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.     

The ultrastructural organization of the mature spermatozoon of Luidia clathrata (Say) (Echinodermata: Asteroidea)

The sperm of Luidia clathrata are morphologically typical of asteroid sperm. The head is spherical and contains the nucleus and acrosomal complex. The nucleus has an anterior indentation in which rests the acrosomal complex. There is no evidence of a centriolar fossa along the posterior border of the nucleus. The acrosome is a cup-shaped structure containing a less electron dense central region. The periacrosomal material is homogeneous in nature, and the subacrosomal specialization of the periacrosomal materials appear as bands of varying electron density. The middle piece is an annular band of mitochondria which surrounds the proximal and distal centrioles. The centrioles exhibit the typical nine triplet arrangement. Both the centrioles and the axoneme project to one side of the middle piece region. Associated with the distal centriole is an elaborate pericentriolar process.     

The structure of the larval nervous system of Pisaster ochraceus (Echinodermata: Asteroidea)

Ultrastructural observations and glyoxilic acid-induced fluorescence of catecholamines indicate that tracts of axons lie at the base of the ciliary bands and run throughout their length in bipinnaria and brachiolaria larvae of Pisaster ochraceus. Two types of nerve cells occur at regular intervals within the ciliary bands. Type I nerve cells are associated with the axonal tracts, and type II nerve cells, which are ciliated, occur along the edge of the ciliary bands. Two prominent ganglia, which appear as accumulations of nerve cells and neuropile, occur on the lower lip of the larval mouth. Smaller ganglia occur irregularly throughout the ciliary band. Synapses were never clearly identified and were assumed to be unspecialized. Nervous tissues were also found associated with the esophageal muscles, the attachment organ, and the larval arms. Organization of the nervous system and its association with effectors suggest it controls swimming and feeding. Several similarities exist between the nervous systems of larval asteroids, larval echinoids, and adult echinoderms.     

Individual recognition and phenotypic variability in Acanthaster planci (Echinodermata: Asteroidea)

Summary A coding method for identifying individuals by utilizing natural attributes (numbers and disposition of arms, madrporites, anuses, spines and deformities) permits individual recognition of crown-of-thorns sea stars Acanthaster planci. Because natural injuries altering parts of the code are frequent in A. planci (present in 20% to 40% of the individuals in a population), the physiognomy of the centrally located periproct, which is much less frequently injured than peripherally located areas, was used in combination with the arrangement of arms and madreporites. Identical madreporite/arm codes occurred in fewer sea stars in Guam (0%) than in Panamá (14%). The greater variability among sea stars at the western Pacific locality could be due in part to the higher incidence of gross deformities (presumed injuries) in Guam (43%) compared with Panamá (17%).     

Catalogue of neritic-benthonic echinoderms (Echinodermata: Asteroidea and Ophiuroidea) from the Cuban Archipelago

This is an updated taxonomic catalogue of neritic-benthonic Echinodermata (Asteroidea and Ophiuroidea) from Cuba, that includes locality, range, habitat and depth where each species is found. The species list is based on the collections of the Oceanology Institute; Havana; the "Felipe Poey" Museum (Havana University) and the National Museum of Natural History, Smithsonian Institution, Washington D.C. Specimens for those collections were captured by a variety of field techniques, narcotized with Cloral hydrate, fixed in formalin and stored in 70-80% alcohol. A total of 75 species have been identified from Cuban waters: Asteroidea (20 species) and Ophiuroidea (55 species).     

Skeletal morphology of the mud star,Ctenodiscus crispatus (Echinodermata: Asteroidea)

The morphology of the following eight major ossicle types is described and illustrated for the goniopectinid asteroid Ctenodiscus crispatus: terminal plates, superomarginal and inferomarginal ossicles, adambulacral and ambulacral ossicles, odontophores, oral intermediate plates, and superambulacral ossicles. Development, variation, and relationships with soft body-parts and with other ossicles are embphasized. Each ossicle type is distinguished by numerous structures related to its function and to articulation with adjoining skeletal elements. Because major structures (such as pustules, alveoli, and articulation surfaces) distinguishing ossicle types develop early during ontogeny, immature ossicles are readily identifiable. However, changes in form and orientation of these structures occur during ossicle growth. Ontogenetic changes are influenced by development of associated skeletal and soft parts. Ambulacral and adambulacral ossicles near the peristome are highly modified but retain the basic characteristics of structure and orientation which define these major types of skeletal elements.     

