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     

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.     

Born to lose. I. Measures of tissue loss and regeneration by the brittlestar Microphiopholis gracillima (Echinodermata: Ophiuroidea)

To measure amounts of tissue lost in natural populations of the burrowing amphiurid ophiuroid Microphiopholis gracillima (Stimpson), individuals were collected from subtidal mud flats in North Inlet, South Carolina, USA, at monthly intervals between February 1985–February 1987 and December 1989–November 1990. Between 20 and 70% of all individuals were regenerating the disc, and 85% of the 2045 arms examined had regeneration scars; >50% had one scar and some arms had up to 4 scars. Fewer individuals were regenerating discs in warmer months, but there was no seasonality in arm-loss frequency. To quantify rates of arm regeneration in the field, individuals which had 1, 2, or 3 arms removed were placed in mud-filled cores in the field in late July and November 1988 and in March and May 1989, and recovered after periods of about one month. Another set of cores was held in a running seawater laboratory during the May 1989 experiment. No regeneration occurred during the cooler times of year (November and March), and rates of regeneration were slower in May (total: 0.13 mg/d; tissue: 0.03 mg/d) than July (total: 0.17 mg/d; tissue: 0.05 mg/d). These rates indicate complete replacement of lost tissue in 100 to 120 d during the growth season. Within experiments, per arm regeneration rates were similar regardless of the number of arms removed. This finding is complicated by small sample size, high variability and low statistical power, but in general individuals which lost 2 or 3 arms regenerated proportionally more tissue than individuals which lost 1 arm. Individuals held in the laboratory regenerated the same amount of tissue but 30% less skeleton than individuals in the field. Sublethal tissue loss is common in this population, and M. gracillima is capable of regenerating at least 50% (each arm=17% of total body weight x 3) of its standing crop in a single growing season. Burrowing brittlestars probably constitute a significant renewable energy source for higher trophic levels in areas where they occur in dense populations.     

Growth cycle and related vertical distribution of the thecosomatous pteropod Spiratella (“Limacina”) helicina in the central Arctic Ocean

The growth cycle and related vertical distribution of the thecosomatous pteropod Spiratella (“Limacina”) helicina (Phipps) were studied. S. helicina has a life cycle of approximately 1.5 to 2 years in the central Arctic Ocean (Canada Basin). It spawns mainly during the spring to summer period, and on a small scale during the winter. The young double their sizes during the winter months of October to May, slow down in growth until late summer, and attain maximum size in early winter. The oldest disappear by late March. Gonadal tissue was first seen in young pteropods of 0.7 mm diameter, the predominant size from February to April. S. helicina 0.8 mm in diameter, the size predominant from May through July, are mature and hermaphroditic. Growth during the winter months suggests that particulate organic matter is available during this period to these obligate ciliary feeders. Vertical distribution is size and season-dependent. The youngest specimens collected (0.2 to 0.4 mm) were found concentrated in the first 50 m. The larger sizes dispersed during the summer months, and tended to concentrate in the top 150 m during the rest of the year. They aggregated in the top 50 m from late winter through early spring, and fall through early winter; then concentrated in the 100 to 50 m level until the end of winter. Numerous environmental factors seem to be involved in determining the vertical distribution of the species in the central Arctic Ocean.     

Dispersal and use of sponges and ascidians as camouflage by Cryptodromia hilgendorfi (Brachyura: Dromiacea)

