Anna Metaxas

Quantifying the “Bio-” Components in Biophysical Models of Larval Transport in Marine Benthic Invertebrates: Advances and Pitfalls

Biophysical models are being used increasingly, both as predictive tools of larval dispersal for a particular system and for general evaluation of the role of different factors in larval transport. In the results of such models, larval duration, mortality, and behavior in the water column have exhibited pronounced effects on larval dispersal of marine benthic invertebrates. The parameterization of these processes has broadly reflected values from laboratory experiments, but the accuracy of these values is unknown. The pelagic larval duration used in models should be determined by laboratory, or preferably field, studies and should incorporate environmentally dependent variability. For mortality, in situ estimates are now feasible and, likely, more accurate than the currently used values. Larval behavior can be measured in the field, by high-frequency sampling of distributional changes relative to features in the water column or by controlled larval releases in tractable systems. To successfully validate the outcomes of these models, we must either improve our techniques for measuring larval abundance at the end of larval transport immediately before settlement, or incorporate components for settlement into the models. We must also address the mismatch in sampling resolution between biological and physical processes. If used with caution, this powerful approach can significantly advance our understanding of larval transport.

Estimating dispersal distance in the deep sea: challenges and applications to marine reserves

Population connectivity refers to the exchange of individuals among populations: it affects gene flow, regulates population size and function, and mitigates recovery from natural or anthropogenic disturbances. Many populations in the deep sea are spatially fragmented, and will become more so with increasing resource exploitation. Understanding population connectivity is critical for spatial management. For most benthic species, connectivity is achieved by the planktonic larval stage, and larval dispersal is, in turn, regulated by complex interactions between biological and oceanographic processes. Coupled biophysical models, incorporating ocean circulation and biological traits, such as planktonic larval duration (PLD), have been used to estimate population connectivity and generate spatial management plans in coastal and shallow waters. In the deep sea, knowledge gaps in both the physical and biological components are delaying the effective use of this approach. Here, we review the current efforts in conservation in the deep sea and evaluate (1) the relevance of using larval dispersal in the design of marine protected areas and (2) the application of biophysical models in the study of population connectivity. Within biophysical models, PLD can be used to estimate dispersal distance. We propose that a PLD that guarantees a minimum dispersal distance for a wide range of species should be used in the planning of marine protected areas in the deep sea. Based on a review of data on species found at depths > 200 m, a PLD of 35 and 69 days ensures a minimum distance for 50% and 75%, respectively, of eurybathic and deep-sea species. We note that more data are required to enhance accuracy and address the high variability in PLD between and within taxonomic groups, limiting generalizations that are often appealing to decision-makers. Given the imminent expansion of resource exploitation in the deep sea, data relevant to spatial management are needed urgently.

Hydrothermal Vents and Methane Seeps: Rethinking the Sphere of Influence

Although initially viewed as oases within a barren deep ocean, hydrothermal vent and methane seep communities are now recognized to interact with surrounding ecosystems on the sea floor and in the water column, and to affect global geochemical cycles. The importance of understanding these interactions is growing as the potential rises for disturbance from oil and gas extraction, seabed mining and bottom trawling. Here we synthesize current knowledge of the nature, extent and time and space scales of vent and seep interactions with background systems. We document an expanded footprint beyond the site of local venting or seepage with respect to elemental cycling and energy flux, habitat use, trophic interactions, and connectivity. Heat and energy are released, global biogeochemical and elemental cycles are modified, and particulates are transported widely in plumes. Hard and biotic substrates produced at vents and seeps are used by “benthic background” fauna for attachment substrata, shelter, and access to food via grazing or through position in the current, while particulates and fluid fluxes modify planktonic microbial communities. Chemosynthetic production provides nutrition to a host of benthic and planktonic heterotrophic background species through multiple horizontal and vertical transfer pathways assisted by flow, gamete release, animal movements, and succession, but these pathways remain poorly known. Shared species, genera and families indicate that ecological and evolutionary connectivity exists among vents, seeps, organic falls and background communities in the deep sea; the genetic linkages with inactive vents and seeps and background assemblages however, are practically unstudied. The waning of venting or seepage activity generates major transitions in space and time that create links to surrounding ecosystems, often with identifiable ecotones or successional stages. The nature of all these interactions is dependent on water depth, as well as regional oceanography and biodiversity. Many ecosystem services are associated with the interactions and transitions between chemosynthetic and background ecosystems, for example carbon cycling and sequestration, fisheries production, and a host of non-market and cultural services. The quantification of the sphere of influence of vents and seeps could be beneficial to better management of deep-sea environments in the face of growing industrialization.

