Stanislas Dubois

Biodiversity associated with Sabellaria alveolata (Polychaeta: Sabellariidae) reefs: effects of human disturbances

Infauna diversity is reported from the Sabellaria alveolata reefs in the bay of Mont Saint-Michel in France, known as the greatest European reef formation. Polychaetes dominate the fauna, but other species also play a role in terms of reef functioning. Species richness of the associated infauna is much higher than that of the surrounding sediments and is concentrated mainly on the reef surface. Species richness clearly varied according to the three stages of reef evolution: the degraded reef stage is different from the ball-shaped structure and platform stages. Multivariate analyses indicate that three species communities can be distinguished, each corresponding to a different stage of reef development. The number of species is very high, but only a few species are restricted to a particular reef stage. Variations of surface topography and spatial heterogeneity can explain unusual associations of species that make infauna associated with S. alveolata reefs very unique. Moreover, the high densities of S. alveolata (up to 60,000 ind m -2 ) raise questions regarding the role of the reefs in the ecosystem of the bay. Anthropogenic influences, notably mechanical disturbances due to fishing activity, show a steady increase and may have a serious impact on the assemblage diversity and the ecological stability of the reef, even though recolonization of S. alveolata is possible in degraded reef areas. An important question that must be addressed is whether human activities could influence the role of S. alveolata reefs in the bay of Mont Saint-Michel.

Efficiency of particle retention and clearance rate in the polychaete Sabellaria alveolata L.

The development of Sabellaria alveolata, a gregarious reef-building polychaete species, is maximal in Mont-Saint-Michel Bay (France), where trophic capacity is now threatened by increasing shellfish farming. As no data are available concerning the ecophysiological response of this species, the purpose of the present study was to obtain clearance rate and retention efficiency values to provide a first order of magnitude for the trophic role of this species. Data were obtained using a flow-through system with novel troughs suitable for 225 cm2 reef blocks containing a mean number of 940 +/- 102 (S.E.) individuals. The experimental diet used consisted of a mixture of two live microalgae, Skeletonema costatum (3800 cell ml-1) and Isochrysis galbana (23,700 cell ml-1), chosen to cover a broad size range (2 to 16 microns equivalent spherical diameter, ESD), as determined by a particle counter. On the basis of a mean clearance rate of 0.7 lh-1 obtained with reef blocks, the mean rate for an individual was estimated at 7.5 x 10(-4) L h-1. Particles larger than 6 microns ESD were cleared with 100% efficiency, but S. alveolata was unable to retain particles smaller than 2 microns ESD. The results are compared with data obtained for other polychaete species, and clearance rate values are extrapolated to an entire reef.

Isotopic Diversity Indices: How Sensitive to Food Web Structure?

Recently revisited, the concept of niche ecology has lead to the formalisation of functional and trophic niches using stable isotope ratios. Isotopic diversity indices (IDI) derived from a set of measures assessing the dispersion/distribution of points in the δ-space were recently suggested and increasingly used in the literature. However, three main critics emerge from the use of these IDI: 1) they fail to account for the isotopic sources overlap, 2) some indices are highly sensitive to the number of species and/or the presence of rare species, and 3) the lack of standardization prevents any spatial and temporal comparisons. Using simulations we investigated the ability of six commonly used IDI to discriminate among different trophic food web structures, with a focus on the first two critics. We tested the sensitivity of the IDI to five food web structures along a gradient of sources overlap, varying from two distinct food chains with differentiated sources to two superimposed food chains sharing two sources. For each of the food web structure we varied the number of species (from 10 to 100 species) and the type of species feeding behaviour (i.e. random or selective feeding). Values of IDI were generally larger in food webs with distinct basal sources and tended to decrease as the superimposition of the food chains increased. This was more pronounced when species displayed food preferences in comparison to food webs where species fed randomly on any prey. The number of species composing the food web also had strong effects on the metrics, including those that were supposedly less sensitive to small sample size. In all cases, computing IDI on food webs with low numbers of species always increases the uncertainty of the metrics. A threshold of ∼20 species was detected above which several metrics can be safely used.

