Andreas Reul

Physical–biological coupling in the Strait of Gibraltar

This study presents a joint analysis of the distributions of some biogeochemical variables and their relation to the hydrodynamics of Gibraltar Strait. It is a synthesis paper that brings together many results obtained during CANIGO project. We show the role of hydrodynamics as a forcing agent for the plankton community structure in the Strait, with emphasis on the two physical processes that we propose as key factors for the coupling: interface position and oscillations, and mixing processes along the Strait. As a general pattern, autotrophic plankton biomass increases at the Strait from southwest to northeast, a tendency that coincides with a gradual elevation of the interface depth in the same direction. The different mechanisms of mixing that take place in the Strait are briefly reviewed: The occurrence of the internal hydraulic jump is an important mechanism of mixing constrained to the spring tide situations, but other processes such as the generation of arrested internal waves of wavelength around 1 km are proposed as a complementary mixing mechanism, particularly during neap tides situations. Both mechanisms, the elevation of the pycnocline and these mixing events, can enhance biological productivity and biomass accumulation on the northeastern sector of the Strait, since phytoplankton cells are there packaged in a water mass with sufficient light and nutrients and smaller advective velocity. There is a clear north–south difference in the biological response to these upwelling episodes in the eastern section, with high nutrient and low chlorophyll in the south and the opposite in the north. The deeper interface and the greater water speed are the proposed reasons for this lower nutrient uptake on the southeastern sector. Finally, the temporal scales of variation of the mixing events, the influence of its periodicity on the productivity of the area and the influence of these upwelling episodes in the nearest Albor! an Sea are discussed. r 2002 Elsevier Science

Surface distribution of chlorophyll, particles and gelbstoff in the Atlantic jet of the Alboran Sea: from submesoscale to subinertial scales of variability

The surface distribution of light attenuation due to particles (c) as well as chlorophyll-a and gelbstoff fluorescence (Fch and Fcd, respectively) were recorded during an OMEGA (EU funded, MAST III project) cruise in the northwestern Alboran Sea through a high spatial (zonally separated by 10 km and virtually meridionally continuous) and temporal (about 3 days between each of the three repeated surveys made in the zone) resolution sampling design. The distributions obtained for these variables were tightly linked to the physical forcing at the different scales that the sampling design was able to resolve. Low values dominate the quasi permanent anticyclonic gyre occupying the western Alboran Sea, whereas the frontal zone directly affected by the entrance of the Atlantic jet depicts much higher records for c, Fch and Fcd. High geostrophic Froude numbers in the jet, and the subsequent increase in turbulence diffusion of nutrients towards the surface, cannot alone justify this spatial distribution. Instead, high phytoplankton concentration at the jet could also result from the entrainment and advection of water from the upwelling zone at the Spanish coast. However, T–S characteristics suggest that this is neither the most important process for the biological enrichment of the jet, so that other mechanisms such as vertical ageostrophic velocities at the edge of the gyre must also be considered. Due to the time needed for phytoplankton growth, the intense horizontal velocities associated to the jet can decouple the sectors where deep nutrient-rich waters reach the surface from sectors where high values of the recorded variables are observed. The decoupling hinders a differentiation of this fertilization mechanism from other possible alternatives as mixing at the sills in the Strait of Gibraltar. In the third survey, the spatial structure of surface warm waters in the gyre and cold waters in the front became less apparent. ADCP data show a southward migration of the jet in a fluctuation probably related to transient states in the Atlantic jet and western Alboran gyre system. The qualitative response of c, Fch and Fcd to these scales of variability was very similar and close to the changes observed in temperature. However, the values of Fcd varied in a much narrower range than c or Fch (a factor of 2 and 10, respectively), which indicates a distinct control for the abundance of Gelbstoff. This control dumps the range of variability in the western Alboran and its origin is discussed in the context of photobleaching or bacterial degradation of these substances.

