The life history responses of the abalone pest,<Emphasis Type="Italic"> Terebrasabella heterouncinata</Emphasis>, under natural and aquaculture conditions

The sabellid, Terebrasabella heterouncinata, is a small (<5 mm) intratubular brooder that lives in burrows within the hosts shell matrix. It is a semi-continuous breeder and despite producing small numbers of large eggs, infestation by this animal has reached epidemic proportions on local abalone farms. The present study compared the morphometrics and reproductive characteristics of worms from farmed and wild abalone, in the Walker Bay area of the south Western Cape Province of South Africa, to gain insights into why this animal has become so successful under aquaculture conditions. The farms designated farm A and farm B each had one on-farm site, and two wild sites, while farm C had two on-farm sites and two wild sites. The wild sites were natural abalone habitats located within 2.5 km of the farms. Our results conclusively showed that environmental conditions prevalent on the farms enhanced the reproductive success of these worms relative to that observed in its natural environment. At farms B and C, worms occurred in significantly higher densities at the on-farm sites than in the corresponding wild samples, but at farm A, density was equally low at the three sites. At all three farms, a greater proportion of the population was reproductively active in the on-farm samples than in the wild samples. Worms on farmed abalone had a higher instantaneous fecundity, brooded more clutches simultaneously and were larger than their conspecifics from the wild. There was a positive correlation between adult size and brood size and the number of clutches brooded simultaneously. Within the three on-farm sites there was a negative correlation between egg volume and brood size, indicating a trade-off between these traits. However, such a trade-off was not apparent between sites, with brood size being higher at the on-farm sites than at the wild sites, irrespective of egg size. This suggests that the stable nutrient-enriched environment on the farm led to an increase in fecundity without compromising the size (and implicitly the quality) of the eggs. Worm density did not have a significant effect on body size or any other reproductive traits at most sites, and the density of T. heterouncinata was unaffected by the density of other shell-infesting polychaetes. The results suggest that the farm environment has selected for larger, more fecund worms that breed rapidly with high recruitment success as a consequence of abundant nutrients, high host density, habitat stability and a possible lack of predation and interspecific competition.Communicated by J.P. Thorpe, Port Erin     

Vertical distribution,population structure and lifecycle of<Emphasis Type="Italic"> Eucalanus bungii</Emphasis> (Copepoda: Calanoida) in the Oyashio region,with notes on its regional variations

Vertical distribution and population structure of Eucalanus bungii were investigated at site H in the Oyashio region (western subarctic Pacific) from September 1996 through October 1997 to evaluate the species lifecycle pattern and associated ontogenetic vertical migration. Additional temporary samplings were also made at several stations covering the entire subarctic Pacific, Okhotsk Sea and Japan Sea, as a basis for regional comparison of lifecycle features of this species. At site H, a marked phytoplankton bloom occurred from mid-March to June, and E. bungii spawned in April/May in the surface layer. Resulting nauplii and copepodite stage 1 (C1) formed a prominent abundance peak in early June. The C1 developed and reached C5 by August. The development of nauplii through C4 occurred in the surface layer. From August onwards, C5 and a small fraction of C3–C4 sank gradually deeper, and entered diapause to overwinter at >500 m depth. The C5 molted to C6 males and females in February and March, respectively. The C6 males and females mated at 250–500 m depth, and only mated C6 females ascended to the surface layer in April for spawning. Judging from the size of lipid droplets in the body, the C3–C5 specimens deposited lipids in the body through the phytoplankton bloom period, and the lipids were consumed gradually during overwintering. Taking account of sampling season, temporal changes in population structure, and vertical distribution, the data collected from the western subarctic Pacific and Okhotsk Sea are consistent with a 1-year lifecycle for the site H population, while the data from the central and eastern subarctic Pacific were consistent with a 2-year lifecycle. The populations from the southern and southeastern Japan Sea did not fit the features of either lifecycle scenario, and because of their very small population size it is suggested that they originated from the northern Japan Sea. Regional comparison of the prosome length of C6 females, including those in the Bering Sea, indicated significantly larger specimens from the Japan Sea and Okhotsk Sea, and smaller specimens in the eastern subarctic Pacific, as compared with those in the western subarctic Pacific (including site H) and Bering Sea. A possible overwintering mechanism of E. bungii is discussed.Communicated by O. Kinne, Oldendorf/Luhe     

