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     

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.     

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.     

Reproduction in Amphiura filiformis (Echinodermata: Ophiuroidea): Seasonality in gonad development

The burrowing ophiuroid Amphiura filiformis (O. F. Müller) (Echinodermata: Ophiuroidea) colonises an extensive area and is a numerically dominant member of the macrofauna in Galway Bay on the west coast of Ireland. There, it has a discrete, relatively short annual breeding period, from June to September, with peak activity in the month of August. The breeding period approximates in timing those of other European populations. The oocytes of a particular year are laid down in the autumn of the previous year; they begin to develop in spring, with the period of fastest growth in May/June. Gonad growth corresponds quite closely to the annual rise in water temperature, with spawning taking place during the warmest months of the year. The species is polytelic, and it is suggested that the long-lived Galway Bay individuals may breed every year for several years. The relationship between spawning and population dynamics is discussed in the light of current, available literature concerning A. filiformis from other parts of Europe. The role of so-called ultimate factors in the process of ophiuroid maturation and spawning is discussed.     

Tolerance to low oxygen and sulfide in Amphiura filiformis and Ophiura albida (Echinodermata: Ophiuroidea)

The brittle stars Amphiura filiformis (Müller) and Ophiura albida (Forbes) were exposed to different oxygen saturations (100, 10, 5, 3, and <1% oxygen saturation) and to physiological anoxia (<1% oxygen saturation) at different total sulfide concentrations (0, 2, 20, 200 μM). The mortality was followed during experiments and the median survival time (LT₅₀) was determined. The infaunal A. filiformis had a significantly higher tolerance to both hypoxia and sulfide than did the epibenthic O. albida. After exposure to 10% oxygen saturation for a month, only 2.0% A. filiformis and 0% O. albida were dead. In oxygen saturations <1% A. filiformis and O. albida had a LT₅₀ of 7.5 and 2.5 d, respectively. The presence of even very small concentrations of sulfide decreased the survival significantly. Sulfide is shown to be the key factor for the survival of the two species. Received: 11 October 1996 / Accepted: 12 November 1996     

Unusual lecithotrophic development of the caribbean brittle star Ophiothrix oerstedi

The ophiuroid Ophiothrix oerstedi Lütken spawned in the laboratory at Barbados, West Indies, from August, 1975 until early December. The embryo passes through a wrinkled blastula stage, and the larva is a reduced lecithotrophic ophiopluteus with a shortened pelagic existence. A larva of this type is unusual for brittle stars in general and unique for ophiothricids for which development has been described. Metamorphosis is completed 4.3 days (24° to 27°C) after fertilization with a single pair of ciliated larval arms, the postero-lateral arms, being retained as a swimming device for the late larva. Settlement, with subsequent separation of the postero-lateral arms from the young brittle-star, begins as early as 4.5 days, but can be delayed for at least one week, at the end of which time midwater separation can occur resulting in the pelagic dispersal of post-larvae. A comparison of gametic and larval characteristics of O. oerstedi with the literature suggests that the larva of this species is most closely allied to the abbreviated developers. The adaptive significance of this larval form is discussed.     

Photoperiodic activity pattern in the brittle star<Emphasis Type="Italic"> Amphiura filiformis</Emphasis>

Diurnal activity pattern of suspension-feeding arms of the infaunal brittle star Amphiura filiformis was recorded in situ by time-lapse photography at 30 m water depth in the Gullmarsfjord, west Sweden. Activity was clearly related to the photoperiod with high activity at night and low or no activity during daytime. In laboratory experiments, the activity of the arms of A. filiformis changed in relation to new manipulated photoperiods, demonstrating exogenous circadian rhythms. The time difference for the greatest change in activity of the arms between control and treatment was correlated with the photoperiodicity. The recently found microlenses and associated neural photoreceptors in ophiuroids may apply also to A. filiformis. The trade-off between efficient feeding and predator avoidance is discussed.Communicated by M. Kühl, Helsingør     

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.     

Non-viable seasonal settlement of larvae of the upper bathyal brittle star Ophiocten gracilis in the Rockall Trough abyssal

Re-examination of ine structure of postlarvae previously identified as the abyssal species Ophiura ljungmani (Lyman) in a time series of samples taken from 1975 to 1980 from a 2900 m deep permanent station in the southern Rockall Trough (Northeast Atlantic Ocean) has shown the majority of the large summertime peak of brittle-star postlarvae to belong to a second species. Most were incomplete and probably corpses when collected. The few intact specimens present were identical to postlarvae collected in benthic samples from the neighbouring continental slope, where a dense population of the bathyal species Ophiocten gracilis (G. O. Sars) is present from 600 to 1 200 m depth. Examination of population size-structures indicates synchronous reproduction in January/February and annual recruitment to the population. The presence both of Ophiopluteus ramosus and intact postlarvae of Ophiocten gracilis in midwater plankton samples from the vicinity of the permanent station in March/April are thought to derive either from the adjacent slope population or, less probably, from larvae transported in the North Atlantic Current from the Flemish Cap area off Newfoundland. In deep water, viability of these benthic postlarvae is low, none surviving longer than the following winter in the Rockall Trough. These observations suggest a large seasonal transfer from the slope of non-viable larvae and postlarvae as food to the abyssal benthic community.Communicated by J. Mauchline, Oban     

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.     

