A dynamic mass balance model for phosphorus fluxes and concentrations in coastal areas

This paper presents a general, process-based mass balance model (CoastMab) for total phosphorus (TP) in defined coastal areas (at the ecosystem scale). The model is based on ordinary differential equations and calculates inflow, outflow and internal fluxes on a monthly basis. It consists of four compartments: surface water, deep water, erosion/transportation areas for fine sediments and accumulation areas for fine sediments. The separation between surface water and deep water is not done based on water temperature, but on sedimentological criteria instead (from the theoretical wave base). There are algorithms for all major internal TP fluxes (sedimentation, resuspension, diffusion, mixing and burial). Validations were performed using data from 21 different Baltic coastal areas. The results show that the model predicts monthly TP in water and chlorophyll a very well (generally within the uncertainty bands of the empirical data). The model has also been put through sensitivity tests, which show that the most important factor regulating the predictions of the model is generally the TP concentration in the sea beyond the coast. The model is simple to apply, since all driving variables may be accessed from maps or monitoring programs. The driving variables include coastal area, section area (between the defined coastal area and the adjacent sea), mean and maximum depths, latitude (used to predict water temperatures, stratification and mixing), salinity and TP concentration in the sea. Many of the model structures are general and could be used for areas other than those included in this study, e.g., for open coasts, estuaries or tidal coasts, as well as for other substances than phosphorus.     

Artificial mixing prevents nuisance blooms of the cyanobacterium Microcystis in Lake Nieuwe Meer, the Netherlands

1. Artificial mixing in the hypertrophic Lake Nieuwe Meer was successful in preventing blooms of the cyanobacterium Microcystis. During the 2 years of artificial, deep mixing the number of colonies of Microcystis per litre and also per m² was lower than in the two preceding control years. Hardly any nuisance scums of Microcystis occurred in the lake.
2. The phytoplankton shifted from a cyanobacteria-dominated community in summer to a mixed community of flagellates, green algae and diatoms. Reduced sedimentation losses in the mixed lake, probably in combination with a lower pH, favoured non-buoyant algae, while the entrainment of cyanobacteria in the turbulent flow nullified their advantage of buoyancy.
3. The chlorophyll concentrations were much lower in the mixed lake, but the euphotic depth did not show clear differences between the years. The chlorophyll content integrated through depth (m^–2) increased in the artificially mixed lake.
4. The deep lake normally stratified in summer, but artificial mixing of the lake in 1993 resulted in a homogeneous temperature and oxygen distribution with depth. In spring 1994, the mixing was applied intermittently with a reduction of 75% of the energy costs, while the mixing was still sufficient to prevent stratification.
5. Determination of the buoyancy state of the colonies on a sunny and calm day showed that the buoyancy loss was low close to the bubble plumes, and high at some distance from these plumes. This suggests that Microcystis could escape the mixing at some distance from the plumes, and could synthesize more carbohydrates during its stay in the upper illuminated layer of the lake than the deep mixed colonies close to the bubble plumes. Determination of the buoyancy state appeared to be a good and simple method to investigate the extent of entrainment of colonies in the turbulent flow.     

Artificial mixing prevents nuisance blooms of the cyanobacterium Microcystis in Lake Nieuwe Meer, the Netherlands

