Rate of regeneration of two arms in the field and its effect on body components in Luidia clathrata (Echinodermata: Asteroidea)

The incidence of arm regeneration, effects of arm regeneration on non-regenerating body components and rate of arm regeneration were examined under field conditions. Approximately 60% of Luidia clathrata found in the upper portion of Tampa Bay, Florida were regenerating one or more arms. Individuals with two arms amputated at the disk edge took 380 days to regenerate completely. Both arms of an individual regenerated at the same rate. Length of regenerating arms initially increased faster than dry mass. No difference was found between non-regenerating and regenerating individuals for the dry mass of the body wall, pyloric caeca, and gonad of a non-regenerating arm. The absolute masses of the gonad and pyloric caeca were low through out the year possibly indicating a low food environment. Based on the rate of regeneration, arm loss just prior to the reproductive season may reduce the reproductive potential of this species.     

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.     

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.     

Arm regeneration in two species of cuttlefish Sepia officinalis and Sepia pharaonis

To provide quantitative information on arm regeneration in cuttlefish, the regenerating arms of two cuttlefish species, Sepia officinalis and Sepia pharaonis, were observed at regular intervals after surgical amputation. The third right arm of each individual was amputated to ~10–20 % starting length. Arm length, suction cup number, presence of chromatophores, and behavioral measures were collected every 2–3 days over a 39-day period and compared to the contralateral control arm. By day 39, the regenerating arm reached a mean 95.5 ± 0.3 % of the control for S. officinalis and 94.9 ± 1.3 % for S. pharaonis. The process of regeneration was divided into five separate stages based on macroscopic morphological events: Stage I (days 0–3 was marked by a frayed leading edge; Stage II (days 4–15) by a smooth hemispherical leading edge; Stage III (days 16–20) by the appearance of a growth bud; Stage IV (days 21–24) by the emergence of an elongated tip; and Stage V (days 25–39) by a tapering of the elongated tip matching the other intact arms. Behavioral deficiencies in swimming, body postures during social communication, and food manipulation were observed immediately after arm amputation and throughout Stages I and II, returning to normal by Stage III. New chromatophores and suction cups in the regenerating arm were observed as early as Stage II and by Stage IV suction cup number equaled that of control arms. New chromatophores were used in the generation of complex body patterns by Stage V. These results show that both species of cuttlefish are capable of fully regenerating lost arms, that the regeneration process is predictable and consistent within and across species, and provide the first quantified data on the rate of arm lengthening and suction cup addition during regeneration.     

Regeneration,predatory–prey interaction,and evolutionary history of articulate crinoids

Regeneration and predatory–prey interaction of crinoids are reviewed. Crinoids have strong powers of regeneration, and arm regeneration is common in fossil and Recent crinoids. Regenerated arms commonly start from the ligamentary articulation called syzygy or cryptosyzygy, where crinoids can autotomize their arms. Therefore, regenerated arms can be formed after loss of arms by autotomy of arms, and such autotomy is commonly the response to predatory attacks. Thus, regenerated arms can be used as the clue to estimate the predatory frequencies. Regeneration of “correct” skeletal morphology as in the original depends on the existence of adoral nerve center. A stalk without the adoral nerve center cannot regenerate the “correct” morphology of the original skeleton, but forms of “callus” as skeletal overgrowth. The strong ability of regeneration is a key factor of the success of articulate crinoids in the geologic history since the Triassic onward.     

Wound healing and arm regeneration in Ophioderma longicaudum and Amphiura filiformis (Ophiuroidea, Echinodermata): comparative morphogenesis and histogenesis

All species of the Ophiuroidea have exceptional regenerative capabilities; in particular, they can replace arms lost following traumatic or self-induced amputation. In order to reconstruct this complex phenomenon, we studied arm regeneration in two different ophiuroids, Ophioderma longicaudum (Retzius, 1805) and Amphiura filiformis O. F. Müller, 1776, which are quite distantly related. These species present contrasting regeneration and differentiation rates and differ in several ecological traits. The aim of this paper is to interpret the primary sequence of morphogenetic and histogenetic events leading to the complete reconstruction of a new arm, comparing the arm regenerative processes of these two ophiuroid species with those described in crinoids. Arm regeneration in ophiuroids is considered an epimorphic process in which new structures develop from a typical blastema formed from an accumulation of presumptive undifferentiated cells. Our results showed that although very different in some respects such as, for instance, the regeneration rate (0.17 mm/week for O. longicaudum and 0.99 mm/week for A. filiformis), morphogenetic and histogenetic aspects are surprisingly similar in both species. The regenerative process presents similar characteristics and follows a developmental scheme which can be subdivided into four phases: a repair phase, an early regenerative phase, an intermediate regenerative phase and an advanced regenerative phase. In terms of histogenesis, the regenerative events involve the development of new structures from migratory pluripotent cells, which proliferate actively, in addition in both cases there is a significant contribution from dedifferentiated cells, in particular dedifferentiating myocytes, although to varying extents. This evidence confirms the plasticity of the regenerative phenomenon in echinoderms, which can apparently follow different pathways in terms of growth and morphogenesis, but nevertheless involve both epimorphic and morphallactic contributions at the cellular level.     

