Scarred limpets at hydrothermal vents: evidence of predation by deep-sea whelks

The chaotic physical and chemical environment at deep-sea hydrothermal vents has been associated with an ecosystem with few predators, arguably allowing the habitat to provide refuge for vulnerable species. The dominance of endemic limpets with thin, open-coiled shells at north Pacific vents may support this view. To test their vulnerability to predation, the incidence of healed repair scars, which are argued to reflect non-lethal encounters with predators, were examined on the shells of over 5,800 vent limpets of Lepetodrilus fucensis McLean (1988) that were collected from 13 to 18 August 1996. Three vent fields on the Juan de Fuca Ridge at ca. 2,200 m depth were sampled, two within 70 m of 47°56.87′N 129°05.91′W, and one at 47°57.85′N 129°05.15′W with the conspicuous potential limpet predators, the zoarcid fish Pachycara gymninium Anderson and Peden (1988), the galatheid crab Munidopsis alvisca Williams (1988), and the buccinid snail Buccinum thermophilum Harasewych and Kantor (2002). Limpets from the predator-rich vent were most often scarred, a significant difference created by the high incidence of scars on small (<4 mm long) limpets in this sample. Collected with the limpets were small (median shell diameter 4.4 mm) buccinids. They, rather than the larger, more conspicuous mobile fishes and crabs are argued to be the shell-damaging predator. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Sperm polymorphism and genetic divergence in the mussel Perumytilus purpuratus

For the first time in a marine bivalve species, geographic intraspecific variation in sperm morphology in the Chilean mussel, Perumytilus purpuratus, is reported. Samples were obtained in December 2010 from the rocky intertidal at three locations: Antofagasta (23o44′01.08′′S, 70o26′26.11′′W), Valparaíso (32o57′23.07′′S, 71o33′01.86′′W), and Valdivia (39°50′44.26′′S, 73°25′51.99′′W). Scanning electron microscopy was used to describe sperm traits, and genetic data were obtained by sequencing a 16S mtDNA 427-bp fragment. The results showed geographic intraspecific variation with two statistically significant patterns of genetically differentiated sperm morphologies. One genetic clade was found for Antofagasta and Valparaíso, displaying a unique sperm morphotype with a short acrosome accounting for 26 % of the head length, and the other from Valdivia, with a larger acrosome occupying 47 % of the head length. These results will lead to new research to distinguish among alternative hypotheses: the presence of cryptic species, incipient speciation, or a polymorphism in sperm traits of P. purpuratus.     

Mantle-to-shell CaCO<Subscript>3</Subscript> transfer during shell repair at different hydrostatic pressures in the deep-sea vent mussel <Emphasis Type="Italic">Bathymodiolus azoricus</Emphasis> (Bivalvia: Mytilidae)

Calcium carbonate transfer was experimentally examined in hydrothermal mussels Bathymodiolus azoricus, which were collected from 850 m depth at Menez Gwen hydrothermal vent site (31°31′W, 37°50′N) on the Mid Atlantic Ridge in May 2007. In each of four 10-day experiments, groups of mussels were maintained at atmospheric pressure or re-pressurised to depths relevant to their site of occurrence, i.e. 850 m depth at Menez Gwen, 1,750 m at Lucky Strike (31°31′W, 37°18′N) and 2,300 m at Rainbow (31°31′W, 36°13′N). The shells of experimental mussels were perforated and mantle tissue was fixed for light and TEM studies at days 7 and 10 following the injury. Simultaneously, haemocytes from the extrapallial fluid (EPF) at the site of induction were studied. At day 7 the response was most intense in the middle fold of the mantle margin and possibly proportional to hydrostatic pressure. At day 10 the epithelial cells on the mantle surface facing the body cavity produced copious organic secretions that avidly bound calcium. Haemocyte migration was noted within the mantle tissue, and the haemocytes at the mantle surface facing the shell had a Ca-positive granular content. Large haemocytes were detected in the EPF at the injury site, and some showed evidence of an immune reaction while others showed Ca-positive granular content. These results suggest that haemocytes are involved in shell repair in these deep-sea mussels just as in some freshwater and shallow marine molluscs.     