From May 1977 to February 1979, the use of sponges and ascidians by Cryptodromia hilgendorfi was studied in Moreton Bay, Queensland, Australia. The aim was to investigate patterns of seasonal use, cap making behaviour, cap turnover, the effect of intraspecific interactions on cap life and the effect of movement of crabs between hosts on background matching. C. hilgendorfi uses 12 (of 16 available) species of sponge and 3 species of ascidians to construct caps, which are carried by the crabs using their last two pairs of legs. Cap area increases non-linearly with crab size, and caps are normally two to three times as large as the crabs. Cap making behaviour is described. It occurs during intermoult periods, with females making most of their caps at night. Caps decrease in size with time, but conceal the crabs which commonly occupy exposed sites on sponges. Cap life is independent of crab size, differs between different cap species and is influenced by the presence of other crabs who can dislodge caps through aggressive behaviour. Caps are made from the sponge Suberites carnosus more often than from other available sponges. S. carnosus caps also decay less rapidly than caps made from other sponges. Use of sponge and ascidian species varies seasonally, with Halichondria sp. and S. carnosus being used in all months. C. hilgendorfi exhibits a preference for certain sponges. The majority of crabs carried caps which matched their host sponge or ascidian, but mis-matches varied seasonally with a winter peak following the breeding season. Young C. hilgendorfi settle only on S. carnosus sponges and disperse from this host to other species in the environment. Males and females differ in their rate of discovery of new hosts. Males, despite their greater mobility, find new hosts slower than females. It is hypothesized that males occupy “home ranges” which females do not. Crabs frequently move between sponges, mostly at night. Sponges and ascidian species grow in intimate association with each other, and sponge crabs act a selective asexual propagation mechanism. Depending upon the nature of the interactions between sponge and ascidian species (co-operative or competitive) and whether competitive hierarchies or networks are involved, the sponge crabs may have either stabilizing or destabilizing effects on the sponge community.     

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     

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.     

Brooding behaviour of the northern sea star Leptasterias polaris

The northern sea star Leptasterias polaris (Müller and Troschel) has a unique mode of brooding its young. The mother curves her arms along the plane of the disc, the whole individual assuming a flattened plate-like shape. In this form she covers the larvae, which are attached to the substratum beneath her. The central disc is not raised to form a brood chamber, as in related species, and there is no tendency to hide while brooding. In the St. Lawrence Estuary, brooding L. polaris are common from February through May and decline in numbers during June and early July. The genus Leptasterias originates in the Arctic and probably all species brood their young.This paper is dedicated to François Dorion who lost his life during benthic studies in the St. Lawrence Estuary. His enthusiasm in studying marine organisms will always be an inspiration to usContribution to the program of GIROQ (Groupe Interuniversitaire de Recherches Océanographiques du Québec)     

Field survey of the occurrence and significance of regeneration in Amphiura chiajei (Echinodermata: Ophiuroidea) from Killary Habrour,west coast of Ireland

The boreo-Mediterranean amphiurid Amphiura chiajei Forbes occurs in high numbers (700 individuals/m²) in Killary Harbour, a fjordic inlet on the west coast of Ireland. 99.1% of the adult individuals show signs of arm regeneration, while 0.5% show regeneration of the disc. A study of the phenomenon commenced in November 1987 and continued on a seasonal basis until July 1989. Specimens were collected by SCUBA. The overall mean length of regenerated arm tissue over the 21 mo sampling period was 33.5±6.5 mm. Regeneration points were most common in the proximal and in the distal one-third of each arm. An overall mean of 4.21±0.3 arms per individual showed evidence of regeneration, while biomass assays revealed that regenerated tissue accounted for up to 57.9% of the total body weight. Anthropogenic activity (e.g. bottom trawling) may contribute to regeneration, although sub-lethal predation appears to be the main causitive factor of regeneration. Gut analyses of potential predators at the sampling station revealed that fish species (especially the Pleuronectidae) are particularly implicated. This predation seems to vary seasonally, being greatest in the summer months. Since the larger individuals showed the greatest evidence of arm regeneration, it is assumed that such sub-lethal damage is a persistent experience. Whatever the causality, the population of A. chiajei in Killary Harbour is deemed to be highly resilient.     

Multivariate analysis of the mud crab Scylla serrata (Brachyura: Portunidae) from four locations in Southeast Asia

The large, edible mud crab Scylla serrata (Forskål) exhibits different phenotypes which are recognised by the fishermen of Southeast Asia and are given local names such as “white”, “green”, “red” or “black” crabs on the basis of their colouration. A preliminary study using multivariate techniques was undertaken in order to examine the degree of dissimilarity between the different morphs of commercially fished stocks of S. serrata. Twenty-two morphometric and 20 meristic characters were measured on male crabs from four locations in three countries in Southeast Asia, including Surat Thani in Thailand where “black” and “white” morphs exist sympatrically. Canonical variate analysis (CVA) on the morphometric data discriminated three groups with no chain-linking. Surat Thani “white” crabs and those from south Vietnam were phenotypically similar and formed one distinct group; a second group contained crabs from Ranong (Thailand) and Sarawak which were phenotypically “black”; the third group contained “black” crabs from Surat Thani. CVA on meristic data confirmed two groupings, but implied that the Surat Thani “black” morph may only be a variant of the Surat Thani/Vietnam “white” form, rather than a third species. These findings are discussed in relation to the ecology, fishery management and aquaculture potenial of S. serrata.     