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.     

Re-evaluation of the Devonian family Helianthasteridae Gregory, 1899 (Asteroidea: Echinodermata)

Because of their taxonomic and morphologic diversity, the asteroids of the Lower Devonian Hunsrück Slate of Germany are important to both an understanding of the history of the class Asteroidea and to the interpretation of community evolution during the Paleozoic. Helianthaster Roemer, 1863, a large multiarmed Hunsrück asteroid, is redescribed. The Helianthasteridae Gregory is restricted to Helianthaster and Arkonaster Kesling, 1982 (Middle Devonian, Canada); Lepidasterella Schuchert is similar to the other two genera but known specimens are of poor quality and as a result the status of the genus is uncertain. Helianthaster is noteworthy in part because its size, multiarmed state, and presence of pedicellariae suggest certain crown-group genera, yet aspects of the arrangement of the ambulacral column are characteristic of the Paleozoic asteroid evolutionary grade. Neither Helianthaster nor other Hunsrück asteroids appear closely linked to the crown group; instead, functional patterns apparently re-emerged through time, although identification of specific behavior of ancient asteroids is difficult to impossible.      

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.     

Timing of fission in the starfish Allostichaster capensis (Echinodermata: Asteroidea) in laboratory

Timing of fission in the fissiparous starfish Allostichaster capensis under laboratory conditions is described. A. capensis generally splits across the disc along the fission plane during the spring, and then each half regenerates the missing arms during the rest of the year. The fission process can last eight hours. The healing process starts immediately after fission. A. capensis has impressive regeneration ability, including a comet, without signs of disc, regenerated two new arms.     

Seven Hox gene sequences from the asterinid starfish Patiriella exigua (Echinodermata: Asteroidea)

Partial homeobox gene fragments were amplified from Patiriella exigua genomic DNA by PCR using degenerate primers. The primers spanned a region of 82 nucleotides within the homeobox, encompassing amino acids 21–47 of the homeodomain. The deduced amino acid sequences of the seven Hox gene sequences plus one Xlox-type gene sequence from P. exigua are presented and compared with similar sequences from other organisms. This work is a preliminary step in the analysis of the evolution of development in the Patiriella species group and the roles of Hox gene expression therein.     

Determination of polyamines in digestive and reproductive tissues of adult Asterias vulgaris (Echinodermata: Asteroidea)

1. The polyamines spermine, spermidine and putrescine were extracted from tissues of Asterias vulgaris and quantitated using thin layer chromatography. 2. In the pyloric caeca, mean (+/- SE) levels of spermine, spermidine and putrescine were 138(15), 86(24) and 415(77) nmol/g wet wt tissue, respectively. In the testes, levels were 95(12), 13(6) and less than 6 nmol/g wet wt. In the ovaries, levels were 105(9), 11(0.8) and 15(8) nmol/g wet wt. 3. High levels of polyamines in the pyloric caeca may be related to secretion or excretion in this tissue. 4. We hypothesize that polyamines will be important in the regulation of cellular activity in these tissues during the annual reproductive cycle.     

Hettangian Asteriidae (Echinodermata: Asteroidea) from southern Germany: taxonomy,phylogeny and life habits

Germanasterias amplipapularia andHystrixasterias hettangiurnus are new genera and species of Asteriidae (Asteroidea: Forcipulatida) described from the Hettangian (lowest Jurassic) of southern Germany. They are among the oldest known relatively complete asteroids assignable to surviving families, yet they are remarkably similar to certain living species. Modern asteriids are efficient predators of sessile and slow-moving stoutly-armored mullusks, however much asteriid morphology is suggested to have originated with suspension-feeding habits. The lineage leading to the modern Forcipulatacea might have separated from that of the remainder of the modern asteroids as early as the Carboniferous.     