Spatiotemporal variation in the population ecology of scaly slipper lobsters <Emphasis Type="Italic">Scyllarides squammosus</Emphasis> in the Northwestern Hawaiian Islands

Scaly slipper lobster (Scyllarides squammosus) population ecology was examined using tag/recapture information at Necker Island (23°30′N; 164°35′W), Gardner Pinnacles (25°00′N; 168°50′W), Maro Reef (25°30′N; 170°45′W), and Laysan Island (25°48′N; 171°45′W) in the Northwestern Hawaiian Islands (NWHI) USA from 2002 to 2008. Although many aspects of S. squammosus life history and population dynamics were similar to those of other scyllarids, somatic growth differed from its congers. Scyllarides squammosus growth abruptly declined at maturity and, because of this, growth at length was better described using the Schnute as opposed to the more commonly von Bertalanffy growth model. Growth varied among locations, and survival varied among years; thereby being the first documentation of variability in the life history of a scyllarid. This study has expanded knowledge of scyllarid biology and documented that spatiotemporal variability in biological characteristics must be considered to understand and describe the population ecology of this species and probably of other scyllarids.     

Historical demography and contemporary spatial genetic structure of an estuarine crab in the northeast Pacific (<Emphasis Type="Italic">Hemigrapsus oregonensis</Emphasis>)

Discrete estuary subpopulations of the mud crab Hemigrapsus oregonensis (Dana, 1851) are connected via larval dispersal. Sequence variation at the mtDNA COI locus was examined in eight populations sampled in 2001–2002 from central California through northern Oregon in the northeast Pacific (36.6–45.8°N) to infer patterns of dispersal and historical connectivity in the region. Strong evidence for persistence since the mid-Pleistocene, with no range truncation resulting from southward shifting temperature isoclines, was provided by a phylogeographic pattern of haplotypes of an older clade distributed throughout the sampled range. A recently derived clade became widespread only north of Cape Blanco after the last glacial maximum. Its clear pattern of restriction to the northern area, in the absence of similarly restricted southern clades, suggests that contemporary dispersal around Cape Blanco is rare (population F _ST = 0.192). Low pairwise differentiation within Oregon and within central California, as well as contrasts between northern and southern groups in the shape of the pairwise mismatch distribution, nucleotide diversity, and Tajima’s D suggest that these regions reflect different demographic histories. Potential mechanisms explaining this latitudinal break include contemporary coastal circulation patterns, selection, and ancient patterns of larval dispersal in the California Current.     

Species boundaries in the starfish genus Linckia

 The genetic basis for species boundaries in the starfish genus Linckia was examined using variation observed in 613 base pairs (bp) of sequence from the cytochrome oxidase I gene of mtDNA and 16 allozyme loci. Five groups within Linckia were clearly genetically differentiated; L. columbiae, L. bouvieri, two clades within L. guildingi, and one clade with two sub-clades consisting of both L. laevigata and L. multifora. Genetic divergence among these groups is consistent with interspecific variation. The two clades within L. guildingi suggest the presence of a cryptic, partially sympatric, species. Genetic divergence between these two clades implies that they have been genetically distinct from each other for at least one million years. In contrast, genetic data suggest that L. laevigata and L. multifora are a single species, despite the fact that live individuals can be distinguished by their colour and colour pattern, number of madreporites and ratio of arm length to breadth. There are probably three closely related groups within the L. laevigata/L. multifora clade –L. multifora, and two groups in L. laevigata defined by biogeographic province. It is difficult to determine boundaries for these three entities, since genetic and morphological differences are complicated by phenotypic differences arising from both environmental variation and population genetic structure. The difficulties encountered in defining species boundaries in Linckia, particularly with respect to variation arising from the overlap of Indian and Pacific biogeographic provinces, may be a general issue for many marine organisms from this region. Received: 24 May 1999 / Accepted: 6 October 1999     

Reproductive traits of pioneer gastropod species colonizing deep-sea hydrothermal vents after an eruption