Biology of euphausiids in the subarctic waters north of Iceland

The seasonal abundance, maturity, spawning, and population dynamics of Thysanoessa inermis (Krøyer, 1846), T. longicaudata (Krøyer, 1846), and Meganyctiphanes norvegica (M. Sars, 1857) were studied in the subarctic waters north of Iceland from February 1993 to February 1994. The material was sampled at approximately monthly intervals along a transect of eight stations extending from 66°16′ to 68°00′N at 18°50′W. Information on temperature and chlorophyll a concentrations is also presented. Spring warming of the water began in March to April and maximum temperatures were recorded in August (3.8?°C). The spring bloom of the phytoplankton started in late March and highest chlorophyll a concentrations were measured during middle to late April (7.0?mg chlorophyll a m^?3). T. inermis was the dominant species in the samples, constituting 77% of juvenile, male and female euphausiids present. The greatest abundance of juvenile, male and female T. inermis and M. norvegica was observed during autumn and winter, with lower abundance in spring and summer. T. longicaudata showed only limited changes in seasonal abundance. Male T. inermis had spermatophores in their ejaculatory ducts from February to May, while mature females had spermatophores attached during April and May. T. longicaudata males bore spermatophores from February to July, whereas females only bore spermatophores in April and May. M. norvegica males had spermatophores from February to April, while the single female with spermatophores was caught in February. Euphausiid eggs were first recorded during the latter part of April; the highest numbers of eggs were observed in the samples taken in late May. Maximum numbers of nauplii of both Thysanoessa spp. and M. norvegica were recorded in late May. The main spawning of the euphausiids coincided with the phytoplankton spring bloom. Most male T. inermis took part in breeding at 1 yr of age while most females appeared not to mature until 2 yr of age. T. inermis has a life span of just over 2 yr, T. longicaudata appears to live just over 1 yr. Limited data did not allow the life span of M. norvegica to be determined.     

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     

One size fits all: stability of metabolic scaling under warming and ocean acidification in echinoderms

Responses by marine species to ocean acidification (OA) have recently been shown to be modulated by external factors including temperature, food supply and salinity. However the role of a fundamental biological parameter relevant to all organisms, that of body size, in governing responses to multiple stressors has been almost entirely overlooked. Recent consensus suggests allometric scaling of metabolism with body size differs between species, the commonly cited ‘universal’ mass scaling exponent (b) of ¾ representing an average of exponents that naturally vary. One model, the Metabolic-Level Boundaries hypothesis, provides a testable prediction: that b will decrease within species under increasing temperature. However, no previous studies have examined how metabolic scaling may be directly affected by OA. We acclimated a wide body-mass range of three common NE Atlantic echinoderms (the sea star Asterias rubens, the brittlestars Ophiothrix fragilis and Amphiura filiformis) to two levels of pCO₂ and three temperatures, and metabolic rates were determined using closed-chamber respirometry. The results show that contrary to some models these echinoderm species possess a notable degree of stability in metabolic scaling under different abiotic conditions; the mass scaling exponent (b) varied in value between species, but not within species under different conditions. Additionally, we found no effect of OA on metabolic rates in any species. These data suggest responses to abiotic stressors are not modulated by body size in these species, as reflected in the stability of the metabolic scaling relationship. Such equivalence in response across ontogenetic size ranges has important implications for the stability of ecological food webs.     