Phylogeography of the Antarctic planktotrophic brittle star Ophionotus victoriae reveals genetic structure inconsistent with early life history

In the marine environment, connectivity is influenced by physical oceanography as well as life history and behavioral traits, which in combination with historical events such as geology, physical oceanography, and climate, determine population structure. The Antarctic brittle star Ophionotus victoriae develops via a feeding planktonic larval stage, and therefore has potential for long-distance dispersal throughout its Antarctic/subantarctic range. To evaluate this hypothesis, phylogeography of this ecologically dominant species was elucidated by sequence analysis of two mtDNA genes from individuals collected throughout the Antarctic Peninsula and from two subantarctic islands. Counter to expectations of genetic homogeneity, mtDNA data revealed substantial levels of genetic differentiation as well as diversity. Although there were some genetically homogeneous populations, such as those throughout Bransfield Strait, we found O. victoriae to have significant population structure throughout much of the Antarctic Peninsula, with evidence of potential cryptic speciation between the western and eastern Antarctic Peninsula. Furthermore, Antarctic Peninsula populations were genetically distinct from subantarctic island populations. The low levels of connectivity implied for O. victoriae contrast with those found for many other Antarctic benthic taxa, and suggest a complex interplay between oceanography, recent climate history, and larval ecology.     

Divide and broadcast: sexual reproduction in the West Indian brittle star Ophiocomella ophiactoides and its relationship to fissiparity

It is known that asexual reproduction by fission is the dominant mode of propagation in the small (disc diameter <5.0 mm) ophiocomid brittle star Ophiocomella ophiactoides (H. L. Clark). This species is, however, able to reproduce sexually as well. Sexual reproduction occurs throughout much or all of the year in a population of O. ophiactoides at Jamaica. Gonads are often present in recently split individuals, indicating that sexual and asexual reproduction could occur simultaneously. Regeneration of new gonads in the new disc half is underway by the time an individual is competent to split again. Unsexable individuals are significantly smaller than males, which in turn are significantly smaller than females. The unsexable individuals appear to be below a threshold size for gonad production. Fecundity is low, with the largest females producing a maximum of about 7 400 eggs of small (mean diameter 80 m) size. Fertilized eggs develop into planktotrophic ophioplutei which are typical of the larvae of other species of ophiocomid brittle stars in morphology, rate of development and duration of planktonic period. Egg size and larval type for O. ophiactoides are characteristic of brittle stars with the planktotrophic mode of development, but egg number is uncharacteristically low. Fissiparity in O. ophiactoides may have evolved in association with small body size, concomitant low fecundity and a high-risk planktotrophic larva as a means of ensuring reliable recruitment. Available data, although sparse, suggest that this hypothesis may be applicable to fissiparous brittle stars in general. Fission in brittle stars has a broad systematic incidence at the familial level which suggests that it has evolved recurrently. The ability both to divide and broadcast appears to be a useful combination for O. ophiactoides: it can recruit reliably at large size to occupy favourable habitats quickly, but, it can still maintain low level dispersal of larvae which could provide escape from deteriorating habitats and the potential to colonize new habitats.Contribution No. 328 of the Discovery Bay Marine Laboratory, University of the West Indies     

Submersible and laboratory observations on Asteroschema tenue,a long-armed euryaline brittle star epizoic on gorgonians

In situ and laboratory obervations of the biology and behaviour of the deep-sea, euryaline brittle star Asteroschema tenue Lyman, from Jamaica, are described. The brittle star is found clinging to the gorgonian Ellisella barbadensis at depths of 168 to 230 m off Jamaica, and is here restricted to this gorgonian. The brittle star, which is usually cryptically coloured, is active principally nocturnally, when it extends arm loops and employs the tube feet for microphagous filter-feeding. Locomotory activity is limited to extension and retraction along the gorgonian, achieved by means of unusual arm-loop patterns. Association with a gorgonian appears to be essential for the survival of the brittle star. The study was conducted in 1985.Order of authors' names is arbitrary     