Exchange of planktonic biomass through the Strait of Gibraltar in late summer conditions

In order to estimate plankton biomass transport through the Gibraltar Strait, plankton biomass and velocity profiles were measured at three stations located in the eastern side of the Strait as a part of the CANIGO project. Abundance and biomass measurements were carried out for autotrophic (Prochlorococcus, Synechococcus and eukaryotic piconano- and microplankton) and heterotrophic (bacteria and nanoflagellates) organisms, in September 1997. Biomass and velocity decreased from the surface to deeper water. Highest biomass concentration was observed at the northern station ð0:12 g C m � 3 ), whereas maximum mean velocities ð80 cm s � 1 ) were found at the central and southern stations. Biomass transport is estimated with a approach with a 10 -m resolution in the vertical and three subareas of approximately 5 km in the horizontal direction. Estimate of plankton biomass transports towards the Mediterranean and the Atlantic are 5570 and 1140 tonnes C day � 1 ; respectively. The former is co-dominated by heterotrophic bacteria (37%) and autotrophic nanoplankton (42%), while the latter is dominated by heterotrophic organisms like bacteria (75%) and heterotrophic nanoflagellates (14%). The variation during a one-day period of the biomass transport estimate at the central part of the Strait was explored. Also, in order to estimate the influence of spatial distribution of both biomass and velocity in the transport estimates, a comparison of our results with other possible estimates performed with less spatial resolution is carried out. The results confirms that both temporal and spatial resolution are key factors for transport estimates of inhomogeneous distributed variables through the Strait. r 2002 Elsevier Science Ltd. All rights reserved.

Effect of Drought Conditions on Plankton Community and on Nutrient Availability in an Oligotrophic High Mountain Lake

Abstract Natural water level fluctuations (WLFs) are an inherent characteristic of Mediterranean inland waters, which are projected to be amplified by global climate change. La Caldera (Sierra Nevada National Park, Spain) is an oligotrophic high mountain lake (3050 m a.s.l.) that has experienced large fluctuations in water volume (13–100%) during the past 20 years due to irregular annual precipitation patterns (371–1816 mm). Because of the lakes cold and dilute abiotic environment, it is likely susceptible to projected increases in global temperature and represents an ideal sentinel of global change. We analyze the effect of WLFs on water quality and on plankton community in La Caldera to better understand the potential effects of recurrent droughts (3 droughts in a 20-year period) on lake ecology. We have found significantly positive effects of WLFs on total phosphorus (TP) concentrations. There was extreme variability in TP concentrations during three recurrent droughts (1995, 1999, and 2005) reflecting sediment resuspension. However, the data also suggest that this was not the only source of phosphorus. Extremely high P-enriched atmospheric dust inputs could have maintained the abnormally high TP in-lake concentrations measured during 2005. The data indicate that recurrent droughts have reduced lake resistance to TP changes but have increased lake resistance to total nitrogen (TN) changes, which supports the idea that a P-enriched atmospheric dust inputs during 2005. An increase in dissolved inorganic nitrogen (DIN)∶TP mass ratio after 2005 was observed, revealing a higher ecosystem homeostasis of this ratio.

Fertilization and connectivity in the Garrucha Canyon (SE-Spain) implications for Marine Spatial Planning

Marine Spatial Planning is usually based on benthic georeferenced information or GPS tracked human activities, whereas the pelagic ecosystem is often ignored because of scarce and limited surface information. However, the 3-D pelagic ecosystem plays a key role connecting all the other ecosystems by physical (currents) and biological (migration) processes. According to remote sensing the Garrucha Canyon is oligotrophic, but 3-D sampling reveals subsurface upwelling, and converts it into the richest area around the Cape of Gata. Vertical connectivity by means of zooplankton migration, measured at two sampling stations, is 40 and 220 times faster than microphytoplankton settling and vertical water velocities respectively. Thus coupled physical-biological connectivity models are necessary to estimate the ecosystem connection and the fate of carbon, but also other substances (e.g. radioactivity), that might accumulate throughout the food-web. This is especially important in the Garrucha Canyon and the Coastal Areas Management Programme Levante de Almería where natural heritage and extractive fishery are important for the local economy.