1. Artificial mixing in the hypertrophic Lake Nieuwe Meer was successful in preventing blooms of the cyanobacterium Microcystis. During the 2 years of artificial, deep mixing the number of colonies of Microcystis per litre and also per m² was lower than in the two preceding control years. Hardly any nuisance scums of Microcystis occurred in the lake.
2. The phytoplankton shifted from a cyanobacteria-dominated community in summer to a mixed community of flagellates, green algae and diatoms. Reduced sedimentation losses in the mixed lake, probably in combination with a lower pH, favoured non-buoyant algae, while the entrainment of cyanobacteria in the turbulent flow nullified their advantage of buoyancy.
3. The chlorophyll concentrations were much lower in the mixed lake, but the euphotic depth did not show clear differences between the years. The chlorophyll content integrated through depth (m^–2) increased in the artificially mixed lake.
4. The deep lake normally stratified in summer, but artificial mixing of the lake in 1993 resulted in a homogeneous temperature and oxygen distribution with depth. In spring 1994, the mixing was applied intermittently with a reduction of 75% of the energy costs, while the mixing was still sufficient to prevent stratification.
5. Determination of the buoyancy state of the colonies on a sunny and calm day showed that the buoyancy loss was low close to the bubble plumes, and high at some distance from these plumes. This suggests that Microcystis could escape the mixing at some distance from the plumes, and could synthesize more carbohydrates during its stay in the upper illuminated layer of the lake than the deep mixed colonies close to the bubble plumes. Determination of the buoyancy state appeared to be a good and simple method to investigate the extent of entrainment of colonies in the turbulent flow.     

The Burgundy-blood phenomenon: a model of buoyancy change explains autumnal waterblooms by Planktothrix rubescens in Lake Zürich

Buoyancy changes of the cyanobacterium Planktothrix rubescens- the Burgundy-blood alga - were modelled from its buoyancy response to light and irradiance changes in Lake Zürich during autumnal mixing. The daily insolation received by filaments at fixed depths and circulating to different depths was calculated from the measured light attenuation and surface irradiance. The active mixing depth, za5, was determined from the vertical turbulent diffusion coefficient, Kz, calculated from the wind speed, heat flux and temperature gradients. The fixed depth resulting in filament buoyancy, zn, decreased from 13 to 2 m between August and December 1998; the critical depth for buoyancy, zq, to which filaments must be circulated to become buoyant, decreased from >60 m in the summer to <10 m in winter. When za5 first exceeded zn, in September, P. rubescens was mixed into the epilimnion. In October, zq > za5: circulating filaments would have lost buoyancy in the high insolation. Often in November and December, after deeper mixing and lower insolation, za5 > zq: filaments would have become buoyant but would have floated to the lake surface (the Burgundy-blood phenomenon) only under subsequent calm conditions, when Kz was low. The model explains the Burgundy-blood phenomenon in deeper lakes; waterblooms near shallow leeward shores arise from populations floating up in deeper regions of the lake.     

Suspended particulate matter dynamics in the shallow mesotidal urdaibai estuary (Bay of Biscay,Spain)

Seston quantity and quality was measured in the Urdaibai Estuary in summer, when river discharge is low and tidal flushing is the main driving force to transport and resuspend particulate matter in the estuary. The highest seston concentrations are found in the upper estuary, where more than 90% of total suspended particulate matter is made up of particles <20 μm. There is a temporal ebb-flood asymmetry and turbulent mixing is stronger at flood tide. During dry calm periods sediment resuspension is observed for a short period of time at floods during spring tides, decreasing drastically before slack waters. In spite of this, particulate organic matter mainly originates from anthropogenic sources (domestic wastewater discharge) and plankton production.     

Calcium sensors in sea urchin sperm flagella

The asymmetry of ATP-reactivated flagellar bending waves of Triton-demembrated sea urchin spermatozoa has been measured over a range of free Ca2+ ion concentrations from 10(-9) to 10(-4) M. Detailed examination of the gradual response of asymmetry to Ca2+ ion concentration over this wide range indicates the presence of two Ca2+ sensors. A high-affinity sensor operates at Ca2+ concentrations near 10(-7.5) M. A lower-affinity sensor operates at Ca2+ concentrations above 10(-6) M, in the typical range for calmodulin-mediated responses. Incubation of demembranated sperm flagella at high Ca2+ concentrations to release calmodulin is required to enable these Ca2+ responses to be observed. This treatment also causes a decrease in the apparent affinity of the flagella for calmodulin, as determined by measuring the increase in asymmetry in response to addition of exogenous calmodulin at low Ca2+ concentration.     

