Deposit-feeding in benthic macrofauna : Tracer studies from the Baltic Sea

A low content of organic matter, which is largely refractory in nature, is characteristic of most sediments, meaning that aquatic deposit-feeders live on a very poor food source. The food is derived mainly from sedimenting phytodetritus, and in temperate waters like the Baltic Sea, from seasonal phytoplankton blooms. Deposit-feeders are either bulk-feeders, or selective feeders, which preferentially ingest the more organic-rich particles in the sediment, including phytodetritus, microbes and meiofauna. The soft-bottom benthos of the Baltic Sea has low species biodiversity and is dominated by a few macrobenthic species, among which the most numerous are the two deposit-feeding amphipods Monoporeia affinis and Pontoporeia femorata, and the bivalve Macoma balthica. This thesis is based on laboratory experiments on the feeding of these three species, and on the priapulid Halicryptus spinulosus. Feeding by benthic animals is often difficult to observe, but can be effectively studied by the use of tracers. Here we used the radioactive isotope 14C to label food items and to trace the organic matter uptake in the animals, while the stable isotopes 13C and 15N were used to follow feeding on aged organic matter in the sediment. The abundance of M. balthica and the amphipods tends to be negatively correlated, i.e., fewer bivalves are found at sites with dense populations of amphipods, with the known explanation that newly settled M. balthica spat are killed by the amphipods. Whether the postlarvae are just accidentally killed, or also ingested after being killed was tested by labelling the postlarvae with 14C and Rhodamine B. Both tracer techniques gave similar evidence for predation on and ingestion of postlarval bivalves. We calculated that this predation was likely to supply less than one percent of the daily carbon requirement for M. affinis, but might nevertheless be an important factor limiting recruitment of M. balthica. The two amphipods M. affinis and P. femorata are partly vertically segregated in the sediment, but whether they also feed at different depths was unknown. By adding fresh 14C-labelled algae either on the sediment surface or mixed into the sediment, we were able to distinguish surface from subsurface feeding. We found M. affinis and P. femorata to be surface and subsurface deposit-feeders, respectively. Whether the amphipods also feed on old organic matter, was studied by adding fresh 14C-labelled algae on the sediment surface, and using aged, one-year-old 13C- and 15N-labelled sediment as deep sediment. Ingestion of old organic matter, traced by the stable isotopes, differed between the two species, with a higher uptake for P. femorata, suggesting that P. femorata utilises the older, deeper-buried organic matter to a greater extent. Feeding studies with juveniles of both M. affinis and P. femorata had not been done previously. In an experiment with the same procedure and treatments as for the adults, juveniles of both amphipod species were found to have similar feeding strategies. They fed on both fresh and old sediment, with no partitioning of food resources, making them likely to be competitors for the same food resource. Oxygen deficiency has become more wide-spread in the Baltic Sea proper in the last half-century, and upwards of 70 000km2 are now devoid of macrofauna, even though part of that area does not have oxygen concentrations low enough to directly kill the macrofauna. We made week-long experiments on the rate of feeding on 14C-labelled diatoms spread on the sediment surface in different oxygen concentrations for both the amphipod species, M. balthica and H. spinulosus. The amphipods were the most sensitive to oxygen deficiency and showed reduced feeding and lower survival at low oxygen concentrations. M. balthica showed reduced feeding at the lowest oxygen concentration, but no mortality increase. The survival of H. spinulosus was unaffected, but it did not feed, showing that it is not a surface deposit-feeder. We conclude that low oxygen concentrations that are not directly lethal, but reduce food intake, may lead to starvation and death in the longer term.

Comparison of epifaunal assemblages between Cymodocea nodosa and Caulerpa prolifera meadows in Gran Canaria (eastern Atlantic)

Máster en Oceanografía

Chlorophyll a concentration and zooplankton abundance and biomass below melting sea ice in the Amundsen Gulf

