Is <Emphasis Type="Italic">Hippolyte williamsi</Emphasis> gonochoric or hermaphroditic? A multi-approach study and a review of sexual systems in <Emphasis Type="Italic">Hippolyte</Emphasis> shrimps

Sexual systems vary considerably among caridean shrimps and while most species are gonochoric, others are described as sequential protandric hermaphrodites or simultaneous hermaphrodites with an early male phase. At present, there is confusion about the sexual system exhibited by several species mostly because those studies attempting to reveal their sexual system draw inferences solely from the distribution of the sexes across size classes. Here we investigated the sexual system of the shrimp Hippolyte williamsi from Chile to determine if the species is protandric or gonochoric with sexual dimorphism (males smaller than females). Morphological identification and size frequency distributions indicated that the population comprised small males, small immature females, and large mature females, which was confirmed by dissections. No transitional individuals were found. Males maintained in the laboratory molted 1–8 times, and many grew up to reach sizes observed in only a small fraction of males in the field. No indication of sex change was recorded. Our results indicate that H. williamsi is a sexually dimorphic gonochoric species and emphasizes the importance of using several kinds of evidence (size measurements, growth experiments, morphological dissections, and histological studies) to reveal the sexual system of Hippolyte species. Whether the observed size dimorphism between males and females in many species of Hippolyte is expression of contrasting sexual and natural selection, and whether divergent sexual fitness functions can contribute to the evolution of hermaphroditism remains to be revealed in future studies.     

Size–frequency dynamics of NE Pacific abyssal ophiuroids (Echinodermata: Ophiuroidea)

The 17-year time-series study at Station M in the NE Pacific has provided one of the longest datasets on deep-sea ophiuroids to date. Station M is an abyssal site characterized by low topographical relief and seasonal and interannual variation in surface-derived food inputs. From 1989 to 2005, over 31,000 ophiuroid specimens were collected. Size–frequency distributions of the four dominant species, Ophiura bathybia, Amphilepis patens, Amphiura carchara and Ophiacantha cosmica, were examined for recruitment and the role of surface-derived food supplies on body size distributions. Juveniles were collected in sediment traps and used to investigate settlement patterns and seasonality. Trawl samples showed no indication of seasonal changes in recruitment to larger size classes; however, there was evidence of seasonal settling of juveniles. Interannual differences in median disk diameters and size distributions of trawl-collected adults are greater than those at the seasonal scale. Three of the four species, O. bathybia, A. patens and O. cosmica, had co-varying monthly median disk diameters, suggesting they may have a similar factor(s) controlling their growth and abundance. Interannual differences in monthly size distributions were generally greater than those between seasons. Cross-correlations between the particulate organic carbon (POC) flux (food supply) and size distribution indices for O. bathybia, A. patens and O. cosmica all were significant indicating that increases in food supply were followed by increases in the proportion of smaller size classes after approximately 17–22 months. These findings suggest that food inputs are indeed an important factor influencing deep-sea ophiuroid populations on interannual time scales, more generally supporting the long-hypothesized connection between food availability and population size structure in the deep sea. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The life cycles of two coexisting copepods, <Emphasis Type="Italic">Calanus chilensis</Emphasis> and <Emphasis Type="Italic">Centropages brachiatu</Emphasis>s,in the upwelling zone off northern Chile (23°S)

A time series study of the copepods Calanus chilensis and Centropages brachiatus was carried out at the coastal upwelling zone of Mejillones (23°S, northern Chile), to analyze their annual life cycles in association with upwelling variation. These species co-exist in the upwelling zone. Weekly sampling of zooplankton and oceanographic variables including Chlorophyll-a and phytoplankton composition were obtained during January–December 2002 at a fixed station (ca. 90 m depth). Stages of abundances, their proportions, changes in body length of adult females, sex ratio and egg production rate (EPR), were used as proxies to examine copepods’ demography. Upwelling, assessed by weekly Ekman transport and oceanographic conditions, was intermittent throughout the year with lack of periodicity components. Populations of both copepod species did not correlate with these non-predictable upwelling events. C. chilensis reproduced year-round and the population showed ca. 15 peaks of adults with an average time interval between peaks of 20 days. C. brachiatus showed a similar life cycle, also having 15 peaks of adults at about 22 days of time intervals. Cross-correlation functions and spectral analysis showed that both populations correlated positively through time, but not in phase, evidencing a time lag for their reproductive cycles. The lag was also evident in their population abundances. Both species differ in their development rates and this may result in non-in phase life cycles. Our findings suggest that species-dependent attributes, such as development rates, modulated by adaptations to temperature, might impose constraints in the species life cycles determining the population cycles. Such attributes must be considered when modeling and understanding population dynamics and secondary production of copepods.     

