Modern development of barrier salinity zones relativity and plurality conception

1. ICES Annual Science ConferenceBaltic Committee meeting Maastricht, September 18-23, 2006 Modern development of barrier salinity zones relativity and plurality conception N.V. Aladin, I.S. Plotnikov, M.B. DianovZoological Institute of Russian Academy of Sciences, St.Petersburg

2. Water salinity is one of major environmental factors influencing hydrobionts. Ascertainment of specificity of the attitude of aquatic animals and plants to this factor is important to understand both autoecological and synecologilal rules. • Conception of relativity and plurality of water barrier salinity zones was formulated more than 20 years before in the frames of V.V. Khlebovich’s school of thought (Aladin, 1986). Its main theses were published in the Journal of General Biology (Aladin, 1988). • Two main theses were stated: • Zones of barrier salinities are relative, on the one hand, to the degree of perfection of hydrobionts osmoregulatory capacities and, on the other hand, to the water chemical composition. • There are several zones of barrier salinities and they are unequal by their importance.

3. Zones of barrier salinities and tolerance ranges of hydrobionts from marine and continental waters : horizontal axis – salinity, ‰; over horizontal axis are given salinity tolerance ranges of hydrobionts from marine waters; below horizontal axis – for those from continental waters. Osmoconformers: 1 – I, 2 – II, 3 – III; confohyperosmotics: 4 – I, 5 – II; 6 – hyperosmoticsI, 7 –hyperosmoticsIIor secondary confohyperosmotics; amphiosmotics: 8 – I, 9 – II, 10 – III, 11 – IV; 12 – hypoosmotics; Barrier salinities of marine waters: IM – first, 5–8‰, IIM – second, 16–20‰, IIIM– third, 26–30‰, IVM – fourth, 36–40‰, VM – fifth, 50–55‰; barrier salinities of continental waters: I c – first, 5–20‰, IIc – second, 50–60‰, IIIc – third, 100–300‰ and more; A– marine brackish waters; A*– before “critical salinity” 5–8‰, A**– after“critical salinity” 5–8‰, B– typical marine waters, C– marine hyperhaline waters, D– fresh waters, E– continental brackish waters, E*– in the “critical salinity” zone 5–20‰, E**– after the the “critical salinity” zone, F– continental hyperhaline waters. Summarized from: Journal of General Biology, 1988, vol. 67(7): 974-982.

4. All the hydrosphere of our planet could be conditionally divided into freshwater, brackish water, marine and hyperhaline zones. Marine zone occupies about 95% of the hydrosphere surface. Freshwater zone occupies about 3%. Brackish water and hyperhaline zones occupy about 0.5% each. Between these four basic zones there are transitional ones occupying less than 0.5% each. Approximate boundaries and corresponding barrier salinities are found for all of these basic and transitional zones of the hydrosphere are defined.

5. Following main principles of conception of relativity and plurality of salinity barrier zones (Aladin, 1986, 1988; Aladin, Plotnikov, 2007) the following salinity zones were suggested for oceanic, Caspian and Aral waters.

6. Following the main principles of conception of relativity and plurality of salinity barrier zones (Aladin, 1986, 1988; Aladin, Plotnikov, 2007) the following salinity zones were suggested for oceanic, Caspian and Aral waters.

7. Revealing barrier salinity zones in the hydrosphere supposes studying osmoregulatory capacities of hydrobionts first of all. It is to reveal types of osmotic relations of internal media with the environment, to find experimentally limits of salinity tolerant ranges, to analyze data on salinity boundaries of hydrobionts distribution in the nature.

8. Classification of osmoconformers and osmoregulators Aquatic organisms Osmoconformers Osmoregulators Osmoconformers I Confohyperosmotics Hypoosmotics Osmoconformers II Amphyosmotics Hyperosmotics Osmoconformers III Amphyosmotics I Hyperosmotics I Confohyperosmotics I Amphyosmotics II Hyperosmotics II Confohyperosmotics II Amphyosmotics III Amphyosmotics IV

9. General table of osmoconformers and osmoregulators I I I I I II II II II III III IV Osmoconformers – majority of recent primary marine hydrobionts: Coeletnterata, Vermes, Mollusca, Arthropoda, Echinodermata, etc. Confohyperosmotics – majority of recent widely euryhaline primary marine hydrobionts: Polychaeta, Gastropoda, Crustacea, etc. Hyperosmotics – majority of recent freshwater hydrobionts: Oligochaeta, Rotatoria, Mollusca, Crustacea, Insecta, freshwater Pisces, etc. Amphyosmotics – some Crustacea, some Insecta, anadrom Pisces. Hypoosmotics – some secondary marine Crustacea, majority of recent secondary marine Pisces.

10.  Evolution of all known types of osmoregulation A0 – Hypothetic ancestral osmoconformer A1  – Stenohaline marine hydrobionts (osmoconformers I) A2  – Marine hydrobionts (osmoconformers II) A3  – Euryhaline marine hydrobionts (osmoconformers III) B1 – Widely euryhaline marine hydrobionts (confohyperosmotics I) B2  – Brackish water hydrobionts of marine origin (confohyperosmotics II) C1  – Freshwater hydrobionts (hyperosmotics I) C2  – Brackish water hydrobionts of freshwater origin (hyperosmotics II or secondary confohyperosmotics) D1  – Some Caspian brackish water hydrobionts (amphiosmotics I) D2  – Some euryhaline Australian hydrobionts of freshwater origin (amphiosmotics II) D3  – Euryhaline hydrobionts of freshwater origin (amphiosmotics III) D4  – Widely euryhaline hydrobionts of freshwater origin (amphiosmotics IV) E  – Euryhaline marine hydrobionts of freshwater origin (hypoosmotics)           

11. Different levels of Ostracoda and Branchiopoda occurrence in marine and continental waters(by: Aladin, 1986)

12. The position and width of barrier salinities ranges cannot depend on water physicochemical properties only. The values of barrier salinities can change following evolution of salinity adaptations and osmoregulation capacities of aquatic plants and animals.

