Diversity dynamics of echinoderms and evolution of marine communities

A compound analysis of two global paleontological databases (Sepkoski??s database (SDB) and The Paleobiology Database) allowed the recognition of a number of previously undescribed trends in the evolution of the phylum Echinodermata. Paleozoic echinoderms, dominated by sessile epibenthic filter feeders, played an important role in benthic communities, especially in the Ordovician and Carboniferous. Paleozoic echinoderms typically showed an increased rate of genus renewal, which significantly decreases in the Meso-Cenozoic. After the P-T crisis the echinoderms became dominated by motile taxa, while the role of infaunal forms increased. During the global turnover in the benthic communities at the K-T boundary, which was accompanied by a sharp increase in the mean alpha-diversity, many marine organisms became inhabitants of much richer (compared to the Mesozoic) communities. However, of all echinoderms, this trend is observed only in crinoids. In contrast to most large taxa, echinoderms do not show positive correlation between the duration of genera and alpha-diversity of communities, which included these genera. During the Phanerozoic the geographical distribution of echinoderms showed a sharp paleolatitudinal gradient, i.e., each period was characterized by one paleolatitudinal zone with the maximum diversity of echinoderms, and the diversity rapidly decreasing to the north and to the south of this zone. The zone of the maximum diversity of echinoderms, like of entire marine biota, during the Phanerozoic gradually moved from the tropics of the southern hemisphere to the middle latitudes of the northern hemisphere.     

The development of an Early Ordovician hard ground community in response to rapid sea-floor calcite precipitation

The Kanosh Shale (Upper Arenig, Lower Ordovician) of west-central Utah. USA. contains abundant carbonate hardgrounds and one of the earliest diverse hardground communities. The hardgrounds were formed through a combination of processes including the development of early digenetic nodules in clay sediments which were exhumed and concentrated as lags by storms. These cobble deposits. together with plentiful biogenic metrical. were cemented by inorganically precipitated calcite on the sea floor. forming intraformational conglomerate hardgrounds. Echinoderms may have -played a critical role in the development of hardground faunas since their disarticulated calcite ossicles were rapidly cemented by syntaxial overgrowths. forming additional cobbles and hardgrounds. The echinoderms thus may have taphonomically facilitated the development of some of the hard substrates they required. A significant portion of the hardground cements may have been derived from the early dissolution of aragonitic mollusk shells. Kanosh hardground species include the earliest bryozoans recorded on hardgrounds and large numbers of stemmed echinoderms. primarily rhipidocystid cocrinoids. Bryozoans and echinoderms covered nearly equal areas of the hardground surfaces. and there was a distinct polarization between species which preferred the upper. exposed portions of the hardgrounds and others which were most common on undercut. overhang surfaces. The Kanosh Shale hardground fossils combine elements of Late Cambrian assemblages and Middle Ordovician faunas, thus confirming predicted trends in hardground community evolution. especially the replacement of cocrinoids by bryozoans and. to a lesser extent, by other stemmed echinoderms, especially crinoids. The Kanosh community marks the transition from the Cambrian Fauna to The Paleozoic Fauna in The hardground ecosystem. *Carbonate hardgrounds, aragonite dissolution, calcite cement, Echinodermara, Trepostomata, Nicholsonclla. Dianulites. Porifpra. taphonomic facilitation, Utah. Pogonip Group, Kanosh Shale. Ordovician.     

Deciphering the early evolution of echinoderms with Cambrian fossils

Echinoderms are a major group of invertebrate deuterostomes that have been an important component of marine ecosystems throughout the Phanerozoic. Their fossil record extends back to the Cambrian, when several disparate groups appear in different palaeocontinents at about the same time. Many of these early forms exhibit character combinations that differ radically from extant taxa, and thus their anatomy and phylogeny have long been controversial. Deciphering the earliest evolution of echinoderms therefore requires a detailed understanding of the morphology of Cambrian fossils, as well as the selection of an appropriate root and the identification of homologies for use in phylogenetic analysis. Based on the sister‐group relationships and ontogeny of modern species and new fossil discoveries, we now know that the first echinoderms were bilaterally symmetrical, represented in the fossil record by Ctenoimbricata and some early ctenocystoids. The next branch in echinoderm phylogeny is represented by the asymmetrical cinctans and solutes, with an echinoderm‐type ambulacral system originating in the more crownward of these groups (solutes). The first radial echinoderms are the helicoplacoids, which possess a triradial body plan with three ambulacra radiating from a lateral mouth. Helicocystoids represent the first pentaradial echinoderms and have the mouth facing upwards with five radiating recumbent ambulacra. Pentaradial echinoderms diversified rapidly from the beginning of their history, and the most significant differences between groups are recorded in the construction of the oral area and ambulacra, as well as the nature of their feeding appendages. Taken together, this provides a clear narrative of the early evolution of the echinoderm body plan.     

