The importance of fine-scale flow processes and food availability in the maintenance of soft-sediment communities

Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy at the Massachusetts Institute of Technology and the Woods Hole Oceanographic Institution January 1993Although the association between soft-sediment invertebrates and a specific sediment type has been documented for many habitats, most studies have been correlative and have failed to convincingly demonstrate any single mechanism to explain this association. Sediment type has generally been characterized by grain size, however, many other potential causal factors correlate with grain size, including organic content, microbial content, stability, food supply, and larval supply. One hypothesis for animal-sediment associations is that settling larvae are transported as passive particles and are sorted into different sedimentary habitats much like sediment grains. To test the hypothesis that near-bed hydrodynamics may modify larval settlement, field and flume experiments were conducted where larval settlement was compared between microdepositional environments (small depressions) and non-trapping environments (flush treatments). Depressions have been observed to trap passive particles, and these experiments were therefore designed to test whether settling larvae would be trapped in depressions like passive particles. Flume flow simulations were carried out with the polychaete Capitella sp. I and the bivalve Mulinia latera/is. Experiments with flush and depression sediment treatments were conducted in the absence of the potentially confounding effects of suspended sediment and organic matter and therefore offered a highly controlled, explicit test of passive hydrodynamic deposition of larvae in depressions. Although larvae of both species were generally able to actively select a high-organic sediment over a low-organic alternative with a comparable grain size, elevated densities of both species were observed in depressions for a given sediment treatment. Thus, both species appeared to be vulnerable to hydrodynamic trapping. M. latera/is larvae, however, often made a "poor choice" by settling in high numbers in depressions containing the low-organic sediment while Capitella sp. I larvae were generally able to "escape" from depressions if the sediment was unsuitable. In field experiments carried out at Station R in Buzzards Bay, Massachusetts, significantly higher densities of Mediomastus ambiseta juveniles, spionid polychaete juveniles, bivalves, gastropod larvae, and nemerteans were observed in depressions compared with flush treatments over 5 relatively short experimental periods (3 or 4 days each) during the summer of 1990. Of the abundant taxa, only Capitella spp. was not significantly more abundant in depressions compared with flush treatments, although numbers tended to be higher in depressions. Experiments were conducted over a short time period to minimize potential biological interactions between taxa and reduce the likelihood that organic material would accumulate in depressions and provide a cue for settling larvae. Thus, higher numbers in depressions suggest that larvae were passively entrained. These flume and field experiments suggest that near-bed hydrodynamics may modify settlement at some scales, and that both active and passive processes may operate in determining larval distributions in shallow-water, muddy habitats. In deep-sea ecosystems, the role of near-bed hydrodynamics is also of interest because of the potential role that larval settlement in organic patches may play in maintaining the immense species diversity characteristic of many deep-sea ecosystems. To try to understand the role of organic patches in deep-sea communities, several investigators have used colonization trays containing sediments that have been treated in different ways. These experiments have been criticized in the past because the sediment surface in the trays was elevated above the bottom and may therefore have interfered with natural boundary layer flow. Flume simulations of flow over these colonization trays revealed serious flow artifacts generated by the trays, and that flow across the sediment surface of the trays was characterized by turbulent eddies, accelerated velocities and boundary layer thickening. These sorts of flow characteristics would not be expected over natural sediments, and an alternative colonization tray was designed to eliminate these artifacts. To test the hypothesis that different types of food patches would result in different types of larval response, and determine how near-bed hydrodynamics may influence larval settlement, flush colonization trays filled with prefrozen sediment were deployed in tandem with artificial depressions south of St. Croix, U.S.V.I at 900 m depth. Colonization trays and artificial depressions were either unenriched or enriched with Thalassiosira sp. and Sargassum sp. two types of algae chosen to mimic natural food patches on the sea floor. Unexpectedly high densities of organisms colonized trays after only 23 days. The Thalassiosira trays were colonized by high densities of a relatively low diversity, opportunistic fauna, Sargassum trays were colonized by lower densities of a higher diversity fauna, and unenriched trays were colonized by very low numbers of a very diverse fauna. All tray faunas were markedly different in composition from the natural, ambient fauna. These fmdings suggest that different patch types did, indeed, result in a specialized faunal response to each of the "patch" types. Depressions on the sea floor provide a natural mechanism for food patch formation because passive particles such as detritus and algae tend to be entrained in the depressions. To determine whether dominant colonizers would be entrained in depressions like passive particles or could differentiate between depression "patch" types in a flow environment that might be expected to make active selection more difficult, artificial depressions were unenriched or enriched with Sargassum sp. or Thalassiosira sp. Total densities of organisms and densities of the most abundant species were substantially lower in artificial depressions than in trays. Densities in Thalassiosira depressions were lower than in Sargassum depressions and densities in unenriched depressions were extremely low, suggesting that dominant colonizers were not passively entrained in depressions and that colonization was specialized and highly active for these taxa. A different fauna was also observed in natural depressions compared with flush sediments, suggesting that natural depressions do contribute to species coexistence. Long-term tray deployments designed to test whether different faunas would be present in "patches" of different ages indicated that time may also play an important part in a deep-sea patch mosaic.This was funded by NSF and ONR, NOAA, NSERC (Canada), WHOI Ocean Ventures Fund and the WHOI Ditty Bag Fund