Ephemeral Bio-engineers or Reef-building Polychaetes: How Stable are Aggregations of the Tube Worm Lanice conchilega (Pallas, 1766)?

Dense aggregations of tube-worms can stabilize sediments and generate oases for benthic communities that are different and often more diverse and abundant than those of the surroundings. If these features are to qualify as biogenic reefs under nature-conservation legislation such as the EC Habitats Directive, a level of stability and longevity is desirable aside from physical and biological attributes. Lanice conchilega (Pallas, 1766) is widely distributed around the European coast and aggregations of this tube-dwelling polychaete are known to have a positive effect on the biodiversity of associated species in inter- and sub-tidal areas. This increases the value of L. conchilega-rich habitats for higher trophic levels such as birds and fish. However, L. conchilega is currently not recognized as a reef builder primarily due to uncertainty about the stability of their aggregations. We carried out three studies on different spatial and temporal scales to explore a number of properties relating to stability: (1) Individual aggregations of L. conchilega of ∼1 m(2) were monitored for up to 1 year, (2) records of L. conchilega from a 258-ha area over a 35-year period were analyzed, (3) the recovery of a population of L. conchilega subjected to disturbances by cultivation of Manila clams (Ruditapes philippinarum) was followed over 3 years. The studies provided evidence about the longevity of L. conchilega aggregations, their resistance to disturbance, their resilience in recovering from negative impact and their large-scale persistence. The results showed that populations of L. conchilega were prone to considerable fluctuation and the stability of aggregations depended on environmental factors and on recruitment. The tube-worms proved to be susceptible to disturbance by cultivation of Manila clams but demonstrated the potential to recover from that impact. The long-term monitoring of a large L. conchilega population in the Bay of Mont Saint Michel (France) indicated that aggregations can persist over many decades with a constant, densely populated core area and an expanding and contracting more thinly populated fringe zone. The stability of aggregations of L. conchilega and the structures they form do not unequivocally fit the currently accepted definition of a reef. However, given L. conchilegas accepted reef-like potential to influence diversity and abundance in benthic communities, we suggest clarifying and expanding the definition of reefs so that species with records of significant persistence in particular areas and which otherwise meet expectations of reefs are included within the definition.

Diversity and composition of macrobenthic community associated with sandy shoals of the Louisiana continental shelf

Along the Louisiana, USA continental shelf, sandy shoals are shallow, possibly oxygen-rich “islands” surrounded by deeper muddy deposits prone to hypoxia. Shoals also contain significant quantities of fine sand that may be mined in the future for coastal restoration. The ecological role of shoals remains poorly understood and we hypothesized that shoals provide critical habitat for benthic invertebrates. Using Ship Shoal as a model system, we assessed the diversity and structure of macrobenthic assemblages and how community structure varies with season and environmental parameters. High biomass (averaging 26.7xa0gxa0m−2) and high diversity (161 species) of macrobenthos was found in 2006. Polychaetes (45%—72 species) and crustaceans comprised most of the species (28%—46 species); spionids and amphipods dominated the polychaete and crustacean groups respectively, both in terms of number of species and abundances. Sharp decreases in diversity, abundance and biomass occurred from spring to autumn. Species diversity and total abundance significantly increased with decreasing sediment grain size and increasing bottom water dissolved oxygen. Across seasons, mole crabs Albunea paretii and amphioxus Branchiostoma floridae typified the community and contributed most of the biomass. The polychaetes Nephtys simoni, Neanthes micromma, Dispio uncinata, Mediomastus californiensis and Magelona sp. A, the amphipod Acanthohautorius sp. A and the burrowing shrimp Ogyrides alphaerostris also contributed to variation in community composition. Cluster analyses quantified seasonal variation, mainly based on sharp decreases in abundance, as well as spatial differences in species composition oriented along both east–west and north–south gradients. Variation in benthic assemblages was correlated with water depth and sediment characteristics (mean grain size and percentage of gravel-sized shell debris). We conclude that Ship Shoal is an unrecognized biodiversity hotspot and a hypoxia refuge compared to the immediate surrounding area where the benthic community is affected by seasonal hypoxia events and we discuss how sand-mining may influence this community.