The relative impacts of grazing by caribbean coral reef fishes and Diadema: effects of habitat and surge

The effects of grazing on fleshy algal turfs by large herbivorous fishes and by the black spiny urchin, Diadema antillarum (Philippi), were investigated in three shallow (<3 m), moderately exposed, reef flat habitats in the San Blas Islands on the Caribbean coast of Panama. Grazing by Diadema was found to be a primary determinant of the biomass of fleshy algae in this shallow reef environment, and its impact was exposure-dependent. Feeding activity of Diadema was inhibited during the protracted periods of heavy wave action that characterize the dry season when strong north winds (24–27 kph) are typical. In the shallowest, most exposed elkhorn bluff habitats urchin grazing was minimal even during calm periods due to the effects of waves generated by unpredictable, light winds. In a relatively protected backreef habitat, water movement was inadequate to discourage grazing significantly, even during the turbulent dry season. As a result, algal biomass (decalcified dry weight) was typically three-fold greater in the elkhorn bluff habitat than on the ackreef. In habitats of intermediate exposure (elkhorn fields) algal biomass was reduced during calm periods, but increased to the level characteristic of the elkhorn bluff habitat during protracted rough periods that inhibited grazing by Diadema.A caging experiment in the backreef habitat demonstrated a relatively minor effect of grazing by herbivorous fishes in comparison to that of grazing by Diadema. Observations on the distributions and abundances of herbivorous fishes also indicated a relatively minor role for these grazers as determinants of algal biomass in the more exposed habitats.     

Colonization of the deep sea by fishes

Analysis of maximum depth of occurrence of 11 952 marine fish species shows a global decrease in species number (N) with depth (x; m): log₁₀N = ?0·000422x + 3·610000 (r² = 0·948). The rate of decrease is close to global estimates for change in pelagic and benthic biomass with depth (?0·000430), indicating that species richness of fishes may be limited by food energy availability in the deep sea. The slopes for the Classes Myxini (?0·000488) and Actinopterygii (?0·000413) follow this trend but Chondrichthyes decrease more rapidly (?0·000731) implying deficiency in ability to colonize the deep sea. Maximum depths attained are 2743, 4156 and 8370 m for Myxini, Chondrichthyes and Actinopterygii, respectively. Endemic species occur in abundance at 7–7800 m depth in hadal trenches but appear to be absent from the deepest parts of the oceans, >9000 m deep. There have been six global oceanic anoxic events (OAE) since the origin of the major fish taxa in the Devonian c. 400 million years ago (mya ). Colonization of the deep sea has taken place largely since the most recent OAE in the Cretaceous 94 mya when the Atlantic Ocean opened up. Patterns of global oceanic circulation oxygenating the deep ocean basins became established coinciding with a period of teleost diversification and appearance of the Acanthopterygii. Within the Actinopterygii, there is a trend for greater invasion of the deep sea by the lower taxa in accordance with the Andriashev paradigm. Here, 31 deep‐sea families of Actinopterygii were identified with mean maximum depth >1000 m and with >10 species. Those with most of their constituent species living shallower than 1000 m are proposed as invasive, with extinctions in the deep being continuously balanced by export of species from shallow seas. Specialized families with most species deeper than 1000 m are termed deep‐sea endemics in this study; these appear to persist in the deep by virtue of global distribution enabling recovery from regional extinctions. Deep‐sea invasive families such as Ophidiidae and Liparidae make the greatest contribution to fish fauna at depths >6000 m.     

A simple model of plankton population dynamics coupled with a LES of the surface mixed layer

