Feeding biology of the Antarctic brittle star Ophionotus victoriae (Echinodermata: Ophiuroidea)

Summary The diet and feeding behavior of the brittle star Ophionotus victoriae were studied in 1981–1983. Frequency-of-occurrence and points methods were used to examine stomach contents of 2475 specimens from 19 stations representing seven general localities along the Antarctic Peninsula and South Sandwich Islands. Results of the two methods were in agreement and indicated this species is an opportunistic generalist with high diet plasticity. The most abundant component of the stomach contents was sediment but the diet derived from at least 13 phyla. Crustaceans, notably euphausiids and amphipods, were most important at some locations. Diet composition varied significantly between locations and years, but did not differ among brittle star size groups. Stomach content data and laboratory studies indicate that O. victoriae utilizes a variety of feeding methods but not suspension feeding.     

Global diversity of brittle stars (Echinodermata: Ophiuroidea)

This review presents a comprehensive overview of the current status regarding the global diversity of the echinoderm class Ophiuroidea, focussing on taxonomy and distribution patterns, with brief introduction to their anatomy, biology, phylogeny, and palaeontological history. A glossary of terms is provided. Species names and taxonomic decisions have been extracted from the literature and compiled in The World Ophiuroidea Database, part of the World Register of Marine Species (WoRMS). Ophiuroidea, with 2064 known species, are the largest class of Echinodermata. A table presents 16 families with numbers of genera and species. The largest are Amphiuridae (467), Ophiuridae (344 species) and Ophiacanthidae (319 species). A biogeographic analysis for all world oceans and all accepted species was performed, based on published distribution records. Approximately similar numbers of species were recorded from the shelf (n = 1313) and bathyal depth strata (1297). The Indo-Pacific region had the highest species richness overall (825 species) and at all depths. Adjacent regions were also relatively species rich, including the North Pacific (398), South Pacific (355) and Indian (316) due to the presence of many Indo-Pacific species that partially extended into these regions. A secondary region of enhanced species richness was found in the West Atlantic (335). Regions of relatively low species richness include the Arctic (73 species), East Atlantic (118), South America (124) and Antarctic (126).     

Lecithotrophic development and metamorphosis in the Indo-West Pacific brittle star Ophiomastix venosa (Echinodermata: Ophiuroidea)

Summary

The larval development of the ophiocomid ophiuroid Ophiomastix venosais described using SEM. The gastrula transforms into a uniformly ciliated early larva which progressively changes into a lecithotrophic late premetamorphic larva with a continuous bilateral ciliated band. This stage is short-lived and equivalent to a highly reduced ophiopluteus. Comparisons between O. venosa and other ophiuroid species whose development has been investigated suggest that, whatever the developmental mode (lecithotrophic or planktotrophic), a pluteus stage always occurs in ophiuroids with planktonic development. Two metamorphic stages were identified, the late metamorphic larva differing from the early one by the closure of the larval mouth. The appearance of the permanent mouth marks the end of the metamorphosis. The postlarva still possesses remnants of larval features. The transformation of the reduced ophiopluteus into a barrel-shaped metamorphic larva with transverse ciliated bands, a vitellaria larva, is followed. The possible occurrence of a unique type of metamorphic larva in non-brooding ophiuroids is discussed. Verification of this, however, needs further SEM investigations on metamorphic larva from species having “regular” planktotrophic development.     