Chaotic genetic patchiness and high relatedness of a poecilogonous polychaete in a heterogeneous estuarine landscape

The genetic structure of benthic marine invertebrates is often described as “chaotic” when genetic structure cannot be explained and barriers to dispersal and gene flow cannot be identified. Here, chaotic patterns of genetic structure for the polychaete Pygospio elegans (Claparède) sampled at 16 locations from the heterogeneous Isefjord–Roskilde Fjord estuary complex in Denmark were found. There was no isolation by distance, and the geography of the estuary complex did not seem to pose a barrier to dispersal and gene flow in this species. We investigated whether characteristics of the environment could be related to the genetic structure and possibly restrict gene flow in this species. Additionally, since P. elegans is poecilogonous, producing larvae with different pelagic developmental periods, we investigated whether observed developmental modes in the samples might clarify the genetic patterns. None of the tested factors explained the population genetic structure. However, a high degree of relatedness among individuals in almost all samples was found. Samples with a larger percentage of young individuals had more related individuals, suggesting that different cohorts could be comprised of individuals with different degrees of relatedness. Relatedness within a site could be increased by limited larval dispersal, collective dispersal of related larvae, sweepstakes reproductive success, or asexual reproduction, but distinguishing between these requires further study. Using a “seascape genetics” approach allowed us to investigate some of the numerous potential factors that could influence population genetic structure in a poecilogonous species.     

Nachweis der epitokie als fortpflanzungsmodus des polychaeten Eunice siciliensis (Polychaeta: Annelida)

Eunice siciliensis (Grube)¹ is a sedentary polychaete with separate sexes; its germ cells develop only in the posterior part of the body. In March and April, females with whitish or even dark-green oocytes, males with only few spermatogonia, and individuals with no germ cells at all were discovered in the “coralligène” of the Banyuls region (European Mediterranean Sea). In august, besides immature and sexually non-differentiated worms, mature individuals of both sexes were captured: females with dark bluish-green oocytes of diameters up to 250μm, and males with milky genital segments containing spermatozoa. Soon after isolation from the substrate, these individuals performed characteristic movements with their posterior body parts, which then autotomized, and released ova and spermatozoa. Evidence is given that the anterior, atokous body parts survive in their tubes in the “coralligène” after separation from the epitokous portions, and regenerate new genetal segments. The development of gametes observed in these regenerated segments indicates the ability of E. siciliensis to reproduce more than once during its lifetime. Not only caudal, but also prostomial regeneration was observed; the formation of a (morphologically different) secondary prostomium appears to be essential for any development of germ cells in worm fragments. E. siciliensis exhibits surprising similarities in habits and in formation of an epitokous form with Eunice viridis (Gray), the “palolo” worm of the South Pacific Ocean, but nevertheless, lacks the prominent criteria of this famous species: the ventral eye spots on each genital segment in males and females, and the paired areas with brown pigmentation on the ventral side of each epitokous segment in the males.     

Sexualité d'Asterina gibbosa,asteride hermaphrodite,des côtes de Marseille

Asterina gibbosa from Marseille, a protandric hermaphrodite is considered belonging to a balanced sexual race similar to the Dinard (France), Plymouth (England) and Banyuls (France) reces. Spermatogenetic activity is seasonal. Sexreversal occurs when animals are 3 years old (average arm length (R)=12 mm). Ultimately, sea-stars with ovogenetic reactions could produce a spermatogenetic resurgence. Few individuals are “permanent males”.     

Supralittoral amphipod abundances across habitats on Mediterranean temperate beaches

Community complexity and abiotic conditions are key components of environmental heterogeneity that affect the abundance and distribution of species. In this study we evaluated how environmental conditions affect abundances of supralittoral amphipods (Talitridae) in four habitats (sandy beach, rivermouth, wrack and lakeshore), along the Italian peninsula in the Mediterranean Sea. All samplings covered a 12 month period, and used the same sampling methodology thereby enabling comparison of abundances and species composition and richness. Four species (Talitrus saltator (Montagu, 1808), Orchestia gammarellus (Pallas, 1766), O. montagui Audouin 1826, O. cf. cavimana Heller 1865) were collected in the different habitats, but most species were found or were abundant in only one of the four habitats. Abundances of talitrids (numbers per sampling hour) differed significantly among the habitats with highest abundances found in the wrack and on the riverbank in proximity to an estuary, and lowest abundances observed on four sandy beach sites. Environmental conditions (temperature, moisture, substrate penetrability) differed among the habitats and were associated with some of the among-site variability in abundances. Our findings demonstrate that talitrids thrive better in some supralittoral habitats than others, and that some habitats could be considered to be “hotspots” of talitrid ecology and biodiversity.     