Autotomy and arm number increase in Oxycomanthus japonicus (Echinodermata, Crinoidea)

Abstract. We have explored the process by which crinoids increase arm number as they grow. Two hypotheses have been proposed: (1) arm autotomy with subsequent bifurcation and regeneration of a pair of arms, and (2) growth of a pinnule into an additional arm. We have traced the development of Oxycomanthus japonicus for about a year after fertilization and provide the first confirmation that the number of arms increases by autotomy, bifurcation, and subsequent regeneration of a pair of arms. The next such addition tends to occur at some distance from the previous pair. Thus, increase of arm number takes place in such a manner that the density of the arms remains relatively constant, and an efficient filtration fan for feeding is maintained. Although arm autotomy in crinoids has been considered to occur only as a response to physical or chemical disturbance, the present results suggest that autotomy also occurs as a specific, intrinsically programmed event during normal development.     

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.     

Cellular and molecular mechanisms of arm regeneration in crinoid echinoderms: the potential of arm explants

 Crinoid echinoderms can provide a valuable experimental model for studying all aspects of regenerative processes from molecular to macroscopic level. Recently we carried out a detailed study into the overall process of arm regeneration in the crinoid Antedon mediterranea and provided an interpretation of its basic mechanisms. However, the problem of the subsequent fate of the amputated arm segment (explant) once isolated from the animal body and of its possible regenerative potential have never been investigated before. The arm explant in fact represents a simplified and controlled regenerating system which may be very useful in regeneration experiments by providing a valuable test of our hypotheses in terms of mechanisms and processes. In the present study we carried out a comprehensive analysis of double-amputated arm explants (i.e. explants reamputated at their distal end immediately after the first proximal amputation) subjected to the same experimental conditions as the regenerating donor animals. Our results showed that the explants undergo similar regenerative processes but with some significant differences to those mechanisms described for normal regenerating arms. For example, whilst the proximal-distal axis of arm growth is maintained, there are differences in terms of the recruitment of cells which contribute to the regenerating tissue. As with normal regenerating arms, the present work focuses on (1) timing and modality of regeneration in the explant; (2) proliferation, migration and contribution of undifferentiated and/or dedifferentiated/transdifferentiated cells; (3) putative role of neural growth factors. These problems were addressed by employing a combination of conventional microscopy and immunocytochemistry. Comparison between arm explants and regenerating arms of normal donor adults indicates an extraordinary potential and regenerative autonomy of crinoid tissues and the cellular plasticity of the phenomenon. Received: 9 March 1998 / Accepted: 5 June 1998     

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.     

An experimental study of the use of arm regeneration in estimating rates of sublethal injury on brittle-stars

The ophiuroids Ophiomyxa flaccida (Say), Ophionereis reticulata (Say), Ophiocoma wendtii Müller & Troschel, O. echinata (Lamarck), O. pumila Lütken, Ophioderma appressum (Say), O. cinereum Müller & Troschel, and Ophiolepis impressa (Lütken) in back-reef populations in Jamaica were found to differ significantly in the number of arms regenerating. The proportion of arms regenerating varied from 74.4% for Ophionereis reticulata to 28.3% for Ophiolepis impressa. Regeneration rates, hitherto unknown except for Ophiocoma pumila, were determined for all the species for a single arm over a 3-month period. O. wendtii had the highest rate, regenerating 55% of its mean maximum arm length in 3 months, and O. impressa the slowest, regenerating only 23% of its mean maximum arm length in the same time.Injury rates, i.e., the number of arms broken and the total percent loss estimated to have occurred in the month prior to collection, were determined by dating wounds with reference to regeneration rates over 1 month. The species differed significantly in injury rates. O. pumila and O. reticulata suffered the highest levels of injury and O. impressa the least. Injury rates were not simply related to either the palatability to predators or the behaviour of the brittle-stars.     

