Phylogeny and classification of the Asteroidea (Echinodermata)

Post-Palaeozoic asteroids share a large number of derived characters of the ambulacral column and the mouth frame, and constitute the crown group of the monophyletic group Asteroidea. This crown group is here called the Neoasteroidea (new subclass). The stem species of the crown group lived in the Permian or early Triassic and so the evolution of the asteroids parallels that of the echinoids. Character distribution within the Neoasteroidea, especially morphology of the skeleton, digestive system, larvae and tube feet, allows subdivision into four orders (Paxillosida, Notomyotida, Valvatida, Forcipulatida). The latter three orders possess the synapomorphy of suckered tube feet and are united as the Surculifera (new superorder); the Paxillosida are their primitive sister group. Palaeozoic asteroids represent the stem group of the class, and may be divided into plesions according to the order of appearance of synapomorphies with the crown group. Classification of Palaeozoic asteroids requires much further study. The appearance of new characters within the crown group asteroids, such as suckered tube feet, implies that these were absent in the stem group. The range of life-habits possible in Palaeozoic asteroids can thus be partly deduced from evidence provided by living asteroids. Palaeozoic asteroids are deduced to have lacked suckered tube feet and were presumably unable to evert the stomach; hence they were precluded from life on hard substrates and extraoral feeding on epifaunal organisms. It is suggested that they lived on soft substrates by deposit feeding, scavenging and predation on small benthos.     

Comptoniaster adamsi nov. sp. (Echinodermata, Asteroidea) from the middle Cretaceous of Texas and its phylogenetic position

Comptoniaster adamsi nov. sp. (Asteroidea, Valvatida, Goniasteridae) is described from the middle Cretaceous (Cenomanian-Turonian) Britton Formation of northcentral Texas, USA. The new species provides the focus for an exploratory cladistic analysis of Mesozoic asteroids of the Comptoniaster-Tylasteria Group sensu G. Breton. The systematics of Mesozoic goniasterids has relied heavily on the morphology of marginal ossicles, which generally are the best-preserved elements of the skeleton. Unfortunately, marginal ossicular data are scanty for most species because ossicles tend to be morphologically simple yet varied even within individuals, and few even partially articulated specimens are available to provide more comprehensive information. Further, both plesiomorphy and homeomorphy have been important. Because of limited available data, phylogenetic reconstruction here is preliminary. Nevertheless, a number of taxon groupings recognized in the literature are recovered, and stratigraphic distribution provides some support for results. Data unfortunately are particularly incomplete for species of Comptoniaster, the focus of the study. Comptoniaster adamsi nov. sp. clustered with three other species assigned to the genus, including the type, all Cretaceous in age. Jurassic species of Comptoniaster are more widely distributed in the analysis, perhaps reflecting an early stage in diversification or perhaps suggesting the need for species assignment reassessment, but also reflecting the limited available data.     

A chemical alarm response in Pycnopodia helianthoides (Echinodermata: Asteroidea)

Adult Pycnopodia helianthoides exhibit a classic alarm response, moving away from fragments and tissue fluid of the tube feet, pyloric caeca, and body wall of conspecific individuals.     

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.      

Salinity effects on arm regeneration in Luidia clathrata (Echinodermata: Asteroidea)

The effect of salinity (20, 25, and 30?psu) on regeneration of two arms of adult starfish Luidia clathrata (n?=?8 per salinity treatment group collected from Pensacola Bay, Florida, USA 30° 20′ N, 87° 13′ W) was examined in a laboratory experiment (November 2009 to March 2010) to test the hypothesis that L. clathrata is adapted to euryhaline conditions found in bays with regards to arm regeneration. Regenerating arm condition (lengths and weights) relative to treatment was 20-psu treatment?<?25-psu treatment?<?30-psu treatment, and non-regenerating arm condition was 20-psu treatment?<?25-psu treatment?=?30-psu treatment. Luidia clathrata is not adapted to lower salinity conditions found in bays with regard to arm regeneration, despite the common occurrence of the species in bays. Lower salinity in bays should reduce the ability of L. clathrata to recover from damage sustained during, or just prior to, low-salinity periods.     

