organism. Ichthyotoxicity assays were conducted on the body wall tissues of thirteen species of asteroids and three species of holothuroids, the tests of three species of echinoids and the arms of three species of ophiuroids and one species of crinoid. Patterns of toxicity were class-specific, with varying degrees of toxicity occurring exclusively among the Asteroidea and Holothuroidea. Seven of the thirteen species of asteroids were mildly to highly toxic (54%), while one of the three holothuroids was mildly and one highly toxic. Toxicity was not related to the level of calcification of the body wall and energy level in asteroids. The only armored asteroid, Notasterias armata, was non-toxic. The occurrence of toxins in the body tissues of antarctic echinoderms is similar in pattern and frequency with that of temperate and tropical species.     

What a New Model of Skeletal Homologies Tells Us About Asteroid Evolution

The Extraxial-Axial Theory (EAT) is applied to the body wallhomologies of asteroids. Attempts to characterize major platesystems of asteroids as axial or extraxial, particularly thosethat are highly organized into series, can be problematic. However,the Optical Plate Rule (OPR) is instrumental in establishingthat ambulacrals and terminals are axial. It is equally clearthat the region aboral to the marginal frame is a part of theperforate extraxial body wall (with the possible exception ofthe centrodorsal, which is likely imperforate extraxial). Previouslyestablished EAT criteria, particularly those strongly rootedin the embryologically expressed boundary between axial andextraxial body wall in larvae, suggest that marginals, and perhapsadambulacrals, are extraxial in origin. We also explore theextraxial nature and phylogenetic significance of the odontophore.Our data from both juveniles and adults show that plate andtube foot addition sequences occur according to the OPR, andshed light on poorly known homologies of the asteroid mouthframe. These data indicate that the mouth angle ossicle mustat least contain the first ambulacral, although we cannot ruleout the possibility that the first adambulacral also contributesto the construction of this ossicle. The interpretations providedby the EAT for all ossicles suggest a synapomorphy scheme forsomasteroids, ophiuroids, and asteroids.     

Records of echinoderms (excluding holothurians) from the Norfolk Ridge and Three Kings Rise north of New Zealand

Abstract

Fifty-eight species of echinoderms (excluding holothurians) are newly recorded from the southern Norfolk Ridge and the Three Kings Rise to the north of New Zealand. Two species of Asteroidea, in the genera Ceramaster and Pseudoceramaster, ceramaster are described as new. A total of 115 species of echinoderms, not counting the holothurians, are now known from this area. They include 14 crinoids, 35 asteroids, 41 ophiuroids, and 25 echinoids.     

Occluding-like junctions at mesaxons of central myelin in Anolis carolinensis are not 'tight'. A freeze-fracture-protein tracer analysis.

The junctional complexes of the myelin sheath of central nervous system axons in the American chameleon, Anolis carolinensis, exhibit an intramembrane ridge and groove construction in freeze-fracture replicas that has usually been interpreted in other organisms as evidence for an occluding or tight intercellular junction. Close examination of PF fracture face ridges, however, shows them to be made up of discontinuous rows of particles of variable length separated by frequent gaps of non-uniform width. Introduction of horseradish peroxidase into the intercellular milieu of the lizard central nervous system is followed by appearance of this protein in interlamellar spaces of the myelin sheath and in the intercellular spaces containing focal membrane fusions that correspond precisely in position and center-to-center spacing to the ridges and grooves in platinum replicas of the same tissue. Since the junctional ridges on PF fracture faces in these mesaxonal junctional complexes are conspicuously discontinuous and since the areas within the myelin sheath where these junctional complexes are located inner and outer mesaxons) are readily permeated by exogenous protein tracer, it is concluded that the junctional complexes of central myelin mesaxons, heretofore incorrectly interpreted as functionally tight, are actually very leaky and probably contribute only to the structural stability of the myelin sheath architecture.     

Phylogenomic analysis of echinoderm class relationships supports Asterozoa

While some aspects of the phylogeny of the five living echinoderm classes are clear, the position of the ophiuroids (brittlestars) relative to asteroids (starfish), echinoids (sea urchins) and holothurians (sea cucumbers) is controversial. Ophiuroids have a pluteus-type larva in common with echinoids giving some support to an ophiuroid/echinoid/holothurian clade named Cryptosyringida. Most molecular phylogenetic studies, however, support an ophiuroid/asteroid clade (Asterozoa) implying either convergent evolution of the pluteus or reversals to an auricularia-type larva in asteroids and holothurians. A recent study of 10 genes from four of the five echinoderm classes used ‘phylogenetic signal dissection’ to separate alignment positions into subsets of (i) suboptimal, heterogeneously evolving sites (invariant plus rapidly changing) and (ii) the remaining optimal, homogeneously evolving sites. Along with most previous molecular phylogenetic studies, their set of heterogeneous sites, expected to be more prone to systematic error, support Asterozoa. The homogeneous sites, in contrast, support an ophiuroid/echinoid grouping, consistent with the cryptosyringid clade, leading them to posit homology of the ophiopluteus and echinopluteus. Our new dataset comprises 219 genes from all echinoderm classes; analyses using probabilistic Bayesian phylogenetic methods strongly support Asterozoa. The most reliable, slowly evolving quartile of genes also gives highest support for Asterozoa; this support diminishes in second and third quartiles and the fastest changing quartile places the ophiuroids close to the root. Using phylogenetic signal dissection, we find heterogenous sites support an unlikely grouping of Ophiuroidea + Holothuria while homogeneous sites again strongly support Asterozoa. Our large and taxonomically complete dataset finds no support for the cryptosyringid hypothesis; in showing strong support for the Asterozoa, our preferred topology leaves the question of homology of pluteus larvae open.     

