Rapid, local adaptation of zooplankton behavior to changes in predation pressure in the absence of neutral genetic changes

Organisms producing resting stages provide unique opportunities for reconstructing the genetic history of natural populations. Diapausing seeds and eggs often are preserved in large numbers, representing entire populations captured in an evolutionary inert state for decades and even centuries. Starting from a natural resting egg bank of the waterflea Daphnia, we compare the evolutionary rates of change in an adaptive quantitative trait with those in selectively neutral DNA markers, thus effectively testing whether the observed genetic changes in the quantitative trait are driven by natural selection. The population studied experienced variable and well documented levels of fish predation over the past 30 years and shows correlated genetic changes in phototactic behavior, a predator-avoidance trait that is related to diel vertical migration. The changes mainly involve an increased plasticity response upon exposure to predator kairomone, the direction of the changes being in agreement with the hypothesis of adaptive evolution. Genetic differentiation through time was an order of magnitude higher for the studied behavioral trait than for neutral markers (DNA microsatellites), providing strong evidence that natural selection was the driving force behind the observed, rapid, evolutionary changes.

Raw data of SCADCP (self-contained Acoustic Doppler Current Profiler) from three moorings in the Eastern Weddell Sea, 2005-2008

The success of any efforts to determine the effects of climate change on marine ecosystems depends on understanding in the first instance the natural variations, which contemporarily occur on the interannual and shorter time scales. Here we present results on the environmental controls of zooplankton distribution patterns and behaviour in the eastern Weddell Sea, Southern Ocean. Zooplankton abundance and vertical migration are derived from the mean volume backscattering strength (MVBS) and the vertical velocity measured by moored acoustic Doppler current profilers (ADCPs), which were deployed simultaneously at 64°S, 66.5°S and 69°S along the Greenwich Meridian from February, 2005, until March, 2008. While these time series span a period of full three years they resolve hourly changes. A highly persistent behavioural pattern found at all three mooring locations is the synchronous diel vertical migration (DVM) of two distinct groups of zooplankton that migrate between a deep residence depth during daytime and a shallow depth during nighttime. The DVM was closely coupled to the astronomical daylight cycles. However, while the DVM was symmetric around local noon, the annual modulation of the DVM was clearly asymmetric around winter solstice or summer solstice, respectively, at all three mooring sites. DVM at our observation sites persisted throughout winter, even at the highest latitude exposed to the polar night. Since the magnitude as well as the relative rate of change of illumination is minimal at this time, we propose that the ultimate causes of DVM separated from the light-mediated proximal cue that coordinates it. In all three years, a marked change in the migration behaviour occurred in late spring (late October/early November), when DVM ceased. The complete suspension of DVM after early November is possibly caused by the combination of two factors: (1) increased availability of food in the surface mixed layer provided by the phytoplankton spring bloom, and (2) vanishing diurnal enhancement of the threat from visually oriented predators when the illumination is quasi-continuous during the polar and subpolar summer. Zooplankton abundance in the water column, estimated as the mean MVBS in the depth range 50-300 m, was highest end of summer and lowest mid to end winter on the average annual cycle. However, zooplankton abundance varied several-fold between years and between locations. Based on satellite and in situ data of chlorophyll and sea ice as well as on hydrographic measurements, the interannual and spatial variations of zooplankton mean abundance can be explained by differences in the magnitude of the phytoplankton spring bloom, which develops during the seasonal sea ice retreat. Whereas the vernal ice melt appears necessary to stimulate the blooming of phytoplankton, it is not the determinator of the blooms magnitude, its areal extent and duration. A possible explanation for the limitation of the phytoplankton bloom in some years is top-down control. We hypothesise that the phytoplankton spring development can be curbed by grazing when the zooplankton had attained high abundance by growth during the preceding summer.