Release of phagocytosis-stimulating factor(s) by morula cells in a colonial ascidian

In vitro yeast phagocytosis by haemocytes of the compound ascidian Botryllus schlosseri was studied, with particular attention to interactions among different immunocyte types. It is demonstrated that the supernatant from haemocyte cultures matched with yeast cells contains factor(s) able to enhance yeast ingestion by Botryllus phagocytes. The increase in phagocytosis is not the consequence of yeast opsonisation, as the phagocytic index does not significantly increase when yeast cells, previously incubated in the culture media, are washed and re-suspended in filtered sea water. When haemocytes were fractionated by density gradient centrifugation and each band was incubated with yeast, the ability to stimulate phagocytosis was found in the supernatants from haemocyte cultures of fractions rich in morula cells (MC). Previous studies have demonstrated that MC express molecules recognised by anti-cytokine antibodies, as a consequence of the recognition of foreign molecules or cells. Our results indicate that molecules immunoreactive with anti-cytokine antibodies are required for modulating phagocyte activity, as the above-reported enhancing effect is completely absent in the presence of anti-IL-1-α and anti-TNF-α, but not of anti-rabbit-IgG antibodies, and they also highlight the presence of ‘cross-talk‘ between MC and phagocytes. A new scenario is therefore sketched, in which MC actively recognise non-self molecular patterns and, upon this recognition, release humoural factor(s) recognised by phagocytes, which modulate phagocytosis.     

Preliminary Measurements of Turbulence and Environmental Parameters in a Sub-Tropical Estuary of Eastern Australia

In natural systems, mixing is driven by turbulence, but current knowledge is limited in estuarine zones where predictions of contaminant dispersion are often inaccurate. A series of detailed field studies was conducted in a small sub-tropical creek in eastern Australia. Hydrodynamic, physio-chemical and ecological measurements were conducted simultaneously to assess the complexity of the estuarine zone, notably the interactions between turbulence and environment. The measurements were typically performed at high frequency over a tidal cycle. The results provide an original data set to complement long-term monitoring and a basis for a more detailed study of mixing in sub-tropical systems. Unlike many long-term observations, velocity and water quality scalars were measured herein with sufficient spatial and temporal resolutions to determine quantities of interest in the study of turbulence, while ecological indicators were sampled systematically and simultaneously. In particular the results yielded contrasted outcomes, and the finding impacts on the selection process for key water quality indicators.     

Innate immune defense of the sponge<Emphasis Type="Italic"> Suberites domuncula</Emphasis> against gram-positive bacteria: induction of lysozyme and AdaPTin

Sponges are filter feeders that are exposed to large amounts of bacteria present in their surrounding aqueous milieu. The characteristic cell wall component of gram-positive bacteria, peptidoglycan (PPG), was used as a model molecule to study the responsiveness of cells from the marine demosponge Suberites domuncula toward gram-positive bacteria. The sponge lysozyme, which hydrolyzes PPG, was isolated from the living sponge; in addition its gene was cloned (SDLYS) and expressed in Escherichia coli. Antibodies were raised against the recombinant protein to demonstrate that in the Western blot both molecules give the same signal. In situ hybridization with SDLYS as a probe showed that cells in the mesohyl, the gray cells, strongly react with SDLYS. Subsequent immunofluorescence studies with antibodies raised against lysozyme revealed that only bacteria react with anti-lysozyme and only those that are scattered within the mesohyl of the tissue. An adaptor gene (AdaPTin-1) was isolated from the same sponge species that encodes a putative protein involved in endosome formation. Based on its differential expression we conclude that sponge cells react to PPG with a rapid activation of endocytosis, followed by the release of lysozyme.Communicated by O. Kinne, Oldendorf/LuheThe cDNA sequences from Suberites domuncula have been deposited (EMBL/GenBank): cDNA for the lysozyme (SDLYS) under the accession number AJ699166 and the AdaPTin-1 cDNA (SDAP1) under AJ699167.     