Relative importance of parental and larval nutrition on larval development and metamorphosis of the sea urchin Strongylocentrotus droebachiensis

Abstract We examined the relative importance of parental nutritional condition and larval food ration on the rates of development, growth and metamorphosis of larvae of Strongylocentrotus droebachiensis (Muller) in a laboratory experiment. Parents were reared for 22 months on either a high ration of kelp ( Laminaria spp., 6 days week −1 ) supplemented with mussel flesh ( Mytilus spp., 1 day week −1 ) (KM), or a low ration of kelp (1 day week −1 ) (KL). Larvae were fed either a high ration (5000 cells ml −1 ) or a low ration (500 cells ml −1 ) of microalgae ( Dunaliella tertiolecta ). Larval food ration had a strong effect on the rates of development, growth, and metamorphosis, which were all significantly greater in larvae fed the high ration. Test diameter of settlers also was significantly greater in the high than the low ration. Parental nutritional condition had little or no effect on the rates of development and growth, and no effect on settler size. The rate of metamorphosis was significantly higher in larvae from the KM than the KL treatment in the high but not the low ration (where rates of metamorphosis were similar). Although parental condition generally had a small effect on larval development, our results suggest that when planktonic food is abundant, larvae of adults from nutritionally rich habitats (such as kelp beds) may metamorphose sooner than those of adults from nutritionally poor habitats (such as barrens).

Predicting habitat for two species of deep-water coral on the Canadian Atlantic continental shelf and slope

Documentation of hundreds of locations for Canadian deep-water corals has been obtained through scientific initiatives and local fishermen’s knowledge. Using these locations, as well as relevant oceanographic data, this study determined areas of suitable habitat for Paragorgia arborea and Primnoa resedaeformis along the Canadian Atlantic continental shelf and shelf break using predictive models. The study area included a band approximately 800 km long x 200 km wide from Cape Breton to the Gulf of Maine, and was chosen based on density of coral sites. Several environmental factors including slope, temperature, chlorophyll a, current speed and substrate may be important in determining suitable coral habitat and were included in the analysis. There are many different techniques used to model habitats, but frequently they are limited by the type of data available. Comparatively, few techniques using presence-only data are available. We utilized BioMapper, a program which uses Ecological Niche Factor Analysis (ENFA), to generate habitat suitability maps by relating data on species presence with background environmental data to determine the species’ niche. We found that habitat requirements differed between the two species of coral. For Paragorgia arborea, the niche was highly specialized, and characterized by steeply sloped environments and rocky substrate. In contrast, for Primnoa resedaeformis, suitable habitat was more broadly distributed in the study area, and located in areas with high current speed, rocky substrates and an approximate temperature range between 5 and 10°C. This is the first study to use predictive modelling to identify suitable habitat for deep-water coral, which may prove an important tool for the conservation of these organisms.

Biogeographic and bathymetric ranges of Atlantic deep-sea echinoderms and ascidians: the role of larval dispersal

Dispersal plays an important role in the establishment and maintenance of biodiversity and, for most deep-sea benthic marine invertebrates, it occurs mainly during the larval stages. Therefore, the mode of reproduction (and thus dispersal ability) will affect greatly the biogeographic and bathymetric distributions of deep-sea organisms. We tested the hypothesis that, for bathyal and abyssal echinoderms and ascidians of the Atlantic Ocean, species with planktotrophic larval development have broader biogeographic and bathymetric ranges than species with lecithotrophic development. In comparing two groups with lecithotrophic development, we found that ascidians, which probably have a shorter larval period and therefore less dispersal potential, were present in fewer geographic regions than elasipod holothurians, which are likely to have longer larval periods. For asteroids and echinoids, both the geographic and bathymetric ranges were greater for lecithotrophic than for planktotrophic species. For these two classes, the relationships of egg diameter with geographic and bathymetric range were either linearly increasing or non-monotonic. We conclude that lecithotrophic development does not necessarily constrain dispersal in the deep sea, probably because species with planktotrophic development may be confined to regions of high detrital input from the sea surface. Our data suggest that more information is necessary on lengths of larval period for different species to accurately assess dispersal in the deep sea.

Environmental Impacts of the Deep-Water Oil and Gas Industry: A Review to Guide Management Strategies

The industrialization of the deep sea is expanding worldwide. Expanding oil and gas exploration activities in the absence of sufficient baseline data in these ecosystems has made environmental management challenging. Here, we review the types of activities that are associated with global offshore oil and gas development in water depths over 200 m, the typical impacts of these activities, some of the more extreme impacts of accidental oil and gas releases, and the current state of management in the major regions of offshore industrial activity including 18 exclusive economic zones. Direct impacts of infrastructure installation, including sediment resuspension and burial by seafloor anchors and pipelines, are typically restricted to a radius of approximately 100 m on from the installation on the seafloor. Discharges of water-based and low-toxicity oil-based drilling muds and produced water can extend over 2 km, while the ecological impacts at the population and community levels on the seafloor are most commonly on the order of 200-300 m from their source. These impacts may persist in the deep sea for many years and likely longer for its more fragile ecosystems, such as cold-water corals. This synthesis of information provides the basis for a series of recommendations for the management of offshore oil and gas development. An effective management strategy, aimed at minimizing risk of significant environmental harm, will typically encompass regulations of the activity itself (e.g. discharge practices, materials used), combined with spatial (e.g. avoidance rules and marine protected areas) and temporal measures (e.g. restricted activities during peak reproductive periods). Spatial management measures that encompass representatives of all of the regional deep-sea community types is important in this context. Implementation of these management strategies should consider minimum buffer zones to displace industrial activity beyond the range of typical impacts: at least 2 km from any discharge points and surface infrastructure and 200 m from seafloor infrastructure with no expected discharges. Although managing natural resources is, arguably, more challenging in deep-water environments, inclusion of these proven conservation tools contributes to robust environmental management strategies for oil and gas extraction in the deep sea.