Early summer in the Arctic with extensive ice melt and break-up represents a dramatic change for sympagic-pelagic fauna below seasonal sea ice. As part of the International Polar Year-Circumpolar Flaw Lead system study (IPY-CFL), this investigation quantified zooplankton in the meltwater layer below landfast ice and remaining ice fauna below melting ice during June (2008) in Franklin Bay and Darnley Bay, Amundsen Gulf, Canada. The ice was in a state of advanced melt, with fully developed melt ponds. Intense melting resulted in a 0.3- to 0.5-m-thick meltwater layer below the ice, with a strong halocline to the Arctic water below. Zooplankton under the ice, in and below the meltwater layer, was sampled by SCUBA divers. Dense concentrations (max. 1,400 ind./m**3) of Calanus glacialis were associated with the meltwater layer, with dominant copepodid stages CIV and CV and high abundance of nauplii. Less abundant species included Pseudocalanus spp., Oithona similis and C. hyperboreus. The copepods were likely feeding on phytoplankton (0.5-2.3 mg Chl-a/m**3) in the meltwater layer. Ice amphipods were present at low abundance (<10 ind./m**2) and wet biomass (<0.2 g/m**2). Onisimus glacialis and Apherusa glacialis made up 64 and 51% of the total ice faunal abundance in Darnley Bay and Franklin Bay, respectively. During early summer, the autochthonous ice fauna becomes gradually replaced by allochthonous zooplankton, with an abundance boom near the meltwater layer. The ice amphipod bust occurs during late stages of melting and break-up, when their sympagic habitat is diminished then lost.

Near-bottom zooplankton aggregations in Kongsfjorden with link to images from the optical zooplankton sensor MOKI

Near-bottom zooplankton communities have rarely been studied despite numerous reports of high zooplankton concentrations, probably due to methodological constraints. In Kongsfjorden, Svalbard, the near-bottom layer was studied for the first time by combining daytime deployments of a remotely operated vehicle (ROV), the optical zooplankton sensor moored on-sight key species investigation (MOKI), and Tucker trawl sampling. ROV data from the fjord entrance and the inner fjord showed high near-bottom abundances of euphausiids with a mean concentration of 17.3 ± 3.5 n/100 m**3. With the MOKI system, we observed varying numbers of euphausiids, amphipods, chaetognaths, and copepods on the seafloor at six stations. Light-induced zooplankton swarms reached densities in the order of 90,000 (euphausiids), 120,000 (amphipods), and 470,000 ind/m**3 (chaetognaths), whereas older copepodids of Calanus hyperboreus and C. glacialis did not respond to light. They were abundant at the seafloor and 5 m above and showed maximum abundance of 65,000 ind/m**3. Tucker trawl data provided an overview of the seasonal vertical distribution of euphausiids. The most abundant species Thysanoessa inermis reached near-bottom concentrations of 270 ind/m**3. Regional distribution was neither related to depth nor to location in the fjord. The taxa observed were all part of the pelagic community. Our observations suggest the presence of near-bottom macrozooplankton also in other regions and challenge the current view of bentho-pelagic coupling. Neglecting this community may cause severe underestimates of the stock of elagic zooplankton, especially predatory species, which link secondary production with higher trophic levels.

Biometrics, fatty acid composition and stable isotopic ratios of five seabird species from Kongsfjorden, Svalbard

Marine birds are important predators in the marine ecosystem, and dietary studies can give useful information about their feeding ecology, food webs and oceanographic variability. The aim of this study was to increase our understanding of the diet and trophic level of the seabirds breeding in Kongsfjorden, Svalbard. We have used fatty acids and stable isotopes, both of which integrate diet information over space and time, to determine trophic relationships in marine food webs. Fatty acid compositions of muscle from Little auk (Alle alle), Brünnich's guillemot (Uria lomvia), Black-legged kittiwake (Rissa tridactyla), Northern fulmar (Fulmarus glacialis) and Glaucous gull (Larus hyperboreus) were determined and compared with their prey species. Canonical analysis (CA) showed that fatty acid composition differed among the five seabird species. Little auk, Black-legged kittiwake and Northern fulmar had high levels of the Calanus markers 20:1n9 and 22:1, indicating that these seabirds are a part of the Calanus food chain. Brünnich's guillemot differed from the other species with much lower levels of 20:1n9 and 22:1. Brünnich's guillemot is a pursuit diver feeding on fish and amphipods deeper in the water column, below 30 m. Glaucous gull also differed from the other seabird species, with a larger variation in the fatty acid composition indicating a more diverse diet. Trophic level analysis placed Little auk at the lowest trophic level, Brünnich's guillemot and Black-legged kittiwake at intermediate levels and Glaucous gull and Northern fulmar at the highest trophic level.

