The benefits of polyandry in the free-spawning polychaete Galeolaria caespitosa

In many species, females are thought to benefit from polyandry due to the reduced risks of fertilization by genetically incompatible sperm. However, few studies that have reported such benefits have directly attributed variation in female reproductive success to the interacting effects of males and females at fertilization. In this paper, we determine whether male x female interactions influence fertilization in vitro in the free-spawning, sessile polychaete Galeolaria caespitosa. Furthermore, we determined whether polyandry results in direct fertilization benefits for females by experimentally manipulating the number of males contributing towards staged spawning events. To test for male x female interaction effects we performed an initial experiment that crossed seven males with six females (in all 42 combinations), enabling us to assess fertilization rates for each specific male-female pairing and attribute variation in fertilization success to males, females and their interaction. This initial experiment revealed a strong interaction between males and females at fertilization, confirming that certain male-female combinations were more compatible than others. A second experiment tested the hypothesis that polyandry enhances female reproductive success by exposing each female's eggs to either a single male's sperm (monandry) or the sperm from three males simultaneously (polyandry). We performed this second experiment at two ecologically relevant sperm concentrations. This latter experiment revealed a strong fertilization benefit of polyandry, independent of the effects of sperm concentration (which were also significant). We suggest that these direct fertilization gains arising from polyandry will constitute an important source of selection on females to mate multiply in nature.

Seawater carbonate chemistry and sperm swimming speed of the polychaete Galeolaria caespitosa in lab experiment

The rapidity of ocean acidification intensifies selection pressure for resilient phenotypes, particularly during sensitive early life stages. The scope for selection is greater in species with greater within-species variation in responses to changing environments, thus enhancing the potential for adaptation. We investigated among-male variation in sperm swimming responses (percent motility and swimming speeds) of the serpulid polychaete Galeolaria caespitosa to near- (delta pH 0.3) and far-future ocean acidification (delta pH 0.5). Responses of sperm swimming to acidification varied significantly among males and were overall negative. Robust sperm swimming behavior under near-future ocean acidification in some males may ameliorate climate change impacts, if traits associated with robustness are heritable, and thereby enhance the potential for adaptation to far-future conditions. Reduced sperm swimming in the majority of male G. caespitosa may decrease their fertilization success in a high CO2 future ocean. Resultant changes in offspring production could affect recruitment success and population fitness downstream.

Does egg competition occur in marine broadcast-spawners?

When the availability of sperm limits female reproductive success, competition for sperm, may be an important broker of sexual selection. This is because sperm limitation can increase the variance in female reproductive success, resulting in strong selection on females to compete for limited fertilization opportunities. Sperm limitation is probably common in broadcast-spawning marine invertebrates, making these excellent candidates for investigating scramble competition between broods of eggs and its consequences for female reproductive success. Here, we report our findings from a series of experiments that investigate egg competition in the sessile, broadcast-spawning polychaete Galeolaria caespitosa. We initially tested whether the order in which eggs encounter sperm affects their fertilization success at two ecologically relevant current regimes. We used a split-clutch-split-ejaculate technique to compare the fertilization success of eggs from individual females that had either first access (competition-free treatment) or second access (egg competition treatment) to a batch of sperm. We found that fertilization success depended on the order in which eggs accessed sperm; eggs that were assigned to the competition-free treatment exhibited significantly higher fertilization rates than those assigned to the egg competition treatment at both current speeds. In subsequent experiments we found that prior exposure of sperm to eggs significantly reduced both the quantity and quality of sperm available to fertilize a second clutch of eggs, resulting in reductions in fertilization success at high and low sperm concentrations. These findings suggest that female traits that increase the likelihood of sperm-egg interactions (e.g. egg size) will respond to selection imposed by egg competition.

Data compilation on the biological response to ocean acidification: environmental and experimental context of data sets and related literature

The exponential growth of studies on the biological response to ocean acidification over the last few decades has generated a large amount of data. To facilitate data comparison, a data compilation hosted at the data publisher PANGAEA was initiated in 2008 and is updated on a regular basis (doi:10.1594/PANGAEA.149999). By January 2015, a total of 581 data sets (over 4 000 000 data points) from 539 papers had been archived. Here we present the developments of this data compilation five years since its first description by Nisumaa et al. (2010). Most of study sites from which data archived are still in the Northern Hemisphere and the number of archived data from studies from the Southern Hemisphere and polar oceans are still relatively low. Data from 60 studies that investigated the response of a mix of organisms or natural communities were all added after 2010, indicating a welcomed shift from the study of individual organisms to communities and ecosystems. The initial imbalance of considerably more data archived on calcification and primary production than on other processes has improved. There is also a clear tendency towards more data archived from multifactorial studies after 2010. For easier and more effective access to ocean acidification data, the ocean acidification community is strongly encouraged to contribute to the data archiving effort, and help develop standard vocabularies describing the variables and define best practices for archiving ocean acidification data.