The colonization dynamics and life histories of pioneer species determine early succession at nascent hydrothermal vents, and their reproductive ecology may provide insight into their dispersal and population connectivity. Studies on the reproductive traits of two pioneer gastropod species, Ctenopelta porifera and Lepetodrilus tevnianus, began within a year after an eruption on the East Pacific Rise (EPR) that eliminated vent communities near 9°50′N from late 2005/early 2006. Standard histology was used to examine gamete release, instantaneous female fecundity, and time to maturation. Both species exhibited two-component oocyte size–frequency distributions indicating quasi-continuous reproduction with high fecundity. In samples collected in December 2006, both C. porifera and L. tevnianus individuals were reproductively mature. The smallest reproducing C. porifera were 4.2 mm (males) and 5.4 mm (females) in shell length, whereas reproductive L. tevnianus were smaller (2.3 and 2.4 mm in males and females, respectively). Most C. porifera were large (>6.0 mm) compared to their size at metamorphosis and reproductively mature. In contrast, most L. tevnianus were small (<1.0 mm) and immature. Reproductive traits of the two species are consistent with opportunistic colonization, but are also similar to those of other Lepetodrilus species and peltospirids at vents and do not fully explain why these particular species were the dominant pioneers. Their larvae were probably in high supply immediately after the eruption, due to oceanographic transport processes from remote source populations.     

Genetic structure of natural populations of California red abalone (<Emphasis Type="Italic">Haliotis rufescens</Emphasis>) using multiple genetic markers

Mitochondrial cytochrome oxidase subunit one (COI) sequence, nuclear microsatellites, and amplified fragment length polymorphisms (AFLPs) were used to evaluate connectivity among nine red abalone (Haliotis rufescens) populations sampled between August 1998 and November 2003 along approximately 1,300 km of California coastline from Crescent City (41°46′N, 124°12′W) to San Miguel Island (34°02′N, 120°22′W). COI sequences and microsatellite genotypes did not show significant genetic divergence among nine sampled populations. A subset of five populations spanning the geographic range of the study was scored for 163 polymorphic AFLP markers. Of these, 41 loci showed significant divergence (P < 0.001) among populations. Still, no AFLP markers were diagnostic for any of the study populations, and assignment tests did not consistently assign individuals to the correct population. Although the AFLP data are the first to suggest there is significant genetic differentiation among California red abalone populations, the discordance between the different genetic markers needs further study before unambiguous conclusions can be drawn with respect to connectivity among the populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

A multi-locus genetic assignment technique to assess sources of <Emphasis Type="Italic">Agaricia agaricites</Emphasis> larvae on coral reefs

Assignment of individuals to populations based upon genetic data is an important ecological problem that requires many polymorphic markers, often more than are available using single locus techniques. To demonstrate the utility of amplified fragment length polymorphisms (AFLP) in studying larval dispersal and recruitment in coral populations, two sets of AFLP primers were used to genotype colonies of the coral Agaricia agaricites Linnaeus from three widely separated geographic locations: the Bahamas (23°28′N, 75°42′W) and Key Largo, Florida (24°55′N, 80°31′W—two sites separated by 12 km) in 1995, and the Flower Garden Banks (FGB) in the Gulf of Mexico (27°55′N,93°36′W) in 1997. In addition to adult samples from each site, recruits were collected from settling plates placed on the East FGB for 1 year (1997–1998). The AFLP technique yielded 45 polymorphic markers. An analysis of molecular variance (AMOVA) showed significant genetic differences among the four adult populations, even between the two Key Largo sites. The recruits were significantly different from all adult populations except those from the FGB. Discriminant function analysis and the program AFLPOP were used to assign individuals to populations. Using the adult AFLP-banding patterns to build the statistical models, both procedures correctly assigned the majority of adults to their respective populations in simulations and assigned all but one of the recruits to the Flower Garden population from where they were collected . The AFLP technique provides a simple and adaptable population assignment method for studying recruitment processes in A. agaricites and other coral species. Electronic Supplementary Material Supplementary material is available for this article at     