Spatial linkages between decapod planktonic and benthic adult stages in a Newfoundland fjordic system

The relative importance of predatory decapod crustaceans in sedimentary communities depends on the spatial variability in their abundance and composition. At the scale of a fjord, such spatial patterns could be related to sill-mediated larval supply. This study examines larval and adult distributions of abundant predatory decapods at six representative sites in a sub-arctic Newfoundland fjord during three consecutive summers. Adult snow crab (Chionoecetes opilio) and toad crab (Hyas coarctatus, H. araneus) characterized outer areas of the fjord, whereas pandalid shrimp (Pandalus montagui) dominated inner areas, and rock crab (Cancer irroratus) showed only minor spatial differences. Multivariate analysis and nonparametric comparisons of larval abundance and composition suggest that the sill separating inner and outer areas of the fjord results in differences in larval supply that correspond to adult abundances for at least two of the species analyzed here: snow crab and pandalid shrimp. Although larval abundance was not related to adult distribution when all zoeal stages were considered, correspondence between larval and adult patterns emerged when only late stages (zoeae ≥ II) were included in the multivariate analyses. Nonparametric comparisons supported these results, indicating significant differences in larval abundance inside and outside the sill for corresponding species and stages. Our results suggest that larval supply may play a critical role in establishing adult spatial patterns at the scale of the entire fjord for some species, but a less relevant role at the finer scale represented by the sites and habitats located at each side of the sill

Adult macrofauna effects on Capitella sp. I larval settlement: A laboratory flume study

The opportunistic, deposit-feeding polychaete Capitella sp. I is the overwhelming numerical dominant in disturbed and enriched sediments and rarely co-occurs in appreciable numbers with other abundant mud-dwelling macrofauna. Rapid colonization and population increase in organicrich sediments is typically followed by subsequent sharp decline. The mechanistic basis for these characteristics was explored in flume-flow experiments that tested whether settling Capitella sp. I larvae avoid sediments inhabited by macrofaunal adults or sediments reworked by them. The first set of experiments consisted of four treatments: conspecific adults or no adults in reworked or non-reworked sediment. Capitella sp. I settlement was significantly altered (depressed) only by pelletized sediment of conspecific adults. The second set of experiments involved similar treatments, but with adults of the deposit-feeding bivalve Tellina agilis. Neither adult presence nor sediment reworking significantly affected settlement of Capitella sp. I larvae. A third set of experiments that compared settlement in sediments with and without the suspension-feeding bivalve Mulinia lateralis demonstrated no significant treatment effect. These results suggest that larval settlement behavior could contribute to population growth in a boom and bust species when a critical limiting resource is overexploited. That is, sediments completely pelletized by Capitella sp. I adults may signal settling larvae that organic matter is depleted. Larvae may therefore settle in smaller numbers and are more likely to be dispersed away from abundant populations of adults. Active avoidance of conspecific adults or adults of other taxa is unimportant for the taxa at the densities tested here