Properties and faunal abundances of level sea ice and sea-ice ridges in the Chukchi/Beaufort Seas and Canada Basin

The abundances and distribution of metazoan within-ice meiofauna (13 stations) and under-ice fauna (12 stations) were investigated in level sea ice and sea-ice ridges in the Chukchi/Beaufort Seas and Canada Basin in June/July 2005 using a combination of ice coring and SCUBA diving. Ice meiofauna abundance was estimated based on live counts in the bottom 30 cm of level sea ice based on triplicate ice core sampling at each location, and in individual ice chunks from ridges at four locations. Under-ice amphipods were counted in situ in replicate (N=24-65 per station) 0.25 m**2 quadrats using SCUBA to a maximum water depth of 12 m. In level sea ice, the most abundant ice meiofauna groups were Turbellaria (46%), Nematoda (35%), and Harpacticoida (19%), with overall low abundances per station that ranged from 0.0 to 10.9 ind/l (median 0.8 ind/l). In level ice, low ice algal pigment concentrations (<0.1-15.8 µg Chl a /l), low brine salinities (1.8-21.7) and flushing from the melting sea ice likely explain the low ice meiofauna concentrations. Higher abundances of Turbellaria, Nematoda and Harpacticoida also were observed in pressure ridges (0-200 ind/l, median 40 ind/l), although values were highly variable and only medians of Turbellaria were significantly higher in ridge ice than in level ice. Median abundances of under-ice amphipods at all ice types (level ice, various ice ridge structures) ranged from 8 to 114 ind/m**2 per station and mainly consisted of Apherusa glacialis (87%), Onisimus spp. (7%) and Gammarus wilkitzkii (6%). Highest amphipod abundances were observed in pressure ridges at depths >3 m where abundances were up to 42-fold higher compared with level ice. We propose that the summer ice melt impacted meiofauna and under-ice amphipod abundance and distribution through (a) flushing, and (b) enhanced salinity stress at thinner level sea ice (less than 3 m thickness). We further suggest that pressure ridges, which extend into deeper, high-salinity water, become accumulation regions for ice meiofauna and under-ice amphipods in summer. Pressure ridges thus might be crucial for faunal survival during periods of enhanced summer ice melt. Previous estimates of Arctic sea ice meiofauna and under-ice amphipods on regional and pan-Arctic scales likely underestimate abundances at least in summer because they typically do not include pressure ridges.

Travel time and speed of gray whales and amphipod abundance along Chukotka Peninsula in 2006

The purpose of this study was to evaluate summer and fall residency and habitat selection by gray whales, Eschrichtius robustus, together with the biomass of benthic amphipod prey on the coastal feeding grounds along the Chukotka Peninsula. Thirteen gray whales were instrumented with satellite transmitters in September 2006 near the Chukotka Peninsula, Russia. Nine transmitters provided positions from whales for up to 81 days. The whales travelled within 5 km of the Chukotka coast for most of the period they were tracked with only occasional movements offshore. The average daily travel speeds were 23 km/day (range 9-53 km/day). Four of the whales had daily average travel speeds <1 km/day suggesting strong fidelity to the study area. The area containing 95% of the locations for individual whales during biweekly periods was on average 13,027 km**2 (range 7,097-15,896 km**2). More than 65% of all locations were in water <30 m, and between 45 and 70% of biweekly kernel home ranges were located in depths between 31 and 50 m. Benthic density of amphipods within the Bering Strait at depths <50 m was on average ~54 g wet wt/m**2 in 2006. It is likely that the abundant benthic biomass is more than sufficient forage to support the current gray whale population. The use of satellite telemetry in this study quantifies space use and movement patterns of gray whales along the Chukotka coast and identifies key feeding areas.

Stomach content and parasite characteristics of M. scorpius from Frobisher Bay, Baffin Island

Shorthorn sculpin (Myoxocephalus scorpius) from Frobisher Bay, Baffin Island, is a slow growing long-lived species. A wide range of diet items were present in the stomachs of the shorthorn sculpins sampled but 2-3 diet items (amphipod species) comprised 99.5 % of total food consumed. These amphipods were present in the stomachs in similar proportions among all age classes of shorthorn sculpin. Several new host records for parasites were reported and mean numbers of parasite species increased with shorthorn sculpin age. The increased diversity of parasite species and higher d15N values in older/larger individuals suggest that their diets were more diverse and the prey items consumed had higher d15N values. By contrast, the value of d13C in dominant diet items masked the d13C values of minor diet items. We conclude that parasites and stable isotope values provide complementary data on feeding patterns of the shorthorn sculpin. The ubiquitous marine acanthocephalan, Echinorhynchus gadi, was found at high prevalences (87-100 %) and mean intensities (28-35), and were localized in the midgut. In contrast to other studies on acanthocephalans, E. gadi did not influence fish condition as measured by condition factor, liver somatic and gonado-somatic indices.