Emergence of an oocytic circumnuclear ring in response to increasing day length in Atlantic herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>)

Understanding physiological and environmental variables that initiate sexual maturity would provide fundamental information on life history dynamics. The aim of this study was to test the usefulness of the common circumnuclear ring (CNR), an oocytic structure similar to the Balbiani body, which appears just prior to oocyte development as a predictor of first maturation in Atlantic herring (Clupea harengus). The relative roles of physiology (e.g. fat) and photoperiod as triggers of maturation were also investigated. Samples were collected in May 2008 (72°26′–73°84′N/11°26′–18°40′E) and February 2009 (56°12′–59°45′N/00°25′–03°06′W). These data suggested that thresholds in body size may influence the decision to mature. We also found that short days (winter solstice) may be the photoperiod trigger for a first-decision window for both Norwegian spring-spawning (NSS) and North Sea autumn-spawning (NSAS) herring. The second-decision window for NSAS herring maturation appears to be triggered by longer days (spring equinox), while a decreasing rate of day lengthening may trigger NSS herring maturation. So, photoperiodic cycle is a key determinate of the timing of maturation in Atlantic herring.     

Social monogamy in the shrimp Pontonia margarita, a symbiont of Pinctada mazatlanica, off the Pacific coast of Panama

A previous study predicted the evolution of monogamy in symbiotic crustaceans inhabiting scarce, relatively small hosts in tropical environments where predation risk away from hosts is high. This prediction was tested in the shrimp Pontonia margarita, which inhabits the pearl oyster Pinctada mazatlanica. A total of 68 oysters were collected from the intertidal and shallow subtidal at two islands (Islas Secas [N 27° 55′, W 82° 03′] and Isla de La Coiba [N 27° 50′, W 97° 03′]) off the eastern tropical Pacific coast on 15 and 17 March 2007, respectively. The population structure, distribution, male-female association pattern, and relative growth of the major claw and pleura of the second abdominal segment of each shrimp retrieved were examined. Shrimps were found as heterosexual pairs in the mantle cavity of hosts more frequently than would be expected by chance alone. Males occurred with females in the same host, independent of the reproductive condition of the female or the stage of development of brooded embryos. This observation, and strong correlations between the host and shrimp body size in both males and females suggest a long-term association between males and females in each host. Sexual dimorphism in body size was minor, with males being just slightly smaller than females. In agreement with predictions for monogamous species, the major claw of males did not display positive allometry, which has been generally reported for polygamous shrimps. In turn, the pleura of the second abdomen presented negative allometry in males but positive allometry in females. All available information suggests that Pontonia margarita has a socially monogamous mating system with males and females forming exclusive pairs in their hosts.     

Domoic acid in benthic flatfish on the continental shelf of Monterey Bay,California, USA

Within Monterey Bay, California, USA, the food web transfer of domoic acid (DA), a neurotoxin produced by diatoms of the genus Pseudo-nitzschia, has led to major mortality events of marine mammals and birds. Less visible, and less well known, is whether invertebrates and fish associated with the benthos are also affected by blooms of DA-producing Pseudo-nitzschia spp. This study examines the presence of DA in benthic flatfish offshore of Davenport, California, (37°0′36″N, 122°13′12″W) and within Monterey Bay, California (36°45′0″N, 122°1′48″W), including species that feed primarily in the sediment (benthic-feeding) and species that feed primarily in the water column (benthopelagic-feeding). Flatfish caught between 10 December 2002 and 17 November 2003 at depths of 30–180 m had concentrations of DA in the viscera ranging from 3 to 26 μg DA g⁻¹ of viscera. Although the DA values reported are relatively low, benthic-feeding flatfish were frequently contaminated with DA, especially as compared with the frequency of contamination of flatfish species that feed in the water column. Furthermore, on days in which both benthic-feeding and benthopelagic-feeding flatfish were collected, the former had significantly higher concentrations of DA in the viscera. Curlfin turbot, Pleuronicthys decurrens, the flatfish with both the highest level and frequency of DA contamination, are reported to feed exclusively on polychaetes, suggesting that these invertebrates may be an important vector of the toxin in benthic communities and may pose a risk to other benthic-feeding organisms. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Recovery after trampling disturbance in a canopy-forming seaweed population