Settling strategy of stalked echinoderms from the Kaili Biota (middle Cambrian), Guizhou Province,South China

A total of 124 articulated echinoderms, including Sinoeocrinus lui (n = 30), “S. globus” (n = 92), one indeterminate gogiid, and one indeterminate edrioasteroid, from the Kaili Formation (Middle Cambrian), eastern Guizhou, China was examined. Among them, 73% of gogiid echinoderms are preserved attached to skeletal substrates. Suitable skeletal substrates for gogiid holdfasts in the Kaili Biota are organophosphatic brachiopods; large trilobite fragments, including cranidia, free cheeks, thoracic segments, and pygidia; hyoliths; and Scenella shells. The high frequency of “S. globus” attached to organophosphatic brachiopods is due to the high richness of organophosphatic brachiopods rather than a host-specific association in the Kaili Biota. However, the possible discrimination against attachment to small trilobites, such as Pagetia, requires further investigation. Based on 11 hand samples, echinoderm population density (EPD) varies from 0.3 to 4.5 echinoderms per 50 grind points (0.06 to 0.96 echinoderms per cm²), depending on the percent coverage of skeletal debris. Crowding effect is evident when EPD is approximately 0.32 echinoderms per cm² and the skeletal coverage is less than 20%.     

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.     

Life history evolution and comparative developmental biology of echinoderms

Evolutionary biologists studying life history variation have used echinoderms in experimental, laboratory, and field studies of life history evolution. This focus on echinoderms grew originally from the tradition of comparative embryology, in which echinoderms were central. The tools for obtaining and manipulating echinoderm gametes and larvae were taken directly from comparative embryological research. In addition, the comparative embryologists employed a diverse array of echinoderms, not a few model species, and this diversity has led to a broad understanding of the development, function, and evolution of echinoderm larvae. As a result, this branch of life history evolution has deep roots in comparative developmental biology of echinoderms. Here two main aspects of this relationship are reviewed. The first is a broad range of studies of fertilization biology, dispersal, population genetics, functional morphology, and asexual reproduction in which developmental biologists might take a keen interest because of the historical origins of this research in echinoderm comparative embryology. The second is a similarly broad variety of topics in life history research in which evolutionary biologists require techniques or data from developmental biology in order to make progress on understanding patterns of life history variation among echinoderm species and higher taxa. Both sets of topics provide opportunities for interaction and collaboration.     

On Mellitella stokesii and Amphipholis geminata (Echinodermata), from an intertidal flat in the upper Gulf of Nicoya estuary, Pacific, Costa Rica

Information on changes over time in the biodiversity and ecology of tropical systems is of increasing importance against the background of local, regional, and global dynamics. This study aimed to contribute with long-term data on the abundance fluctuations of two species of echinoderms from a tropical tidal flat. From February 1984 to December 1986, March 1985 to April 1987, and July 1994 to September 1996 (total: 76 dates), core samples (17.7 cm2, 15 cm deep) were collected at low tide at a mud-sand flat in the mid upper Gulf of Nicoya estuary, Costa Rica, as part of a survey of the benthic community. Among more than 100 taxa, the burrowing brittle star, Amphipholis geminata, and the sand dollar, Mellitella stokesii, consistently appeared in the samples over the study period. A total of 63 M. stokesii and 149 A. geminata were collected. The 43 sand dollars found during 1984-1985, give an approximate density of 35 ind./m2, which is within the range reported for this species. M. stokesii was almost absent from 1994 to 1996, while A. geminata had its peak of abundance at the end of 1995. Low abundances of the sand dollar during the rainy seasons (May-November) and slight increments in dry seasons (December-April) cores, also agrees with a report from the region. A. geminata also presented an irregular pattern of abundance, with slight increases at the end of the rainy seasons or during the dry seasons, when higher salinities are more suitable for echinoderms. The patchy spatial distribution of both species makes difficult the detection of patterns with a corer. Nevertheless, this information is unique due to its extensive time coverage and provides a baseline for future surveys designed specifically for the study of tropical intertidal estuarine echinoderms.     

Diseases of echinoderms

Diseases of echinoderms are poorly documented. Most reports concern biotic diseases caused by animal agents. While parasites on echinoderms have been described in increasing numbers of papers for more than one century, the host-parasite relationship and the effects of parasitism on echinoderm life-cycles were rarely considered. The parasitic fauna differs markedly according to the echinoderm group concerned, depending on various factors such as feeding-habits or symbiogenesis. Microorganismic diseases undoubtedly occur in echinoderms but they were not investigated until recently. Microorganisms have frequently been assumed to act as agents causing mass mortalities. As for stress-caused diseases, the only — and very preliminary — data available concern almost exclusively those induced by pollution. Since echinoderms are major components of benthic ecosystems, echinoderm diseases may be expected to exert prominent ecological effects.     