Contrasts between spawning times of <Emphasis Type="Italic">Anguilla</Emphasis> species estimated from larval sampling at sea and from otolith analysis of recruiting glass eels

This study reviewed literature on spawning times for three north temperate species of anguillid eels estimated by sampling for small leptocephali (larvae) at sea and for several temperate and tropical species by back-calculating from putative daily ages derived from otolith increment analysis of glass eels that recruited to coastal waters. Estimates from otoliths of European eels, Anguilla anguilla, American eels, Anguilla rostrata, and Japanese eels, Anguilla japonica, imply much more protracted spawning seasons than are indicated by sampling at sea during various times of year. European eels are inferred to spawn year-round from otolith analysis, but the smallest, recently hatched leptocephali are found only in late winter and spring. From otoliths, the spawning times of these three species are all estimated to occur much later in the year than when small leptocephali are found at sea, indicating that ages appear to be underestimated. For these and other temperate and tropical eels, there are inconsistencies in assigned ages among various studies, which are most extreme for the European eel. This species has the longest larval migration and often has an opaque zone in the glass eels’ otoliths where it is difficult to discern growth increments. These inconsistencies suggest that interpretation of otolith growth increments is incorrect at least in some studies, and the apparently consistent mismatch between otolith and sea-sampling studies suggests that increments may not always be formed at some period during the unusual early life history of anguillids. Because daily increments may be formed in eels during most of their early life history, future research is needed to determine the cause of the mismatch of glass eel aging studies and the apparent spawning times of eels offshore. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.

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Nutrient transfer in a marine mutualism: patterns of ammonia excretion by anemonefish and uptake by giant sea anemones

Many symbioses involve multiple partners in complex, multi-level associations, yet little is known concerning patterns of nutrient transfer in multi-level marine mutualisms. We used the anemonefish symbiosis as a model system to create a balance sheet for nitrogen production and transfer within a three-way symbiotic system. We quantified diel patterns in excretion of ammonia by anemonefish and subsequent absorption by host sea anemones and zooxanthellae under laboratory conditions. Rates of ammonia excretion by the anemonefish Amphiprion bicinctus varied from a high of 1.84 μmole g⁻¹ h⁻¹ at 2 h after feeding, to a basal rate of 0.50 μmole g⁻¹ h⁻¹ at 24–36 h since the last meal. Conversely, host sea anemones Entacmaea quadricolor absorbed ammonia at a rate of 0.10 μmole g⁻¹ h⁻¹ during the daytime in ammonia-enriched seawater, but during the night reduced their absorption rate to near zero, indicating that ammonia uptake was driven by zooxanthella photosynthesis. When incubated together, net ammonia excretion was virturally zero, indicating that host anemones absorbed most of the ammonia produced by resident fish. Adult anemonefish weighed about 11 g under laboratory conditions, but on the coral reef may reach up to 64 g, resulting in a maximal potential ammonia load of >200 μmole h⁻¹ produced by two adult fish during daylight hours. In contrast, host sea anemones weighed about 47 g in the laboratory, but under field conditions, large individuals may reach 680 g, so their maximal ammonia clearance rates may reach about 70 μmole h⁻¹ during the daytime. As such, the ammonia load produced by adult anemonefish far exceeds the clearance rate of host anemones and zooxanthellae. Ammonia transfer likely occurs mainly during the daytime, when anemonefish consume zooplankton and excrete rapidly, and in turn the zooxanthellae are photosynthetically active and drive rapid ammonia uptake. We conclude that zooplanktivorous fishes that form mutualisms with coral reef cnidarians may serve as an important link between open water and benthic ecosystems, through the transfer of large quantities of nutrients to zooxanthellate hosts, thus enhancing coral reef productivity.     

