Sulfide assimilation by ectosymbionts of the sessile ciliate, <Emphasis Type="Italic">Zoothamnium niveum</Emphasis>

We investigated the constraints on sulfide uptake by bacterial ectosymbionts on the marine peritrich ciliate Zoothamnium niveum by a combination of experimental and numerical methods. Protists with symbionts were collected on large blocks of mangrove-peat. The blocks were placed in a flow cell with flow adjusted to in situ velocity. The water motion around the colonies was then characterized by particle tracking velocimetry. This shows that the feather-shaped colony of Z. niveum generates a unidirectional flow of seawater through the colony with no recirculation. The source of the feeding current was the free-flowing water although the size of the colonies suggests that they live partly submerged in the diffusive boundary layer. We showed that the filtered volume allows Z. niveum to assimilate sufficient sulfide to sustain the symbiosis at a few micromoles per liter in ambient concentration. Numerical modeling shows that sulfide oxidizing bacteria on the surfaces of Z. niveum can sustain 100-times higher sulfide uptake than bacteria on flat surfaces, such as microbial mats. The study demonstrates that the filter feeding zooids of Z. niveum are preadapted to be prime habitats for sulfide oxidizing bacteria due to Z. niveum’s habitat preference and due to the feeding current. Z. niveum is capable of exploiting low concentrations of sulfide in near norm-oxic seawater. This links its otherwise dissimilar habitats and makes it functionally similar to invertebrates with thiotrophic symbionts in filtering organs. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Endemism and dispersal: comparative phylogeography of three surgeonfishes across the Hawaiian Archipelago

To evaluate the hypothesis that a general correlation exists between species range size and dispersal ability, we surveyed mitochondrial cytochrome b sequence variation in three surgeonfish species with vastly different ranges: Ctenochaetus strigosus, Hawaiian endemic, N = 531; Zebrasoma flavescens, North Pacific, N = 560; Acanthurus nigrofuscus, Indo-Pacific, N = 305. Collections were made throughout the 2,500 km expanse of the Hawaiian Archipelago and adjacent Johnston Atoll. Analyses of molecular variance demonstrate that all three species are capable of maintaining population connectivity on a scale of thousands of km (all species global Φ_ST = NS). However, rank order comparison of pairwise Φ_ST results and Exact test P-values revealed modest but significantly different patterns of gene flow among the three species surveyed, with the degree of genetic structure increasing as range size decreases (P = 0.001). These results are consistent with mtDNA surveys of four additional Hawaiian reef fauna in which a wide-spread Indo-Pacific species exhibited genetic homogeneity across the archipelago, while three endemics had significant population subdivision over the same range. Taken together, these seven cases invoke the hypothesis that Hawaii’s endemic reef fishes evolved from species with reduced dispersal ability that, after initial colonization, could not maintain contact with parent populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Anomalies de structure des radioles de Heterocentrotus mammillatus (Echinodermata: Echinoidea) en microcosme in vitro

After a 3 mo period in a confined (closed-circuit) microcosm, all speciment of the echinoid Heterocentrotus mammillatus (L.), originally collected from the fringing coral reef of the Gulf of Aqaba (Jordan), exhibited anomalies of their long primary spines, both aboral and adoral, especially at the distal extremities. The abnormality was confined to the top portion of the spine (1 to 2 cm in length) and consisted of (1) the formation of a callosity, and (2) a change in the terminal shape of the spine to either a very sharp thorn, or to a triple carination. The surface of the spine became rough to the touch, and its original colour notably faded. The very marked structural degeneration of the surface microstructure progresses from the callosity to the tip of the spine in 4 phases: (1) the surface of the grooves, originally smooth, becomes perforated; (2) the perforations become increasingly larger (both in width and depth): (3) the roughness increases; (4) the overall appearance of the microstructure becomes necrotic. Transverse sections of the distal extremity of such spines revealed a disorganized internal microstructure with a network of finer meshes which partially disappear in some zones. The normal microstructural arrangement of the spine-in ordered, concentric and radial lines-is replaced by a disordered network, and tomographs of the spines reveal a decalcification of the whole portion of the spine distal to the callosity. Some illustrations of the different phases of the progressive disorganisation of the characteristic microstructure of apices in abnormal spines are quite similar to illustrations by authors investigating regeneration of broken spines, but the process of necrosis proceeds by stages which are in an inverse order to those observed in regeneration. Such abnormalities of shape and structure of the spine have hitherto never been recorded for natural populations of H. mammillatus from any coral reefs in the Indo-Pacific region. Certainly, the abnormal spines we observed after a 3 mo period within a closed microcosm might be ascribable to a malfunction of the calcification mechanism, since the characteristic callosity never appeared in regeneration experiments. The biotope of H. mammillatus is the upper levels of very exposed coral reefs, and its food chiefly consists of Corallinacea which are abundant therein. In contrast, specimens transferred to this sheltered and confined environment, where the diet is impoverished, display an insufficient calcification rate in the extremities of their spines, as if the calcium content in the whole body is so depleted that this element can not reach the tips of the spines. In contrast, the thickness of the test plates appears normal. The study was carried out in 1978.     