Regeneration from Injury and Resource Allocation in Sponges and Corals – a Review

The ability of bottom‐dwelling marine epifauna to regenerate injured or lost body parts is critical to the survival of individuals from disturbances that inflict wounds. Numerous studies on marine sponges (Phlyum Porifera) and corals (of the orders Scleractinia and Alcyonacea) suggest that regeneration is limited by many intrinsic (individual‐dependent) and extrinsic (environment‐dependent) factors, and that other life history processes may compete with regeneration for energetic and cellular resources. We review how intrinsic (size, age, morphology, genotype) and extrinsic (wound characteristics, water temperature, food availability, sedimentation, disturbance history, selection) factors limit regeneration in sponges and corals. We then review the evidence for impaired somatic growth and sexual reproduction, and altered outcomes of interactions (anti‐predator defenses, competitive abilities, self‐ and non‐self recognition abilities) with other organisms in regenerating sponges and corals. We demonstrate that smaller, older sponges and corals of decreasing morphological complexities tend to regenerate less well than others, and that regeneration can be modulated by genotype. Large wounds with small perimeters inflicted away from areas where resources are located tend to be regenerated less well than others, as are injuries inflicted when food is limited and when the animal has been previously or recently injured. We also demonstrate that regeneration strongly impairs somatic growth, reduces aspects of sexual reproduction, and decreases the ability for sponges and corals to defend themselves against predators, to compete, and to recognize conspecifics. Effects of limited regeneration and impaired life histories may manifest themselves in higher levels of biological assembly e.g., reduced accretion of epifaunal biomass, reduced recruitment and altered biotic associations, and thus affect marine community and ecosystem recovery from disturbances. (© 2005 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Asteroid and Ophiuroid Trace Fossils from the Lower Cretaceous of Chile

Abundant asterozoan trace fossils in the Lower Cretaceous Apeleg Formation of southern Chile were produced by the infilling of traces made by asteroids and ophiuroids on a muddy surface. Most are preserved as hypichnial ridges formed as casts on the bottom of fine-grained sandstone laminae. Star-shaped Asteriacites lumbricalis are interpreted as the infillings of shallow excavations made by asteroids. Hook-shaped and sinuous ridges of the newly defined ichnogenus and ichnospecies Ophioichnus aysenensis are interpreted as the casts of imprints made by the walking action of the arms of ophiuroids. Deposition probably took place in an offshore tide-swept marine shelf environment. In life the animals formed an assemblage equivalent to echinoderm aggregations on the modern sea floor.     

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.     

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.      

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 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)))].     

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.     

Taphonomy of dense ophiuroid accumulations from the Middle Triassic of Poland

The taphonomy of three Middle Triassic (Muschelkalk) monospecific ophiuroid taphocoenoses, comprising Aspiduriella similis (Eck), from different regions in Poland (Holy Cross Mountains, Upper Silesia and North-Sudetic Basin) has been investigated. The majority of specimens (88.5% of a total of 428 individuals) are partially disarticulated, having only proximal and median portions of their arms preserved (Taphonomic Group 2). Pristine specimens, with only faint traces of disarticulation (Taphonomic Group 1), as well as those that preserve the disc only, or which have only proximal stumps of arms preserved (Taphonomic Group 3), are much rarer (6.5% and 5%, respectively). Moreover, most specimens (76.4%) are oral side up. Only 19.2% of specimens are preserved in life position, and a small fraction (4.4%) are preserved oblique to bedding. All ophiuroids studied occur in thin, pelitic layers devoid of any trace fossils. Associated body fossils, such as bivalves, gastropods or crinoids, are very rare. Taking into account ophiuroid taphonomy, as well as sedimentological characteristics of the thin layer (burial layer), it is clear that all assemblages were transported prior to burial. The predominance of articulated skeletons indicates that the burial event (obrution event) was not only rapid, but also single – subsequent events would have destroyed the previously buried ophiuroids. Storm-related resuspension of fine-grained material from nearshore environments which covered the ophiuroids is the most likely burial agent. The fact that in all regions the ophiuroid taphofacies studied is identical suggests that the three assemblages underwent the same burial history, and that a single event might have been responsible.     Muschelkalk , ophiuroids , Poland , taphonomy , Triassic .