Large-scale estimate of somatic growth in Amphiura filiformis (Echinodermata: Ophiuroidea)

Quantitative data on size structure of the ophiuroid Amphiura filiformis (O.F. Müller) from 35 benthic stations in the Kattegat sampled twice with a 143 d interval, June and October 1991, were used to estimate somatic growth. The material was objectively divided into cohorts, and cohorts, from the two occasions were paired to give estimates of growth. The growth constant, K, in the Von Bertalanffy equation, was estimated from a Ford-Walford plot to 0.54 yr^-1. Results were in agreement with previous estimates from a few single sites, and suggest that the main part of dise growth occurs within the first 5 to 7 yr of living. Size specific growth in oral width was density independent despite high densities of A. filiformis (>3000 ind m^-2) and high total benthic biomass (up to 1000 g wet wt m^-2) in some areas. Growth was uncorrelated with plant pigment concentrations in the sediment and showed weak positive correlation with sediment carbon and nitrogen, as well as water depth. Growth was higher in fine sediments. This is the first attempt to estimate growth in this important species over a large area and to relate growth in the field to environmental factors.     

Reproductive biology of a deep-sea brittle star Amphiura carchara (Echinodermata: Ophiuroidea)

 New information on the brooding reproduction of Amphiura carchara, and previous studies of several deep-sea congeners, contradict the prevailing notion that direct development is exceptional among deep-sea echinoderms. Over 500 specimens of A. carchara from 2,850 to 4,100-m depths off the coast of California, USA, were studied. The species was found to be gonochoric, although most brooding species of ophiuroids are hermaphroditic. Females each brooded up to 72 embryos at a time, with up to 10 in a single bursa. The embryos of individual adults were often at different stages of development, but those brooded in a single bursa tended to be at the same stage. Thus, the species has characteristics that breach the distinctions between sequential and simultaneous brooding exhibited by shallow-water ophiuroids. The embryos developed from yolky eggs that are large for an ophiuroid, with the mean largest oocytes 0.45 mm, and ranging to 1.28 mm in diameter. Almost all late-stage embryos were positioned with their mouth and arms pressed against the wall of the bursa, possibly to facilitate the uptake of nutrients from the parent. Thus, A. carchara may be matrotrophic. The largest embryos examined had a rudimentary disk skeleton, and arms with four joints and a terminal plate. Emerging juveniles probably differ in disk diameter. The species appears to brood year round, although differences in gonad size, the incidence of brooding, and the relative numbers of early developmental stages in summer and winter samples indicated that there are seasonal trends in reproduction. Received: 14 February 2000 / Accepted: 3 July 2000     

Field survey of the occurrence and significance of regeneration in Amphiura filiformis (Echinodermata: Ophiuroidea) from Galway Bay,west coast of Ireland

The Boreo-Mediterranean amphiurid Amphiura filiformis (O. F. Müller) occurs in high densities within Galway Bay. This burrowing species consistently shows signs of recent regeneration (of both arms and disc “cap”) at the above locality. Specimens were collected by SCUBA in March, April and May 1980, May 1981, and April 1982. Biomass assays have revealed that individuals with up to 50% (mean ca. 25%) of their entire body weight consisting of regenerated tissue are not uncommon. The occurrence of arm breakage and regrowth was found to be more frequent in the proximal (basal) and mid-arm regions than in the exposed arm tip portions. Roughly 3% of the population were found to be in the process of regenerating the disc “cap” or covering. Preliminary results from gut analyses of potential predators at the sampling station indicate that young flatfish (plaice and dab) actively “crop” the arms of A. filiformis. Records from north western Europe of this species occurring in the diets of fish and invertebrates are considered. In this long-lived species, the continual necessity to regenerate is deemed to be a major drain on bodily resources.     