Penetration and storage of sponge spicules in tissues and coelom of spongivorous echinoids

When echinoids feed on sponges, silicate spicules of the sponge were found to enter their body either by penetration through the wall of the food canal into coelomic cavities or by penetration into skeletal plates and spines. The spicules, which have penetrated into the coelom, obviously evoke a kind of protective answer. They were found to be entangled by clusters of cell remnants, the so-called brown bodies. The brown bodies contain melanin and gather at special sites of the echinoid body; these are the Stewart Organs, the gills and the inner side of the ambulacral plates. Sometimes the silicate spicule becomes surrounded by a calcareous sheath. The length of the sponge spicules makes their removal impossible, so that they are stored. The spicules penetrating into the plates are partly incorporated into the stereom. The four species examined in this study were Asthenosoma ijimai, Araeosoma owstoni, Diadema setosum (collected in Sagami Bay, Japan in 1991) and Hapalosoma gemmiferum (collected in Suruga Bay, Japan in 1991).     

Embryonic,larval, and post-metamorphic development of the sea urchin <Emphasis Type="Italic">Cassidulus mitis</Emphasis> (Echinoidea; Cassiduloida): an endemic brooding species from Rio de Janeiro,Brazil

Cassiduloids are currently rare irregular echinoids with a highly conserved adult morphology. Aristotle’s lantern is present only during the post-metamorphic stage, and little is known about the early development of species in this group. Cassidulus mitis produces eggs of about 375 μm in diameter, lecithotrophic larvae with four reduced arms with skeletal fenestrated rods, cilia along the body surface, and a ciliated band on arms and lobes. Offspring is brooded among the female spines from embryo to settler’s stage. The echinopluteus larval stage is reached 6 days after fertilization, and the settler’s stage is formed at the age of 17 days. Aristotle’s lantern appears around the thirteenth day of development. The lantern is well developed and functional in settlers. It remains until at least 62 days after fertilization and can be used to acquire food from the environment. The early development of C. mitis is unusual concerning features of typical lecithotrophic larvae (such as reduced arms), but retains some features of planktotrophic larvae (such as skeletal rods and a ciliated band). Regarding egg size, early development in C. mitis seems to be transitioning from facultative lecithotrophic to typical obligate lecithotrophic pattern in echinoid larval evolution.     

Genetic diversity of Nymphon (Arthropoda: Pycnogonida: Nymphonidae) along the Antarctic Peninsula with a focus on Nymphon australe Hodgson 1902