Re-appraisal of age composition,growth and survivorship of the deep-sea brittle star Ophiura ljungmani from size structure in a sample time series from the Rockall Trough

Samples were taken from a 2 900 m deep permanent station (centred on 54°40N; 12°16W) in the southern Rockall Trough, from November 1975 to September 1980, inclusive. Revision of juvenile size frequencies was necessary after removal of postlarvae of an annual non-viable settlement of Ophiocten gracilis (G. O. Sars) that were mistakenly included in a previous analysis. The present study of the revised data shows that settlement of Ophiura ljungmani (Lyman) occurs in summer when disk-size frequencies are dominated by the newly metamorphosed postlarvae. The likely age composition in larger sizes was suggested from the progression of modes along the size axis in juvenile size classes in consecutive were developed from Gaussian mixtures fitted by constrained computer search using, as starting values, component parameter values suggested from the frequency histograms. Choice between competing solutions was made on the basis both of goodness-of-fit and biological plausibility. However, identification of year-class structure amongst sizes >3 to 4 mm disk diameter remained conjectural owing to the low frequencies of adults in the samples. Comparison of the fitted component-means in the time series suggested a marked seasonal growth pattern, with maximal growth in spring. Probable age composition amongst adult sizes was extrapolated from von Bertalanffy growth parameters estimated from the means of juvenile year classes.Communicated by J. Mauchline, Oban     

Diets of some deep-sea brittle stars in the Rockall Trough

The stomachs of 1 165 individuals of the species Ophiacantha bidentata (Retzius), Ophiactis abyssicola (M. Sars), Ophiocten gracilis (O. Sars), Ophiura irrorata (Lyman), O. ljungmani (Lyman) and Ophiomusium lymani Wyville Thomson, collected from the general area of the Rockall Trough from 1973 to 1983, were examined; 47% contained material. This varied in nature and in volume between individuals, but overall, the diets of the 6 species showed quite high similarity. The few differences evident are interpreted as reflecting different lifestyles. Except for Ophiocten gracilis, the probably motile, epifaunal species (Ophiura irrorata, O. ljungmani and Ophiomusium lymani) showed the greatest variety in items, and the lowest similarity with the remaining species; they are probably unselective omnivores, eating whatever small prey or organic detritus they are able to find. Ophiacantha bidentata and Ophiactis abyssicola showed both the least dietary variety, and the highest similarity to each other. This probably reflects their more sedentary lifestyle, feeding on current-borne particles and small prey from perches on sessile fauna. The stomach contents of Ophiocten gracilis consisted mainly of amorphous organic material and mineral particles, indicating a microphagous feeding mechanism. With O. gracilis, the proportion of stomachs with food decreased markedly in summer, this coinciding with seasonal gonadal development when stomachs frequently appeared ruptured or compressed. The possible importance of seasonal sedimentation of fast-sinking particulates from the surface is discussed in relation to finds of flocculent organic material, often containing diatom frustules, in stomachs of all 6 species. Some doubts remain as to whether our results are biased by specimens egesting stomach contents on capture. As in shallow water, deep-sea ophiuroids seem to be trophic generalists lacking in dietary specialization. Brief notes on parasites encountered are also given.     

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.     

Male response to an aggressive visual signal, the wing wave display, in swamp sparrows

Signaling often involves complex suites of behaviors that incorporate different sensory modalities. Whatever modality is used to establish that a signal functions in communication researchers must demonstrate that receivers respond to it. The territory defense response of male swamp sparrows involves a variety of behaviors that includes both vocal and visual displays. One of these, the “wing wave” display, is a distinctive movement that predicts physical attack. Here, we use robotic taxidermic mounts paired with song to test the hypothesis that wing waving is a signal and, specifically, that male receivers respond to wing waving as a signal of aggressive intent. As predicted, subjects responded more aggressively to the mount during wing waving trials than during stationary trials. A second experiment demonstrated that this effect cannot be attributed simply to increased attention to movement. Less expectedly, subjects did not alter their own display behavior in response to wing waving as compared to a static mount. We conclude that the wing wave display in the context of singing is a signal that functions in male–male aggressive communication. Questions remain, including whether wing waving functions as a signal in the absence of singing and whether wing waving and song are redundant signals or communicate different information.     

Dislocation experiments in the migrating sea star Astropecten jonstoni

Movements of tagged specimens of the sea star Astropecten jonstoni (D. Ch.), both spontaneous and after displacement, were studied by means of SCUBA diving in shallow coastal waters of Sardinia (Mediterranean Sea). In spring, the entire population nigrates towards the shore. Displaced sea stars were able to return to their original depth zone. Both migration and return to the original depth zone occur in a unidirectional way.