Stability of eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis> L.) depth limits: influence of habitat type

Seagrass meadows are generally considered stable although few studies have specified and tested this statement. On the basis of a large monitoring dataset from Danish coastal waters, we aimed to test whether the stability of deep eelgrass populations changes along a eutrophication gradient extending from inner bays over outer bays to open coastal waters. We defined stability in terms of the stability properties of constancy, resilience, and/or persistence. Our data allowed us to investigate the stability property constancy expressed as temporal variability in eelgrass depth limits over a time scale of 10 years. We hypothesised that colonisation depths were large and relatively constant along open coasts because of low and relatively constant levels of nutrients and turbidity. Conversely, colonisation depths were hypothesised to be low and variable in protected bays due to higher and more variable levels of nutrients and turbidity. We found that depth limits increased from inner bays towards open coastal waters, matching declines in nutrient concentration and increases in water clarity and oxygen concentration. Stability expressed as constancy of depth limits did not differ significantly between habitat types, and neither did stability of physicochemical variables. However, when data from all habitat types were analysed together, they showed that eelgrass populations at the depth limit were significantly more constant and thus, in this respect, more stable when occurring in deep waters as compared to shallow waters. Areas of good water quality may thus obtain the double benefit of deeper-growing and more stable eelgrass populations. The most likely reason why this pattern did not appear at habitat-type level is that the habitat types studied represented wide spatial variation in water quality and depth limit. We conclude that the question of whether eelgrass populations are stable depends on the stability property and the ecological situation in question. Populations may be considered unstable in terms of inter-annual variation but stable in terms of long-term persistence. Therefore, the common statement that eelgrass populations are stable is not universally true.Communicated by M. Kühl, Helsingør     

Is benthic trophic structure affected by food quality? The Dogger Bank example

The aim of this study was to evaluate the relationship between food quality and trophic structure of macrobenthic communities of the Dogger Bank. Therefore, 21 stations along different depth gradients (a transect across the central Dogger Bank and at a station cross in the north-eastern part) were sampled and analysed for macrofauna trophic structure and food supply, indicated by phytopigment composition and organic matter contents of the sediment. The study revealed differences in the trophic structure of the benthic community in areas with contrasting food supply. The trophic structure reflected differences in relative quality of organic matter. The sandy sediments on top of the bank receiving intermediate quantities of high food quality were inhabited by high abundances of interface feeders and sand-licking amphipods of the genus Bathyporeia. The southern stations characterised by a comparatively high amount of intermediate food quality were dominated by interface feeders such as Amphiura filiformis. The northern and north-eastern stations showing the highest diversity of feeding types were characterised by a comparatively low, but highly variable input of intermediate-quality organic matter consisting of both fresh and degraded organic matter. Results are discussed with a special emphasis on specific Dogger Bank features such as benthic primary production and frontal system characteristics.Communicated by O. Kinne, Oldendorf/Luhe     

Crinoid phylogeny: a preliminary analysis (Echinodermata: Crinoidea)