Population dynamics of Amphiura chiajei (Echinodermata: Ophiuroidea) in Killary Harbour,on the west coast of Ireland

The infaunal ophiuroid Amphiura chiajei Forbes is a dominant member of the A. chiajei community in Killary Harbour, a fjord-like inlet on the west coast of Ireland. High density populations (700 individuals/ m²) occur in sediments with a silt/clay content of 80 to 90% and organic carbon levels of 5 to 7%. A study of the population dynamics of this species was carried out from November 1985 to October 1988. At the outset, the population consisted of approximately equal numbers of adults and juveniles. By October 1988, the percentage of adults had increased to 95%, with little or no change in the mean annual dersities. However, there was some degree of variability between the mean monthly densities. Such variability may have been due to very low bottom temperatures, resulting in mortality among the older members of the population. Observations suggest that in Killary Harbour the species may attain an age of 10 yr, with an early annual growth rate of 0.5 mm (oral width). Annual recruitment was variable and low, due presumably to intraspecific competition with the adults; recruitment success may be very closely linked to adult mortality. The Killary Harbour findings are considered in the light of what is currently known of similar amphiurid populations in different geographical locations.     

Effect of arm regeneration on energy storage and gonad production in Ophiocoma echinata (Echinodermata: Ophiuroidea)

Two laboratory experiments were conducted on Ophiocoma echinata (Lamarck, 1816) collected from Missouri Key, Florida, USA, to ascertain the effects of arm regeneration on energy storage and gonad production. In each experiment individuals were divided into non-regenerating (all arms intact) and regenerating (three arms autotomized) groups and maintained together in a re-circulating seawater system for 2 mo. In Experiment 1, individuals were fed once a week, and in Experiment 2 once every 2 wk. The amount of food energy required for maintenance was estimated to be 0.172 kJ d⁻¹. The lower feeding level (Experiment 2) was below maintenance ration and both groups lost material relative to an initial group. At the higher feeding level (Experiment 1), regeneration of three arms resulted in less storage of organic material, mainly lipid in the stomach, and less gonad production. Thus, regeneration may reduce energy reserves and reproductive output. Received: 23 April 1998 / Accepted: 11 January 1999     

Reproduction and population dynamics of the fissiparous brittle star Ophiactis savignyi (Echinodermata: Ophiuroidea)

Reproductive cycle, asexual reproduction, and population dynamics of the fissiparous brittle star, Ophiactis savignyi, which inhabits the exhalant passages of the sponge Haliclona sp. were examined monthly from February 1991 to January 1992 at Wanlitung, southern Taiwan (22°N; 120°E). Mature gametes were found from March to December, but release was mainly in May and June. Sexual recruits were found from May to December, with the highest frequency (14.1% of population) in June. Sexual recruits composed 2.4% of the 1-yr sample. Sex ratio of male to female was 24:1. Fission occurred throughout the year, although the frequency of recently split individuals was lower from January to June (6 to 31%), and higher from July to December (42 to 52%). The occurrence of fission was highest after spawning. Regenerating individuals composed 48.2% of the 1-yr sample. Population density fluctuated greatly during summer due to recruitment by fission and mortality or dispersal due to the stressful environmental conditions. Both sexual and asexual reproduction of O. savignyi were successful at this site.     

Comparative study of sub-cuticular bacteria in brittlestars (Echinodermata: Ophiuroidea)

Arm tissues from species of brittlestars from the British Isles and North America were examined by light and electron microscopy for the presence of sub-cuticular bacteria (SCB) which were recorded in 17 of the 19 species studied. Three major groups of SCB could be distinguished on the basis of their morphology. The bacterial morphology was not correlated with the host ecology but did appear to be related to the phylogeny of the hosts. Ophiuroids of the same genera had morphologically similar SCB and this was also generally true of species from the same family. This indicates the possibility of co-evolution between the SCB and their hosts. Estimates of SCB load were made for 9 of the species by direct counting and by quantifying the amount of bacterial endotoxin present in the host tissues. SCB numbers varied interspecifically from 3.7×10⁸ to 4.6×10⁹ SCB per gram of ash-free brittlestar arm tissue. Endotoxin values ranged from 5.97 to 285 g g^–1 ash-free arm tissue. Calculations suggest that SCB form at least 0.1 to 1% of the total biomass of the arm tissues. There was considerable intraspecific variation in SCB load. The possible role and significance of SCB is discussed.     

