Carlos M. García

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

Morphological and physiological differences between two morphotypes of Zostera noltii Hornem. from the south-western Iberian Peninsula

Abstract The morphological and physiological differences between two morphotypes of Z. noltii Hornem. were studied in the intertidal meadows on the south-western Iberian Peninsula (Palmones river estuary and Ria Formosa). A small-leaved morphotype (SM) grows mainly at high intertidal sites, meadow edges or in recently deposited sandbanks, whereas a large-leaved morphotype (LM) generally thrives in well-structured beds or in deeper places. This study deals with the morphological, biochemical and physiological differences between these morphotypes as well as the ecological implications of the occurrence of different morphotypes in the same meadow. Shoot length, leaf width, rhizome internode length, roots per node, root length, leaf nutrient and pigment contents, and photosynthetic rates of both morphotypes were compared. The below-ground architecture (root and rhizome complex) of both morphotypes was more developed in sites characterized by higher hydrodynamics and/or a lower nitrogen content in sediments. Both morphotypes showed similar values for photosynthetic efficiency, dark respiration rate and compensation irradiance. On the other hand, the net photosynthetic capacity was much greater (5-fold) for the SM. This difference could explain the greater growth rate and faster leaf turnover rate of the SM compared with the LM. The occurrence of the SM in newly settled areas (and in the meadow edges) could be explained on the basis of its higher growth rate, which would allow a faster spreading of the meadow and/or better recovery after burial resulting from stormy weathers.

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.

Hydrological cycle and interannual variability of the aquatic community in a temporary saline lake (Fuente de Piedra, Southern Spain)

Fuente de Piedra is a shallow, temporary saline lake whoseseasonal behavior is strongly dependent on the annual hydrologicalbudget. In this study, we outline the characteristics of Fuente dePiedra Lake for two years that had different hydrological budgets.The high precipitations in 1989–90 caused the lake not to dry asusual, and decreased both salinity and the amplitude of changes.There were also differences in nutrient dynamics, with generallylower concentrations of soluble reactive phosphorus and ammonium,whereas in the more humid year nitrate showed a distinct maximum inwinter. Winter bloom chlorophyll a concentrations were alsomuch higher in 1989–90 (>600 μg l-1) but there wasalso a winter productive phase that was presumably poorly coupledwith consumption processes that predominate in spring. Planktonicassemblages were different between years. Highly halotolerantphytoplankton species (Dunaliella salina and D. viridis) became scarcer, and especially two previouslyunrecorded diatoms (Cyclotella sp. and Chaetoceros sp.)became dominant in the bloom time in the wet year. The speciesrichness of the zooplankton increased in the wet year, with newspecies appearing that were not collected during 1987–88(Branchinectella media, Daphnia mediterranea, Macrothrix sp.,Arctodiaptomus salinus, Cyclops sp., Sigara sp...).There was also a much higher development of macrophytes (Ruppiadrepanensis, Althenia orientalis, Lamprothamnium papulosum)and bird populations, especially flamingoes (Phoenicopterusruber).Important interannual variations in this sort of system pointto the need for long term studies to observe the whole range ofstates that define the lake as an entity.

Phytoplankton pigment distribution in the northwestern Alboran Sea and meteorological forcing: A remote sensing study

A set of weekly-composite SeaWiFS (Sea-viewing Wide Field of view Sensor) imagery (from 1998 to 2004) has been used to asses the temporal and spatial variability of the surface chlorophyll distribution in the Northwestern Alboran Sea. Empirical Orthogonal Function (EOF) analysis of the images series shows the existence of three different regions in which surface chlorophyll presents a homogeneous temporal behavior in the neighborhood of the Strait of Gibraltar. The temporal patterns of surface chlorophyll concentrations in each region have been characterized by studying the role of several forcing factors, mainly the zonal component of the wind field. The first identified region occupies the coastal area in front of Estepona and Malaga and shows the highest chlorophyll concentration in the entire region of study in absence of easterly winds. A second region is located far offshore, in the zone usually occupied by the Atlantic jet entering the Alboran Sea, showing high concentrations of chlorophyll when the easterlies blow. Finally, a third region has been situated in the normal location of the Western Alboran Gyre (WAG). The zonal wind field arises as a main forcing function in the modulation of the intensity and location of the upwelling processes in the area, while the entrance of the Atlantic jet seems to be the main agent feeding of all these processes.

Seasonal change in a saline temporary lake (Fuente de Piedra, southern Spain)

Fuente de Piedra saline lake is located in an endorheic basin in the south of Spain. This lake is very shallow (0.5 m max. depth during 1987–88) and relatively large (± 1350 ha). It is a temporary playa lake, showing irregular cycles, with frequent seasonal drought and a high degree of unpredictability. The lake was sampled monthly during a relatively rainy year (1987–88, 10.5 months permanence). The result of combined analyses for environmental variables (salinity, temperature and soluble inorganic forms of nitrogen and phosphorus), variables related to biological activity (chlorophyll ‘a’, sediment organic matter and redox potential) and the direct analysis of the planktonic community, shows the existence of two periods of dominance by autotrophs. The first occurs during winter, exhibits a progressively higher surface to volume ratio for phytoplankton and is followed in the spring by high zooplankton densities (Moina salina, Fabrea salina) and very low phytoplankton densities, suggesting the existence of a period with a detritus-based food web. The summer period coincides with a community better adapted to high salinities that is dominated by Dunaliella salina, D. viridis, diatoms and the ciliate Fabrea salina, and associated with high ammonium concentrations. A new period of organic matter accumulation could be facilitated, in the last moments before the lake dries, by a progressive decrease in zooplankton abundance.