Thorium-234, POC and PON fluxes during POLARSTERN cruise ANT-XXVIII/3, January-March 2012

Carbon fixation by phytoplankton plays a key role in the uptake of atmospheric CO2 in the Southern Ocean. Yet, it still remains unclear how efficiently the particulate organic carbon (POC) is exported and transferred from ocean surface waters to depth during phytoplankton blooms. In addition, little is known about the processes that control the flux attenuation within the upper twilight zone. Here, we present results of downward POC and particulate organic nitrogen fluxes during the decline of a vast diatom bloom in the Atlantic sector of the Southern Ocean in summer 2012. We used thorium-234 (234Th) as a particle tracer in combination with drifting sediment traps (ST). Their simultaneous use evidenced a sustained high export rate of 234Th at 100 m depth in the weeks prior to and during the sampling period. The entire study area, of approximately 8000 km**2, showed similar vertical export fluxes in spite of the heterogeneity in phytoplankton standing stocks and productivity, indicating a decoupling between production and export. The POC fluxes at 100 m were high, averaging 26 ± 15 mmol C/m**2/d, although the strength of the biological pump was generally low. Only <20% of the daily primary production reached 100 m, presumably due to an active recycling of carbon and nutrients. Pigment analyses indicated that direct sinking of diatoms likely caused the high POC transfer efficiencies (~60%) observed between 100 and 300 m, although faecal pellets and transport of POC linked to zooplankton vertical migration might have also contributed to downward fluxes.

Effects of physical disturbance on infaunal and epifaunal assemblages in subtropical, intertidal seagrass beds

We assessed the impact of large-scale commercial and recreational harvesting of polychaete worms Marphysa spp. on macrobenthic assemblages in a subtropical estuary in Queensland, Australia, by examining: (1) the spatial extent of harvesting activities and the rate of recovery of the seagrass habitat over an 18 to 20 mo period; (2) the recovery of infauna in and around commercial pits of known age; (3) the indirect effects of physical disturbance from trampling and deposition of sediments during harvesting on epibenthos in areas adjacent to commercial and recreational pits; (4) impacts of potential indirect effects through manipulative experimentation. Harvesting caused a loss of seagrass, changes to the topography and compaction of the sediments associated with the creation of walls around commercial pits, and the deposition of rubble dug from within the pit. The walls and rubble were still evident after 1.8 to 20 mo, but comprised only a small proportion of the total area on the intertidal banks. There was a shift from an intertidal area dominated by Zostera capricorni to one with a mixture of Z. capricorni, Halophila spp. and Halodule uninervis, but there was no overall decline in the biomass of seagrass in these areas. There were distinct impacts from harvesting on the abundance of benthic infauna, especially amphipods, polychaetes and gastropods, and these effects were still detectable after 4 mo of potential recovery. After 12 me, there were no detectable differences in the abundances of these infauna between dug areas and reference areas, which suggested that infauna had recovered from impacts of harvesting; however, an extensive bloom of toxic fireweed Lyngbya majsucula may have masked any remaining impacts. There were no detectable impacts of harvesting on epifauna living in the seagrass immediately around commercial or recreational pits.

Robust spatially resolved pressure measurements using MRI with novel buoyant advection-free preparations of stable microbubbles in polysaccharide gels

MRI of fluids containing lipid coated microbubbles has been shown to be an effective toot for measuring the local fluid pressure. However, the intrinsically buoyant nature of these microbubbles precludes lengthy measurements due to their vertical migration under gravity and pressure-induced coalescence. A novel preparation is presented which is shown to minimize both these effects for at least 25 min. By using a 2% polysaccharide gel base with a small concentration of glycerol and 1,2-distearoyl-sn-glycero-3-phosphocholine coated gas microbubbles, MR measurements are made for pressures between 0.95 and 1.44 bar. The signal drifts due to migration and amalgamation are shown to be minimized for such an experiment whilst yielding very high NMR sensitivities up to 38% signal change per bar.