Sea spiders are conspicuous, and often abundant, members of the Antarctic benthic community. Nymphonidae (Pycnogonida) in Southern Ocean waters comprise over 240 species which are often difficult to assign due to their intraspecific ‘highly variable’ morphology. In particular, Nymphon australe, the numerically dominant species in Antarctic waters is known to have a high level of phenotypic variation in external morphology and is also reported to have a circumpolar distribution. Circumpolarity seems contradictory to the pycnogonid’s brooding lifestyle and presumably limited dispersal. Here we examine the genetic diversity of several Nymphon species collected in the Antarctic Peninsular region. Concomitantly, we assess the genetic structure of N. australe to gain insight into Nymphon dispersal capacity. Cytochrome c oxidase subunit I (COI) and 16S ribosomal gene data suggest a recent common history and/or recent gene-flow of N. australe populations across nearly 800 km of the Antarctic Peninsula. Furthermore, these data support that the Antarctic Peninsula region may hold two previously unrecognized species of Nymphon. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Particularités microstructurales du squelette de Paracentrotus lividus et Arbacia lixula: Rapports avec l'écologie et l'éthologie de ces échinoïdes

Scanning-electron-microscope investigations on the test microstructure of two regular sea urchins, Paracentrotus lividus (Lamarck) and Arbacia lixula Linné have been especially devoted to the sutures between test plates, and the spines. Some features of both these components may be related to ecological and ethological differences between the two species. It is well documented that the structure of the sea urchin's test plates consists of a meshwork of calcareous trabecules embedded within the mesenchyme. The suture between the two lines of plates of each radial or intertradial zone exhibits a gap which plays some part in the process of the plates' growth. However, the gap may also constitute a complementary stress-breaker of mechanical forces (such as waves) which are exerted upon the test in the natural environment. In A. lixula this gap is so wide and the height: diameter ratio so low, that this species is particularly well fitted to bear physical stress and force on its apical region: since A. lixula almost exclusively inhabits vertical or subvertical rocky substrates at 2 to 15 m depth, where wave action is mainly exerted perpendicular to the substrate, these particular features of sutures and test shape may be a morphofunctional adaptation to this habitat. The sutures of P. lividus exhibit a narrower gap, making this species less able to bear strong apical pressure; consequently, P. lividus usually occurs on exposed horizontal or gently inclined substrates, but also inhabits sea-grass beds as deep as 15 to 20 m. The spines of P. lividus bear deep longitudinal grooves with lateral teeth, which seem especially fitted for collecting and transporting organic particles from the top of the spines to the apical region of the test where they are digested and assimilated by coelomocytes and epithelial cells. In the natural environment, the slightest water motion provides the spines with suspended particles; in extremely sheltered places or in aquaria however, the spines actively collect particles. Therefore, P. lividus populations can thrive in places where food resources other than suspended particles are scarce. The grooves on the spines of A. lixula are less marked, and thus unsuitable for collection of suspended particles. This species can therefore fulfil its energy requirements only by grazing and absorption of dissolved material. It appears that although belonging to the same biocoenosis, P. lividus and A. lixula do not occupy the same ecological niche. Their potential utilization in either urban or chemical pollution monitoring studies is discussed.     

Laboratory culture of the aeolid nudibranch <Emphasis Type="Italic">Spurilla neapolitana</Emphasis> (Mollusca,Opisthobranchia): life history aspects

Spurilla neapolitana from the Eastern Mediterranean was cultured in a self-sustained, temperature-controlled laboratory culture system, and its life cycle is described. Adults were collected from three field sites situated 120 km apart, along the Israeli Mediterranean coastline, between March 2006 and August 2007. Cultures of the life-cycle stages were raised at 24°C. S. neapolitana deposited white, coiled, spiral egg masses containing zygotes. Veliger larvae hatched 3.0 ± 0.4 days post oviposition. The veliger larvae are obligatory planktotrophs, with a minimal larval phase of 22 days. In the lab, larvae settled and metamorphosed following exposure to metabolites derived from distinct prey sea anemone species. Reproductive maturity was reached 42 ± 5 days post metamorphosis, resulting in a laboratory generation time of 67 days (egg to egg). The average life span of reproductive specimens in this study was 157 ± 13 days post-oviposition and they reached a length of 7–10 cm. During this period, an average adult deposited ca. 40 × 10⁶ zygotes. This species has several characteristics that suggest it will be a useful model for laboratory-oriented research. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Comparative phylogeography of marine cladocerans