Microvascularization on collared peccary placenta: a microvascular cast study [corrected] in late pregnancy

The microvascularization of the collared peccary (Tayassu tajacu) placenta was studied by vascular casts and immunolocalization of α-smooth muscle actin and vimentin, to identify the three dimensional organization and vascular flow interrelation in the microvasculature between the maternal and fetal compartments of the placentae. The immunolocalization of vimentin in the vascular endothelium and in the smooth muscle cells of blood vessels showed indented capillaries along the uterine epithelium and the trophoblast at the sides of complementary maternal and fetal microfolds, or rugae. This confers the three-dimensional structure observed in vascular casts. On the maternal side, casts demonstrated uterine folds coated by with primary and secondary ridges, and by areolae dispersed between these ridges. The arteriole runs through the center/middle of ridges, branching at the top into a microvascular network wall in a basket-like fashion. At the base of these baskets venules were formed. On the fetal side, arterioles branched centrally in the fetal rugae into a capillary network in a bulbous form, complementary to the opposite maternal depressions forming the baskets. At the base of the bulbous protrusions, the fetal venules arise. The blood vessel orientation in the materno-fetal interface of the placentae of collared peccaries suggests a blood flow pattern of the type countercurrent to cross current. The same pattern has been reported in domestic swine demonstrating that, even after 38 million years, the Tayassuidae and Suidae families exhibit similar placental morphology, which is here characterized at the microvascular level.     

Energetic costs of loss and regeneration of arms in stellate echinoderms

Loss of arms has energetic consequences for stellate echinoderms (crinoids, ophiuroids, and asteroids). The energetic cost of losing an arm includes loss of investment, decrease in ability to obtain nutrients and allocation of nutrients to regeneration of the lost arms at a cost to other body compartments. The cost to other body compartments is low when food availability is very low or very high. The cost becomes apparent when food availability is sufficient to support production but not high enough that the cost of regeneration has no effect on production of other body compartments. Loss of investment is greater in asteroids than in crinoids and ophiuroids because of greater development of the body wall and presence of gonads and pyloric caeca in the arms. The cost of regeneration of organic matter in an arm can be estimated from the amount of organic matter present in intact arms and the cost of anabolism. Protein production is the primary cost of regeneration of an arm because of the high concentration of protein in the regenerated arm and the high anabolic cost of protein production. A major energetic cost of loss of arms that affects regeneration is decrease in food consumption. It is necessary to separate cost of decrease in consumption from cost of regeneration. Comparison of intact and regenerating individuals requires they consume the same amount of food. The cost of regeneration will also be affected by the quality of food because of the nutrient requirements for growth. Because the quantity and quality of the food ingested is not known, it is not possible to quantify the cost of regeneration in the field. Asteroids appear to be a good model for the study of regeneration in the laboratory because it is possible to control the quantity and quality of food they ingest. They are also a good model for the study of the evolutionary significance of regeneration by comparing individuals that have lost arms and are regenerating them to those that have lost arms and are not. The difference in the frequency of loss of arms of species is related to the difference in availability of food and the ability to feed that affect the capacity for re-investment in the lost arm. This is important in considering life-history strategies.     

Molecular phylogenetic analyses indicate multiple independent emergences of parasitism in Myzostomida (Protostomia)

The fossil record indicates that Myzostomida, an enigmatic group of marine worms, traditionally considered as annelids, have exhibited a symbiotic relationship with echinoderms, especially crinoids, for nearly 350 million years. All known extant myzostomids are associated with echinoderms and infest their integument, gonads, celom, or digestive system. Using nuclear (18S rDNA) and mitochondrial (16S and COI) DNA sequence data from 37 myzostomid species representing nine genera, we report here the first molecular phylogeny of the Myzostomida and investigate the evolution of their various symbiotic associations. Our analyses indicate that the two orders Proboscidea and Pharyngidea do not constitute natural groupings. Character reconstruction analyses strongly suggest that (1) the ancestor of all extant myzostomids was an ectocommensal that first infested crinoids, and then asteroids and ophiuroids, and (2) parasitism in myzostomids emerged multiple times independently.