Polychaete assemblage of an impacted estuary, Guanabara Bay, Rio de Janeiro, Brazil

Thirty-eight stations were sampled in Guanabara Bay, Rio de Janeiro, Brazil, to assess the spatio-temporal diversity and biomass of sublittoral polychaetes. Samples were collected during the dry (September 2000) and rainy season (May 2001) in shallow sublittoral sediments. The polychaete spatial composition showed a heterogeneous distribution throughout the bay. A negative gradient of diversity and biomass was observed towards the inner parts of the bay and sheltered areas. A wide azoic area was found inside the bay. Some high-biomass and low-diversity spots were found near a sewage-discharge point. In these areas, the polychaete biomass increased after the rainy season. A diversified polychaete community was identified around the bay mouth, with no dramatic changes of this pattern between the two sampling periods. Deposit-feeders were dominant in the entire study area. The relative importance of carnivores and omnivores increased towards the outer sector, at stations with coarse sediment fractions. Guanabara Bay can be divided into three main zones with respect to environmental conditions and polychaete diversity and biomass patterns: A) High polychaete diversity, hydrodynamically exposed areas composed of sandy, oxidized or moderately reduced sediments with normoxic conditions in the water column. B) Low diversity and high biomass of deposit and suspension-feeding polychaete species in the middle part of the bay near continental inflows, comprising stations sharing similar proportions of silt, clay and fine sands. C) Azoic area or an impoverished polychaete community in hydrodynamically low-energy areas of silt and clay with extremely reduced sediments, high total organic matter content and hypoxic conditions in the water column, located essentially from the mid-bay towards the north sector. High total organic matter content and hypoxic conditions combined with slow water renewal in the inner bay seemed to play a key role in the polychaete diversity and biomass. Sedimentation processes and organic load coming from untreated sewage into the bay may have negatively affected the survivorship of the fauna.

The polychaete worm Nereis diversicolor increases mercury lability and methylation in intertidal mudflats

The polychaete worm Nereis diversicolor engineers its environment by creating oxygenated burrows in anoxic intertidal sediments. The authors carried out a laboratory microcosm experiment to test the impact of polychaete burrowing and feeding activity on the lability and methylation of mercury in sediments from the Bay of Fundy, Canada. The concentration of labile inorganic mercury and methylmercury in burrow walls was elevated compared to worm-free sediments. Mucus secretions and organic detritus in worm burrows increased labile mercury concentrations. Worms decreased sulfide concentrations, which increased Hg bioavailability to sulfate-reducing bacteria and increased methylmercury concentrations in burrow linings. Because the walls of polychaete burrows have a greater interaction with organisms, and the overlying water, the concentrations of mercury and methylmercury they contain is more toxicologically relevant to the base of a coastal food web than bulk samples. The authors recommend that researchers examining Hg in marine environments account for sediment dwelling invertebrate activity to more fully assess mercury bioavailability.

Mercury and methylmercury bioaccumulation by polychaete worms is governed by both feeding ecology and mercury bioavailability in coastal mudflats

Polychaete worms are abundant in many mudflats but their importance to coastal food web Hg biomagnification is not known. We sampled sediments and polychaete worms from mudflats in the Bay of Fundy to investigate the bioaccumulation of mercury (Hg) and methylmercury (MeHg) in the coastal invertebrate food web. Hg concentrations in the sediments were low (<20 μg kg−1). Labile Hg (methanol/KOH sediment extraction) in surface sediments (0–1 cm) was positively correlated with Hg bioaccumulation by surface sediment-ingesting polychaetes but, surprisingly, there was a negative correlation between δ15N (i.e. trophic level) and THg bioaccumulation factors in polychaete worms. Worms feeding on deeper sediments contained the greatest MeHg concentrations (69.6 μg kg−1). Polychaetes are an important vector for Hg biomagnification to the coastal avian food web. This research demonstrates that feeding depth and method of feeding are more important than trophic position or sediment Hg concentrations for predicting Hg bioaccumulation.

Burrow ventilation and associated porewater irrigation by the polychaete Marenzelleria viridis