We examined the population genetics of six species of marine cladocerans, using a ~600 bp fragment of the cytochrome oxidase subunit I gene sequence. Phylogenetic analysis revealed significant intraspecific, semi-allopatric phylogenetic breaks in four out of five species belonging to the Podonidae, supporting an ancient radiation and oceanic expansion for this group. By contrast, Penilia avirostris (Sididae) displayed no phylogeographic structure across a global sampling, suggesting a recent worldwide expansion. Our results also show a transoceanic distribution of identical or very similar haplotypes in several species of marine Cladocera, which may be interpreted as either natural transport or evidence of recent anthropogenic transport. If the latter is the case, marine cladocerans represent one of the first genetically documented cases of exotic or invasive marine zooplankton, likely an underreported group. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Sequence data from this article have been deposited with the EMBL/GenBank Data Libraries under accession nos. EU675871–EU675924.     

Latitudinal variation in the Dungeness crab,Cancer magister: zoeal morphology explained by incubation temperature

The difference in morphology between zoeae of Cancer magister Dana from Alaskan and Californian waters was documented to determine if the morphological variation is attributable to environmental influences. First-stage zoeae from Alaska have significantly longer carapace spines than zoeae from central California. The dorsal, rostral and lateral carapace spines were 14, 14 and 29% longer, respectively, in the Alaskan zoeae. The effect of temperature was tested on zoeal morphology as it is an obvious environmental difference between Alaskan and Californian waters. Ovigerous female crabs collected in southeastern Alaska in 1984 were held at 1°, 5°, 10° and 15° C until hatching occurred. Eggs were sampled seven times during the incubation period, and relative mortality, egg diameter and development stage were measured. All of the crabs and eggs at 1° C died before hatching occurred. Egg mortality averaged less than 2% in the other temperature treatments. Egg diameter increased significantly over the incubation period for all temperatures. Developmental rate of the embryos was inversely related to temperature. Hatching first occurred in 42 d at 15° C, 60 at 10° C and 160 d at 5° C. Newly hatched zoeae were collected and body length, dorsal, rostral and lateral carapace spines were measured. Significant differences existed between all temperatures for all spine lengths, with longer spines occurring at lower temperatures. Zoeal body lengths were also significantly different between the three temperatures. The results of this study question the use of spine lengths to distinguish similar larval species.     

Function of the shell spine in the predaceous rocky intertidal snail Acanthina spirata (Prosobranchia: Muricacea)

Feeding observations revealed that Acanthina spirata (Blainville, 1832) uses the shell spine in a ramming and prying motion, resulting in the fracturing and/or forcing open of opercular plates in the barnacles Balanus glandula (Darwin, 1854) and Chthamalus fissus (Darwin, 1854). Attacking barnacles by using the shell spine may represent a second widespread feeding mechanism (in addition to drilling) in muricacean gastropods. Specialization of the shell spine for attacking barnacles is advantageous over drilling within the context of an optimal foraging theory. Rockey intertidal field and laboratory experiments at Palos Verdes and Los Angeles, California, respectively, in 1981–1982 revealed, that snails with a shell spine have lower handling times and feed at a greater rate than spineless snails which drill their prey. Spine penetration of prey as opposed to drilling takes considerably less time than a tidal cycle. This is expected to increase the success rate for completion of feeding, since during tidal flux snails risk being dislodged. Plasticity in attack mechanisms allows A. spirata with broken spines to penetrate prey by drilling while undergoing spine repair. Differences exist in spine-feeding by A. spirata on B. glandula and C. fissus. The attack process takes more than twice as long when snails attack B. glandula versus C. fissus. However, there is no difference in the mean number of spine thrusts required to penetrate the opercular plates of the two species. When feeding on the larger barnacle B. glandula, A. spirata ingests significantly greater dry weight per unit handling time than with C. fissus.     