Water chemistry, and abundance and carbon biomass of under-ice fauna during POLARSTERN cruise ARK-XIX/1 to the Storfjord area in 2003

The under-ice habitat and fauna were studied during a typical winter situation at three stations in the western Barents Sea. Dense pack ice (7-10/10) prevailed and ice thickness ranged over <0.1-1.6 m covered by <0.1-0.6 m of snow. Air temperatures ranged between -1.8 and -27.5°C. The ice undersides were level, white and smooth. Temperature and salinity profiles in the under-ice water (0-5 m depth) were not stratified (T=-1.9 to -2.0°C and S=34.2-34.7). Concentrations of inorganic nutrients were high and concentrations of algal pigments were very low (0.02 µg chlorophyll a/l), indicating the state of biological winter. Contents of particulate organic carbon and nitrogen ranged over 84.2-241.3 and 5.3-16.4 µg/l, respectively, the C/N ratio over 11.2-15.5 pointing to the dominance of detritus in the under-ice water. Abundances of amphipods at the ice underside were lower than in other seasons: 0-1.8 ind/m**2 for Apherusa glacialis, 0-0.7 ind/m**2 for Onisimus spp., and 0-0.8 ind/m**2 for Gammarus wilkitzkii. A total of 22 metazoan taxa were found in the under-ice water, with copepods as the most diverse and numerous group. Total abundances ranged over 181-2,487 ind/m**3 (biomass: 70-2,439 µg C/m**3), showing lower values than in spring, summer and autumn. The dominant species was the calanoid copepod Pseudocalanus minutus (34-1,485 ind/m**3), contributing 19-65% to total abundances, followed by copepod nauplii (85-548 ind/m**3) and the cyclopoid copepod Oithona similis (44-262 ind/m**3). Sympagic (ice-associated) organisms occurred only rarely in the under-ice water layer.

Haemolymph inorganic ion composition of crustaceans from the Southern Ocean

Brachyuran and anomuran decapod crabs do not occur in the extremely cold waters of the Antarctic continental shelf whereas caridean and other shrimp-like decapods, amphipods and isopods are highly abundant. Differing capacities for extracellular ion regulation, especially concerning magnesium, have been hypothesised to determine cold tolerance and by that the biogeography of Antarctic crustaceans. Magnesium is known to have a paralysing effect, which is even more distinct in the cold. As only few or no data exist on haemolymph ionic composition of Sub-Antarctic and Antarctic crustaceans, haemolymph samples of 12 species from these regions were analysed for the concentrations of major inorganic ions (Na+, K+, Ca2+, Mg2+, Cl-, SO4 2-) by ion chromatography. Cation relationships guaranteed neuromuscular excitability in all species. Sulphate and potassium correlated positively with magnesium concentration. The Antarctic caridean decapod as well as the amphipods maintained low (6-20% of ambient sea water magnesium concentration), Sub-Antarctic brachyuran and anomuran crabs as well as the Antarctic isopods high (54-96% of ambient sea water magnesium concentration) haemolymph magnesium levels. In conclusion, magnesium regulation may explain the biogeography of decapods, but not that of the peracarids.

Seawater carbonate chemistry and processes during experiments with amphipod Gammarus locusta, 2009

We report an investigation of the effects of increases in pCO2 on the survival, growth and molecular physiology of the neritic amphipod Gammarus locusta which has a cosmopolitan distribution in estuaries. Amphipods were reared from juvenile to mature adult in laboratory microcosms at three different levels of pH in nominal range 8.1-7.6. Growth rate was estimated from weekly measures of body length. At sexual maturity the amphipods were sacrificed and assayed for changes in the expression of genes coding for a heat shock protein (hsp70 gene) and the metabolic enzyme glyceraldehyde-3-phosphate dehydrogenase (gapdh gene). The data show that the growth and survival of this species is not significantly impacted by a decrease in sea water pH of up to 0.5 units. Quantitative real-time PCR analysis indicated that there was no significant effect of growth in acidified sea water on the sustained expression of the hsp70 gene. There was a consistent and significant increase in the expression of the gapdh gene at a pH of ~7.5 which, when combined with observations from other workers, suggests that metabolic changes may occur in response to acidification. It is concluded that sensitive assays of tissue physiology and molecular biology should be routinely employed in future studies of the impacts of sea water acidification as subtle effects on the physiology and metabolism of coastal marine species may be overlooked in conventional gross "end-point" studies of organism growth or mortality.

Studying the effect of CO2-induced acidification on sediment toxicity using acute amphipod toxicity test

Carbon capture and storage is increasingly being considered one of the most efficient approaches to mitigate the increase of CO2 in the atmosphere associated with anthropogenic emissions. However, the environmental effects of potential CO2 leaks remain largely unknown. The amphipod Ampelisca brevicornis was exposed to environmental sediments collected in different areas of the Gulf of Cádiz and subjected to several pH treatments to study the effects of CO2-induced acidification on sediment toxicity. After 10 days of exposure, the results obtained indicated that high lethal effects were associated with the lowest pH treatments, except for the Ría of Huelva sediment test. The mobility of metals from sediment to the overlying seawater was correlated to a pH decrease. The data obtained revealed that CO2-related acidification would lead to lethal effects on amphipods as well as the mobility of metals, which could increase sediment toxicity.