Functional and biochemical properties of the hemoglobins of the burrowing brittle star Hemipholis elongata say (Echinodermata,Ophiuroidea)

The burrowing brittle star Hemipholis elongata (Say) possesses hemoglobin-containing coelomocytes (RBCs) in its water vascular system. The RBCs, which circulate between the arms and body, are thought to play a role in oxygen transport. The hemoglobin of adult animals has a moderate affinity for oxygen (P(50) = 11.4 mm Hg at pH 8, 20 degrees C, measured in cellulo) and exhibits cooperativity (Hill coefficient > 1.7). The hemoglobin of juveniles has a higher affinity (P(50) = 2.3 mmHg at pH 8.0, 20 degrees C) and also exhibits cooperativity. The oxygen-binding properties of the hemoglobin are relatively insensitive to pH, temperature, and hydrogen sulfide. Adult hemoglobin is a heterogeneous mixture composed of three major fractions. The combined results of electrospray mass spectrometry and oxygen-binding experiments performed on purified fractions indicate that the native hemoglobin is in the form of homopolymers. A partial amino acid sequence (about 40 amino acids) of adult hemoglobin reveals little homology with holothurian hemoglobins.     

Two new species of Heterokrohnia (Chaetognatha) from Antarctic waters

Summary Two new species of the genus Heterokrohnia, H. longidentata and H. fragilis, are described and compared with the other three known Heterokrohnia species, H. mirabilis Ritter-Záhony 1911; H. bathybia Marumo and Kitou 1966 and H. involucrum Dawson 1968. The species have been found at great depths (1,000 m–2,000 m) near Elephant Island, north of the Antarctic Peninsula.     

Defining reproductively isolated units in a cryptic and syntopic species complex using mitochondrial and nuclear markers: the brooding brittle star, Amphipholis squamata (Ophiuroidea)

At a time when biodiversity is threatened, we are still discovering new species, and particularly in the marine realm. Delimiting species boundaries is the first step to get a precise idea of diversity. For sympatric species which are morphologically undistinguishable, using a combination of independent molecular markers is a necessary step to define separate species. Amphipholis squamata , a cosmopolitan brittle star, includes several very divergent mitochondrial lineages. These lineages appear totally intermixed in the field and studies on morphology and colour polymorphism failed to find any diagnostic character. Therefore, these mitochondrial lineages may be totally interbreeding presently. To test this hypothesis, we characterized the genetic structure of the complex in the French Mediterranean coast using sequences of mitochondrial DNA (16S) and for the first time, several nuclear DNA markers (introns and microsatellites). The data revealed six phylogenetic lineages corresponding to at least four biological species. These sibling species seem to live in syntopy. However, they seem to display contrasted levels of genetic diversity, suggesting they have distinct demographic histories and/or life-history traits. Genetic differentiation and isolation-by-distance within the French Mediterranean coasts are revealed in three lineages, as expected for a species without a free larval phase. Finally, although recombinant nuclear genotypes are common within mitochondrial lineages, the data set displays a total lack of heterozygotes, suggesting a very high selfing rate, a feature likely to have favoured the formation of the species complex.     

A new brittle star (Ophiuroidea: Ophiodermatina) from the Early Triassic Paris Biota (Bear Lake County,Idaho, USA)

Knowledge on the morphology of Early Triassic ophiuroids is very limited, in spite of the relatively high number of described species. This hampers attempts to use morphology-based phylogenetic analyses in order to explore the early diversification of the ophiuroid modern clades between the latest Paleozoic and the early Mesozoic. Here, a new ophiuroid from the Early Triassic of southern Idaho (USA) is described and assigned to the new taxon Shoshonura brayardi Thuy nov. gen., nov. sp. It is part of an exceptional Early Triassic faunal assemblage recently described as the Paris Biota. On account of the arm morphology and the shape of the arm spines and arm spine articulations, Shoshonura brayardi nov. gen., nov. sp. is considered a basal member of the suborder Ophiodermatina.     