Distribution of a marginal population of <Emphasis Type="Italic">Mytilus edulis</Emphasis>: responses to biotic and abiotic processes at different spatial scales

Physical and biological processes interact to produce pattern in nature. Pattern is scale dependent as processes generating pattern are heterogeneous in time and space. We tested some causes of variation in abundance and distribution of three marginal populations of sublittoral blue mussels, Mytilus edulis, in the non-tidal northeastern Baltic Sea. We studied the role of substrate inclination, perennial algae and siltation along local wave exposure gradients on mussel distribution over a regional salinity gradient. We found marked differences on regional scales (p < 0.001) with lower densities and biomasses of mussels with declining salinity. Along local gradients, mussel densities increased with increasing exposure (p < 0.001) and declining slope and sedimentation (p < 0.01). Site specifically, densities of blue mussels and the perennial red algae, Furcellaria lumbricalis, were positively related, results supported by a colonisation experiment. Also, young post-recruits showed significant relations to adult biomass, wave exposure, algal biomass, bottom slope and sediment cover. Findings showed that the relative importance of the determinants affecting blue mussels at the edge of their range vary with scale and are affected by the density and size structure of mussel populations. The study provides an indication of the types of factors that may be invoked as causes of spatial variation in marginal blue mussel populations and reinforces the need to consider multiple aspects when distributional patterns are assessed.     

Differences in morphology and habitat use among the native mussel <Emphasis Type="Italic">Mytilus trossulus</Emphasis>, the non-native <Emphasis Type="Italic">M. galloprovincialis,</Emphasis> and their hybrids in Puget Sound,Washington

Mytilus galloprovincialis (Mg), the Mediterranean blue mussel, is sympatric with the native M. trossulus (Mt) throughout much of the north Pacific, likely as the result of human introduction. We investigated the distribution of the two species and their hybrids (Mgt) in Puget Sound, Washington, to determine whether differences occur in habitat preference between the two species and hybrids. In addition, we investigated whether there were consistent size and shape differences between the native and introduced mussels and hybrids. Measurements of over 6,000 mussels from 30 sites, of which 1,460 were genotyped for a species-specific genetic marker, revealed that Mg and Mgt can be found throughout Puget Sound. Mg and Mgt were larger and exhibited a greater height:length ratio than Mt. Frequencies of Mg and Mgt were higher in subtidal habitats, such as docks, than on intertidal rocks, walls or pilings. Within intertidal habitats, Mg and Mgt were more frequent than Mt in the lower reaches of the intertidal. At slightly more than half the sites the frequency of the three genotypes accorded with random mating expectations suggesting no consistent barriers to gene flow between species. The standardized random sampling methods and simple morphometric identification techniques described here can be used to test whether the frequency of invasive mussels changes over time and space in Puget Sound.     

Feeding behavior of adult <Emphasis Type="Italic">Vinciguerria nimbaria</Emphasis> (Phosichthyidae), in the tropical Atlantic (0°–4°N, 15°W)

Adult Vinciguerria nimbaria are the main prey of tuna during the tuna fishing season (late autumn and winter) in the equatorial Atlantic (0–4°N, and ~15°W). V. nimbaria trophic behavior in the fishing grounds was studied in relation to hydrobiological factors to determine its role in the trophic food web. Sampling stations spaced by 20 nautical miles were set up along a 15°W north–south transect from 4°N to 0°40S. At each station, the temperature and vertical fluorescence profiles were recorded. Nitrate and chlorophyll a analyses were performed on water sampled at different levels in the euphotic zone. Vertical plankton hauls were carried out at depths of 0–100 and 0–200 m using a standard WP2 net fitted with a 200-μm mesh gauze. Vinciguerria nimbaria adults were collected using a young-fish mid-water trawl net (10 × 15 m opening mouth, 10 mm cod end mesh). The weight of the stomach contents, the stomach fullness index, the number of prey, the frequency of occurrence and the prey preponderance were recorded for 20 fish from each haul. An oligotrophic typical tropical structure (TTS) was found between 1° and 4°N where small zooplankton was relatively abundant above or near the thermocline. In the TTS, V. nimbaria behaved as an epipelagic fish, feeding on the dominant small prey during the daytime. In turn, it was a prey for tuna. In the equatorial zone, where zooplankton was more abundant than in the north equatorial zone, V. nimbaria behaved as a mesopelagic fish and as an opportunistic mesozooplankton feeder. It consumed a wide range of sizes of food, feeding on the most abundant species of zooplankton as well as the largest zooplankton species, possibly while migrating towards the surface in the late afternoon or in the deep layer.     