A NEW SPECIES OF BRITTLESTAR (OPHIUROIDEA, ECHINODERMATA) FROM THE HUNSRÜCK SLATE (LOWER EMSIAN, LOWER DEVONIAN) OF GERMANY

Abstract:  Lapworthura lehmanni , a new species of ophiuroid, is described from four specimens from the Lower Devonian Hunsrück Slate of Germany. It is the only known ophiuroid in the Hunsrück Slate with paired but unfused ambulacrals, and it exhibits unique rows of spine-bearing dorsal arm ossicles. The genus Lapworthura Gregory was previously known only from the Ordovician of Scotland and the Silurian of England and Australia.     

New arthropods from the Lower Devonian Hunsru¨ck Slate (Lower Emsian, Rhenish Massif, western Germany)

Four new genera and species of arthropod, Cambronatus brasseli Wingertshellicus backesi Eschenbachiellus wuttkensis and Magnoculus blindi , are described from the Hunsru¨ck Slate (Lower Emsian) of Germany. All four occur in the Wingertshell Member in the vicinity of Bundenbach. They preserve remarkable details of the ventral morphology, including the appendages, as a result of pyritization. In each case the body consists of just two tagmata, a cephalon and a large number of similar trunk somites. Both Cambronatus Wingertshellicus have fluke-like appendages making up a tail fan. The affinities of these arthropods lie with the Crustacea, apart from Magnoculus , which is an arachnomorph, but they do not fall within those clades with modern representatives. They show that morphologies other than those represented by trilobites and modern arthropod groups persisted long after the Cambrian, at least in muddy bottom marine settings.     

Lysosomes and yolk formation in the oocytes of Asterina gibbosa (Echinodermata,Asteroidea)

Summary

Primary lysosomes appear in the oocytes of A. gibbosa at the end of previtellogenesis. The lysosomes fuse with the cisternae of the endoplasmic reticulum and give rise to yolk globules containing acid phosphatase. The yolk globules then grow by fusion.     

Tracing the evolution of the holothurian body plan through stem‐group fossils

The fossil echinoderm Palaeocucumaria, from the early Devonian Hunsrück Slate of southwestern Germany, has been studied using both traditional techniques and X‐ray microtomography, and its anatomy clarified. Phylogenetic analysis shows that it is a stem‐group holothurian with a combination of characters that help understand how the modern (crown‐group) holothurian body plan developed. Echinoids and holothurians have evolved along different paths, by differential growth of the larval‐ and rudment‐derived body regions. Palaeocucumaria shows that late stem‐group holothurians had a water vascular organization with a single external madreporite and calcified stone canal leading to the aboral end of the peripharyngeal coelom, and five primary radial water vessels that gave rise to tentacle‐like tube‐feet. This fossil data, in combination with a molecular phylogeny based on 18 s‐like rRNA gene sequence data, is used to order evolutionary steps in the making of the crown‐group holothurian body plan. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 670–681.     

Acrosomal reaction and early events at fertilization in Marthasterias glacialis (Echinodermata: Asteroidea)

When the spermatozoon of M glacialis contacts the mature oocyte jelly it adheres to it. Following this, there is a slight tumefaction of the acrosome, which is followed by the disruption of the apical acrosomal vesicle and cytoplasmic membranes. Acrosomal vesicle contents are liberated and spread along the outer surface of the oocyte jelly. Meanwhile, the acrosomal process begins to extend, penetrates all the jelly extension, then the vitelline layer, and finally contacts the cytoplasmic egg membrane. Nevertheless, the sperm cell continues lying at the outer border of the jelly. From the beginning of the acrosome reaction the dense and finely fibrillar subacrosomal material is connected, by some expansions, to the basal acrosomal vesicle membrane. Both nuclear and mitochondrial diameters have diminished.     