Brittle stars from Southern Ocean (Echinodermata: Ophiuroidea)

The present biogeographic study on the ophiuroid fauna from the Southern Ocean (SO) contains an updated checklist, based on a compilation of all the published information provided for the Antarctic and sub-Antarctic regions as well as the information available in SCAR-MarBIN database. Faunal composition and geographical and bathymetric distribution are included. So far, 219 species have been recorded, of which 126 are endemic to the SO, 76 are exclusive to Antarctic waters, and 30 are exclusive to sub-Antarctic waters. This study corroborated the circumpolar and eurybathic character of the ophiuroid fauna of the SO, but some differences are discussed when considering shelf and deep-sea fauna in the whole SO, or in the Antarctic and sub-Antarctic regions separately. The biogeographic affinities of 17 areas considered in the SO are revised, based on a presence/absence datamatrix of the 219 species. This similarity analysis shows three main groups, two of them including sub-Antarctic areas and one for Antarctic areas. The faunal movement patterns between the main geographical connections have been based on historical site records of each species. These movements have a level of faunal exchange that exceeds that of other Antarctic benthic groups. Such movements are mainly from Antarctic and sub-Antarctic regions to the subtropical waters of South America, and from New Zealand and southern Australian waters to sub-Antarctic areas. In this context, the origin of the ophiuroid Antarctic fauna is discussed.     

A new genus Squamophis of Asteroschematidae (Echinodermata, Ophiuroidea, Euryalida) from Australia

Squamophis, a new genus of brittle star is described. Two species are included in the genus: Squamophis amamiensis (Okanishi & Fujita, 2009) from south-western Japan and Squamophis albozosteressp. n. from north-western Australia. Squamophisgen. n. is distinguished from the other genera of the family Asteroschematidae by the following characters: each radial shield is single-layered and is completely covered by plate-shaped epidermal ossicles, and the relative length of the longest arm spine throughout the arms is as long as the length of the corresponding arm segment. Squamophis albozosteressp. n. is distinguished from Squamophis amamiensis in having white, slightly domed, plate-shaped epidermal ossicles on the aboral side of the body, the ossicles on aboral and lateral portion of the arms form transverse rows, and the other part of aboral side of disc and basal to middle portion of arms are brown but tip of the arms are light purple.     

An escape response behaviour in the brittle star ophiopteris papillosa (echinodermata: Ophiuroidea)

This study describes a behaviour of the brittle star Ophiopteris papillosa and tests two hypotheses concerning its possible function. During this behaviour an individual rapidly raises all five arms above its disk as the brittle star falls through the water column. The final position, with the five arms held tightly together above the disk, is termed the tulip position. Individuals in the tulip position fell faster, thus reaching the bottom sooner, than individuals not in the tulip position when tested in the laboratory. Individuals in the tulip position were consumed less frequently than individuals not in the tulip position when tested in the field. I conclude that the tulip position increases the likelihood of a brittle star reaching the protective cover of the bottom and escaping fatal predation.     

Two new species of Ammothea (Pycnogonida, Ammotheidae) from Antarctic waters

Two new species of the genus Ammothea are described from Elephant Island and the South Shetlands Islands, Antarctica. The material was captured during the Polarstern cruise XXIII/8 to the Antarctic Peninsula area. The main features of Ammothea pseudospinosa n. sp. are a proboscis distinctly trilobulated distally with a constriction at 2/3 of its length and dimorphism between the propodi of the anterior (first and second) and posterior (third and fourth) legs, and a trunk: proboscis length ratio of about 1.5. The main features of Ammothea childi n. sp. are a cylindrical proboscis, longer than trunk length, and adults with functional chelifores. These species are compared with their closest congeners from the Southern Ocean: A. pseudospinosa n. sp. with Ammothea spinosa and Ammothea allopodes; A. childi n. sp. with Ammothea gigantea, Ammothea bicorniculata and Ammothea hesperidensis.     