The recovery of Ascophyllum nodosum, and the associated benthic community, was followed after repeated trampling disturbance at different intensities, from 2005 to 2010, at its southern limit (41°41′27″N; 8°50′57″W). The cover of A. nodosum and associated organisms and the demography of A. nodosum individuals in the most disturbed areas were followed over a recovery period of 54 months. Recovery of A. nodosum assemblages was negatively affected by increasing trampling intensity. After 54 months, the highest intensity plots were still significantly different from control plots. Growth and mortality rates of A. nodosum increased shortly after disturbance but progressively recovered to levels close to control plots. The results of this study demonstrate the sensitivity of this habitat-building species to human trampling and also show its plasticity to cope with disturbance events.     

Geographic and ontogenetic variation in the diet and daily ration of the bonnethead shark, <Emphasis Type="Italic">Sphyrna tiburo</Emphasis>, from the eastern Gulf of Mexico

To examine variation in diet and daily ration of the bonnethead shark, Sphyrna tiburo (Linnaeus 1758), animals were collected from three areas in the eastern Gulf of Mexico: northwest Florida (∼29°40′N, 85°13′W), Tampa Bay near Anclote Key (∼28°10′N, 82°42.5′W), and Florida Bay (∼24°50′N, 80°48′W) from March through September, 1998–2000. In each area, diet was assessed by life stage (young-of-the year, juveniles, and adults) and quantified using five indices: percent by number (%N), percent by weight (%W), frequency of occurrence (%O), index of relative importance expressed on a percent basis (%IRI), and %IRI based on diet category (%IRI_DC). Diet could not be assessed for young-of-the-year in Tampa Bay or Florida Bay owing to low sample size. Diet analysis showed an ontogenetic shift in northwest Florida. Young-of-the-year stomachs from northwest Florida (n = 68, 1 empty) contained a mix of seagrass and crustaceans while juvenile stomachs (n = 82, 0 empty) contained a mix of crabs and seagrass and adult stomachs (n = 39, 1 empty) contained almost exclusively crabs. Crabs made up the majority of both juvenile and adult diet in Tampa Bay (n = 79, 2 empty, and n = 88, 1 empty, respectively). Juvenile stomachs from Florida Bay (n = 72, 0 empty) contained seagrass and a mix of crustaceans while adult stomachs contained more shrimp and cephalopods (n = 82, 3 empty). Diets in northwest Florida and Tampa Bay were similar. The diet in Florida Bay was different from those in the other two areas, consisting of fewer crabs and more cephalopods and lobsters. Plant material was found in large quantities in all stomachs examined from all locations (>15 %IRI_DC in 6 of the 7 life stage-area combinations, >30 %IRI_DC in 4 of the 7 combinations, and 62 %IRI_DC in young-of-the-year diet in northwest Florida). Using species- and area-specific inputs, a bioenergetic model was constructed to estimate daily ration. Models were constructed under two scenarios: assuming plant material was and was not part of the diet. Overall, daily ration was significantly different by sex, life stage, and region. The bioenergetic model predicted increasing daily ration with decreasing latitude and decreasing daily ration with ontogeny regardless of the inclusion or exclusion of plant material. These results provide evidence that bonnetheads continuously exposed to warmer temperatures have elevated metabolism and require additional energy consumption to maintain growth and reproduction. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Differential ingestion of zooplankton by four species of bivalves (Mollusca) in the Mali Ston Bay,Croatia

This study provides information about differences in composition of ingested zooplankton amongst bivalve species coexisting in the same area in a period from May 2009 to December 2010. The study was conducted at the Mali Ston Bay (42°51′ N, 17°40′ E)—the most important bivalve aquaculture area in the eastern Adriatic Sea. Stomach content analysis was performed on cultured species—Ostrea edulis and Mytilus galloprovincialis, and commercially important bivalve species from their natural environment—Modiolus barbatus and Arca noae. Results confirmed carnivory in bivalves, both from natural and cultured populations, but cultured species had higher numbers of zooplankters than those living on the seabed. The most abundant taxa were bivalve larvae, followed by tintinnids, copepods, unidentified eggs and gastropod larvae. Recorded numbers of bivalve larvae in M. galloprovincialis stomach were the highest so far reported and show that mussels impact the availability of natural spat.     

