Vertical distribution of bivalve larvae along a cross-shelf transect during summer upwelling and downwelling

Previous time-series studies of meroplankton abundances in the LEO-15 research area off Tuckerton, New Jersey, USA (39°28′N, 74°15′W) indicated short-lived (6–12 h) pulses in larval surfclam (Spisula solidissima Dillwyn) concentration often associated with the initiation of downwelling. To examine possible larval surfclam (and other bivalve) concentrating mechanisms during upwelling and downwelling, six sets of adaptive mobile zooplankton pump samples were taken in July 1998 at different depths at five to six stations along a 25-km transect perpendicular to the coastline and crossing Beach Haven Ridge at LEO-15. Sampling was guided by near real-time, satellite imagery of sea surface temperature overlain by sea surface currents from a shore-based ocean surface current radar (OSCR) unit. A Seabird CTD on the mobile pump frame near the intake provided information on thermocline depth, and sampling depths were adjusted according to the temperature profiles. Near shore, the thermocline was tilted down during downwelling, and up during upwelling. The highest concentrations of surfclam larvae occurred near the bottom at a station near Beach Haven Ridge during downwelling, and just above the thermocline 3 km further offshore during well-developed upwelling. For other bivalve taxa, the larvae were concentrated near the thermocline (Anomia simplex Orbigny and Pholadidae spp.) or concentrated upslope near the bottom (Mytilidae spp.) during upwelling, and the larvae were concentrated near the bottom or were moved downslope during downwelling. Donax fossor Say larvae were found near the surface or above the thermocline during upwelling and downwelling. The general patterns of larval bivalve distribution appear to be influenced by water mass movement during upwelling and downwelling. The larval concentration patterns of individual species are likely a consequence of advection due to upwelling and downwelling circulation, vertical shear in the front region, species-specific larval behaviors, and larval sources.     

Settlement-competency periods of larvae of three species of scleractinian corals

The pattern of settlement over time of three broadcast spawning coral species (Cyphastrea serailia, Acanthastrea lordhowensis, and Goniastrea australensis) from the Solitary Islands (30°00′S; 153°20′E) was studied in 1995 and 1996 in order to determine the maximum length of time these larvae could remain in the water column and still retain the ability to settle and metamorphose. Larvae were maintained in aquaria and the number which had settled on biologically-conditioned tile pairs was monitored every 5 to 10 d. While the majority of larvae settled quickly after becoming competent, some larvae survived and settled for extended periods after spawning. Competency periods ranged from 26 d for C. serailia to 56 d for G. australensis and 78 d for A. lordhowensis. These data greatly extend the known competency periods for larvae of broadcast-spawning corals and indicate the potential for transport of broadcast-spawned coral larvae over large distances. Medium to long-distance larval dispersal of the species studied provides a mechanism for their widespread distribution in subtropical regions, on reefs which are often widely spaced and relatively isolated. Received: 27 May 1997 / Accepted: 27 November 1997     

Spatial–temporal variability of epibenthic assemblages on subtidal biogenic reefs in the northern Adriatic Sea

Spatial distribution and temporal variation of epibenthic assemblages of coralligenous biogenic rocky outcrops occurring in the northern Adriatic Sea (45°04′–45°24′N; 12°23′–12°43′E) were investigated by photographic sampling from 2003 to 2006 at 12 randomly selected sites. The dominant reef-forming organisms were the encrusting calcareous algae (Lithophyllum stictaeforme, Lithothamnion minervae and Peyssonnelia polymorpha), while the main bioeroders were boring sponges (Cliona viridis, C. celata, C. thoosina, C. rhodensis, Piona vastifica) and the bivalve Gastrochaena dubia. Composition of the assemblages varied thorough years and among sites. Spatial heterogeneity, at local and regional scale, prevailed over temporal variation. This variability was related both to the geo-morphological features of the outcrops and to environmental variables. Sites clearly differed in the percent cover of reef builder and bioeroder species while only limited temporal variation within site was found. Some taxa revealed complex intra-site temporal trends. These results provide valuable information on the diversity and variability of epibenthic assemblages of the northern Adriatic coralligenous reefs, essential for the management and conservation of these unique biogenic habitats.     

