Short-term climatic variation in the Black Sea region is significantly influenced by the operation of the North Atlantic oscillation, the climatic mechanisms resulting from the interaction between the north Atlantic and mid-latitude air masses. [1] While the exact mechanisms causing the North Atlantic Oscillation remain unclear, [2] it is thought the climate conditions established in western Europe mediate the heat and precipitation fluxes reaching Central Europe and Eurasia, regulating the formation of winter cyclones, which are largely responsible for regional precipitation inputs [3] and influence Mediterranean Sea surface temperatures (SST's). [4]
The relative strength of these systems also limits the amount of cold air arriving from northern regions during winter. [5] Other influencing factors include the regional topography, as depressions and storms systems arriving from the Mediterranean are funneled through the low land around the Bosporus, Pontic and Caucasus mountain ranges acting as waveguides, limiting the speed and paths of cyclones passing through the region. [6]
The Black Sea supports an active and dynamic marine ecosystem, dominated by species suited to the brackish, nutrient-rich, conditions. As with all marine food webs, the Black Sea features a range of trophic groups, with autotrophic algae, including diatoms and dinoflagellates, acting as primary producers. The fluvial systems draining Eurasia and central Europe introduce large volumes of sediment and dissolved nutrients into the Black Sea, but the distribution of these nutrients is controlled by the degree of physiochemical stratification, which is, in turn, dictated by seasonal physiographic development. [7]
During winter, strong wind promotes convective overturning and upwelling of nutrients, while high summer temperatures result in a marked vertical stratification and a warm, shallow mixed layer. [8] Day length and insolation intensity also controls the extent of the photic zone. Subsurface productivity is limited by nutrient availability, as the anoxic bottom waters act as a sink for reduced nitrate, in the form of ammonia. The benthic zone also plays an important role in Black Sea nutrient cycling, as chemosynthetic organisms and anoxic geochemical pathways recycle nutrients which can be upwelled to the photic zone, enhancing productivity. [9]
In total, Black Sea's biodiversity contains around one-third of Mediterranean's and is experiencing natural and artificial invasions or Mediterranizations. [10] [11]
The main phytoplankton groups present in the Black Sea are dinoflagellates, diatoms, coccolithophores and cyanobacteria. Generally, the annual cycle of phytoplankton development comprises significant diatom and dinoflagellate-dominated spring production, followed by a weaker mixed assemblage of community development below the seasonal thermocline during summer months and surface-intensified autumn production. [8] [12] This pattern of productivity is also augmented by an Emiliania huxleyi bloom during the late spring and summer months.
Since the 1960s, rapid industrial expansion along the Black Sea coast line and the construction of a major dam has significantly increased annual variability in the N:P:Si ratio in the basin. In coastal areas, the biological effect of these changes has been an increase in the frequency of monospecific phytoplankton blooms, with diatom bloom frequency increasing by a factor of 2.5 and non-diatom bloom frequency increasing by a factor of 6. The non-diatoms, such as the prymnesiophytes Emiliania huxleyi (coccolithophore), Chromulina sp., and the Euglenophyte Eutreptia lanowii are able to out-compete diatom species because of the limited availability of Si, a necessary constituent of diatom frustules. [31] As a consequence of these blooms, benthic macrophyte populations were deprived of light, while anoxia caused mass mortality in marine animals. [32] [33]
The decline in macrophytes was further compounded by overfishing during the 1970s, while the invasive ctenophore Mnemiopsis reduced the biomass of copepods and other zooplankton in the late 1980s. Additionally, an alien species—the warty comb jelly (Mnemiopsis leidyi)—was able to establish itself in the basin, exploding from a few individuals to estimated biomass of one billion metric tons. [34] The change in species composition in Black Sea waters also has consequences for hydrochemistry, as Ca-producing coccolithophores influence salinity and pH, although these ramifications have yet to be fully quantified. In central Black Sea waters, Si levels were also significantly reduced, due to a decrease in the flux of Si associated with advection across isopycnal surfaces. This phenomenon demonstrates the potential for localized alterations in Black Sea nutrient input to have basin-wide effects.