The concentration of phytoplankton in the sea is affected by biological processes, such as growth/mortality rates, predatory zooplankton concentrations and nutrient levels. Phytoplankton concentrations are also influenced by physical processes, in particular the mixing properties of the local fluid environment. On planktonic scales (approximately 10-1000 microm) one can assume the local turbulent flow is isotropic, with no distinction between horizontal and vertical mixing. However, agglomerations of phytoplankton into patches are observed on larger scales of up to hundreds of metres, whose formation will be influenced by the anisotropic advection/mixing properties and large-eddy structures prevalent in the surface mixed layer. This paper presents the results of the coupling of a large-eddy simulation (LES) model of the mixed layer with an advection-diffusion system of coupled equations for nitrate-phytoplankton-zooplankton (NPZ) concentration, incorporating sub-grid parameterizations of the biological processes. Typically these include phytoplankton growth due to light levels and ambient nitrate concentration, offset by grazing losses due to the presence of zooplankton. The primary goal of this work is to investigate how the characteristics of the mixed layer turbulence influence the observed distribution of phytoplankton. One novel feature is the incorporation of a 'vortex-force' term in the LES code in order to generate Langmuir circulations. It has been speculated that the enhanced mixing rates associated with 'Langmuir turbulence' play a significant role in regulating planktonic activity. Results derived from the coupled LES-NPZ model, run with and without the presence of Langmuir circulations, are presented in order to investigate these ideas.     

Wave and sediment dynamics along a shallow subtidal sandy beach inhabited by modern stromatolites

To help define the habitat of modern marine stromatolites, wave-dominated flow and sediment transport were studied in the shallow subtidal region (1-2 m depth) along the slightly concave, windward face of Highborne Cay, Exuma, Bahamas - the only face of the cay that includes a population of stromatolites concentrated near the region of highest curvature of the beach. Wave energy impacting this island's most exposed beach was driven by local wind forcing which increases largely in response to the passage of atmospheric disturbances that typically affect the region for periods of a few days. Although some wave energy is almost always noted (maximum horizontal orbital speeds at the bottom are rarely <10 cm s(-1)), wave conditions remain comparatively calm until local winds increase above speeds of approximately 3-4 m s(-1) at which point maximum wave speeds rapidly increase to 50-80 cm s(-1). Stromatolites, which are largely restricted to the shoreward side of a shallow platform reef, are sheltered by the reef beyond which wave speeds are one to four times higher (depending on tidal stage). Moreover, stromatolite populations are predominantly found along a region of this wave-exposed beach that experiences comparatively reduced wave energy because of the curved morphology of the island's face. Maximum wave speeds are 1.4 to 2 times higher along more northern sections of the beach just beyond the locus of stromatolite populations. A quantitative model of sediment transport was developed that accurately predicted accumulation of suspended sediment in sediment traps deployed in the shallow subtidal zone along this beach. This model, coupled with in situ wave records, indicates that gross rates of suspended sediment deposition should be two to three times higher northward of the main stromatolite populations. Regions of the beach containing stromatolites nevertheless should experience significant rates of gross suspended sediment deposition averaging 7-10 g cm(-2) day(-1) ( approximately 4-6 cm day(-1)). Results suggest that one axis of the habitat of modern marine stromatolites may be defined by a comparatively narrow range of flow energy and sediment transport conditions.     

The effect of wind on shore gastropods

A preliminary investigation into the effect of wind on shore gastropods has been undertaken. Moderate winds displace some Monodonta lineata (da Costa) from boulders on the shore and some individuals detach within six hours in weaker winds in the laboratory (5–10 km/h, 10°C, 94% humidity). Sheltered aspects of the rock are used to minimize the effects of the wind on shores, and maintenance movements on the shore are extensive.  

Summary:

In calm conditions and in light winds Monodonta lineata is more widely dispersed about the rocks than in moderate winds when the animals are either sheltering in cracks or displaced.
Providing the rock surface is damp then animals continue to move about during emersion in light winds. On a small boulder beach animals moved as much as 4 m upshore during one tidal cycle. Only slight reorientation movements were recorded during moderate winds.
Animals withstand a windspeed of 4 km/h for six hours in a wind tunnel in a humidity of 94% at 10°C, but are displaced within an hour by 10 km/h winds in similar conditions.     