Presence of Alexandrium catenella and paralytic shellfish toxins in finfish, shellfish and rock crabs in Monterey Bay, California, USA

The central California coast is a highly productive, biodiverse region that is frequently affected by the toxin-producing dinoflagellate Alexandrium catenella. Despite the consistent presence of A. catenella along our coast, very little is known about the movement of its toxins through local marine food webs. In the present study, we investigated 13 species of commercial finfish and rock crabs harvested in Monterey Bay, California for the presence of paralytic shellfish toxins (PSTs) and compared them to the presence of A. catenella and PSTs in sentinel shellfish over a 3-year period. Between 2003 and 2005, A. catenella was noted in 55% of surface water samples (n = 307) and reached a maximum concentration of 17,387 cells L⁻¹ at our nearshore site in Monterey Bay. Peak cell densities occurred in the month of July and were associated with elevated shellfish toxicity in the summers of 2004 and 2005. When A. catenella was present, particulate PSTs were detected 71% of the time and reached a maximum concentration of 962 ng STXeq L⁻¹. Of the 13 species tested, we frequently detected PSTs in Pacific sardines (Sardinops sagax; maximum 250 μg STXeq 100 g⁻¹), northern anchovies (Engraulis mordax; maximum 23.2 μg STXeq 100 g⁻¹), brown rock crabs (Cancer antennarius; maximum 49.3 μg STXeq 100 g⁻¹) and red rock crabs (C. productus; 23.8 μg STXeq 100 g⁻¹). PSTs were also present in one sample of Pacific herring (Clupea pallas; 13.3 μg STXeq 100 g⁻¹) and one sample of English sole (Pleuronectes vetulus; 4.5 μg STXeq 100 g⁻¹), and not detected in seven other species of flatfish tested. The presence of PSTs in several of these organisms reveals that toxins produced by A. catenella are more prevalent in California food webs than previously thought and also indicates potential routes of toxin transfer to higher trophic levels. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Multivariate characterisation of the habitats of seven species of Malayan mudskippers (Gobiidae: Oxudercinae)

Mudskippers (Gobiidae: Oxudercinae) are amphibious gobies. The occurrence and habitats of seven sympatric species of Malayan mudskippers were recorded: Boleophthalmus boddarti, Boleophthalmus pectinirostris, Periophthalmodon schlosseri, Periophthalmus chrysospilos, Periophthalmus gracilis, Periophthalmus variabilis and Scartelaos histophorus. A total of 14 surveys were made at six localities along 120 km of the west coast of the Malay Peninsula in August 1996 and September 2006. A multivariate set of ordinal parameters were used to measure the quantity of environmental water in mudskippers’ habitats, and three guilds of species were discriminated. The resulting pattern mirrored the adaptive scenario depicted by physiological and anatomical studies of mudskippers’ terrestriality, although in one case a terminal taxon was less terrestrial than other more basal ones. Differentiation of trophic niches may also play a role in habitat selection by the herbivorous Boleophthalmus spp., which were only found in unvegetated, illuminated areas and on wet mud at low tide. With the more terrestrial guild, a significant negative co-occurrence suggested direct interspecific competition. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Ultrastructural examination of chamber morphogenesis and biomineralization in the planktonic foraminiferOrbulina universa

Pre-sphere, trochospiralOrbulina universa (d'Orbigny) were hand-collected between June and September, 1982 to 1984, from surface waters of the Pacific Ocean ca. 2 km off Santa Catalina Island, California, USA. The formation of a spherical chamber by this planktonic foraminifer was studied with light and electron microscopy. Chamber morphogenesis is preceded by the secretion of a cytoplasmic layer rich in mitochondria. The layer expands away from the pre-existing foraminiferal shell surface in a spherical pattern. Full cytoplasm expansion is followed by the secretion of an organic matrix (OM) approximately 200 m above the original shell surface. Cytoplasm, mitochondria and/or vesicles may play a role in OM secretion. Calcification and chamber thickening, new spine growth and pore development are documented and continue for a period of 1–7 d. Approximately 24 h prior to gamete release, the foraminiferal spines are resorbed and a veneer of calcite is deposited on the shell surface. An extensive intracellular organelle unique to foraminifera, the fibrillar system, is examined and is hypothesized to play a role in the biomineralization process. The taxonomic and functional significance of theO. universa spherical chamber is discussed with respect to other foraminiferal species.     