Combined effects of temperature and salinity on fed and starved larvae of the mediterranean mussel Mytilus galloprovincialis and the Japanese oyster Crassostrea gigas

The combined effects of temperature, salinity and nutrition on survival and growth of larvae of the Mediterranean mussel Mytilus galloprovincialis and the Japanese oyster Crassostrea gigas were studied over a period of 7 d in the laboratory. Ripe adults, collected in spring and summer 1987 from natural populations in the Bay of Arcachon, France, were induced to spawn. Larvae of both species were cultured at four temperatures (15°, 20°, 25° and 30°C), four salinities (20, 25, 30 and 35S) per temperature, and two levels of nutrition (fed and unfed) per temperature/salinity combination. The fed larvae received a mixed algal diet of 50 cells each of Isochrysis galbana and Chaetoceros calcitrans forma pumilum per microlitre. In both bivalve species, larvae survived over a wide range of temperature and salinity, with the exception of mussel larvae, which died at 30°C. Statistical analysis indicated that nutrition had the greatest effect on larval development, explaining 64 to 75% of the variance in growth of M. galloprovincialis and 54 to 70% in growth of Crassostrea gigas. Unfed mussel larvae displayed little growth. Compared with temperature, the effect of salinity was very slight. M. galloprovincialis larvae exhibited best growth at 20°C and 35S and C. gigas at 30°C and 30S.     

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.     

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     

Single-step species identification of bivalve larvae using multiplex polymerase chain reaction

One of the biggest obstacles to studying recruitment variation in marine bivalves is the need to collect and process large numbers of plankton samples. Larval bivalves are notoriously difficult, if not impossible, to identify to species using morphological criteria alone. Remote time-series collections could satisfy the sampling challenge, but efficient identification techniques must be developed to obtain species-specific data. Thus, we have developed a multiplex polymerase chain reaction (PCR) identification assay in which a single reaction is capable of accurate and efficient discrimination of five target bivalve species based on the size of cytochrome oxidase I products. The assay was tested with cultured and field-sampled larvae as well as adult genomic DNAs. Using a single whole larva as template, multiplex PCR reactions were capable of discriminating among the commercially important bivalves: Mercenaria mercenaria, Argopecten irradians, Mulinia lateralis, Spisula solidissima and Mya arenaria. Overall accuracy was 92%, including very few false positives. The efficiency of this assay stems from its ability to discriminate multiple target species with a single molecular step that ultimately can be automated to process large numbers of larvae. Received: 30 March 2000 / Accepted: 10 July 2000     

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.     

Physiological tolerance predicts species composition at different scales in a barnacle guild

This study examined how the species composition of an intertidal barnacle guild varied according to physical gradients in the environment at small scales governed by microclimates, medium scales of wave exposure and large scales of latitude. Barnacle distributions at small and medium scales were sampled in Ireland between 51°29′ and 52°44′N and 6°50′ and 10°08′W. Sampling on European shores spanned ~18° latitude from 37°05′ to 55°16′N. Barnacle surveys mainly took place in 2003–2004. An index of wave fetch was calculated along the wave exposure gradient using a digital coastline-based model that was supported by a biological exposure scale. A ‘dryness’ index was defined according to mean monthly wind speed, fetch along the average wind direction and mean monthly air or sea surface temperatures for 2 years (January 2001–December 2002) which is the period when the most recent adults in the barnacle community would have settled and grown to adulthood. The proportion of the dry-loving barnacle Chthamalus montagui Southward increased within the barnacle guild at all scales as the habitat became warmer and drier. Barnacle densities were high in all habitats, mean densities ranged from a minimum of 4.16 cm⁻² on moderately exposed shores to a maximum of 6.27 cm⁻² in sunlit or south-facing microclimates. Percentage cover of barnacles across the gradient of latitudes was usually >70%. The results suggest that the distribution and abundance of interacting barnacle species on European coasts is strongly controlled by abiotic factors, most likely temperature and desiccation.     