We describe the first molecular and morphological analysis of extant crinoid high-level inter-relationships. Nuclear and mitochondrial gene sequences and a cladistically coded matrix of 30 morphological characters are presented, and analysed by phylogenetic methods. The molecular data were compiled from concatenated nuclear-encoded 18S rDNA, internal transcribed spacer 1, 5.8S rDNA, and internal transcribed spacer 2, together with part of mitochondrial 16S rDNA, and comprised 3,593 sites, of which 313 were parsimony-informative. The molecular and morphological analyses include data from the bourgueticrinid Bathycrinus; the antedonid comatulids Dorometra and Florometra; the cyrtocrinids Cyathidium, Gymnocrinus, and Holopus; the isocrinids Endoxocrinus, and two species of Metacrinus; as well as from Guillecrinus and Caledonicrinus, whose ordinal relationships are uncertain, together with morphological data from Proisocrinus. Because the molecular data include indel-rich regions, special attention was given to alignment procedure, and it was found that relatively low, gene-specific, gap penalties gave alignments from which congruent phylogenetic information was obtained from both well-aligned, indel-poor and potentially misaligned, indel-rich regions. The different sequence data partitions also gave essentially congruent results. The overall direction of evolution in the gene trees remains uncertain: an asteroid outgroup places the root on the branch adjacent to the slowly evolving isocrinids (consistent with palaeontological order of first appearances), but maximum likelihood analysis with a molecular clock places it elsewhere. Despite lineage-specific rate differences, the clock model was not excluded by a likelihood ratio test. Morphological analyses were unrooted. All analyses identified three clades, two of them generally well-supported. One well-supported clade (BCG) unites Bathycrinus and Guillecrinus with the representative (chimaeric) comatulid in a derived position, suggesting that comatulids originated from a sessile, stalked ancestor. In this connection it is noted that because the comatulid centrodorsal ossicle originates ontogenetically from the column, it is not strictly correct to describe comatulids as unstalked crinoids. A second, uniformly well-supported clade contains members of the Isocrinida, while the third clade contains Gymnocrinus, a well-established member of the Cyrtocrinida, together with the problematic taxon Caledonicrinus, currently classified as a bourgueticrinid. Another cyrtocrinid, Holopus, joins this clade with only weak molecular, but strong morphological support. In one morphological analysis Proisocrinus is weakly attached to the isocrinid clade. Only an unusual, divergent 18S rDNA sequence was obtained from the morphologically strange cyrtocrinid Cyathidium. Although not analysed in detail, features of this sequence suggested that it may be a PCR artefact, so that the apparently basal position of this taxon requires confirmation. If not an artefact, Cyathidium either diverged from the crinoid stem much earlier than has been recognised hitherto (i.e., it may be a Palaeozoic relic), or it has an atypically high rate of molecular evolution.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by J.P. Thorpe, Port Erin     

The effect of feeding behavior of the gastropods<Emphasis Type="Italic"> Batillaria zonalis</Emphasis> and<Emphasis Type="Italic"> Cerithideopsilla cingulata</Emphasis> on their ambient environment

Feeding behaviors of the gastropods Batillaria zonalis, a suspension and deposit feeder, and Cerithideopsilla cingulata, an obligate deposit feeder, were studied to examine their effect on dynamics of suspended materials, total nitrogen (TN) and total organic carbon (TOC) in sediments. Suspension feeding in B. zonalis was observed in detail visually, as it had been previously unreported. An experimental system where B. zonalis and C. cingulata were cultured for 10 weeks, using previously frozen microalgae Nannochloropsis oculata as food, was then constructed. During feeding observations, the suspension-feeding B. zonalis formed a mucus food cord to entangle particulate materials, which were subsequently ingested. The feeding mode of B. zonalis is hence categorized as ctenidial filter feeding. For the culture experiments, decreases in suspended materials were seen only in the B. zonalis cultures, while the control (no gastropods) and C. cingulata cultures remained nearly unchanged. Sediment TN and TOC showed no significant differences between B. zonalis (with mean TN at 0.0345% and mean TOC at 0.261%) and control cultures (with TN at 0.0389% and TOC at 0.331%), but the sediments in C. cingulata cultures had lower levels (with TN at 0.0204% and TOC at 0.156%). The C/N ratios were similar for both B. zonalis (7.55) and C. cingulata (7.68) cultures, and both were lower than the control cultures (8.55). The filtration rate for B. zonalis was lower than that previously observed in bivalves inhabiting the same intertidal flat (e.g. Cyclina sinensis, Grafrarium tumidum and Barbatia virescens). However, Batillaria zonalis occurs at higher abundances than these bivalves. Therefore, it is expected that this species has a large affect upon the transport of suspended materials to the sediments. The addition of TN and TOC to sediments in B. zonalis cultures was probably caused by biodeposition, but deposit feeding by B. zonalis may have restrained the accumulation of those components. The impact of deposit feeding in Cerithideopsilla cingulata cultures was most probably stronger than sedimentation and biodeposition, because of the lower sediment TN and TOC. Bioturbation by both B. zonalis and C. cingulata yields the same effect on sediment quality, as indicated by the low C/N in the culture sediment of both treatments, despite difference in feeding modes. This paper demonstrates, for the first time, the importance of gastropods in bioturbation and removal of suspended materials in subtropical tidal flat habitats.Communicated by T. Ikeda, Hakodate