Variation in colony geometry modulates internal light levels in branching corals, <Emphasis Type="Italic">Acropora humilis</Emphasis> and <Emphasis Type="Italic">Stylophora pistillata</Emphasis>

Colonial photosynthetic marine organisms often exhibit morphological phenotypic plasticity. Where such plasticity leads to an improved balance between rates of photosynthesis and maintenance costs, it is likely to have adaptive significance. To explore whether such phenotypic plasticity leads to more favourable within-colony irradiance for reef-building branching corals, this relationship was investigated for two coral species Acropora humilis and Stylophora pistillata, along a depth gradient representing light habitats ranging from 500 to 25 μmol photons m⁻² s⁻¹, during 2006 at Heron Island, Great Barrier Reef (23.44°S, 151.91°E). In the present study changes in flow-modulated mass transfer co-varied with light as a function of depth. In low-light (deep) habitats, branch spacing (colony openness) in A. humilis and S. pistillata was 40–50% greater than for conspecifics in high-light environments. Also, branches of A. humilis in deep water were 40–60% shorter than in shallow water. Phenotypic changes in these two variables lead to steeper within-colony light attenuation resulting in 38% higher mean internal irradiance (at the tissue surface) in deep colonies compared to shallow colonies. The pattern of branch spacing was similar for S. pistillata, but this species displayed an alternate strategy with respect to branch length: shade adapted deep and cave colonies developed longer and thinner branches, allowing access to higher mass transfer and irradiance. Corals in cave habitats allowed 20% more irradiance compared to colonies found in the deep, and had a 47% greater proportion of irradiance compared to colonies in the shallow high-light environment. Such phenotypic regulation of internal light levels on branch surfaces partly explains the broad light niches of many branching coral species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Production, hatching success and surface ornamentation of eggs of calanoid copepods during a winter at 57°N

Close to 50 species of marine Calanoid copepods have been reported to produce diapause eggs (Engel and Hirche in J Plankton Res 26:1083–1093, 2004); eggs that are viable but require a refractory phase before they hatch, sometimes after months. Diapause eggs are often described as morphologically different with respect to egg membrane ultrastructure and having a thicker egg shell with surface ornamentation as opposed to the smooth shell found in subitaneous eggs that hatch within days (Belmonte in J Mar Syst 15:35–39, 1998; Chen and Marcus in Mar Biol 127:587–597, 1997; Castro-Longoria in Crustaceana 74:225–236, 2001). Egg production rates, egg surface ornamentation, and hatching success were monitored in large aquaculture fish enclosures during winter with close to zero water temperatures (N57°). Surprisingly, all female copepods (Acartia spp.—presumably A. tonsa, and Centropages hamatus) produced eggs all through the winter with no obvious pattern with respect to light, temperature and food availability, and no diapause eggs were observed. However, individual females produced several categories of eggs with or without surface spines even within the same egg batch as evidenced by scanning electron microscopy (SEM). Four egg categories were distinguishable: ‘no spines’, smooth eggs; ‘short spines’, 5–15 μm long; ‘truncated spines’, with the spine tips cut-off <10 μm long; and ‘long spines’, up to 30 μm long. All egg categories remained unchanged with respect to surface structures from when we took them out of the incubation bottles until they hatched. In general, the frequency of ‘no spines’ was 10–40%, and most eggs were ornamented with ‘short-’ or ‘long spines’. Further, a given egg can be ornamented with all types of surface spines simultaneously, which might even be a fifth egg category. The different egg categories were all able to hatch within days when exposed to normoxic conditions suggesting that they were subitaneous.     