Potential flow artifacts associated with benthic experimental gear: Deep-sea mudbox examples

In response to the growing recognition of the potential effects of near-bed hydrodynamics on various benthic processes, flume studies were conducted to document fine-scale flow patterns over several types of mudboxes that have been used to study colonization by deep-sea organisms. Mudboxes are typically filled with natural sediments or sediment treatments and placed in the field to observe how timing, larval supply and sediment composition may affect larval settlement. This study addresses potential hydrodynamic biases of mudbox structures as obstructions to the near-bed flow. Detailed velocity profiles were made over two types of “free vehicle” mudboxes that could be deployed and recovered from a surface vessel. One of these (“Old Free Vehicle”) was not designed with regard for potential hydrodynamic biases whereas the other (“New Free Vehicle”) was designed specifically to minimize flow disturbances and maintain a realistic boundary-layer flow over the mudbox sediments. Flume velocity profiles also were made over two smaller mudboxes designed to be deployed by a submersible, one (“Flush Sediment Tray”) which was designed to be placed flush with the ocean bottom, thus minimizing flow disturbance, and another (“Raised Mudbox”) which was not. Flume simulations indicated that the Old Free Vehicle and the Raised Mudbox cause considerable disturbance to the near-bed flow regime; flows over the mudbox sediment surface differed markedly from those predicted for the natural seabed and those observed over the flume bed in the absence of the mudboxes. Flow accelerations, growing secondary boundary layers and eddy formation were observed over these mudbox sediments, and vertical velocity profiles varied considerably in the along-channel direction. The alternative mudbox designs (New Free Vehicle and Flush Sediment Tray) were largely successful in reducing or eliminating these flow artifacts. Boundary-layer flows over both the New Free Vehicle and the Flush Sediment Tray were much more uniform, and velocity profiles over the sediment surfaces were very similar to those in the empty flume channel and those predicted for a natural deep-sea habitat. In addition, there was no evidence of eddy formation and other major flow disturbances. These flume studies underscore the benefit of considering potential hydrodynamic effects in designing benthic experimental sampling gear to reduce potential flow disturbances that may bias data collections and confound data interpretation

Environmentally mediated trends in otolith composition of juvenile Atlantic cod (Gadus morhua)

Author Posting. © The Author(s), 2015. This is the author's version of the work. It is posted here by permission of Oxford University Press for personal use, not for redistribution. The definitive version was published in ICES Journal of Marine Science 72 (2015): 2350-2363, doi: 10.1093/icesjms/fsv070.We evaluated the influence of environmental exposure of juvenile Atlantic cod (Gadus morua) to inform interpretations of natal origins and movement patterns using otolith geochemistry. Laboratory rearing experiments were conducted with a variety of temperature (~ 5, 8.5 and 12 °C) and salinity (~ 25, 28.5 and 32 PSU) combinations. We measured magnesium (Mg), manganese (Mn), strontium (Sr) and barium (Ba), expressed as a ratio to calcium (Ca), using laser ablation inductively coupled plasma mass spectrometry (ICP-MS), and stable carbon (δ13C) and oxygen (δ18O) isotopes using isotope ratio monitoring mass spectrometry. Temperature and salinity significantly affected all elements and isotopes measured, with the exception of salinity on Mg:Ca. We detected significant interactions among temperature and salinity for Mn:Ca and Ba:Ca partition coefficients (ratio of otolith chemistry to water chemistry), with significant temperature effects only detected in the 32 and 28.5 PSU salinity treatments. Similarly, we detected a significant interaction between temperature and salinity in incorporation of δ13C, with a significant temperature effect except at intermediate salinity. These results support the contention that environmental mediation of otolith composition varies among species, thus limiting the ability of generalized models to infer life history patterns from chemistry. Our results provide essential baseline information detailing environmental influence on juvenile Atlantic cod otolith composition, punctuating the importance of laboratory validations to translate species-specific otolith composition when inferring in situ life histories and movements.Research funding and support was provided by a Natural Sciences and Engineering Research Council of Canada (NSERC) Strategic Grant on Connectivity in Marine Fishes. R. Stanley was supported by an NSERC Postgraduate Scholarship and a Research and Development Corporation of Newfoundland (RDC) student fellowship.2016-04-2

