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.     

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.     

Phosphagen kinase evolution. Expression in echinoderms

Arginine kinase and creatine kinase that catalyze the transfer of a phosphate group between ATP and arginine and creatine, respectively, play an important role in cellular energetics. In contrast to most animals which exhibit a single phosphagen kinase activity (creatine kinase in chordates and arginine kinase in protostomians), echinoderms exhibit both arginine kinase and creatine kinase activities, sometimes in the same tissue. In contrast to chordates in which creatine kinases are dimers (consisting of two subunits of 40 kDa) and protostomians in which arginine kinases are usually monomers (40 kDa), echinoids contain specific phosphagen kinases: a dimeric arginine kinase (consisting of two subunits of 42 kDa) in eggs and a monomeric creatine kinase (145 kDa) in sperm. We have examined echinoderms from the five existing classes (echinoids, asteroids, ophiuroids, holothurians and crinoids) for the expression of these specific phosphagen kinases in different tissues. Gel filtration was used to determine the molecular masses of the native enzymes. Antibodies specific for arginine kinase or for creatine kinase were used to characterize the subunit composition of arginine kinase and creatine kinase after SDS/PAGE and transfer. In all echinoderms analyzed, arginine kinase always occurred as an enzyme of about 81 kDa consisting of two subunits of 42 kDa and creatine kinase as a monomeric enzyme of 140-155 kDa. The occurrence in echinoderms of both phosphagen kinases with distinct specificities and specific molecular structures is discussed from both a developmental and evolutionary point of view.     

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.     

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.     

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.     

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.     

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)))].     

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.     

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.     

The interrelationships of the echinoderm classes: morphological and molecular evidence

The relationship between the five echinoderm classes has perplexed phylogeneticists for some time. Although each of the crinoids (C), asteroids (A), ophiuroids (O), echinoids (E) and holothuroids (H) are morphologically distinct, evidence from larval and adult morphology, molecular biology, and stratigraphy have failed to provide a single consensus solution. We have reviewed all available morphological and molecular data, added new data and reanalysed independent data sets individually and in combination, in order to resolve echinoderm class relationships. In total, we present 21 larval and 50 adult morphological characters, partial 28S-like large subunit rRNA gene data for 39 taxa and complete 18S-like small subunit rRNA gene data for 37 taxa. For a 5 taxon problem there are 105 possible rooted tree topologies, and yet we were consistently presented with three competing hypotheses when data sets were analysed both individually and in combination. The total evidence solution favoured (outgroup(C(A(O(E, H))))) although the alternative tree topology, (outgroup(C(O(A(E, H))))) was only one step longer and (outgroup(C((A, O),(E, H)))) was two steps longer. Only these three trees are serious contenders and the distribution of morphological characters suggests we should discount the solution placing ophiuroids as sister group to an asteroid+echinoid + holothurian clade. Thus we are left with (outgroup(C(A(O(E, H))))) and (outgroup(C((A, O),(E, H)))) as the two most plausible phylogenetic hypotheses. Our data showed high levels of phylogenetic signal and these trees best fit the available data.     

Diversity and abundance of echinoderms from the reef lagoon at Cahuita National Park, Caribbean from Costa Rica

A total of 15 species of echinoderms (one asteroid, seven ophiuroids, five echinoids and two holothurians) were recorded at the Cahuita National Park reef lagoon, between September and October 2003, using a 1 m2 quadrant. The sites with coral substrate and algae were the most diverse, while those with seagrass and sand were the least. Ophiuroids were the most abundant group with 170 individuals, asteroids were the least abundant. Adding other studies and reports of echinoderms to this study, a total of 23 species have been found at Cahuita National Park, which is the most diverse area on the Caribbean of Costa Rica. We report nine new echinoderm records for Costa Rica's Caribbean.     

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.     

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.     

Field sampling bias, museum collections and completeness of the fossil record

Museum specimens, particularly old collections, typically lack comprehensive field data and determination of substrate, sampling biases, etc., is problematic. Diversity at the generic level of all identifiable latest Cretaceous (Campanian–Maastrichtian) echinoderm remains in major museum collections from the Mons (southern Belgium) and Danish (Jylland (Jutland) and Sjælland (Zealand)) basins were compared to those of the Liège-Limburg Basin. The last-named has been studied in detail, including microscopical analysis of ossicles picked from bulk samples. Echinoids of the Mons Basin show similarities to those of the Liège-Limburg Basin, but crinoids, asteroids and ophiuroids remain poorly known from the former. Echinoderms of the Danish Basin resemble those of similar chalk lithofacies in the Liège-Limburg Basin, despite significant geographical separation. These disparities can be explained, at least in part, by collector bias in sampling methodology, although differences in substrate presumably also had an influence.     

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.     

Development of the nervous system in the brittle star Amphipholis kochii

There are several studies of neural development in various echinoderms, but few on ophiuroids, which develop indirectly via the production of pluteus larvae, as do echinoids. To determine the extent of similarity of neuroanatomy and neural development in the ophiuroids with other echinoderm larvae, we investigated the development of the nervous system in the brittle star Amphipholis kochii (Echinodermata: Ophiuroidea) by immunohistochemistry. Immunoreactive cells first appeared bilaterally in the animal pole at the late gastrula stage, and there was little migration of the neural precursors during A. kochii ontogeny, as is also the case in echinoids and holothuroids. On the other hand, neural specification in the presumptive ciliary band near the base of the arms does occur in ophiuroid larvae and is a feature they share with echinoids and ophiuroids. The ophiopluteus larval nervous system is similar to that of auricularia larvae on the whole, including the lack of a fine network of neurites in the epidermis and the presence of neural connections across the oral epidermis. Ophioplutei possess a pair of bilateral apical organs that differ from those of echinoid echinoplutei in terms of relative position. They also possess coiled cilia, which may possess a sensory function, but in the same location as the serotonergic apical ganglia. These coiled cilia are thought to be a derived structure in pluteus-like larvae. Our results suggest that the neural specification in the animal plate in ophiuroids, holothuroids, and echinoids is a plesiomorphic feature of the Ambulacraria, whereas neural specification at the base of the larval arms may be a more derived state restricted to pluteus-like larvae.     

Morphologically Specialized Sperm from the Ovary of Isometra vivipara (Echinodermata — Crinoidea)

The morphology of the sperm of Isometra vivipara is markedly different from the spherical-headed sperm typical of crinoids in general. The sperm head of Isometra has a flattened, oval shape. In addition to a nucleus, the head includes a lateral acrosome on one flat surface and a lateral mitochondrion on the opposite flat surface. From the latter surface of the sperm head, a tail flagellum arises adjacent to the mitochondrion. The discussion considers the relationships between reproductive habits and sperm morphology in Isometra and other crinoids.