New early crown-group asteroids (Echinodermata; Triassic of Germany)

Migmaster angularis n. gen. n. sp. (Trichasteropsiidae, Forcipulatacea, Asteroidea) is described from the German Triassic Lower Muschelkalk of Anisian (Pelsonian) age, and new specimens ofTrichasteropsis bielertorum broaden understanding of this species. Phylogenetic events linking the stem groups, known only from Paleozoic rocks, with the crown group, known only from post-Paleozoic rocks, are problematic; the new fossils partially constrain phylogenetic changes associated with emergence of crown-group asteroids.      

Comparative histological and immunohistochemical study of sea star tube feet (Echinodermata, Asteroidea)

Adhesion in sea stars is the function of specialized structures, the tube feet or podia, which are the external appendages of the water-vascular system. Adhesive secretions allow asteroid tube feet to perform multiple functions. Indeed, according to the sea star species considered, the tube feet may be involved in locomotion, fixation, or burrowing. Different tube foot shapes usually correspond to this variety of function. In this study, we investigated the variability of the morphology of sea star tube feet as well as the variability of the composition of their adhesive secretions. This second aspect was addressed by a comparative immunohistochemical study using antibodies raised against the adhesive material of the forcipulatid Asterias rubens. The tube feet from 14 sea star species representing five orders and 10 families of the Class Asteroidea were examined. The histological study revealed three main tube foot morphotypes, i.e., knob-ending, simple disc-ending, and reinforced disc-ending. Analysis of the results suggests that tube foot morphology is influenced by species habitat, but within limits imposed by the evolutionary lineage. In immunohistochemistry, on the other hand, the results were very homogeneous. In every species investigated there was a very strong immunolabeling of the adhesive cells, independently of the taxon considered, of the tube foot morphotype or function, or of the species habitat. This indicates that the adhesives in all the species considered are closely related, probably sharing many identical molecules or, at least, many identical epitopes on their constituents.     

Antarctic Starfish (Echinodermata, Asteroidea) from the ANDEEP3 expedition

This dataset includes information on sea stars collected during the ANDEEP3 expedition, which took place in 2005. The expedition focused on deep-sea stations in the Powell Basin and Weddell Sea.Sea stars were collected using an Agassiz trawl (3m, mesh-size 500µm), deployed in 16 stations during the ANTXXII/3 (ANDEEP3, PS72) expedition of the RV Polarstern. Sampling depth ranged from 1047 to 4931m. Trawling distance ranged from 731 to 3841m. The sampling area ranges from -41°S to -71°S (latitude) and from 0 to -65°W (longitude). A complete list of stations is available from the PANGAEA data system (http://www.pangaea.de/PHP/CruiseReports.php?b=Polarstern), including a cruise report (http://epic-reports.awi.de/3694/1/PE_72.pdf).The dataset includes 50 records, with individual counts ranging from 1-10, reaching a total of 132 specimens.The andeep3-Asteroidea is a unique dataset as it covers an under-explored region of the Southern Ocean, and that very little information was available regarding Antarctic deep-sea starfish. Before this study, most of the information available focused on starfish from shallower depths than 1000m. This dataset allowed to make unique observations, such as the fact that some species were only present at very high depths (Hymenaster crucifer, Hymenaster pellucidus, Hymenaster praecoquis, Psilaster charcoti, Freyella attenuata, Freyastera tuberculata, Styrachaster chuni and Vemaster sudatlanticus were all found below -3770m), while others displayed remarkable eurybathy, with very high depths amplitudes (Bathybiaster loripes (4842m), Lysasterias adeliae (4832m), Lophaster stellans (4752m), Cheiraster planeta (4708m), Eremicaster crassus (4626m), Lophaster gaini (4560m) and Ctenodiscus australis (4489m)).Even if the number of records is relatively small, the data bring many new insights on the taxonomic, bathymetric and geographic distributions of Southern starfish, covering a very large sampling zone. The dataset also brings to light six species, newly reported in the Southern Ocean.The quality of the data was controlled very thoroughly, by means of on-board Polarstern GPS systems, checking of identification by a renowned specialist (Prof. Michel Jangoux, Université Libre de Bruxelles), and matching to the Register of Antarctic Marine Species (RAMS) and World Register of Marine Species (WoRMS). The data is therefore fit for completing checklists, for inclusion in biodiversity patterns analysis, or niche modeling. It also nicely fills an information gap regarding deep-sea starfish from the Southern Ocean, for which data is very scarce at this time. The authors may be contacted if any additional information is needed before carrying out detailed biodiversity or biogeographic studies.     