Geophysical exploration of an active pockmark field in the Bay of Concarneau, southern Brittany, and implications for resident suspension feeders

About a decade ago, a large field of pockmarks (individual features up to 30 m in diameter and <2 m deep) was discovered in water depths of 15–40 m in the Bay of Concarneau in southern Brittany along the French Atlantic coast, covering an overall area of 36 km2 and characterised by unusually high pockmark densities in places reaching 2,500 per square kilometre. As revealed by geophysical swath and subbottom profile data ground-truthed by sediment cores collected during two campaigns in 2005 and 2009, the confines of the pockmark field show a spectacular spatial association with those of a vast expanse of tube mats formed by a benthic community of the suspension-feeding amphipod Haploops nirae. The present study complements those findings with subbottom chirp profiles, seabed sonar imagery and ultrasonic backscatter data from the water column acquired in April 2011. Results show that pockmark distribution is influenced by the thickness of Holocene deposits covering an Oligocene palaeo-valley system. Two groups of pockmarks were identified: (1) a group of large (>10 m diameter), more widely scattered pockmarks deeply rooted (up to 8 ms two-way travel time, TWTT) in the Holocene palaeo-valley infills, and (2) a group of smaller, more densely spaced pockmarks shallowly rooted (up to 2 ms TWTT) in interfluve deposits. Pockmark pore water analyses revealed high methane concentrations peaking at ca. 400 μl/l at 22 and 30 cm core depth in silty sediments immediately above Haploops-bearing layers. Water column data indicate acoustic plumes above pockmarks, implying ongoing pockmark activity. Pockmark gas and/or fluid expulsion resulting in increased turbidity (resuspension of, amongst others, freshly settled phytoplankton) could at least partly account for the strong spatial association with the phytoplankton-feeding H. nirae in the Bay of Concarneau, exacerbating impacts of anthropogenically induced eutrophication and growing offshore trawling activities. Tidally driven hydraulic pumping in gas-charged pockmarks represents a good candidate as large-scale short-term triggering mechanism of pockmark activation, in addition to episodic regional seismic activity.

Investigating isotopic functional indices to reveal changes in the structure and functioning of benthic communities

With the use of stable isotopes, new concepts have emerged based on the idea that the ecological niche can be approximated by the isotopic niche defined as a δ-space area with isotopic δ values as coordinates. This study aims to (i) redefine functional indices originally based on quantitative biological traits of species and demonstrate the ecological significance of newly defined isotopic functional indices (IFI) in a δ-isotopic space, (ii) compare IFI using biomass data with existing unweighted isotopic indices using only isotopic compositions. nUsing a community-wide approach, we tested IFI using isotopic compositions of a large set of associated species from two marine benthic communities widely reported in coastal shallow waters: the common Amphiura filiformis muddy-sand community and the engineered Haploops nirae sandy-mud community. Biomass and isotopic composition (13C and 15N) of all species were measured during four seasons. n IFI were calculated in the isotopic space defined by the two communities, and variations were analysed: (i) isotopic functional richness indices measure the overall extent of the community trophic niche. They are higher in the Haploops community due to a higher diversity in food sources but also to longer food chains. (ii) isotopic functional evenness indices quantify the regularity in species distribution and the density in species packing. They showed that the biomass is concentrated at the edges of the food web in the Haploops community, outside the isotopic range of the main food source. (iii) isotopic functional divergence indices quantify the degree to which species distribution maximizes the divergence. They showed a larger utilization of secondary food sources in the Haploops community. n The IFI variations responded according to expectations overall, based on the extensive knowledge of those communities. Results highlighted that IFI weighted with species biomass provide new insights into how the structure of energy accumulation as biomass between species is likely to underpin community structure and the interplay between structural components of richness, diversity and evenness of biomass distribution.

Food-Web Complexity in Guaymas Basin Hydrothermal Vents and Cold Seeps.