Alarm signal response in the brittle star Amphiura filiformis

 Flat fishes, such as the dab Limanda limanda, commonly prey on arms of Amphiura filiformis. We demonstrate in flume experiments that A. filiformis showed a more or less simultaneous response to an up-stream predation by dab on conspecifics: the arms, which are stretched up into the water column when filter feeding, responded by bending down to the sediment surface, and some arms submerged, at least partly, into the sediment. We interpret this as an alarm response. A similar, but significantly weaker response in A. filiformis was also recorded when dab ate Amphiura chiajei. Homogenates of the two Amphiura species and arm pinching – to simulate partial predation – elicited a similar response in A. filiformis, the arms bent to the sediment surface. In contrast, however, the behavioural response of the arms to the homogenates was to protrude further out of the sediment and sweep the surface vigorously. We interpret this as a feeding behaviour. Response to arm pinching was less clear. The ecological consequences and origin of alarm signal response in A. filiformis are discussed. Received: 8 February 1999 / Accepted: 16 July 1999     

Effects of salinity and calcium concentration on arm regeneration byOphiothrix angulata (Echinodermata: Ophiuroidea)

At 33 salinity a tissue stump formed 2 to 3 d after autotomy and developing ossicles were present by the fourth day inOphiothrix angulata (Say). Regeneration proceeded rapidly from the sixth day until the thirteenth day, when the rate decreased greatly. The length of the regenerated arm and the number of ossicles formed did not vary over a salinity range of 28 to 38 S, but were significantly less at 23 S. The number of ossicles regenerated increased linearly (y=1.9 x-7.7;r=0.9089) with the calcium concentrations ranging from 3.8 to 9.5 mM. No ossicle formation occurred at 3.8 mM calcium concentration. Rate of net uptake of calcium-45 into the ossicles of intact individuals in salinities of 28 and 33 was significantly greater than that in 23 and 38 S. However, net uptake rate of calcium into the soft tissues of the arms was significantly higher at 18S than at the lower two salinities.     

Post-metamorphic growth of the arms in Ophiophragmus filograneus (Echinodermata: Ophiuroidea) from Tampa Bay,Florida (USA)

Juveniles of the burrowing amphiurid Ophiophragmus filograneus from Tampa Bay, Florida (USA) exhibit a growth pattern unreported in the ophiuroid literature. Two nonadjacent arms grow at a greater rate than the other three arms. This phenomenon might be a developmental adaptation to avoid salinity and temperature fluctuations in surface waters of the shallow-water environment which O. filograneus inhabits. It is proposed that concentration of growth into 2 of the 5 arms permits earlier descent of the disc into the substratum with continued ability to feed on the surface with the arm tips. Larger specimens show a gradual equalization of arm lengths. Changes in growth rates and cropping of the longer arms by breakage and predation might account for such eventual equalization.     

The early embryology of the basket star Gorgonocephalus caryi (Echinodermata,Ophiuroidea)

The changes in morphology of Gorgonocephalus caryi embryos over the 8 1/2 day period of their survival are described. Fertilized eggs are 220 in diameter. Division is equal and holoblastic, and the blastula stage is reached within 24 h. Gastrulation begins at about 40 h. At 68 h, the anterior end of the embryo begins to elongate. The embryos are pear-shaped at 72 h. At about 84 h the anterior end begins to bend towards one side. By 98 h the entire embryo has flattened perpendicular to the anterior-posterior axis, and the embryo is a flattened, trilobed structure. At 4 1/2 days the anterior tip twists around to lie with its axis in a plane perpendicular to the other 2 lobes. They, meanwhile, shift in position so that the resulting embryo resembles an umbrella which is flat on top and has a curved handle. Meanwhile, the hydrocoel forms as an out-pouching from the archenteron in the anterior lobe. The general coelom develops by schizocoely. Between 5 and 5 1/2 days, the anterior lobe coelesces with the flattened area as it becomes pentagonal. By 5 1/2 days, many embryos are starshaped. At 6 1/2 days the mouth begins to form. By 8 days the embryos were dying, and none lived beyond 10 1/2 days. At no time did cilia or other locomotory structures develop. The development of G. caryi does not resemble that of any other ophiuroid studied. There is not trace of a pluteus, and it is postulated that the pluteus evolved after the Phrynophiurida and the Ophiurida were differentiated.     