Burrow ventilation of benthic infauna generates water currents that irrigate the interstices of the sediments surrounding the burrow walls. Such activities have associated effects on biogeochemical processes affecting ultimately important ecosystem processes. In this study, the ventilation and irrigation behavior of Marenzelleria viridis, an invasive polychaete species in Europe, was analyzed using different approaches. M. viridis showed to perform two types of ventilation: (1) muscular pumping of water out of the burrow and (2) cilia pumping of water into the burrow. Flowmeter measurements presented muscular pumping in time averaged rates of 0.15 ml min(-1). Oxygen needle electrodes positioned above the burrow openings revealed that muscular undulation of the worm body pumps anoxic water out of the burrow. On the other hand, microscope observations of the animal showed that ventilation of oxygen-rich water in the burrow occurs by ciliary action. The volume of water irrigated by M. viridis appears to vary linearly within the first 24 h incubation, with rates ranging from 0.003 to 0.01 ml min(-1). From those rates we could estimate that the time averaged rate of cilia ventilation should be about 0.16 ml min(-1). Since the cilia pumping into the burrow occurs in periods of 24 +/- 12 min and at 50-70% of the measured time, considerable amounts of water from deeper sediments may percolate upwards to the sediment surface. This water is rich in reduced compounds and nutrients and may have important associated ecological implications in the ecosystem (e.g. affecting redox conditions, organic matter degradation, benthic recruitment and primary production). (C) 2010 Elsevier B.V. All rights reserved.

POLYCHAETE ASSEMBLAGE OF AN IMPACTED ESTUARY, GUANABARA BAY, RIO DE JANEIRO, BRAZIL

Thirty-eight stations were sampled in Guanabara Bay, Rio de Janeiro, Brazil, to assess the spatio-temporal diversity and biomass of sublittoral polychaetes. Samples were collected during the dry (September 2000) and rainy season (May 2001) in shallow sublittoral sediments. The polychaete spatial composition showed a heterogeneous distribution throughout the bay. A negative gradient of diversity and biomass was observed towards the inner parts of the bay and sheltered areas. A wide azoic area was found inside the bay. Some high-biomass and low-diversity spots were found near a sewage-discharge point. In these areas, the polychaete biomass increased after the rainy season. A diversified polychaete community was identified around the bay mouth, with no dramatic changes of this pattern between the two sampling periods. Deposit-feeders were dominant in the entire study area. The relative importance of carnivores and omnivores increased towards the outer sector, at stations with coarse sediment fractions. Guanabara Bay can be divided into three main zones with respect to environmental conditions and polychaete diversity and biomass patterns: A) High polychaete diversity, hydrodynamically exposed areas composed of sandy, oxidized or moderately reduced sediments with normoxic conditions in the water column. B) Low diversity and high biomass of deposit and suspension-feeding polychaete species in the middle part of the bay near continental inflows, comprising stations sharing similar proportions of silt, clay and fine sands. C) Azoic area or an impoverished polychaete community in hydrodynamically low-energy areas of silt and clay with extremely reduced sediments, high total organic matter content and hypoxic conditions in the water column, located essentially from the mid-bay towards the north sector. High total organic matter content and hypoxic conditions combined with slow water renewal in the inner bay seemed to play a key role in the polychaete diversity and biomass. Sedimentation processes and organic load coming from untreated sewage into the bay may have negatively affected the survivorship of the fauna.

Taxonomic sufficiency of polychaete taxocenes for estuary monitoring

The polychaetes assemblage structure was used in order to investigate taxonomic sufficiency in a heavily polluted tropical bay. Species abundance was aggregated into progressively higher taxa matrices (genus, family, order) and was analyzed using univariate and multivariate techniques. Polychaetes distribution in Guanabara Bay (GB) was in accordance with a pollution gradient, probably ruled by the organic enrichment, consequent effects of hypoxia and altered redox conditions coupled with prevailing patterns of circulation. Within the sectors of GB, an increasing gradient in species richness and occurrence was observed, ranging from the azoic and impoverished stations in the inner sector to a well-structured community in terms of species composition and abundance inhabiting the outer sector. Multivariate statistical analysis showed similar results when species were aggregated into genera and families, while greater difference occurred at coarser taxonomic identification (order). The literature about taxonomic sufficiency has demonstrated that faunal patterns at different taxonomic levels tend to become similar with increased pollution. In GB, an analysis carried out solely at family level is perfectly adequate to describe the environmental gradient, considered a useful tool for a quick environmental assessment. (C) 2011 Elsevier Ltd. All rights reserved.

Polychaete fauna of the Northwest Portuguese Coastal Shelf: ecology, diversity and distribution