Carbon dioxide use by chemoautotrophic endosymbionts of hydrothermal vent vestimentiferans: affinities for carbon dioxide, absence of carboxysomes, and δ13C values

The hydrothermal vent vestimentiferans Riftia pachyptila Jones, 1981 and Ridgeia piscesae Jones, 1985 live in habitats with different abundances of external CO₂. R. pachyptila is found in areas with a high input of hydrothermal fluid, and therefore with a high [CO₂]. R. piscesae is found in a range of habitats with low to high levels of hydrothermal fluid input, with a correspondingly broad range of CO₂ concentrations. We examined the strategies for dissolved inorganic carbon (DIC) use by the symbionts from these two species. R. pachyptila were collected from the East Pacific Rise (9°50′N; 104°20′W) in March 1996, and R. piscesae were collected from the Juan de Fuca Ridge (47°57′N; 129°07′W) during September of 1996 and 1997. The differences in the hosts' habitats were reflected by the internal pools of DIC in these organisms. The concentrations of DIC in coelomic fluid from R. piscesae were 3.1 to 10.5 mM, lower than those previously reported for R. pachyptila, which often exceed 30 mM. When symbionts from both hosts were incubated at in situ pressures, their carbon fixation rates increased with the extracellular concentration of CO₂, and not HCO₃ ⁻, and symbionts from R. piscesae had a higher affinity for CO₂ than those from R. pachyptila (K _1/2 of 7.6 μM versus 49 μM). Transmission electron micrographs showed that symbionts from R. piscesae lack carboxysomes, irrespective of the coelomic fluid [DIC] of their host. This suggests that the higher affinity for CO₂ of R. piscesae symbionts may be their sole means of compensating for lower DIC concentrations. The δ¹³C values of tissues from R. piscesae with higher [DIC] in the coelomic fluid were more positive, opposite to the trend previously described for other autotrophs. Factors which may contribute to this trend are discussed. Received: 24 September 1998 / Accepted: 12 May 1999     

Spatial variation in dimethylsulfoniopropionate (DMSP) production in <Emphasis Type="Italic">Ulva lactuca</Emphasis> (Chlorophyta) from the Northeast Pacific

Although dimethylsulfoniopropionate (DMSP) has a variety of functions in marine macroalgae including that of a cryoprotectant, an osmolyte, a way to remove excess sulfur and energy, an antioxidant, and an allelopathic precursor, the latter two functions are believed to be the most important in Ulva lactuca L. (=U. fenestrata) in intertidal populations on the coast of Washington state, USA. The present study found significant variation in DMSP concentrations among U. lactuca collected in May 2005 from six sites ranging from 47°54.45′N (Possession Point, Whidbey Island, WA, USA) to 48°30.55′N (Shannon Point Beach, Anacortes, WA, USA), and also among individuals within sites, and among tissues (basal tissues near the holdfast, middle of the blades, and tips). Concentrations ranged from 37 to 224 μmol g⁻¹ fresh mass (FM). In several 10-day experiments between July 2001 and August 2004 with U. lactuca collected from several places on the coast of Washington, the effects of nutrient level (DIN), light intensity and wavelength, and grazing by the herbivorous gastropod Lacuna vincta, were examined. None of these manipulations resulted in DMSP concentrations that differed significantly from controls, and variance in DMSP concentrations within each experiment was very low. Although DMSP concentrations in U. lactuca may be affected by factors not tested in these experiments, it is also possible that the observed spatial differences reflect constitutive genotypic or phenotypic differences among geographically separated U. lactuca populations or among cryptic Ulva species. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.