Pollution reduction and regulation efforts have led to a partial recovery of the Black Sea ecosystem during the 1990s, and an EU monitoring exercise, 'EROS21', revealed decreased N and P values, relative to the 1989 peak. [35] Recently, scientists have noted signs of ecological recovery, in part due to the construction of new sewage treatment plants in Slovakia, Hungary, Romania, and Bulgaria in connection with membership in the European Union. Mnemiopsis leidyi populations have been checked with the arrival of another alien species which feeds on them. [36]
In the past, the range of the Asiatic lion extended from South Asia to the Balkans, possibly up to the Danube. Places like Turkey and the Trans-Caucasus were in this range. The Caspian tiger occurred in eastern Turkey and the Caucasus, at least. The lyuti zver (Old East Slavic for "fierce animal") that was encountered by Vladimir II Monomakh, Velikiy Kniaz of Kievan Rus' (which ranged to the Black Sea in the south), [37] may have been a tiger or leopard, rather than a wolf or lynx, due to the way it behaved towards him and his horse. [38]
The Black Sea is a marginal mediterranean sea of the Atlantic Ocean lying between Europe and Asia; east of the Balkan Peninsula, south of the East European Plain in Eastern Europe, west of the Caucasus, and north of Anatolia in Western Asia. It is supplied by major rivers, principally the Danube, Dnieper, and Don. The watersheds of many countries drain into the sea beyond the six that share its coast.
Plankton are the diverse collection of organisms found in water that are unable to propel themselves against a current. The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucial source of food to many small and large aquatic organisms, such as bivalves, fish and whales.
An algal bloom or algae bloom is a rapid increase or accumulation in the population of algae in freshwater or marine water systems. It is often recognized by the discoloration in the water from the algae's pigments. The term algae encompasses many types of aquatic photosynthetic organisms, both macroscopic multicellular organisms like seaweed and microscopic unicellular organisms like cyanobacteria. Algal bloom commonly refers to the rapid growth of microscopic unicellular algae, not macroscopic algae. An example of a macroscopic algal bloom is a kelp forest.
A coccolithophore is a unicellular, eukaryotic phytoplankton (alga). They belong either to the kingdom Protista, according to Robert Whittaker's Five kingdom classification, or clade Hacrobia, according to the newer biological classification system. Within the Hacrobia, the coccolithophorids are in the phylum or division Haptophyta, class Prymnesiophyceae. Coccolithophorids are distinguished by special calcium carbonate plates of uncertain function called coccoliths, which are also important microfossils. However, there are Prymnesiophyceae species lacking coccoliths, so not every member of Prymnesiophyceae is a coccolithophorid. Coccolithophores are almost exclusively marine, are photosynthetic, and exist in large numbers throughout the sunlight zone of the ocean.
Phytoplankton are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words φυτόν, meaning 'plant', and πλαγκτός, meaning 'wanderer' or 'drifter'.
The biological pump, also known as the marine carbon pump, is, in its simplest form, the ocean's biologically driven sequestration of carbon from the atmosphere and land runoff to the ocean interior and seafloor sediments. It is the part of the oceanic carbon cycle responsible for the cycling of organic matter formed mainly by phytoplankton during photosynthesis (soft-tissue pump), as well as the cycling of calcium carbonate (CaCO3) formed into shells by certain organisms such as plankton and mollusks (carbonate pump).
Coccoliths are individual plates of calcium carbonate formed by coccolithophores which are arranged around them in a coccosphere.
The spring bloom is a strong increase in phytoplankton abundance that typically occurs in the early spring and lasts until late spring or early summer. This seasonal event is characteristic of temperate North Atlantic, sub-polar, and coastal waters. Phytoplankton blooms occur when growth exceeds losses, however there is no universally accepted definition of the magnitude of change or the threshold of abundance that constitutes a bloom. The magnitude, spatial extent and duration of a bloom depends on a variety of abiotic and biotic factors. Abiotic factors include light availability, nutrients, temperature, and physical processes that influence light availability, and biotic factors include grazing, viral lysis, and phytoplankton physiology. The factors that lead to bloom initiation are still actively debated.
Emiliania huxleyi is a species of coccolithophore found in almost all ocean ecosystems from the equator to sub-polar regions, and from nutrient rich upwelling zones to nutrient poor oligotrophic waters. It is one of thousands of different photosynthetic plankton that freely drift in the euphotic zone of the ocean, forming the basis of virtually all marine food webs. It is studied for the extensive blooms it forms in nutrient-depleted waters after the reformation of the summer thermocline. Like other coccolithophores, E. huxleyi is a single-celled phytoplankton covered with uniquely ornamented calcite disks called coccoliths. Individual coccoliths are abundant in marine sediments although complete coccospheres are more unusual. In the case of E. huxleyi, not only the shell, but also the soft part of the organism may be recorded in sediments. It produces a group of chemical compounds that are very resistant to decomposition. These chemical compounds, known as alkenones, can be found in marine sediments long after other soft parts of the organisms have decomposed. Alkenones are most commonly used by earth scientists as a means to estimate past sea surface temperatures.