Where are larvae of the vermetid gastropod <Emphasis Type="Italic">Dendropoma maximum</Emphasis> on the continuum of larval nutritional strategies?

This study evaluated whether larvae of the Indo-Pacific vermetid gastropod Dendropoma maximum are obligate planktotrophs, or whether they exhibit an intermediate feeding strategy. Experiments were conducted in Moorea, French Polynesia (149°50′W, 17°30′S), Sep–Oct 2009, to examine D. maximum larval growth and metamorphic responses to different diets and amounts of food. Dendropoma maximum larvae required particulate food to undergo metamorphosis, but were able to survive and grow in the absence of food for up to 20 days. Larvae in Low and Unfed food treatments exhibited phenotypic plasticity by growing a larger velum (the larval feeding structure) compared with those in high food. Unfed D. maximum larvae had a slower initial growth rate; however, by 11-day post-hatch fed and unfed larvae had converged on the same mean shell height (553 μm), which was only 10% larger than the initial size at hatching. Therefore, although the nutritional strategy of D. maximum larvae is best described as obligate planktotrophy, it appears to approach an intermediate feeding strategy.     

PCR-based detection of mtDNA haplotypes of native and invading mussels on the northeastern Pacific coast: latitudinal pattern of invasion

Frequencies of mitochondrial haplotypes characteristic of native Mytilus trossulus and introduced M. galloprovincialis were determined in populations along the west coast of North America from San Diego, California, to the Aleutian Islands, Alaska. We also identified the haplotypes of mussels cultured from larvae arriving in Coos Bay, Oregon, during 1988–1990 from sites in Japan in the seawater ballast of ocean-going ships. All mussels from ballast-water samples were M. galloprovincialis. We found that sampled populations north of San Francisco Bay, including Coos Bay, were entirely composed of mussels with the M. trossulus haplotype, while haplotypes of both species were present in all sites in and south of San Francisco Bay. The presence of M. trossulus in southern sites is contrary to evidence from allozyme studies, and we suggest that mtDNA introgression from M. trossulus to M. galloprovincialis may explain this discordance. This study demonstrates that, despite continued transport and release, M. galloprovincialis has not become established in northern sites. Failure to invade the north coast of North America may reflect environmental unsuitability for M. galloprovincialis. However, invasion success may be probabilistic, and the continuing release of M. galloprovincialis larvae may foreshadow a future successful invasion.     

Interpreting seawater temperature range using oxygen isotopes and zooid size variation in Pentapora foliacea (Bryozoa)

This is the first investigation of how two independent proxies for seawater temperature inference (zooid size variation and oxygen isotope ratios of skeletal carbonate) relate to the actual measured ranges of temperature experienced by cheilostome bryozoan colonies. Nine specimens of the bimineralic marine cheilostome bryozoan Pentapora foliacea (Ellis and Solander, 1786) were analysed, collected from ~18-m depth at two localities in Wales, UK—four from Skomer Island (51°42′510″N, 5°13′42.60″W) and five from Porth Ysgaden, Lleyn Peninsula (52°54′6.75″N, 4°38′47.34″W). The annual range of temperature implied by zooid size variability provides a good approximation of the actual range of temperature recorded by a datalogger. However, annual ranges of temperature reconstructed from skeletal oxygen isotope ratios were narrower, typically not showing the lowest temperatures experienced by the colonies. This can be explained by progressive thickening of zooid skeletal walls during the life of the colony that homogenises the temperature signal by time-averaging over the lifetime of the colonies. Our study provides evidence that a combined morphological isotope approach has great potential in the reconstruction of annual ranges in seawater temperatures from historical and fossil bryozoans, particularly for species that lack ontogenetic skeletal wall thickening and bimineralic skeletal composition. As cheilostome bryozoans have been common in benthic communities since the Late Cretaceous, they represent a valuable and underutilised resource for the interpretation of environmental regimes.     