Some effects of turbulence on the activity of the sea urchin Centrostephanus coronatus Verrill

Effects of turbulence on activity of the sea urchin Centrostephanus coronatus Verrill were studied using a new method referred to as the trip-thread method which recorded incidences of emergence from burrows during activity periods. During calm conditions, the number of C. coronatus emerging at 5 m depth was not significantly different from the number emerging at 15 m depth, representing 97% and 92% activity in the populations, respectively; however, during turbulent conditions, the number of individuals emerging at 5 m was significantly less than the number emerging at 15 m, representing 48% and 92% activity, respectively. These results suggest an inverse relationship between turbulence and activity. Although behavioral adaptations, such as reduced activity, allow urchins to occupy turbulent areas, extreme conditions may exclude them entirely. Differences in activity may affect grazing by sea urchins which can significantly affect the species composition and abundance of benthic communities.     

Stormy oceans are associated with declines in sea turtle hatching

Many sea turtle populations are below 10% of their pre-Columbian numbers [1-4]. Though historic and systematic over-exploitation is the principal cause of these declines, sea turtles face similar threats today. Adults and juveniles are actively hunted and commercial fisheries catch them incidentally. Nesting suffers from beach development, egg poaching and the poaching of nesting females. Accompanying these familiar hazards is the largely unknown consequences of recent climate change. Here we report monitoring surveys from the Dry Tortugas National Park (DTNP, 24.64N 82.86W), Florida, and show that hurricanes and other storm events are an additional and increasing threat to loggerhead turtle (Caretta caretta) and green sea turtle (Chelonia mydas) nesting. Both species are listed by the US Endangered Species Act and the IUCN considers them 'endangered'.     

Nutrient fluxes from upwelling and enhanced turbulence at the top of the pycnocline in Mono Lake, California

Time series measurements of temperature at 15 depths and profiles of temperature-gradient microstructure were obtained during a period with strong wind forcing and subsequent calm in Mono Lake, California. The wind forcing increased the amplitude of basin-scale internal waves and energy at all wave frequencies relative to the calm period. Rates of dissipation of turbulent kinetic energy, , were high ( > 10^–6 m² s^–3) at the top of the pycnocline at both an inshore and an offshore site on a day when winds reached 10 m s^–1 and on the following two days at an inshore site ( > 10^–7 m² s^–3). The enhanced turbulence occurred at the depth of a subsurface temperature maximum (z _TM) and coincidentally with elevated concentrations of NH₄, reduced concentrations of chlorophyll a and particulate carbon, and increased abundance of the macrozooplankter Artemia monica. The NH₄ at z _TM was more dispersed and of lower concentration inshore than offshore and indicated greater turbulent transport inshore. Over the course of 4 days, chlorophyll a concentrations increased in the upper mixed layer, and C:N and C:Chl ratios decreased. Offshore, the change in C:N ratio indicated a relaxation of moderate nutrient deficiency. We hypothesize that excretion by A. monica and turbulent transport of the NH₄ from the subsurface temperature maximum led to improved physiological status of phytoplankton in the upper mixed layer.     

If you build it,will they come? Use of restored dunes by beach mice

Restoration of coastal habitat fragmented, degraded, or destroyed by development and climate‐related processes such as sea level rise and storm surge usually involves planting native plants to restore habitat structure, but whether and how restored areas benefit taxa other than plants is rarely reported. Installing restoration plantings is one method used to build habitat such as beach dunes where dunes have been lost, potentially creating habitat for dune‐dependent species. We compared use of natural vegetated dunes, open sand gaps, and restoration plantings (habitat treatment) by Perdido Key beach mice (Peromyscus polionotus trissyllepsis) over 3 years using tracking tubes to assess the value of restoration plantings for beach mice. Tubes were monitored in two seasons (early and mid‐summer), and under new and full moon conditions. Mice used restoration plantings less than natural vegetated dunes but more than open sand gaps, which suggests restoration plantings may facilitate movement of mice across fragmented areas. Both season and moon phase influenced the effect of habitat treatment, interactions which may be attributable to perceived risk associated with movement under a combination of different conditions of ambient light, vegetation cover, and habitat novelty. Our results show restoration plantings provide habitat for movement and foraging, and may ameliorate some consequences of sea level rise and storms for beach mice and potentially other dune‐dependent species into the future.     