Effects of disc and arm loss on regeneration byMicrophiopholis gracillima (Echinodermata: Ophiuroidea) in nutrient-free seawater

Clibanarius longitarsus (De Haan), a hermit crab, feeds by means of both micro- and macrofeeding. Microfeeding is of two types: filter feeding and deposit feeding. In macrofeeding, the animal depends mainly on green algae, barnacles and some worms, etc. In filter feeding, the hermit crab uses its paired antennules for trapping microorganisms drawn into the water current created by the second and third maxillipeds; the antennae do not play any role in filter feeding. In deposit feeding, the hermit crab uses its chelae as well as pereiopods for toughing and obtaining detritus food material. In macrofeeding, the organism employs sharp chelae to collect small pieces of muscle from barnacles and then transfer them to the inner mouth parts. In algal feeding, the chelae as well as maxillipeds help in collecting and moving food material towards the mouth. In this investigation functional organisation ofC. longitarsus mouth parts was studied in detail with reference to the various feeding mechanisms, in specimens collected from the intertidal region of Bheemunipatnam, Andra Pradesh, India, in 1988 and 1989.     

Genetic variance in Ophiomusium lymani (Ophiuroidea: Echinodermata) from lower bathyal depths in the Rocklall Trough (northeast Atlantic)

Specimens of the deep-sea brittle-star Ophiomusium lymani were collected from six sites in the Rockall Trough (northeast Atlantic_. Four monomorphic and four polymorphic loci were detected, with up to 75 individuals screened at any one locus. The results showed little difference in allele frequencies between sites and, consequently, estimates of genetic identity indicated no significant genetic differentiation between the populations sampled. Deficits of heterozygotes were observed at all polymorphic loci, but the deficiencies were only significant at the phosphoglucose isomerase locus from two locations. The deficit of heterozygotes was not the same across loci, suggesting that inbreeding is not the cause of the excess homozygosity. No relationship between heterozygosity and depth was observed.     

Reproduction of<Emphasis Type="Italic"> Ophiactis resiliens</Emphasis> (Echinodermata: Ophiuroidea) in New South Wales with observations on recruitment

Reproduction of the ophiuroid Ophiactis resiliens Lyman 1879, a common species in the shallow waters around Australia, was examined over 1 year in two populations near Sydney, New South Wales. Gametogenesis was documented by histological examination of the gonads. Vitellogenesis in females and spermatocyte proliferation in males started in February and March. O. resiliens had an annual spawning period from May to September/October with peak gamete release from May to July. Thereafter, spawning occurred in some specimens through November. Gravid gonads were present for approximately 2 months longer in O. resiliens from Clovelly Bay than in those from Little Bay. At both sites spawning was episodic and gametogenesis continued through most of the spawning season. Most specimens (50–100%) collected from November to March had indeterminate/recovery stage gonads, the sex of which, could not be determined. The timing of gametogenic renewal during these months varied among individuals. O. resiliens has a 3-month planktonic stage and the presence of juveniles, 0.5 –1.5 mm disk diameter, in the coralline algal turf at Little Bay from February to May, suggests that settlement occurs in summer and autumn at this site. In contrast, juveniles were rarely encountered in coralline turf at Clovelly.     