<Emphasis Type="Italic">Cochlodinium polykrikoides</Emphasis> blooms and clonal isolates from the northwest Atlantic coast cause rapid mortality in larvae of multiple bivalve species

Globally, many commercial bivalve populations have declined in recent decades. In addition to overharvesting and habitat loss, the increasing frequency and intensity of harmful algal blooms (HABs) are likely to contribute to bivalve losses, particularly in cases where blooms negatively impact larval stages. This paper reports on the lethal effects of clonal cultures and blooms of Cochlodinium polykrikoides from the US Atlantic coast on the larvae of three species of commercially and ecologically valuable bivalves: the Eastern oyster (Crassostrea virginica), the bay scallop (Argopecten irradians), and the Northern quahog (hard clam; Mercenaria mercenaria). Both cultures and blooms of C. polykrikoides were highly toxic to all three species of bivalve larvae causing 80–100% mortality during 24- to 72-h exposures at concentrations of 1–2 × 10³ cells ml⁻¹. Toxicity was dependent on cell densities, growth stage of C. polykrikoides (i.e. cultures in exponential stage growth were more toxic than later stages), exposure time of larvae to cells (i.e. longer exposure caused higher mortality), the age of larvae (i.e. younger larvae were more sensitive), and the relative abundance of C. polykrikoides (i.e. the presence of other microalgae decreased toxicity). Free radical-scavenging enzymes (peroxidase and catalase) and the removal of C. polykrikoides cells (i.e. culture filtrate) significantly increased larval survival suggesting toxicity is maximized by contact with live cells and may involve labile toxins bound by these compounds including e.g. reactive oxygen species. The toxicity of C. polykrikoides to bivalve larvae was generally more severe than other HAB species (e.g. Karenia brevis, Karlodinium veneficum, Alexandrium tamarense, Prorocentrum minimum). Since the bivalves in this study spawn in the months when C. polykrikoides blooms on the east coast of North America, these results suggest that these blooms may have detrimental effects on efforts to restore these already diminished populations.     

Growth of the scyllarid lobsters Ibacus peronii and I. chacei

The growth rates of the morphologically similar scyllarid lobsters Ibacus peronii (Leach, 1815) and I. chacei (Brown and Holthuis, 1998) are described using data from a tag/recapture study and from tagged lobsters kept in captivity. Within particular size classes, we found no differences in moult increments between male and female I. peronii nor between male and female I. chacei. Small individuals of both species always had larger moult increments than larger individuals. For I. peronii, females moulted more frequently than males, and smaller size classes moulted more frequently than larger size classes. Female I. peronii therefore grew more quickly than males and reached their estimated size at sexual maturity (51 mm carapace length) after ∼2 yr. Moulting of I. peronii was seasonal, with most lobsters (96.3%) moulting between October and January. We found no differences in growth rates of I. peronii at two locations along the east coast of Australia: Coffs Harbour in New South Wales (30°18′S; 153°08′E), and Lakes Entrance in Victoria (37°53′S; 148°00′E). For I. chacei, we found no differences in the frequency of moulting between males and females and, because we also found no differences in the moult increments between males and females, the growth rates of both sexes were the same. Received: 14 August 1999 / Accepted: 20 January 2000     

Ecological gradients and relative abundance of native (Mytilus trossulus) and invasive (Mytilus galloprovincialis) blue mussels in the California hybrid zone