Effects of Light and Wind Speed on the Vertical Distribution of Microcystis aeruginosa Colonies of Different Sizes during a Summer Bloom

Different dynamics, of the vertical distribution of Microcystis aeruginosa colonies of different sizes, were examined in a large shallow lake (Taihu Lake) during a summer bloom. During the observation the Microcystis colonies, larger than 120 μm, were mainly concentrated in the upper layer both under calm and windy conditions. Colonies from 36 μm to 120 μm showed a clear relationship to wind conditions whereas colonies less than 36 μm showed a nearly uniform vertical distribution at the different depths. There was no clear relationship between the vertical distribution of colonies larger than 36 μm and light intensity. However, at solar noon, colonies less than 36 μm were mainly distributed in the middle layers of the water column. The results indicate that small colonies are easily affected by the wind‐induced mixing, and they show an ability for active movement in response to light intensity under calm conditions. In contrast, large colonies tend to show little diurnal repositioning to light intensity and thereby are mainly concentrated in the surface layer. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Decision between small and large prey: reduced energy acquisition by pre-migratory purple sandpipers,Calidris maritima,on Svalbard

On Svalbard, purple sandpipers forage on the shores when preparing for autumn migration to the European continent after a period of food shortage on the tundra in summer. Their food choice was studied on a beach when they were faced with the choice between two abundant prey species: a small collembolan (springtail) and a much larger amphipod. In spite of several times higher gain of energy when foraging on amphipops, the birds did not show a distinct preference for either of the two prey species in this situation. Alternative explanations for this behaviour are discussed. We suggest that this could be a consequence of adaptations to their summer diet on Svalbard, when they rely on Collembola as food to a considerable extent. Higher variance in energy gain rate when foraging on amphipods might also be important. Collembolan densities high enough to trigger predation from birds occurred only on some days, with cloudy calm weather. This, together with the possible existence of a dietary bottleneck, may reduce the cost associated with the switch to Collembola. Accepted: 25 April 1999     

A synthesis of genetic connectivity in deep‐sea fauna and implications for marine reserve design

With anthropogenic impacts rapidly advancing into deeper waters, there is growing interest in establishing deep‐sea marine protected areas (MPAs) or reserves. Reserve design depends on estimates of connectivity and scales of dispersal for the taxa of interest. Deep‐sea taxa are hypothesized to disperse greater distances than shallow‐water taxa, which implies that reserves would need to be larger in size and networks could be more widely spaced; however, this paradigm has not been tested. We compiled population genetic studies of deep‐sea fauna and estimated dispersal distances for 51 studies using a method based on isolation‐by‐distance slopes. Estimates of dispersal distance ranged from 0.24 km to 2028 km with a geometric mean of 33.2 km and differed in relation to taxonomic and life‐history factors as well as several study parameters. Dispersal distances were generally greater for fishes than invertebrates with the Mollusca being the least dispersive sampled phylum. Species that are pelagic as adults were more dispersive than those with sessile or sedentary lifestyles. Benthic species from soft‐substrate habitats were generally less dispersive than species from hard substrate, demersal or pelagic habitats. As expected, species with pelagic and/or feeding (planktotrophic) larvae were more dispersive than other larval types. Many of these comparisons were confounded by taxonomic or other life‐history differences (e.g. fishes being more dispersive than invertebrates) making any simple interpretation difficult. Our results provide the first rough estimate of the range of dispersal distances in the deep sea and allow comparisons to shallow‐water assemblages. Overall, dispersal distances were greater for deeper taxa, although the differences were not large (0.3–0.6 orders of magnitude between means), and imbalanced sampling of shallow and deep taxa complicates any simple interpretation. Our analyses suggest the scales of dispersal and connectivity for reserve design in the deep sea might be comparable to or slightly larger than those in shallow water. Deep‐sea reserve design will need to consider the enormous variety of taxa, life histories, hydrodynamics, spatial configuration of habitats and patterns of species distributions. The many caveats of our analyses provide a strong impetus for substantial future efforts to assess connectivity of deep‐sea species from a variety of habitats, taxonomic groups and depth zones.