Post-larval development of the common north-west European brittle stars Ophiura ophiura,O. albida and Acrocnida brachiata (Echinodermata: Ophiuroidea)

The post-larval skeletal plate ontogeny of the ophiuroids Ophiura ophiura (Linnaeus) and Acrocnida brachiata (Montagu) from the recently metamorphosed 0.3 mm disc diameter size to a juvenile size of 2.0 mm disc diameter has been examined using scanning electron microscopy. Post-larval development of O. albida Forbes is briefly described for sizes between 1.0 and 2.0 mm disc diameter. Early post-larval development in the 0.2 to 1.0 mm discsize range involves considerable morphogenetic change and necessitates the use of a developmental series in the identification of newly settled ophiuroids. Comparison of these ophiuroids with other closely related species reveals that the post-larvae are readily distinguishable at even the smallest sizes by their transient post-larval morphologies. Abbreviated larval development in A. brachiata is suggested from the egg size and post-larval form. The ophiuroid post-larvae were collected from the shallow sublittoral of Oxwich Bay, Bristol Channel, UK, during the years 1980 to 1982.     

Growth patterns of Amphiura filiformis support the hypothesis of organic enrichment in the Skagerrak-Kattegat area

Population size structure, abundance and biomass of the burrowing ophiuroid Amphiura filiformis were measured in the period 1970 to 1989 at three widely spaced sites at 55 to 100 m depth in the Skagerrak-Kattegat area. This species is a dominant component of the fauna in the area, and also one of the species accounting for a great part of a recent general increase in benthic biomass. Abundance at two of the sites and biomass at all three sites increased significantly with time from the 1970s onwards. Maximum individual size and growth of the cohort of adults showed a marked increase in the period 1980 to 1983 at the three sites. In the 1980s annual absolute growth increments of the recruits were clearly greater than in the 1970s, and size-specific growth rates were virtually unchanged. These results are consistent with the hypothesis that recent increases in benthic biomass in this area result from increased input of food to the benthos. A. filiformis has the potential for rapid growth; estimated maximum weight-specific growth rates were more than 1.5% d^-1 in the disc diameter interval 3 to 9 mm, which is comparable to rates found in shallow water filtering bivalves. The maximum disc diameter growth rate was ca. 0.4 mm mo^-1.     

Reproduction of the intraovarian brooding apodid Leptosynapta clarki (Echinodermata: Holothuroidea) in British Columbia

Reproduction in the viviparous apodid sea cucumber Leptosynapta clarki was examined in Grappler Inlet, Bamfield, Vancouver Island, Canada from July 1989 through July 1991. The reproductive cycle of L. clarki is annual with gonad growth during the summer months. Male sea cucumbers spawn during November and December and internally fertilize, through as yet undetermined means, the oocytes within the ovary. Embryonic development is rapid, so that within 2 wk fertilized eggs reach the pentactula stage. Pentactulae have an open mouth and anus and actively move in the ovarian tubules feeding on droplets and fluid derived from the resorption of unfertilized eggs. Histological examination of gametogenesis shows similar development to that found in non-brooding holothurians. However, the protandric sex change observed in some male L. clarki after spawning and changes to the overian wall structure prior to fertilization are previously undescribed in sea cucumbers.     

Growth,production and ecological significance of Ophiura albida and O. ophiura (Echinodermata: Ophiuroidea) in the German Bight

Growth and production of the shallow-water ophiuroids Ophiura albida and O. ophiura were investigated at two stations in the German Bight from 1988 to 1991. Growth rings visible on the vertebral ossicles of the ophiuroid arms were interpreted as annual age markers. A correction for overgrown first rings allows for more exact estimations of growth and age. In both species growth could be described by Von Bertalanffy growth functions with the asymptotic disc diameter D =10.1 mm, K=0.229 and t _o=-0.192 in O. albida and D =27.7 mm, K=0.084 and t _o=0.042 in O. ophiura. Somatic production was calculated from mass specific growth rates. Annual production:biomass (P:B) ratios were estimated at 0.32 for O. albida and 0.43 for O. ophiura.AWI Publication Number: 618