A large new species of the genus Ptilocrinus (Echinodermata, Crinoidea, Hyocrinidae) from Antarctic seamounts

Ptilocrinus amezianeae n. sp. is a new species of stalked crinoid attributed to the family Hyocrinidae. Forty-five specimens were collected from seamounts north of the Ross Sea, and one specimen from the Kerguelen Plateau at depths ranging from 450 to 1,680 m. The collection from Admiralty and Scott seamounts constitutes the first example of a hyocrinid population known both from in situ photographs and from numerous collected specimens ranging from small juvenile to large adult. Variation in theca and stalk articulation characters throughout ontogeny is congruent with the molecular data and indicates that all the specimens examined belong to a single species. Tegmen and pinnule architecture, brachial arrangement, and stalk articular facets indicate that Ptilocrinus amezianeae n. sp. has close affinities with P. clarki and P. pinnatus from the northeastern Pacific and displays the most derived characters among these three species. Two cases of true arm division into two unequal branches suggest that Ptilocrinus and Calamocrinus are closely related. The picture and video transects on Admiralty seamount show a patchy distribution of living specimens with patches of mean density ca. 2.6 individuals m-2. In situ photographs also document predation by a sea urchin and a sea star on tegmen and proximal arms. The COI gene sequences analyzed in 25 specimens from Admiralty and Scott seamounts display low pairwise distances, low nucleotidic diversity, and intermediate haplotype diversity. These results, together with disarticulated ossicles and attachment disks observed on in situ photographs, indicate that the population investigated here is in decline.     

Arm autotomy in brittlestars (Echinodermata: Ophiuroidea)

Although ophiuroid arm shedding has long been accepted as an example of autotomy, there has been little investigation of the phenomenon to substantiate this. This paper describes the outwardly visible aspects of autotomy and the function of the internal components of the arm during detachment. Observations are focussed on Ophiocomina nigra, some comparisons being made with eight other species.
Ophiuroid autotomy is characterized by its occurrence close to the point of stimulation, its rapidity, and by the pattern of intervertebral muscle separation at the insertions which is constant for a given species. Evidence is presented showing the important role played by the intervertebral ligament. Both this and the muscle insertions are collagenous, and it is suggested that they and the other intersegmental connective tissues facilitate autotomy by undergoing a drastic, nervously mediated loss in tensile strength which enables the animal to part from its arm with a minimum of effort. Comparable properties have been ascribed to other echinoderm connective tissues, and their role in asteroid and holothurian autotomy has been acknowledged, but such a mechanism has not previously been suggested for ophiuroid arm autotomy.     

A new octocoral genus (Cnidaria: Anthozoa) from Antarctic waters

Sphaeralcyon weddellensis gen. nov. et sp. nov. is described and illustrated from three colonies collected in the eastern Weddell Sea on the Polarstern cruises ANT V/3 and ANT VI/3. The new genus is compared with three closely related alcyoniid genera, all having capitate or mushroom-shaped colonies and dimorphic polyps. Sphaeralcyon is readily distinguishable from Malacacanthus by the presence of sclerites (absent in the latter genus), and from Anthomastus by sclerite shape, mainly in the form of tuberculate spheroids, and by the horny periderm on the stalk (absent in Anthomastus). Sphaeralcyon also differs from Verseveldtia in the set of sclerites, which are elongate spiny rods and a different type of tuberculate spheroid, and in the presence of eight-points as anthocodial armature (absent in Verseveldtia). The discovery of this new genus lends further support to the notion that diversity in benthic communities in Antarctica is higher than suggested by previous studies. Accepted: 6 December 1999     

Reproduction and development of the conspicuously dimorphic brittle star Ophiodaphne formata (Ophiuroidea)

Ophiodaphne formata is a conspicuously dimorphic ophiuroid; the disk diameters are approximately 1 mm for males and 5 mm for females. The dwarf male clings to the larger female, with the oral surfaces and bursae of the paired ophiuroids closely appressed. Moreover, the female of each pair adheres aborally to the oral surface of a host sand dollar, Astriclypeus manni. Spawning and external fertilization occur in August, at Tsuruga Bay, Sea of Japan. Development of the dimorphic brittle star O. formata is described for the first time, from spawning through metamorphosis, with special attention to the formation of the skeletal system and the external morphology of early juveniles. Fertilized eggs are about 90 microm in diameter, pale pink, and negatively buoyant. The embryos undergo equal, total, and radial cleavage, and the larval skeleton first forms as a pair of tetraradiate spicules. Larval development proceeds to an 8-armed planktotrophic ophiopluteus, with skeletal elements that consist of a body rod and two recurrent rods. Three weeks after fertilization, all the pluteal arms, except for the postero-lateral arms, are absorbed, and the metamorphosing larvae sink to the bottom. Metamorphosis is completed 21.5 days after fertilization, and the resulting juvenile is pentagonal and approximately 270 microm in diameter. The smallest specimen (480 microm in disk diameter) collected by field sampling exhibited male features on the skeletal plates of the jaw and disk. Sexual dimorphism, the peculiar pairing behavior, and the close relationship with the host sand dollar may have evolved as distinct reproductive characteristics in this ophiuroid with its typical ophiopluteus larvae.     