In the Guaymas Basin, the presence of cold seeps and hydrothermal vents in close proximity, similar sedimentary settings and comparable depths offers a unique opportunity to assess and compare the functioning of these deep-sea chemosynthetic ecosystems. The food webs of five seep and four vent assemblages were studied using stable carbon and nitrogen isotope analyses. Although the two ecosystems shared similar potential basal sources, their food webs differed: seeps relied predominantly on methanotrophy and thiotrophy via the Calvin-Benson-Bassham (CBB) cycle and vents on petroleum-derived organic matter and thiotrophy via the CBB and reductive tricarboxylic acid (rTCA) cycles. In contrast to symbiotic species, the heterotrophic fauna exhibited high trophic flexibility among assemblages, suggesting weak trophic links to the metabolic diversity of chemosynthetic primary producers. At both ecosystems, food webs did not appear to be organised through predator-prey links but rather through weak trophic relationships among co-occurring species. Examples of trophic or spatial niche differentiation highlighted the importance of species-sorting processes within chemosynthetic ecosystems. Variability in food web structure, addressed through Bayesian metrics, revealed consistent trends across ecosystems. Food-web complexity significantly decreased with increasing methane concentrations, a common proxy for the intensity of seep and vent fluid fluxes. Although high fluid-fluxes have the potential to enhance primary productivity, they generate environmental constraints that may limit microbial diversity, colonisation of consumers and the structuring role of competitive interactions, leading to an overall reduction of food-web complexity and an increase in trophic redundancy. Heterogeneity provided by foundation species was identified as an additional structuring factor. According to their biological activities, foundation species may have the potential to partly release the competitive pressure within communities of low fluid-flux habitats. Finally, ecosystem functioning in vents and seeps was highly similar despite environmental differences (e.g. physico-chemistry, dominant basal sources) suggesting that ecological niches are not specifically linked to the nature of fluids. This comparison of seep and vent functioning in the Guaymas basin thus provides further supports to the hypothesis of continuity among deep-sea chemosynthetic ecosystems.

Population structure and spread of the polychaete Diopatra biscayensis along the French Atlantic coast: Human-assisted transport by-passes larval dispersal

Intertidal populations of the ecosystem engineering polychaete, Diopatra biscayensis, were analyzed on the French Atlantic coast for three years with individual size estimated from tube-cap aperture. All but the northernmost population along the Bay of Biscay have yearly recruitment. Individuals live 3-5 years and are likely reproductive as one year olds. Simulations indicate dispersal distances are <50xa0km; yet, populations also exist within the Normano-Breton Gulf in the western English Channel, more than 450xa0km from the northernmost Bay of Biscay population at La Trinité-sur-Mer. Three of the four populations in the Normano-Breton Gulf have no young of the year, but are near to active mussel culture where mussel seed is transported on ropes from dense D.xa0biscayensis areas in the Vendée-Charente region in the Bay of Biscay. The majority of D.xa0biscayensis were adjacent to the likely source, mussel seed ropes. Transport assisted by aquaculture is the likely explanation for the populations in the Normano-Breton Gulf.

Are food web structures well represented in isotopic spaces

Isotopic analyses are increasingly used to assess the structure of food webs and a series of isotopic functional indices have been proposed in the last decade to characterize this structure. These indices are based on the foundational assumption that proximity in the isotopic space informs on trophic similarity between species. While it has been recognized for long that this simplifying assumption should be used with caution, no formal evaluation of its domain of validity has been performed to date. We here simulate a large number (15,000) of food webs with varying characteristics to assess i) whether isotopic distance is a good proxy of trophic dissimilarity; ii) whether isotopic functional indices are good proxies of trophic functional properties; and iii) how the quality of these two proxies depend on various species and food web properties. We first demonstrate that isotopic distance is moderately correlated with trophic similarity in simulated food webs. We find however that two isotopic community metrics investigated (Raos index of functional divergence and NND index of functional originality) are good indicators of community trophic structure, especially in food webs with large connectance, and when predators have low levels of diet specialization. We also find that isotopic functional indices are decreasingly reliable as they aim at characterizing more subtle functional patterns (such as sdNND, an index of between species variation in originality). Taken together, our results suggest that the reliability of the isotopic mapping of consumers decreases with their number of potential prey species, but increases with their number of realized prey species. Consequently, isotopic reliability decreases in species-rich food webs with low connectance, large vertical diversity and large incidence of omnivory. Our study highlights that researchers should not hope to grasp subtle patterns of food web structure based solely on widely used isotopic indices...