Calanus finmarchicus is a species of copepods and a part of zooplankton, which is found in enormous amounts in the northern Atlantic Ocean.
The genus Ceratium is restricted to a small number of freshwater dinoflagellate species. Previously the genus contained also a large number of marine dinoflagellate species. However, these marine species have now been assigned to a new genus called Tripos. Ceratium dinoflagellates are characterized by their armored plates, two flagella, and horns. They are found worldwide and are of concern due to their blooms.
Siliceous ooze is a type of biogenic pelagic sediment located on the deep ocean floor. Siliceous oozes are the least common of the deep sea sediments, and make up approximately 15% of the ocean floor. Oozes are defined as sediments which contain at least 30% skeletal remains of pelagic microorganisms. Siliceous oozes are largely composed of the silica based skeletons of microscopic marine organisms such as diatoms and radiolarians. Other components of siliceous oozes near continental margins may include terrestrially derived silica particles and sponge spicules. Siliceous oozes are composed of skeletons made from opal silica Si(O2), as opposed to calcareous oozes, which are made from skeletons of calcium carbonate organisms (i.e. coccolithophores). Silica (Si) is a bioessential element and is efficiently recycled in the marine environment through the silica cycle. Distance from land masses, water depth and ocean fertility are all factors that affect the opal silica content in seawater and the presence of siliceous oozes.
The deep chlorophyll maximum (DCM), also called the subsurface chlorophyll maximum, is the region below the surface of water with the maximum concentration of chlorophyll. A DCM is not always present - sometimes there is more chlorophyll at the surface than at any greater depth - but it is a common feature of most aquatic ecosystems, especially in regions of strong thermal stratification. The depth, thickness, intensity, composition, and persistence of DCMs vary widely. The DCM generally exists at the same depth as the nutricline, the region of the ocean where the greatest change in the nutrient concentration occurs with depth.
Marine microorganisms are defined by their habitat as microorganisms living in a marine environment, that is, in the saltwater of a sea or ocean or the brackish water of a coastal estuary. A microorganism is any microscopic living organism or virus, that is too small to see with the unaided human eye without magnification. Microorganisms are very diverse. They can be single-celled or multicellular and include bacteria, archaea, viruses and most protozoa, as well as some fungi, algae, and animals, such as rotifers and copepods. Many macroscopic animals and plants have microscopic juvenile stages. Some microbiologists also classify biologically active entities such as viruses and viroids as microorganisms, but others consider these as non-living.
Marine primary production is the chemical synthesis in the ocean of organic compounds from atmospheric or dissolved carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through chemosynthesis, which uses the oxidation or reduction of inorganic chemical compounds as its source of energy. Almost all life on Earth relies directly or indirectly on primary production. The organisms responsible for primary production are called primary producers or autotrophs.
Marine protists are defined by their habitat as protists that live in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. Life originated as single-celled prokaryotes and later evolved into more complex eukaryotes. Eukaryotes are the more developed life forms known as plants, animals, fungi and protists. Protists are the eukaryotes that cannot be classified as plants, fungi or animals. They are usually single-celled and microscopic. The term protist came into use historically as a term of convenience for eukaryotes that cannot be strictly classified as plants, animals or fungi. They are not a part of modern cladistics, because they are paraphyletic.
Many protists have protective shells or tests, usually made from silica (glass) or calcium carbonate (chalk). Protists are mostly single-celled and microscopic. Their shells are often tough mineralised forms that resist degradation, and can survive the death of the protist as a microfossil. Although protists are very small, they are ubiquitous. Their numbers are such that their shells play a huge part in the formation of ocean sediments, and in the global cycling of elements and nutrients.
Particulate inorganic carbon (PIC) can be contrasted with dissolved inorganic carbon (DIC), the other form of inorganic carbon found in the ocean. These distinctions are important in chemical oceanography. Particulate inorganic carbon is sometimes called suspended inorganic carbon. In operational terms, it is defined as the inorganic carbon in particulate form that is too large to pass through the filter used to separate dissolved inorganic carbon.
The Great Calcite Belt (GCB) of the Southern Ocean is a region of elevated summertime upper ocean calcite concentration derived from coccolithophores, despite the region being known for its diatom predominance. The overlap of two major phytoplankton groups, coccolithophores and diatoms, in the dynamic frontal systems characteristic of this region provides an ideal setting to study environmental influences on the distribution of different species within these taxonomic groups.
Linda Karen Medlin is a molecular biologist known for her work on diatoms. She is an elected member of the Norwegian Academy of Science and Letters.