Coulonia parva n. sp., a new species of Astropectinidae (Asteroidea; Echinodermata) from the Santonian (Upper Cretaceous) of Sierra de Montsech (southern Pyrenees, Spain)

A new astropectinid sea star,Coulonia parva n. sp., is described from the Santonian of the Sierra de Montsech, northern Spain. The single specimen shows part of a rather weathered ventral side. It is characterized by comparatively long arms (R/r approx. 3/1), with one row of actinals reaching far into the arms; inferomarginals on the interbrachial arc have deep fasciolar grooves that are less developed on the arms; there are no larger spines on adambulacrals and probably also on inferomarginals. Based on presence of deep marginal fascioles,C. parva n. sp. is inferred to have shared the capability for self-burial with the relatedC. platyspina Hess &Blake.      

An ophiuroid trackway from the Lower Devonian Hunsrück Slate, Germany

A new trace fossil, Arcichnus saltatus , from the Lower Devonian Hunsrück Slate, Germany, is interpreted as the trackway of a protasterid ophiuroid, Taeniaster. The trackway consists of a series of horseshoe-shaped impressions, produced by a pair of arms, a feature characteristic of some ophiuroid trackways. This new trace fossil demonstrates that protasterids lived both infaunally and epifaunally.     

Mass mortality of an asteriid starfish (Forcipulatida,Asteroidea, Echinodermata) from the late Maastrichtian (Late Cretaceous) of Morocco

Abstract: A mass occurrence, numbering tens of thousands of individuals in a single lens, belonging to a single species of asteroid, is described from the late Maastrichtian (Late Cetaceous) of Morocco. The lens of partially silicified asteroidal limestone is made up largely of fully articulated specimens of similar size and probably represents the mass mortality of a single recruitment. By comparison with mass strandings of the present‐day species Asterias rubens (Linnaeus), it can be inferred that a feeding swarm of individuals was swept into a submarine channel by either a storm or an exceptionally strong tidal current, and permanently buried. The genus and species are herein described as Cretasterias reticulatus gen. et sp. nov. The exceptional preservation of the material enables the identification of wreath organs (clusters of crossed pedicellariae set in a dermal pad around spines) for the first time in the fossil record. Comparison between extant Asteriidae, putative fossil asteriids and C. reticulatus provides ambiguous evidence of its affinities; it appears to display a combination of plesiomorphic and derived characters. It is shown that all Mesozoic forcipulatid asteroids described so far share a very simple arm construction (single row of adradial ossicles) unknown in adult extant Forcipulatida.     

<Emphasis Type="Italic">Compsaster formosus</Emphasis><Emphasis Type="SmallCaps">Worthen &; Miller</Emphasis> (Asteroidea; Echinodermata): A Carboniferous homeomorph of the post-Paleozoic Asteriidae