Evaluation and assessment of the efficacy of an abatement strategy in a former lead smelter community,Boolaroo, Australia

This study examines the recent soil Lead Abatement Strategy (LAS) in Boolaroo, New South Wales, Australia, that was designed to “achieve a reduction in human exposure to lead dust contamination in surface soils”. The abatement programme addressed legacy contamination of residential areas following closure of lead smelting operations in 2003 at the Pasminco Cockle Creek Smelter (PCCS). The principal objective of the LAS was to “cap and cover” lead-contaminated soils within the urban environment surrounding the PCCS. Soil lead concentrations of 2500–5000 mg/kg were scheduled for removal and replacement, while concentrations between 1500 and 2500 mg/kg were replaced only under limited circumstances. To date, there has been no industry, government or independent assessment of the clean-up programme that involved >2000 homes in the township of Boolaroo. Thus, by measuring post-abatement soil lead concentrations in Boolaroo, this study addresses this knowledge gap and evaluates the effectiveness of the LAS for reducing the potential for lead exposure. Soil lead concentrations above the Australian residential soil health investigation level value for residential soils (300 mg/kg) were identified at all but one of the residential properties examined (n = 19). Vacuum dust samples (n = 17) from the same homes had a mean lead concentration of 495 mg/kg (median 380 mg/kg). Bio-accessibility testing revealed that lead in household vacuum dust was readily accessible (% bio-accessible) (mean = 92 %, median = 90 %), demonstrating that the risk of exposure via this pathway remains. Assessment of a limited number of properties (n = 8) where pre-abatement soil lead levels were available for comparison showed they were not statistically different to post-abatement. Although the LAS did not include treatment of non-residential properties, sampling of community areas including public sports fields, playgrounds and schools (n = 32) was undertaken to determine the contamination legacy in these areas. Elevated mean soil lead concentrations were found across public lands: sports fields = 5130 mg/kg (median = 1275 mg/kg), playgrounds and schools = 812 mg/kg (median = 920 mg/kg) and open space = 778 mg/kg (median = 620 mg/kg). Overall, the study results show that the LAS programme that was dominated by a “cap and cover” approach to address widespread lead contamination was inadequate for mitigating current and future risk of lead exposures.     

Rythmes nutritionnels et organisation trophique d'une population de crustacés pélagiques (Euphausiacea)

The nature of the food (animal, plant or mixed) and the fullness of the stomachs at different times of the day have been studied through dissection of 18620 specimens representative of almost all the euphausiid species of the Central and Western Tropical Pacific Ocean. Animal food predominates in 22 of the 28 species studied, while 12 can be considered as omnivorous; only 4 are mainly phytophagous. The trophic level of a given species is more or less the same in different zones, but scarcity of phytoplankton in oligotrophic tropical regions results in an increase of the trophic level of herbivorous and omnivorous species. There is no correlation between trophic level and vertical distribution of a species. Each species follows a clearly defined feeding rhythm, usually characteristic for each genus: nutrition most active by night in Euphausia, continuous in Thysanopoda, restricted to light hours in Stylocheiron, mainly from noon to midnight in Nematoscelis and Nematobrachion. As a rule, it is obvious that the smaller the daily vertical migration, the more pronounced the feeding rhythm: the range of fluctuations in fullness of stomachs over 24 h is weak or non-existent in migrating species, maximum in non-migrating ones. Nevertheless, daily vertical migration does not appear to be “advantageous” from the point of view of trophic efficiency: assuming that the stomachal transit is the same for all species (a speculative proposition), it is shown that the ratio “total biomass of species: food consumed during 24 h”, i.e., “biomass permanently available for the upper link: biomass eaten daily at the expense of the lower link” is 4 times higher in non-migrating than in migrating species. It is thus considered that daily vertical migration is an expensive manner to transfer energy from link to link, and therefore fulfills other functions, amongst which diffusion throughout the whole water column of the biomass produced in the upper levels is probably one of the most important.