Polychaetes are one of the larger groups of macroinvertebrates with more than 9000 species recognised, distributed worldwide. Thanks to the broad ecological adaptability and high abundaces, this taxon plays a leading role and is considered an important component of all benthic assemblages. Our knowledge about the West Iberian Coast polychaete fauna are scarce, and the only studies are recent. In this sense, the aim of this work was to investigate the composition and the spatial distribution of the polychaete fauna along the NW Portuguese Coastal Shelf, focusing on their relationship to environmental factors (depth, grain size, longitude and latitude) and to add new data to the existing biological dataset. A total of 39 sites were analysed, collected in an area of about 5665 km², between 20 and 150 m depth, distributed in a way to cover the overall grain size gradient. A total of 9352 specimens belonging to 41 families were found, and the analysis based on the abundance of polychaete species revealed five affinity groups: (a) nearshore medium sand characterised by Pisione parapari and Hesionura elongata; (b) very coarse sand that showed the highest abundance of Syllidae and was characterised by Protodorvillea kefersteini and Syllis garciai; (c) fine sand dominated by Spiophanes bombyx and Glycera tridactyla; (d) very fine sand with Nepthys assimilis and Amage sp. and the highest abundance of Paraonidae; (d) mud characterised by Labioleanira yhleni and Ampharete finmarchica. The combination of the environmental variables and the biological data, done with BIOENV routine, demonstrated that depth, grain size and fine contents were the best related with the biological data (rho=0.598). In general, the results agree with the composition and the spatial distribution of the polychaete fauna in other parts of the world; further polychaete assemblages related to mud sediments were firstly recorded in the Northwestern Portuguese Coastal Shelf.

Contributo dei coralliti sub-fossili di Cladocora caespitosa (L.,1767)(Cnidaria, Scleractinia) ai sedimenti dei fondali circostanti le isole del Golfo di La Spezia.

I sedimenti superficiali dei fondali circostanti l’arcipelago del Golfo di La Spezia sono stati analizzati dal punto di vista granulometrico e composizionale al fine di ottenere la mappatura delle concentrazioni di coralliti sub-fossili di Cladocora caespitosa nel sedimento. Mediante lo studio del sedimento campionato in trentacinque stazioni, sono state individuate tre zone di accumulo di coralliti: (i) in corrispondenza del capo occidentale dell’Isola Palmaria con le concentrazioni più elevate comprese tra il 25 e 55% (ii) sul lato sud-orientale della stessa isola con concentrazioni tra il 10 e 12% e (iii) una fascia contornante l’Isola del Tinetto con quantità inferiori al 3%. La concentrazione anomala di coralliti è il risultato dello scarico di materiali di dragaggio provenienti dal porto di La Spezia, scaricati al largo delle coste occidentali dell’arcipelago tra gli anni ‘50 e ’70 e progressivamente ridistribuiti verso sud-est dalla deriva litorale.

Structures and Concentrations of Surfactants in Gut Fluid of the Marine Polychaete Arenicola Marina

Marine invertebrate deposit feeders secrete surfactants into their gut fluid in concentrations sufficient to induce micelle formation, enhancing solubilization of sedimentary lipids. We isolated and identified 3 related surfactant molecules from the deposit-feeding polychaete lugworm Arenicola marina. Surfactants were isolated and separated by a combination of solvent extraction and thin-layer and gas chromatography. Identification was performed using mass and infrared spectrometry, coupled to various derivatization and hydrolysis reactions. A. marina produces a mixture of related yet distinct anionic surfactants composed of branched, C9, saturated and unsaturated fatty acids that are amide linked to leucine or glycine residues, showing some similarity to crustacean surfactants. The critical micelle concentration of the mixture of these surfactants in gut fluid was about 2 mM, and total concentrations ranged from 5.5 to 19.5 mM. The hydrophilic amide linkage helps to explain previous observations that gut surfactants do not adsorb onto sediment transiting the gut.

Reproductive Biology of the Deep-Sea Polychaete Gorgoniapolynoe Caeciliae (Polynoidae), a Commensal Species Associated with Octocorals

Some aspects of the reproductive biology of the polychaete Gorgoniapolynoe caeciliae have been described for the first time. Gorgoniapolynoe caeciliae is a deep-sea commensal species associated with Candidella imbricala, all octocoral that populates the New England Seamount chain. Gorgoniapolynoe caeciliae is a dioccious species with an equal sex ratio and fertile segments throughout most of the adult body. The gonads of both sexes are associated with genital blood vessels emerging from the posterior surface of most intersegmental septa. In the female, oogenesis is intraovarian with oocytes being retained within the ovary until vitellogenesis is completed. The largest female examined contained over 3000 eggs with a maximum diameter of 80-90 mu m. In the male, the testes are repeated in numerous segments and consist of small clusters of spermatogonia, spermatocytes and early spermatids associated with the walls of the genital blood vessels. Early spermatids are shed into the coelom where they complete differentiation into mature ect-aquasperm with a spherical head (4 mu m), a small cap-like acrosome, and a short mid-piece with four mitochondria. Indirect evidence suggests that this species is an annual breeder that releases its gametes into seawater and produces a planktotrophic larva following fertilization. The reproductive biology of G. caeciliae is consistent with that of most other polynoids including many shallow water species suggesting that phylogenetic history strongly shapes its biology.

Blattformen : 1. Alchemilla vulgaris - 2. Trollius europaeus - 3. Aconitum napellus - 4. Saxifraga - 5. Saxifraga caespitosa - 6. Hydrophyllum virginicum - 7. Alchemilla alpina - 8. Artemisia absinthium

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