Structure and anticoagulant activity of sulfated fucans. Comparison between the regular, repetitive, and linear fucans from echinoderms with the more heterogeneous and branched polymers from brown algae

Sulfated fucans are among the most widely studied of all the sulfated polysaccharides of non-mammalian origin that exhibit biological activities in mammalian systems. Examples of these polysaccharides extracted from echinoderms have simple structures, composed of oligosaccharide repeating units within which the residues differ by specific patterns of sulfation among different species. In contrast the algal fucans may have some regular repeating structure but are clearly more heterogeneous when compared with the echinoderm fucans. The structures of the sulfated fucans from brown algae also vary from species to species. We compared the anticoagulant activity of the regular and repetitive fucans from echinoderms with that of the more heterogeneous fucans from three species of brown algae. Our results indicate that different structural features determine not only the anticoagulant potency of the sulfated fucans but also the mechanism by which they exert this activity. Thus, the branched fucans from brown algae are direct inhibitors of thrombin, whereas the linear fucans from echinoderms require the presence of antithrombin or heparin cofactor II for inhibition of thrombin, as reported for mammalian glycosaminoglycans. The linear sulfated fucans from echinoderms have an anticoagulant action resembling that of mammalian dermatan sulfate and a modest action through antithrombin. A single difference of one sulfate ester per tetrasaccharide repeating unit modifies the anticoagulant activity of the polysaccharide markedly. Possibly the spatial arrangements of sulfate esters in the repeating tetrasaccharide unit of the echinoderm fucan mimics the site in dermatan sulfate with high affinity for heparin cofactor II.     

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.     

Evolutionsmodelle für den Ursprung der Echinodermen: Zusammenfassung und Kritik

The origin of echinoderms is one of the most crucial questions within the evolutionary history of deuterostomes. An ancestral position was suggested byGarstang, Romer andNichols. They also assumed that hemichordates and chordates are sistergroups. In all other hypotheses the echinoderms took a more derived position.Gislén, Jefferies andHolland viewed the hemichordates as basal to the deuterostomes and postulated that echinoderms and chordates are sistergroups. According toJollie, Peterson et al. andMooi &; David echinoderms and hemichordates are sistergroups.Gudo andGutmann adopted the view ofMetschnikoff who combined the hemichordates and echinoderms in the Ambulacraria; they supposed that echinoderms were derived from pterobranchs. This variety of views is linked with different approaches to phylogenetic reconstruction utilized by each of the authors.Garstang, Romer, Jefferies andGislén compared morphological features, in the case ofGislén andJefferies with some attention to fossil evidence, whereasJollie, Holland andGislén also considerd embryological aspects.Mooi &;David as well asPeterson et al. used modern embryological (epigenetical) approaches.Nichols combined functional morphology and comparative anatomy. Evolutionary scenarios were reconstructed only by a few authors.Holland associated the development of echinoderms from pterobranch-like ancestors with repeated changes in feeding modes.Nichols envisioned that echinoderms had evolved from sipunculids that gained protection from predators through skeletal armor. In our own investigations based on constructional morphology echinoderms are interpreted as highly derived chordates.     

Plated Cambrian bilaterians reveal the earliest stages of echinoderm evolution

Echinoderms are unique in being pentaradiate, having diverged from the ancestral bilaterian body plan more radically than any other animal phylum. This transformation arises during ontogeny, as echinoderm larvae are initially bilateral, then pass through an asymmetric phase, before giving rise to the pentaradiate adult. Many fossil echinoderms are radial and a few are asymmetric, but until now none have been described that show the original bilaterian stage in echinoderm evolution. Here we report new fossils from the early middle Cambrian of southern Europe that are the first echinoderms with a fully bilaterian body plan as adults. Morphologically they are intermediate between two of the most basal classes, the Ctenocystoidea and Cincta. This provides a root for all echinoderms and confirms that the earliest members were deposit feeders not suspension feeders.     

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.     

Dumbbell-shaped gogiid clusters: the oldest evidence of secondary tiering for stalked echinoderms

Among 381 specimens of Cambrian stalked echinoderms from eastern Guizhou, China examined, several slabs ( n  = 19) contain either dumbbell-shaped or v-shaped echinoderm clusters. Four slabs of Globoeocrinus globules Zhao, Parsley & Peng, 2008 from the middle-upper part (Cambrian Series 3 portion) of the Kaili Formation are prepared to reveal the attachment sites. Articulated gogiid echinoderms are reported to be attached to both sides of inarticulate (organophosphatic) brachiopods; thus, allowing me to interpret that the larvae of these gogiids were capable of attaching to live benthic brachiopods. This study documents the one of the earliest examples of echinoderms employing secondary tiering, which elevates an organism higher into the benthic boundary layer. Many of the gogiid echinoderm pairs attached to a live brachiopod are similar in size, indicating they were from a single larval spatfall.