Vertical patterns of phytoplankton size distribution in the Cantabric and Balearic Seas

Abstract In this paper we analyze plankton size distribution (2–100 μm of equivalent spherical diameter) under different hydrographic structures, such as the seasonal thermocline of the Cantabric and Balearic seas and the oceanographic front present in the Balearic Sea. Nannoplanktonic fraction (2–20 μm) tends to present biomass maxima at shallower depths than the chlorophyll maximum in stratified waters, as well as in the more productive waters near a front. The slope of the normalized size-biomass spectrum was more negative in these zones of phytoplankton biomass maxima indicating a higher proportion of smaller cells. The main hypotheses to explain chlorophyll and biomass maxima as well as phytoplankton accumulation in productive areas are reviewed. According to the observed results, the most suitable mechanism to explain subsurface biomass maxima is the active net growth of the smaller phytoplanktonic cells. The importance of the methodology employed for analyzing the size distribution of plankton communities and its relationship with hydrographic variability are also discussed.

Effect of nutrient input frequency on the structure and dynamics of the marine pelagic community: A modeling approach

An analysis of the induced changes in both total productivity and community structure as a consequence of different pulsed nutrient inputs has been made by using two different ecosystem modeling tools. One was a modified version of the N-based model proposed by Fasham et al. (1990) for pelagic ecosystems and the other the plankton functional type model by Vichi et al. (2007). Both models lead to higher total biomass production with a pulsed nutrient input compared to a continuous supply, affecting both phytoplankton and zooplankton. Detailed responses are, however, different among plankton groups depending on the frequency of pulsed nutrient input and the complexity of the model. The differences in biomass production were higher for an oscillation period of 150–170 days following a sinusoidal supply pattern. A tidal mixing-induced nutrient supply was also simulated by using both models and a larger biomass increment was observed compared to that obtained with the sinusoidal pattern. Finally, a theoretical application of this approach to the pelagic ecosystem of the North Western Alboran Sea shows a preferential selection of diatoms and larger zooplankton under discontinuous nutrient inputs.

Potential polyunsaturated aldehydes in the Strait of Gibraltar under two tidal regimes.

Diatoms, a major component of the large-sized phytoplankton, are able to produce and release polyunsaturated aldehydes after cell disruption (potential PUAs or pPUA). These organisms are dominant in the large phytoplankton fraction (>10 µm) in the Strait of Gibraltar, the only connection between the Mediterranean Sea and the Atlantic Ocean. In this area, the hydrodynamics exerts a strong control on the composition and physiological state of the phytoplankton. This environment offers a great opportunity to analyze and compare the little known distribution of larger sized PUA producers in nature and, moreover, to study how environmental variables could affect the ranges and potential distribution of these compounds. Our results showed that, at both tidal regimes studied (Spring and Neap tides), diatoms in the Strait of Gibraltar are able to produce three aldehydes: Heptadienal, Octadienal and Decadienal, with a significant dominance of Decadienal production. The PUA released by mechanical cell disruption of large-sized collected cells (pPUA) ranged from 0.01 to 12.3 pmol from cells in 1 L, and from 0.1 to 9.8 fmol cell−1. Tidal regime affected the abundance, distribution and the level of physiological stress of diatoms in the Strait. During Spring tides, diatoms were more abundant, usually grouped nearer the coastal basin and showed less physiological stress than during Neap tides. Our results suggest a significant general increase in the pPUA productivity with increasing physiological stress for the cell also significantly associated to low nitrate availability.

Burrowing beetles of the genus Bledius (Staphylinidae) as agents of bioturbation in the emergent areas and shores of an athalassic inland lake (Fuente de Piedra, southern of Spain)

Bledius (Elbidus) bicornis (Germ.) and B. (Eucerotobledius) furcatus (Oliv.) (Coleoptera: Staphylinidae) are the most important burrowing species in the emergent areas and shores in the athalassic lake of Fuente de Piedra (Málaga, S. of Spain).A first estimate of the importance of these organisms in this system is presented. These insects kick out sediment during their burrowing activity, which accumulates on the surface near the burrows as tumuli which can be easily eroded. The lake perimeter (17 km) is densely colonized (usual densities from 1700 to 2500 ind m−2). The amount of granulated material that can be potentially kicked out was 46.22 g dry wt m−2 day−1. At the same time, the material that constitutes the tumuli shows different characteristics from the compact ground below the surface. Thus, it is relatively enriched with organic matter (6.15 g per square meter), soluble phosphate (406.5 µg m−2) and ammonium (4856 µg m−2), whereas it lacks nitrate. Results of a transect from uninhabited areas to zones of maximum population density also show a similarity between the higher ground level of ammonium and phosphate concentrations and population density.