Regeneration of the epithelial lining of the stomach after autotomy of a disk in the brittle star Amphipholis kochii (Lütken) (Echinodermata: Ophiuroidea)

The regeneration of the epithelial lining of the stomach of the brittle star Amphipholis kochii after autotomy of the aboral part of the disk was studied. It was shown that a part of the stomach epithelium remained after autotomy. Its cells participated in regeneration of the epithelium during restoration of the lost digestive system. There was a partial dedifferentiation of the epithelium cells of stomach, their migration and proliferation; cells of the stomach retained some specialized cytoplasmic structures, secretory vacuoles, and pinocytic vesicles. Migration of ectodermal cells of the esophagus in the damaged area was also recorded.     

Mitogenomes provide insights into the phylogeny and evolution of brittle stars (Echinodermata,Ophiuroidea)

Ophiuroidea is the most speciose of all classes of Echinoderma. It is an important component in benthic ecosystems, occurring in almost all ecological niches of modern seas. To date, the phylogeny and complete evolutionary history of the ophiuroids have not yet been fully resolved. In this study, we sequenced the complete mitochondrial genomes (mitogenomes) of Ophiothrix (Ophiothrix) exigua and two deep-sea species Histampica sp. CS049 and Ophioplinthaca sp. M5261. These two deep-sea ophiuroids displayed reversed strand-compositional bias and rearranged gene orders. Thirteen distinct patterns of mitochondrial gene order among ophiuroid mitogenomes were detected, with two gene order newly found in Ophiuroidea. Our data supported the gene order found in all sampled Ophiuridae as the most likely ancestral order of all Ophiuroidea. To improve phylogenetic accuracy based on nucleotide differences, two different criteria were used for the analyses: (i) nucleotide sequence from all codon positions (PCG₁₂₃); (ii) the NTE method (“Neutral Transitions Excluded”) for ameliorating the misleading effects of a reverse strand bias in the data. The two methods confirmed the polyphyly of the orders Ophiacanthida and Amphilepidia. At family and genus level, Ophiuridae, Ophionotus and Ophioplinthus were not monophyletic. The most notable exception was that the NTE phylogeny showed low variation of branch length. NTE dataset generated younger age for most lower-level nodes than that from PCG123 dataset. All analyses suggested that the ophiuroids radiation occurred around the Permian–Triassic mass extinction event, and the divergence time of the deep-sea lineages was during the Cretaceous.     

Leucon parasiphonatus,a new species (Crustacea: Cumacea: Leuconidae) from Antarctic waters

Six specimens ofLeucon parasiphonatus n. sp. were collected at depths ranging from 15 to 424 m in the vicinity of King George Island (South Shetland Islands, Antarctica) and the south eastern Weddell Sea.Leucon parasiphonatus belongs to the subgenusLeucon and differs from the other already known antarctic and Subantarctic species of the genus, in the absence of a serrated dorsomedian line and in the presence of a long pseudorostrum with several fine setae at its tip, surrounding the very long branchial siphon. The surface of the carapace is granulated; the carapace displays no teeth except for a few at its antero-lateral margin and at its ventral margin. The species most similar toLeucon parasiphonatus isLeucon siphonatus, reported from Mediterranean and North Atlantic waters.