Marine communities are experiencing unprecedented rates of species homogenization due to the increasing success of invasive species, but little is known about the mechanisms that allow a species to invade and persist in a new habitat. In central California, native (Mytilus trossulus Gould 1850) and invasive (Mytilus galloprovincialis Lamarck 1819) blue mussels and their hybrids co-exist, providing an opportunity to analyze the mechanisms that determine the distributions of these taxa. Spatial and temporal variation in temperature and salinity and the relative frequencies of these mussel taxa were examined between 2000 and 2004 at four sites in San Francisco Bay and four in Monterey Bay, which were chosen for their different positions along inferred estuarine/oceanic gradients in the hybrid zone. Mussels were genetically identified as the parent species or hybrids by amplifying regions of two species-specific loci: the adhesive byssal thread protein (Glu-5′) and the internal transcribed spacer region of ribosomal DNA (ITS 1). The proportion of M. trossulus at the eight hybrid zone sites correlated negatively with average salinity (R ²=0.60) and positively with maximal temperature (R ²≥0.72), a somewhat unexpected result given what is known about the phylogeography of this species. The proportion of M. galloprovincialis showed the opposite pattern. The proportion of hybrids was correlated neither with habitat temperature nor salinity. Genotypes of mussel populations at an additional 13 sites from Coos Bay, Oregon (latitude 43.35°N) to Long Beach, California (latitude 33.72°N), sampled at various intervals between 2000 and 2004, were also determined. This survey confirmed previous reports that the hybrid zone lies between Monterey and the Cape Mendocino region (latitudes 36.63°N–40.5°N). Within Monterey and San Francisco Bays, however, the temporal comparisons (1990s vs. 2000s) revealed abrupt changes in the proportions of the two parent species and their hybrids on annual and decadal time scales. These changes indicate that the blue mussel populations are in a highly dynamic state. The survey also showed that, regardless of habitat, M. trossulus is consistently of smaller average size than either M. galloprovincialis or hybrids.

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Tidal height,rather than habitat selection for conspecifics,controls settlement in mussels

Settlement is a major determinant of intertidal populations. However, the energy costs of lost larvae are very high. Accordingly, arrival and attachment on suitable substrata are essential requirements for species’ survival. On the intertidal, the presence of cues left by adult or juvenile conspecifics could be vital for the successful establishment of larvae arriving on the shore. Two mussel species, the indigenous Perna perna and the invasive Mytilus galloprovincialis, co-occur on the lower eulittoral zone on the south coast of South Africa. P. perna dominates the low and M. galloprovincialis the high mussel zones, with co-existence in the mid mussel zone. This study tested the hypothesis of settlement selectivity for conspecifics in these two mussel species, to understand whether the final adult distribution of mussels on the shores is determined by active behavioural and chemical mechanisms. Preferential selection by larvae for conspecifics was tested in the field during the peak settlement period in 2004 in natural mussel beds across zones and through manipulative experiments in the mid-zone where the species co-exist. On natural beds, settlement was determined by counts of settlers attached over 48 h onto artificial collectors. Collectors were placed on beds of P. perna and M. galloprovincialis present at both high- and low-adult densities, as well as in mixed beds. On such natural beds, settlers of both species consistently favored low-zone P. perna beds. Settlement patterns over 24 h onto experimentally created mussel patches consisting of P. perna, M. galloprovincialis or the two species combined beds, set in the mixed zone, did not conform with the results of the natural beds study: settlers of both species settled with no discrimination among different patches. The results indicate that mussels, which are sedentary, lack attraction to conspecifics at settlement. This highlights the importance of tidal height in setting settlement rates, and of post-settlement events in shaping populations of these broadcast spawners.     

Tracking hawksbill sea turtles (<Emphasis Type="Italic">Eretmochelys imbricata</Emphasis>) during inter-nesting intervals around Barbados

Hawksbill sea turtles (Eretmochelys imbricata) nesting in Barbados (Needham’s Point, 13° 04′ 41.33′′ N, 59° 36′ 32.69′′W) were outfitted with GPS dataloggers over three breeding seasons (2008–2010) to track movement during inter-nesting intervals. Most females established spatially restricted resident areas up current and within 7 km of the nesting beach where they spent the majority of the inter-nesting interval. Females nesting earlier in the season settled on shallower sites. Only experienced remigrant turtles occupied the most distant resident areas. Females tracked for multiple inter-nesting intervals exhibited site fidelity, but the area contracted and the activity of females decreased with each successive interval. Hawksbills may trade off site characteristics with distance from the nesting beach and reduce activity over the course of the breeding season to optimise energy reserves during inter-nesting intervals.     

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.