Patterns of asexual reproduction in the fissiparous seastar Coscinasterias acutispina (Asteroidea: Echinodermata) in Japan

The morphological and anatomical characteristics of the fissiparous seastar Coscinasterias acutispina were investigated at 23 sites around Japan in order to clarify the environmental factors determining asexual reproduction. The frequency of recent fission among the sites differed, although no significant differences were evident among the three major sea areas around Japan (the Sea of Japan, Seto Inland Sea, and Pacific Ocean), indicating that the spatial variability in incidence of fission might be determined on a local scale. Morphological observations revealed that the seastars endoparasitized by Dendrogaster okadai had not undergone fission recently, suggesting that fission was suppressed by parasitism. Stepwise multiple regression analysis of the frequency of seastars that had recently undergone fission revealed that nutrient conditions affected the incidence of fission. The relationships between the pyloric caeca index and arm length ratio at the individual level suggested that good nutritional conditions were required before the occurrence of fission. Additionally, regression analysis showed that high temperature was one of the triggers of fission in C. acutispina. Thus, the process of asexual reproduction in C. acutispina appears to be governed by intricate interactions among several factors, including parasitism by D. okadai, nutrient conditions, and high temperature.     

Ultrastructural evidence of the excretory function in the asteroid axial organ (Asteroidea,Echinodermata)

The ultrastructure of the axial organ of Asterias amurensis has been studied The organ is a network of canals of the axial coelom separated by haemocoelic spaces. The axial coelom is lined with two types of monociliary cells: podocytes and musculo-epithelial cells. Podocytes form numerous basal processes adjacent to the basal lamina on the coelomic side. Musculo-epithelial cells form processes running along the basal lamina. Some bundles of these processes wrapped in the basal lamina pass through haemocoelic spaces between neighboring coelomic canals. It is hypothesized that the axial organ serves for filtration of fluid from haemocoelic spaces into the axial coelom cavity, from which urine is excreted through the madreporite to the exterior.     

Daily activity pattern ofAstropecten aranciacus (Echinodermata: Asteroidea) and two related species under natural conditions

Daily activity patterns ofAstropecten aranciacus, A. bispinosus andA. jonstoni were investigated by means of SCUBA diving. The three species generally showed two peaks of activity, at dawn and dusk. However, the activity pattern was variable, depending upon site, depth and season. Two populations ofA. aranciacus showed only one daily peak at certain times of the year. These peaks were at night in one case, at noon in the other. Light appears to be an important factor governing the activity pattern.     

Assemblages of sea stars (Echinodermata: Asteroidea) and brittle stars (Echinodermata: Ophiuroidea) in the Weddell Sea (Antarctica) and off Northeast Greenland (Arctic): a comparison of diversity and abundance

Composition and distribution of asteroid and ophiuroid assemblages were investigated by means of Agassiz trawl catches at 34 stations in 220- to 1,200-m depth in the Weddell Sea and at 17 stations in 90- and 830-m depth off Northeast Greenland. A total of 86 species (48 sea stars, 38 brittle stars) were identified in the Weddell Sea whereas off Northeast Greenland a total of 26 species (16 sea stars, 10 brittle stars) were recorded. In both study areas, brittle stars were numerically more important than sea stars, and abundances generally decreased with water depths. Multivariate analyses revealed a conspicuous depth zonation of sea and brittle stars off Greenland. Very high abundances of Ophiocten sericeum and Ophiura robusta characterized the assemblages on shallow shelf banks whereas in greater depths Ophiopleura borealis, Ophioscolex glacialis and Ophiacantha bidentata became dominant, albeit at significantly lower densities. Mass occurrences of brittle stars, such as those recorded on Greenlandic shelf banks, have not been discovered in the Weddell Sea, where distinct assemblages were discriminated in deep shelf trenches as well as on the eastern and southern shelf. Ophioplocus incipiens, Ophiurolepis martensi and Ophiurolepis brevirima were the most prominent species on the eastern shelf. Ophiacantha antarctica, Ophiurolepis gelida and Ophionotus victoriae on the southern shelf, and Ophiosparte gigas as well as the asteriod Hymenaster sp., in the shelf trenches. Overall, the Weddell Sea housed conspicuously more asterozoan species than the waters off Greenland. Higher species diversity was also evident at both a regional and local scale, especially for the eastern Weddell Sea shelf. However, because many species from the Weddell Sea are closely related, the Weddell Sea assemblages were not significantly different from the Greenland ones in terms of taxonomic diversity and distinctness. Received: 29 April 1996/Accepted: 10 June 1996