Megafauna associated with assemblages of deep-water gorgonian corals in Northeast Channel, off Nova Scotia, Canada

The distribution and abundance of benthic megatauna in areas known to be inhabited by dense gorgonian coral assemblages were examined at Northeast Channel, off Nova Scotia, Canada, in August 2001. Using a remotely operated vehicle, 1 5 video transects during each of 1-2 dives at each of four sites (Rips, Middle Canyon, Hell Hole West and Hell Hole East) were conducted. The relationships in the structure of biological assemblages at three spatial scales: within transects (10s of metres); between dive locations (100s of metres); and among sites (10s to 100s of kilometres) were explored. The most abundant epibenthic taxa included the gorgonians Primnoa resedaeformis and Paragorgia arborea, several suspension feeders (Actinauge verrilli, Bolocera tudiae, an unidentified anemone and encrusting sponge, Ophiacantha abyssicola), the deposit feeder Porania pulvillus insignus and the predatory Solaster endeca. The basket star Gorgonocephalus arcticus was present only on colonies of Paragorgia arborea. Despite large variability in abundance and assemblage composition among transects and dive locations, clear patterns were observed among sites. Mean abundance of most cnidarians and echinoderms was greatest at Hell Hole West. No gorgonians were found at Hell Hole East. The encrusting sponge was most abundant at the Rips and least abundant at Hell Hole East. Cluster analysis and multidimensional sealing (MDS) indicated that, when abundance was averaged across transects for each dive, the megafaunal assemblages fall into groups of dives that separated by site. These differences, among sites are most likely related to variability in the physical environment. The epibenthic megafaunal assemblages were as diverse in the presence as in the absence of gorgonian corals, at least at the abundances that we observed. However, the apparent low recruitment and abundance, combined with small population size make these assemblages particularly vulnerable to perturbations.

Early life history of deep-water gorgonian corals may limit their abundance.

Deep-water gorgonian corals are long-lived organisms found worldwide off continental margins and seamounts, usually occurring at depths of ∼200–1,000 m. Most corals undergo sexual reproduction by releasing a planktonic larval stage that disperses; however, recruitment rates and the environmental and biological factors influencing recruitment in deep-sea species are poorly known. Here, we present results from a 4-year field experiment conducted in the Gulf of Maine (northwest Atlantic) at depths >650 m that document recruitment for 2 species of deep-water gorgonian corals, Primnoa resedaeformis and Paragorgia arborea. The abundance of P. resedaeformis recruits was high, and influenced by the structural complexity of the recipient habitat, but very few recruits of P. arborea were found. We suggest that divergent reproductive modes (P. resedaeformis as a broadcast spawner and P. arborea as a brooder) may explain this pattern. Despite the high recruitment of P. resedaeformis, severe mortality early on in the benthic stage of this species may limit the abundance of adult colonies. Most recruits of this species (∼80%) were at the primary polyp stage, and less than 1% of recruits were at stage of 4 polyps or more. We propose that biological disturbance, possibly by the presence of suspension-feeding brittle stars, and limited food supply in the deep sea may cause this mortality. Our findings reinforce the vulnerability of these corals to anthropogenic disturbances, such as trawling with mobile gear, and the importance of incorporating knowledge on processes during the early life history stages in conservation decisions.

Can Salinity-Induced Mortality Explain Larval Vertical Distribution With Respect to a Halocline?

For the larvae of two echinoderm species that coexist in Atlantic Canada (bipinnaria of the sea star Asterias rubens and 4- and 6-arm echinoplutei of the sea urchin Strongylocentrotus droebachiensis), we examined the effect of short- and long-term exposure to salinity (ranging from 18 to 35) on the probability of larval survival in laboratory experiments. We also related larval vertical distributions in response to sharp haloclines generated in the laboratory to survival probability in the salinity of different layers in the water column. For both species and developmental stages, survival probability decreased with decreasing salinity, and a salinity range of 24–27 emerged as the critical threshold for larval tolerance. The relationship between the proportion of larvae that crossed a halocline into the top water layer and the survival probability of larvae in the salinity of that layer was significant for both species. Interestingly, the shape of this response was species-specific but not stage-specific for S. droebachiensis. Our findings suggest that larval avoidance of low-salinity water layers may be an adaptive behavior that increases survival and indirectly influences larval distribution.