All adequately known post-Paleozoic asteroids are either assignable to surviving families or closely related families whereas no Paleozoic species assignable to a surviving order has been recognized. The Mississippian speciesCompsaster formosus is similar enough to various Recent taxa in overall form as well as in the form and arrangement of body wall ossicles to raise the issue of affinities: IsC. formosus nested within a phylogenetic branch hitherto known only from post-Paleozoic strata or is it only homeomorphic?The nature of the ambulacral system is critical to interpretation of echinoderms, and post-Paleozoic asteroids share three fundamental ambulacral characters or character suites: dorsal podial pores, staggered positioning of ambulacrals and adambulacrals, and complex articular structures between these two ossicular types.Calliasterella americana, a Carboniferous asteroid, shares the three ambulacral features, although it is distinctive from post-Paleozoic asteroids in other ways.Compsaster formosus exhibits at least two of the three ambulacral characters, although presence of staggering has not been finally established. LikeC. americana, C. formosus differs from post-Paleozoic species in details of ambulacral anatomy as well as aspects of ventral body surface ossicular arrangement. Although approaching crown-group organization,C. formosus nevertheless represents a branch basal to the crown group.Because asteroids are generalists, understanding of life habits ofCompsaster is sketchy in spite of morphological similarities between it and younger genera.Compsaster probably was epifaunal and its overall form is strongly reminiscent of that of Recent predatory asteriids but it is also similar to small-particle feeding echinasterids.     

Systematics and palaeoecology of middle Toarcian Asteroidea (Echinodermata) from the “Seuil du Poitou”, Western France

Complete, articulated starfish fossils are rare. However, more frequently encountered dissociated skeletal elements (ossicles) permit reliable taxonomic identification, making them a valuable data source for diversity estimates. Nearly 300 asteroid ossicles, collected from the middle Toarcian marls in western France can be assigned to five species. Four species and two genera are described: Comptoniaster vrinensis nov. sp. (Goniasteridae), Poncetaster crateri nov. gen. nov. sp. (Stauranderasteridae), Galbaster recurrans nov. gen. nov. sp. (Goniasteridae) and Pentasteria? liasica nov. sp. (Astropectinidae). The known diversity of Early Jurassic asteroids is increased from 12 to 16 species. These taxa illustrate the diversification of crown-group asteroids early in the Jurassic, following the Permo-Triassic crisis. They also reflect bias of the fossil record, and imply the existence of numerous ghost lineages in the evolutionary trees of extant groups. Variation in asteroid diversity across the “Seuil du Poitou” was driven by ecological constraints. The relative frequency (abundance of ossicles and diversity) of goniasterids and stauranderasterids increases in shallower environments. The Benthopectinidae, represented by Plesiastropecten hallovensis, occurred primarily from deep-shelf sediments. Similar ecological patterns are observed for more recent fossil and extant relatives, which further supports the idea of conservative evolution in post-Palaeozoic starfishes since the Early Jurassic.     

Fine structure of the spermatozoon of Marthasterias glacialis (Linnaeus) (Echinodermata; Asteroidea), with special reference to acrosomal morphology

The sperm of Marthasterias glacialis (Linnaeus) was studied by light and electron microscopy. It is a long uniflagellated cell of the “primitive” type. The head has a spherical shape and contains a nucleus with a spheroid acrosome lying in a cup-shaped anterior fossa. The acrosome is formed by an acrosomal vesicle surrounded by the periacrosomal material. The basal specializations of the acrosomal vesicle show a clear differentiation of its constituents resembling the structure of membrane. The midpiece contains a very large annular mitochondrion which encircles two perpendicular centrioles. The distal centriole is in close association with a pericentriolar radial complex. The tail, containing a common microtubular axoneme, is projected to a variable position.     

Occurrence of a microcanalicular system within the ossicles and dermal tissue of Asterias rubens and Marthasterias glacialis (Echinodermata: Asteroidea)

Summary A microcanalicular network is demonstrated within the ossicle stroma and the dermal tissue of two asteroid species. Microcanaliculi are presumed to be mesodermal structures. They consist of convoluted tubular ducts lined by epithelial cells associated with scattered basiepithelial nervous processes. Such a microcanalicular system has not been reported previously from any echinoderm species. Its discovery in asteroids entails some conceptual changes, especially considering the physiology of the body wall.Research assistants of the National Fund for Scientific Research (NFSR, Belgium)