The role of colonization in establishing patterns of community composition and diversity in shallow-water sedimentary communities

To determine whether pattern and diversity in benthic sedimentary communities are set primarily at colonization or by post-settlement biological interactions, we collected faunal cores and conducted reciprocal sediment transplant experiments at a sandy and a muddy site at 12 m depth, ~3 km apart off New Jersey. Multivariate analyses of cores collected at these sites in September 1994 indicated differences in the taxa determining local pattern, with the bivalve Spisula solidissima and the polychaete Polygordius sp. being dominant at the sandy site, and oligochaetes, several polychaete species and the bivalve Nucula annulata dominant at the muddy site. Individual cores from the sandy site were significantly less diverse than those at the muddy site. Short-term experiments (3-5 d) were deployed by divers at three different times (August-September, 1994). Replicate trays (100 cm2) filled with azoic sand or mud were placed flush with the ambient seafloor at both sites. Multivariate comparisons indicated that sediment treatment in trays played a greater role in determining colonization patterns in the first experiment, site played a greater role in the second, and both variables contributed in the third. This pattern suggests that larval settlement and habitat choice played an important role in the first and third experiments, and that local transport of recently settled juveniles from the surrounding sediments was important in the second and third experiments. Sandy-site trays had significantly lower diversity than muddy-site trays, but there was no effect of sediment type in trays on diversity of colonizers. These experiments focused on small spatial scales and three short time periods, but they demonstrate that species patterns in some environments may be set by habitat selection by larvae and by juvenile colonization from the surrounding community. Post-colonization processes such as predation and competition likely play a major role for some species, but patterns of initial colonization corresponded well with those in the local community

Resuspension by fish facilitates the transport and redistribution of coastal sediments

Author Posting. © Association for the Sciences of Limnology and Oceanography, 2012. This article is posted here by permission of Association for the Sciences of Limnology and Oceanography for personal use, not for redistribution. The definitive version was published in Limnology and Oceanography 57 (2012): 945-958, doi:10.4319/lo.2012.57.4.0945.Oxygen availability restricts groundfish to the oxygenated, shallow margins of Saanich Inlet, an intermittently anoxic fjord in British Columbia, Canada. New and previously reported 210Pb measurements in sediment cores compared with flux data from sediment traps indicate major focusing of sediments from the oxygenated margins to the anoxic basin seafloor. We present environmental and experimental evidence that groundfish activity in the margins is the major contributor to this focusing. Fine particles resuspended by groundfish are advected offshore by weak bottom currents, eventually settling in the anoxic basin. Transmittance and sediment trap data from the water column show that this transport process maintains an intermediate nepheloid layer (INL) in the center of the Inlet. This INL is located above the redox interface and is unrelated to water density shifts in the water column. We propose that this INL is shaped by the distribution of groundfish (as resuspension sources) along the slope and hence by oxygen availability to these fish. We support this conclusion with a conceptual model of the resuspension and offshore transport of sediment. This fish-induced transport mechanism for sediments is likely to enhance organic matter decomposition in oxygenated sediments and its sequestration in anoxic seafloors.The VENUS Project and University of Victoria supported the ship and submersible time for field experiments, and the U.S. Geological Survey and Coastal and Marine Geological Program generously supported J.C. The project was supported by Discovery Grants from the Natural Sciences and Engineering Research Council of Canada to V.T. and P.S. and a Yohay Ben-Nun fellowship and Moshe Shilo Center for Marine Biogeochemistry Fund award to T.K