Brooding of the sub-Antarctic heart urchin, <Emphasis Type="Italic">Abatus cavernosus</Emphasis> (Spatangoida: Schizasteridae), in southern Patagonia

The reproductive cycle of the sub-Antarctic spatangoid sea urchin, Abatus cavernosus, was examined during a 2-year period in southern Patagonia, Argentina. The population studied is the northernmost known coastal population in the austral oceans, and is influenced by a greater annual range of seawater temperature than other Abatus species. The sex ratio of the overall A. cavernosus population did not differ significantly from 1:1, but was not homogeneous across size classes. A clearly defined annual reproductive cycle was found. Spawning occurred from May to July and was synchronous between sexes. Females were observed to be brooding within a 9-month period, from May to February. Juveniles were released during the austral summer, from January to February. The length at which 50% of the females were brooding occurred at a test length of 25.9 mm. A. cavernosus had a large egg size (mean diameter = 1.4 mm) and low fecundity (maximum = 57 eggs per female) compared to closely related species. The number of eggs within each brood pouch was highest in larger anterior brood pouch, which is close to two gonopores, as opposed to the other anterior and two smaller posterior pouches. Significant interannual variation was observed in gonad cycles, fecundity, and embryo development such as: brood size decreased during 2001; adjusted gonad dry weight and fecundity were higher during 2003. Hypotheses concerning the gonadal and brooding cycles and fecundity of sub-Antarctic and Antarctic Schizasteridae are discussed. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Spawning and development in <Emphasis Type="Italic">Osedax</Emphasis> boneworms (Siboglinidae,Annelida)

We report observations on spawning and early development in bone-eating worms of the genus Osedax. Individual females of Osedax rubiplumus were observed at 1820 m depth freely spawning hundreds of oocytes, and females of an undescribed species, Osedax “orange collar”, were observed spawning in laboratory aquaria. Cytological and molecular analysis of the spawned oocytes of two Osedax species revealed no evidence for the bacterial endosymbionts that the female worms require for their nutrition, suggesting that the bacteria must be acquired later from the environment, as they are in other siboglinids. Individual O. “orange collar” females released an average of 335 (±130) eggs per day, but the number of oocytes spawned per day varied greatly, suggesting that not all the females spawned daily. Fertilization rates of the spawned oocytes varied from 0 to 100%, though most females showed nearly 100% fertilization rates. Oocytes spawned in the laboratory at 4–6°C were negatively buoyant. If fertilized, these oocytes extruded polar bodies and then after at least four hours cleaved unequally. Subsequent cleavages occurred in a spiral pattern at roughly 2-h intervals, resulting in free-swimming trochophore larvae after 24 h. These lecithotrophic trochophores swam for 9–16 days before settling with several hooked chaetae, similar to those of dwarf Osedax males. The larval life span of the Osedax species studied in the laboratory appears to be shorter than in closely related Vestimentifera. Osedax rubiplumus, on the other hand, has much larger oocytes and so may have greater dispersal potential than these other Osedax species. The high fecundity and apparently continuous reproduction of Osedax boneworms permits the opportunistic exploitation of sunken vertebrate bones. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

<Emphasis Type="Italic">Sympagohydra tuuli</Emphasis> (Cnidaria,Hydrozoa): first report from sea ice of the central Arctic Ocean and insights into histology,reproduction and locomotion

Various cnidarians have adapted their life style to interstitial habitats of marine sediments. Recently, for the first time a hydroid was reported from the interstitial brine channel system of Arctic fast ice. Due to its derived features, the new genus and species Sympagohydra tuuli was introduced. Here we describe findings of S. tuuli in sea ice at several sites within the central Arctic Ocean. In our view the results of this study do not allow assignment of Sympagohydra to any known family and we, therefore, suggest the introduction of a new family Sympagohydridae which is placed within the hydrozoan subclass Hydroidolina, order Anthomedusae, suborder Capitata. A first detailed histological analysis of S. tuuli is presented. In vivo analysis of locomotion and reproduction revealed a remarkable convergent evolution in S. tuuli and distant meiobenthic relatives. Shared traits are a flagellated epidermis enabling the animals to glide within small interstices by means of flagellar beating as well as an internalised embryogenesis. In S. tuuli gametogenesis occurs in the absence of gonophores inbetween gastro- and epidermis clearly separated from the epidermis. Budding was observed as the vegetative mode of reproduction. Documentation of feeding behaviour identified copepod nauplii and rotifers as prey items and demonstrates a high trophical position of the hydroids within the sympagic food web. Occurrence of reproducing individuals and pronounced tolerances towards changing temperatures and salinities indicate S. tuuli as a truly sympagic species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.