Morphological and molecular data suggest a cosmopolitan distribution of the polychaete <Emphasis Type="Italic">Proscoloplos cygnochaetus</Emphasis> Day, 1954 (Annelida,Orbiniidae)

The discovery of a possibly invasive Proscoloplos species on the French Atlantic coast led to a detailed morphological and molecular investigation. Proscoloplos consists of three nominal species, but molecular analyses of the variable ITS1 and ITS2 region revealed no clade support for samples from South Africa, France and Australia, representing at least two of these nominal species. We found no unambiguous diagnostic characters for the three different species even with scanning electron microscopy. Two main characters—first appearance of branchiae and hooks—that were used for species delineation show considerable intrapopulational variation and do not withstand critical evaluation. The obtained data and the observed regenerative capabilities point to a cautious use of the position of branchiae and hooked chaetae as taxonomic marker in Proscoloplos. After experimental bisection of animals, the recovery led to a phenotype without recognizable signs of regeneration, but neither branchiae nor hooked chaetae reoccurred at the same initial segmental position. Summarising the molecular and morphological data we suggest the synonymy of the known Proscoloplos species under the senior synonym Proscoloplos cygnochaetus Day, 1967. The enormous geographic range of this species comprises the temperate waters of the whole southern hemisphere and includes the French population. We discuss a dispersal enabled through the effective anchorage on vessels using mucous glands and chaetae possibly combined with architomic reproduction.     

Strong variability in bacterioplankton abundance and production in central and western Bay of Bengal

With large influx of freshwater that decreases sea-surface salinities, weak wind forcing of <10 m s⁻¹ and almost always warm (>28°C) sea-surface temperature that stratifies and shallows the mixed layer leading to low or no nutrient injections into the surface, primary production in Bay of Bengal is reportedly low. As a consequence, the Bay of Bengal is considered as a region of low biological productivity. Along with many biological parameters, bacterioplankton abundance and production were measured in the Bay of Bengal during post monsoon (September–October 2002) along an open ocean transect, in the central Bay (CB, 88°E) and the other transect in the western Bay (WB). The latter representing the coastal influenced shelf/slope waters. Bacterioplankton abundances (<2 × 10⁹cells l⁻¹) were similar to those reported from the HNLC equatorial Pacific and the highly productive northern Arabian Sea. Yet, the thymidine uptake rates along CB (average of 1.46 pM h⁻¹) and WB (average of 1.40 pM h⁻¹) were less than those from the northwestern Indian Ocean. These abundances and uptake rates were higher than those in the oligotrophic northwestern Sargasso Sea (<7 × 10⁸ cells l⁻¹; av 1.0 pM h⁻¹). Concentrations of chlorophyll a (chl a), primary production rates and total organic carbon (TOC) were also measured for a comparison of heterotrophic and autotrophic production. In the WB, bacterioplankton carbon biomass equaled ∼ 95% of chl a carbon than just 31% in the CB. Average bacterial:primary production (BP:PP) ratios accounted for 29% in the CB and 31% in the WB. This is mainly due to lower primary productivity (PP) in the WB (281 mg C m⁻² d⁻¹) than in the CB (306 mg C m⁻² day⁻¹). This study indicates that bacteria–phytoplankton relationship differs in the open (CB) and coastal waters (WB). Higher abundance and contrastingly low bacterial production (BP) in WB may be because of the riverine bacteria, brought in through discharges, becoming dormant and unable to reproduce in salinities of 28 or more psu. Heterotrophic bacteria appear to utilize in situ DOC rather rapidly and their carbon demand is ∼50% of daily primary production. It is also apparent that allochthonous organic matter, in particular in the western Bay, is important for meeting their carbon demand.