C1-C8 hydrocarbons in marine sediments

Light hydrocarbon (C1-C8) profiles are compared for three wells of varying maturities: two immature DSDP wells (Site 397 near the Canary Islands and Site 530A near the Walvis Ridge in the south-east Atlantic) and a mature well, the East Cameron well in the Texas Gulf Coast. Primary migration of C1 and C2 appears to be occurring in all of the sedimentary rocks examined. Primary migration of C3+ components becomes important only as fine-grained sedimentary rocks enter the catagenetic hydrocarbon generation zone or over short distances in more permeable sections. Lateral migration along bedding planes was more important than vertical migration in sedimentary rocks of all maturities. The lightest (methane, ethane and propane gases) hydrocarbon show greater fractionation than do the C4-C8 alkanes which generally show minimal fractionation during the migrational process. Subsurface diffusion coefficients for these p.p.b. quantities of C2-C5 alkanes from immature sediments from DSDP Site 530 are estimated to be several orders of magnitude less than values reported in the literature for diffusion of much larger amounts of these compounds from mature water wet sediments into air or sandstones. Since our calculations suggest light hydrocarbons are present in amounts less than their reported solubilities in pure water at 25°C, we postulate that the sediment organic matter has a substantial effect on retarding the movement of these light hydrocarbons.

Meiofauna and the structure of the benthic community in the Iberian deep sea basin

1. On the cruises 3 and 15 of R.V. "Meteor" 6 grab samples, and 6 hauls with the 6 m Agassiztrawl were taken and at 2 stations the deep sea camera was lowered. This material gave quantitative results on the meiofauna and minimum counts of the macrofauna. 2. The nematodes constitute nearly 95% of the meiofauna, the copepoda only 2%. With increasing sediment depth the density of animals decrease gradually. In the uppermost centimeter of sediment 42.6% of the meiofauna are found while only 3.7% live in layer 6-7 cm. Meiofauna weight ranges from 0.6-5.7 mg/25 m**2 surface i.e. 0.24-2.8 g/m**2. 3. Mean numbers of individuals and weights show standard errors of 20-30 %. As an approximate average values for further considerations the weight of the meiofauna in the area was taken as 1 g/m**2 4. Quantitative information on the macrofauna is derived from the trawls and the photographs for the actinia Chitonanthus abyssorum only, which is found in the rate of 1 individual/36-72 m**2, but seems to be less abundant generally. 5. Animal density does not decrease steadily from nearshore to offshore biocoenoses, i.e. generally with increasing depth. The decrease is more pronounced for macro- than for meiofauna. For the deep sea the weight proportion of macrofauna : meiofauna is of the order of 1 : 1. 6. With the assumption, that adaptation of metabolism to deep sea conditions is similar in macro- and meiofauna total metabolism of invertebrates is ascribed to meiofauna to more than 80%. 7. The structure of the biocoenosis of the deep sea floor is characterized by the meiofauna living on and in the sediment and by the dominance of sediment feeders in the macrofauna. 8. Considering the large numbets and high partition rates of bacteria a comparative large part of the metabolism in the deep sea sediment must be ascribed to bacteria. This favours the hypothesis, that with increasing depth and decreasing addition of organic material to the sediment, the importance of meiofauna and microorganisms for total metabolism increases. 9. Considering the different modes of food transport to the deep sea environment, i.e. sinking of dead particles, transport by vertical migration of organisms, aggregation of organic particles, adsorption of dissoloved organic substance to inorganic particles, and heterotrophy, the sediment may be assumed to contain more food for invertebrates than the water above the bottom. 10. Suspensions feeders of macrofauna are fixed to hard substrates in the sediment surface. Some of them are shown to bend themselves down to the bottom in underwater photographs. This suggests the idea that some deep sea suspension feeders partly depend on food from the sediment surface, on which they feed directly.

Physical oceanography from CTD during Maria S. Merian cruise MSM20/4 in spring 2012