Respiration rate of Themisto libellula determined experimentally

Daily ingestion rates of the pelagic hyperiid amphipod Themisto libellula were studied in the marginal ice zone of the Arctic Fram Strait by feeding experiments, respiration measurements and an allometric approach based on body mass. Amphipods were collected by stratified multiple opening/closing net hauls and Rectangular Midwater Trawl (RMT 8) in August 2000 during the expedition ARK XVI/2 of R/V "Polarstern". T. libellula occurred with abundances of 0.043 and 0.015 ind/m**3 in the upper 30 m of the water column at two RMT 8 stations. Based on respiration data, the daily ingestion necessary to cover metabolic energy demands measured 1.9±0.6% of body carbon per day. Actual prey consumption during feeding experiments with Calanus copepodids as prey was very similar and accounted for 1.9±1.5%/day, indicating that feeding on Calanus can meet the energy demands of T. libellula. In general, experimental results were slightly lower than the maximum potential ingestion (2%/day for an individual of median body dry mass of 32 mg) estimated by an allometric equation based on body mass, but feeding experiments showed a strong variability. Reduced metabolism and low ingestion rates of T. libellula are consistent with low ambient temperature, large body size, slow growth and long life span of this polar species. The effect of the active pelagic life style of T. libellula on metabolism and ingestion rate is discussed in comparison to the sympagic (i.e. ice-associated) amphipod Gammarus wilkitzkii of similar body size living in the same environment. In relation to the mesozooplankton biomass in the investigation area, the predation impact by T. libellula was low. However, high-Arctic conditions also limit the secondary production of principal prey species, such as Calanus glacialis and Calanus hyperboreus, so that even low predation rates may affect the growth of prey populations.

Amphipoda assemblages in a disturbed area (Alicante, Spain, Western Mediterranean)

The distribution and composition of Amphipoda assemblages were analysed off the coasts of Alicante (Spain, Western Mediterranean), a disturbed area affected by several co-occurring anthropogenic impacts. Although differences among sampled stations were mainly related to natural parameters, anthropogenic activities were linked with changes in amphipod assemblages. Expansion of the Port of Alicante, a sewage outfall and a high salinity brine discharge could be causing the disappearance of amphipods at stations closer to these disturbances. However, the completion of port enlargement works and mitigatory dilution of the brine discharge has led to the recovery of the amphipod assemblage. Among the natural parameters, depth determines the distribution of some of the species. While Siphonoecetes sabatieri was abundant at shallow stations, Ampelisca spp., Photis longipes, Pseudolirius kroyeri, Apherusa chiereghinii and Phtisica marina were more abundant at deeper stations. Grain size and percentage of organic matter also influenced amphipod distribution, resulting in changes in species composition and in the relative percentages of different trophic groups. Species such as Ampelisca brevicornis, Perioculodes longimanus, Urothoe hesperiae and Urothoe elegans were more abundant at stations with a high content of fine sand. Carnivorous species, mainly of the Oedicerotidae family, were more abundant at those stations with a low organic matter content, while detritivorous species were more abundant at stations with a higher mud content. Among 62 identified species, three were reported for the first time from the Spanish Mediterranean coast, two species were recorded for the second time and a new species of Siphonoecetes was found, Siphonoecetes (Centraloecetes) bulborostrum. These results confirm the need for further data on amphipods from the Mediterranean Spanish coast.

Bioindicators as metrics for environmental monitoring of desalination plant discharges

Development of desalination projects requires simple methodologies and tools for cost-effective and environmentally-sensitive management. Sentinel taxa and biotic indices are easily interpreted in the perspective of environment management. Echinoderms are potential sentinel taxon to gauge the impact produced by brine discharge and the BOPA index is considered an effective tool for monitoring different types of impact. Salinity increase due to desalination brine discharge was evaluated in terms of these two indicators. They reflected the environmental impact and recovery after implementation of a mitigation measure. Echinoderms disappeared at the station closest to the discharge during the years with highest salinity and then recovered their abundance after installation of a diffuser reduced the salinity increase. In the same period, BOPA responded due to the decrease in sensitive amphipods and the increase in tolerant polychaete families when salinities rose. Although salinity changes explained most of the observed variability in both indicators, other abiotic parameters were also significant in explaining this variability.