North Atlantic Oscillation drives the annual occurrence of an isolated, peripheral population of the brown seaweed Fucus guiryi in the Western Mediterranean Sea

The canopy-forming, intertidal brown (Phaeophyceae) seaweed Fucus guiryi is distributed along the cold-temperate and warm-temperate coasts of Europe and North Africa. Curiously, an isolated population develops at Punta Calaburras (Alboran Sea, Western Mediterranean) but thalli are not present in midsummer every year, unlike the closest (ca. 80 km), perennial populations at the Strait of Gibraltar. The persistence of the alga at Punta Calaburras could be due to the growth of resilient, microscopic stages as well as the arrival of few–celled stages originating from neighbouring localities, and transported by the permanent Atlantic Jet flowing from the Atlantic Ocean into the Mediterranean. A twenty-six year time series (from 1990 to 2015) of midsummer occurrence of F. guiryi thalli at Punta Calaburras has been analysed by correlating with oceanographic (sea surface temperature, an estimator of the Atlantic Jet power) and climatic factors (air temperature, rainfall, and North Atlantic Oscillation –NAO-, and Arctic Oscillation –AO- indexes). The midsummer occurrence of thalli clustered from 1990–1994 and 1999–2004, with sporadic occurrences in 2006 and 2011. Binary logistic regression showed that the occurrence of thalli at Punta Calaburras in midsummer is favoured under positive NAO index from April to June. It has been hypothesized that isolated population of F. guiryi should show greater stress than their congeners of permanent populations, and to this end, two approaches were used to evaluate stress: one based on the integrated response during ontogeny (developmental instability, based on measurements of the fractal branching pattern of algal thalli) and another based on the photosynthetic response. Although significant differences were detected in photosynthetic quantum yield and water loss under emersion conditions, with thalli from Punta Calaburras being more affected by emersion than those from Tarifa, the developmental instability showed that the population from Tarifa suffers higher stress during ontogeny than that from Punta Calaburras. In conclusion, this study demonstrates the teleconnection between atmospheric oscillations and survival and proliferation of marine macroalgae.

Sulphide Resistance in the Cyanobacterium Microcystis aeruginosa: a Comparative Study of Morphology and Photosynthetic Performance Between the Sulphide-Resistant Mutant and the Wild-Type Strain.

The cyanobacterium Microcystis aeruginosa is a mesophilic freshwater organism, which cannot tolerate sulphide. However, it was possible to isolate a sulphide-resistant (Sr) mutant strain that was able to survive in a normally lethal medium sulphide. In order to evaluate the cost of the mutation conferring sulphide resistance in the Sr strain of M. aeruginosa, the morphology and the photosynthetic performance were compared to that found in the wild-type, sulphide-sensitive (Ss) strain. An increase in size and a disrupted morphology was observed in Sr cells in comparison to the Ss counterpart. Phycoerythrin and phycocyanin levels were higher in the Sr than in the Ss cells, whereas a higher carotenoid content, per unit volume, was found in the Ss strain. The irradiance-saturated photosynthetic oxygen-production rate (GPRmax) and the photosynthetic efficiency (measured both by oxygen production and fluorescence, αGPR and αETR) were lower in the Sr strain than in the wild-type. These results appear to be the result of package effect. On the other hand, the Sr strain showed higher quantum yield of non-photochemical quenching, especially those regulated mechanisms (estimated throughout qN and Y(NPQ)) and a significantly lower slope in the maximum quantum yield of light-adapted samples (Fv′/Fm′) compared to the Ss strain. These findings point to a change in the regulation of the quenching of the transition states (qT) in the Sr strain which may be generated by a change in the distribution of thylakoidal membranes, which somehow could protect metalloenzymes of the electron transport chain from the lethal effect of sulphide.