With an extension of > 40 km**2 the recently discovered Campeche cold-water coral province located at the northeastern rim of the Campeche Bank in the southern Gulf of Mexico belongs to the largest coherent cold-water coral areas discovered so far. The Campeche province consists of numerous 20-40 m-high elongated coral mounds that are developed in intermediate water depths of 500 to 600 m. The mounds are colonized by a vivid cold-water coral ecosystem that covers the upper flanks and summits. The rich coral community is dominated by the framework-building Scleractinia Enallopsammia profunda and Lophelia pertusa, while the associated benthic megafauna shows a rather scarce occurrence. The recent environmental setting is characterized by a high surface water production caused by a local upwelling center and a dynamic bottom-water regime comprising vigorous bottom currents, obvious temporal variability, and strong density contrasts, which all together provide optimal conditions for the growth of cold-water corals. This setting - potentially supported by the diel vertical migration of zooplankton in the Campeche area - controls the delivering of food particles to the corals. The Campeche cold-water coral province is, thus, an excellent example highlighting the importance of the oceanographic setting in securing the food supply for the development of large and vivid cold-water coral ecosystems.

Raw data of SCADCP (self-contained Acoustic Doppler Current Profiler) from three moorings in the Eastern Weddell Sea in 2005

Ten-month time series of mean volume backscattering strength (MVBS) and vertical velocity obtained from three moored acoustic Doppler current profilers (ADCPs) deployed from February until December 2005 at 64°S, 66.5°S and 69°S along the Greenwich Meridian were used to analyse the diel vertical zooplankton migration (DVM) and its seasonality and regional variability in the Lazarev Sea. The estimated MVBS exhibited distinct patterns of DVM at all three mooring sites. Between February and October, the timing of the DVM and the residence time of zooplankton at depth were clearly governed by the day-night rhythm. Mean daily cycles of the ADCP-derived vertical velocity were calculated for successive months and showed maximum ascent and descent velocities of 16 and -15 mm/s. However, a change of the MVBS pattern occurred in late spring/early austral summer (October/November), when the zooplankton communities ceased their synchronous vertical migration at all three mooring sites. Elevated MVBS values were then concentrated in the uppermost layers (<50 m) at 66.5°S. This period coincided with the decay of sea ice coverage at 64°S and 66.5°S between early November and mid-December. Elevated chlorophyll concentrations, which were measured at the end of the deployment, extended from 67°S to 65°S and indicated a phytoplankton bloom in the upper 50 m. Thus, we propose that the increased food supply associated with an ice edge bloom caused the zooplankton communities to cease their DVM in favour of feeding.

Particle fluxes of the bathypelagic sediment trap CVOO-3 in the eastern tropical North Atlantic for the period December 2009 until May 2011

Particle fluxes at the Cape Verde Ocean Observatory (CVOO) in the eastern tropical North Atlantic for the period December 2009 until May 2011 are discussed based on bathypelagic sediment trap time-series data collected at 1290 and 3439 m water depth. The typically oligotrophic particle flux pattern with weak seasonality is modified by the appearance of a highly productive and low oxygen (minimum concentration below 2 µmol kg**-1 at 40 m depth) anticyclonic modewater eddy (ACME) in winter 2010. The eddy passage was accompanied by unusually high mass fluxes of up to 151 mg m**-2 d**-1, lasting from December 2009 to May 2010. Distinct biogenic silica (BSi) and organic carbon flux peaks of ~15 and 13.3 mg m**-2 d**-1, respectively, were observed in February-March 2010 when the eddy approached the CVOO. The flux of the lithogenic component, mostly mineral dust, was well correlated with that of organic carbon, in particular in the deep trap samples, suggesting a tight coupling. The lithogenic ballasting obviously resulted in high particle settling rates and, thus, a fast transfer of epi-/meso-pelagic signatures to the bathypelagic traps. We suspect that the two- to three-fold increase in particle fluxes with depth as well as the tight coupling of mineral dust and organic carbon in the deep trap samples might be explained by particle focusing processes within the deeper part of the eddy. Molar C : N ratios of organic matter during the ACME passage were around 18 and 25 for the upper and lower trap samples, respectively. This suggests that some productivity under nutrient (nitrate) limitation occurred in the euphotic zone of the eddy in the beginning of 2010 or that a local nitrogen recycling took place. The d15N record showed a decrease from 5.21 to 3.11 per mil from January to March 2010, while the organic carbon and nitrogen fluxes increased. The causes of enhanced sedimentation from the eddy in February/March 2010 remain elusive, but nutrient depletion and/or an increased availability of dust as a ballast mineral for organic-rich aggregates might have contributed. Rapid remineralisation of sinking organic-rich particles could have contributed to oxygen depletion at shallow depth. Although the eddy formed in the West African coastal area in summer 2009, no indications of coastal flux signatures (e.g. from diatoms) were found in the sediment trap samples, confirming the assumption that the suboxia developed within the eddy en route. However, we could not detect biomarkers indicative of the presence of anammox (anaerobic ammonia oxidation) bacteria or green sulfur bacteria thriving in photic zone suboxia/hypoxia, i.e. ladderane fatty acids and isorenieratene derivatives, respectively. This could indicate that suboxic conditions in the eddy had recently developed and/or the respective bacterial stocks had not yet reached detection thresholds. Another explanation is that the fast-sinking organic-rich particles produced in the surface layer did not interact with bacteria from the suboxic zone below. Carbonate fluxes dropped from -52 to 21.4 mg m**-2 d**-1 from January to February 2010, respectively, mainly due to reduced contribution of shallow-dwelling planktonic foraminifera and pteropods. The deep-dwelling foraminifera Globorotalia menardii, however, showed a major flux peak in February 2010, most probably due to the suboxia/hypoxia. The low oxygen conditions forced at least some zooplankton to reduce diel vertical migration. Reduced "flux feeding" by zooplankton in the epipelagic could have contributed to the enhanced fluxes of organic materials to the bathypelagic traps during the eddy passage. Further studies are required on eddy-induced particle production and preservation processes and particle focusing.