Bridging the gap between policy and science in assessing the health status of marine ecosystems

Human activities, both established and emerging, increasingly affect the provision of marine ecosystem services that deliver societal and economic benefits. Monitoring the status of marine ecosystems and determining how human activities change their capacity to sustain benefits for society requires an evidence-based Integrated Ecosystem Assessment approach that incorporates knowledge of ecosystem functioning and services). Although, there are diverse methods to assess the status of individual ecosystem components, none assesses the health of marine ecosystems holistically, integrating information from multiple ecosystem components. Similarly, while acknowledging the availability of several methods to measure single pressures and assess their impacts, evaluation of cumulative effects of multiple pressures remains scarce. Therefore, an integrative assessment requires us to first understand the response of marine ecosystems to human activities and their pressures and then develop innovative, cost-effective monitoring tools that enable collection of data to assess the health status of large marine areas. Conceptually, combining this knowledge of effective monitoring methods with cost-benefit analyses will help identify appropriate management measures to improve environmental status economically and efficiently. The European project DEVOTES (DEVelopment Of innovative Tools for understanding marine biodiversity and assessing good Environmental Status) specifically addressed t hese topics in order to support policy makers and managers in implementing the European Marine Strategy Framework Directive. Here, we synthesize our main innovative findings, placing these within the context of recent wider research, and identifying gaps and the major future challenges

Seafloor biodiversity of Canada's three oceans: patterns, hotspots and potential drivers

Aim We examined the relationships between bathymetry, latitude and energy and the diversity of marine benthic invertebrates across wide environmental ranges of Canada's three oceans. Location Canadian Pacific, Arctic and Atlantic Oceans from the intertidal zone to upper bathyal depths, encompassing 13 marine ecoregions. Methods We compiled 35 benthic datasets that encompass 3,337 taxa (70% identified to species and 21% to genus) from 13,172 samples spanning 6,117 sites. Partitioning the analyses by different gear types, ecoregions or sites, we used Hill numbers to examine spatial patterns in α‐diversity. We used resampling and extrapolation to standardized sampling effort and examined the effects of depth, latitude, chemical energy (export particulate organic carbon [POC] flux), thermal energy (bottom temperature) and seasonality of primary production on the benthic biodiversity. Results The Canadian Arctic harboured the highest benthic diversity (e.g. epifauna and common and dominant infauna species), whereas the lowest diversity was found in the Atlantic. The Puget Trough (Pacific), Beaufort Sea, Arctic Archipelago, Hudson Bay, Northern Labrador and Southern Grand Bank (Atlantic) were the “hotspots" of diversity among the ecoregions. The infauna and epifauna both exhibited hump‐shaped diversity–depth relationships, with peak diversity near shelf breaks; latitude (positively) predicted infaunal diversity, albeit weakly. Food supply, as inferred from primary production and depth, was more important than thermal energy in controlling diversity patterns. Limitations with respect to calculating POC flux in coastal (e.g. terrestrial runoff) and ice‐covered regions or biological interactions may explain the negative POC flux–infaunal diversity relationship. Main Conclusions We show previously unreported diversity hotspots in the Canadian Arctic and in other ecoregions. Our analyses reveal potential controlling mechanisms of large‐scale benthic biodiversity patterns in Canada's three oceans, which are inconsistent with the prevailing view of seafloor energy–diversity relationships. These results provide insightful information for conservation that can help to implement further MPA networks

Resolving fine-scale population structure and fishery exploitation using sequenced microsatellites in a northern fish

Funding Information Natural Sciences and Engineering Research Council of Canada (NSERC) Strategic Project Atlantic Canada Opportunities Agency and Department of Tourism, Culture, Industry and Innovation grants allocated to the Labrador Institute (MC) Natural Sciences and Engineering Research Council of Canada (NSERC) Discovery Genomics Research and Development Initiative (GRDI) Weston Family AwardPeer reviewedPublisher PD