Winter distribution and size structure of Antarctic krill Euphausia superba populations in-shore along the West Antarctic Peninsula

Antarctic krill Euphausia superba are a key component of food webs in the maritime West Antarctic Peninsula, and their life history is tied to the seasonal cycles of sea ice and primary production in the region. Previous work has shown a general in-shore migration of krill in winter in this region; however, the very near-shore has not often been sampled as part of these surveys. We investigated distribution, abundance, and size structure of krill in 3 fjordic bays along the peninsula, and in the adjacent Gerlache Strait area using vertically stratified MOCNESS net tows and ADCP acoustic biomass estimates. Krill abundance was high within bays, with net estimated densities exceeding 60 krill m-3, while acoustic estimates were an order of magnitude higher. Krill within bays were larger than krill in the Gerlache Strait. Within bays, krill aggregations were observed near the seafloor during the day with aggregations extending to the sediment interface, and exhibited diel vertical migration higher into the water column at night. We suggest these high winter krill abundances within fjords are indicative of an active seasonal migration by krill in the peninsula region. Potential drivers for such a migration include reduced advective losses and costs, and availability of sediment food resources within fjords. Seasonally near-shore krill may also affect stock and recruitment assessments and may have implications for managing the krill fishery in this area.

Winter distribution and size structure of Antarctic krill Euphausia superba populations in-shore along the West Antarctic Peninsula

Antarctic krill Euphausia superba are a key component of food webs in the maritime West Antarctic Peninsula, and their life history is tied to the seasonal cycles of sea ice and primary production in the region. Previous work has shown a general in-shore migration of krill in winter in this region; however, the very near-shore has not often been sampled as part of these surveys. We investigated distribution, abundance, and size structure of krill in 3 fjordic bays along the peninsula, and in the adjacent Gerlache Strait area using vertically stratified MOCNESS net tows and ADCP acoustic biomass estimates. Krill abundance was high within bays, with net estimated densities exceeding 60 krill m-3, while acoustic estimates were an order of magnitude higher. Krill within bays were larger than krill in the Gerlache Strait. Within bays, krill aggregations were observed near the seafloor during the day with aggregations extending to the sediment interface, and exhibited diel vertical migration higher into the water column at night. We suggest these high winter krill abundances within fjords are indicative of an active seasonal migration by krill in the peninsula region. Potential drivers for such a migration include reduced advective losses and costs, and availability of sediment food resources within fjords. Seasonally near-shore krill may also affect stock and recruitment assessments and may have implications for managing the krill fishery in this area.