13. α-, β- and g- horohalinicums in the waters classified by salinity (by: Aladin, 1986, 1988; Khlebovich, 1989 )

14. Special attention needs to be given on comparative analysis of oceanic (thalassic) zones of hydrosphere and those continental (athalassic).

15. Critical salinity shift to higher concentrations in water of Caspian and Aral seas as compared with oceanic water (by: Aladin, 1986, 1989)

16. We shall review some water bodies where we had our studies.

17. Aral Seabefore 1960  - Freshwater ecosystems  - Transitional freshwater-brackishwater ecosystems - Brackishwater ecosystems  - Transitionalbrackishwater-marine ecosystems

18. Aral Seain 1989  - Freshwater ecosystems  - Transitional freshwater-brackishwater ecosystems - Brackishwater ecosystems  - Transitionalbrackishwater-marine ecosystems  - Marine ecosystems

19. Aral Seain 2006  - Freshwater ecosystems  - Transitional freshwater-brackishwater ecosystems - Brackishwater ecosystems  - Transitionalbrackishwater-marine ecosystems  - Hyperhaline ecosystems

20. Caspian Sea  - Freshwater ecosystems  - Transitional freshwater-brackishwater ecosystems - Brackishwater ecosystems  - Transitionalbrackishwater-marine ecosystems  - Marine ecosystems  - Transitionalmarine-hyperhaline ecosystems  - Hyperhaline ecosystems

21.  - Freshwater ecosystems  - Transitional freshwater-brackishwater ecosystems - Brackishwater ecosystems  - Transitionalbrackishwater-marine ecosystems  - Marine ecosystems  - Transitionalmarine-hyperhaline ecosystems  - Hyperhaline ecosystems Sea of Azov

22. Black Sea  - Freshwater ecosystems  - Transitional freshwater-brackishwater ecosystems - Brackishwater ecosystems  - Transitionalbrackishwater-marine ecosystems

23.  - Freshwater ecosystems  - Transitional freshwater-brackishwater ecosystems - Brackishwater ecosystems  - Transitionalbrackishwater-marine ecosystems  - Marine ecosystems Baltic Sea

24.  - Freshwater ecosystems  - Transitional freshwater-brackishwater ecosystems - Brackishwater ecosystems  - Transitionalbrackishwater-marine ecosystems  - Marine ecosystems WhiteSea

25.  - Freshwater ecosystems  - Transitional freshwater-brackishwater ecosystems - Brackishwater ecosystems  - Transitionalbrackishwater-marine ecosystems  - Marine ecosystems BarentzSea

26. Sea of Japan  - Marine ecosystems

27. Ladoga Lake  - Freshwater ecosystems

28. Baltic Sea Sea of Azov Aral Seaprior 1960 Aral Seain 2006 Caspian Sea

29. Area of different salinity zones in different brackish water seas and lakes

30. Percentage of water areas of different salinity zones in different brackish water seas and lakes At present Baltic Sea is the only sea where basic brackishwater zone is occupying more than half of its area (> 60%)

31. There are offered some hypotheses on which basis position of paleo-barrier salinities in water bodies of Parathetys and ancient Baltic Sea is discussed. An attempt is made to apply main principles of the conception to forecast future scenarios of evolution of water bodies anew formed on the place of divided Aral Sea.

32. 1 2 3 4 5 6 7 8 Water bodies of Paleo-Baltic Sea(by Zenkevich, 1963, with corrections and additions)1 – maximal phase of the last glaciation; 2- Danish glaciation (15 ths B.P.); 3 – Baltic Glacial Lake (14 ths B.P.); 4 – Yoldia Sea (12 ths B.P.); 5 – Ancylus Lake-Sea (7 ths B.P.); 6 – last phase of Ancylus Lake-Sea (5 ths B.P.); 7 – Littorina Sea (4 ths B.P.); 8 – modern phase (since 2 ths B.P.)Indicatedonly average salinity without salinity gradient in the Baltic Sea

33. Water bodies of the PalaeocaspianA — Balakhansky(5 mil B.P.); B — Akchagylsky(3 mil B.P.); C — Postakchagylsky(> 2 mil B.P.); D — Apsheronsky(2 mil B.P.); E — Tyurkyansky(< 2 mil B.P.); F — Bakinsky(1.7 mil B.P.); G — Venedsky or Ushtalsky(0.5 mil B.P.); H — the Early Khazarsky (400 ths B.P.); I — the Late Khazarsky; J — Atelsky (> 50 ths B.P.); K — the Early Khvalynsky; L — Enotaevsky; M — the Late Khvalynsky; N — Mangyshlaksky (7.5 ths B.P.); O — the New Caspian (5 ths B.P.); P — the present. Indicatedonly average salinity without salinity gradient

34. Aral Sea: from9000 to1600 years BP

35. Aral Sea: from 450 years BP, till now and in the future

36. Changing of the species number in the Baltic Sea following salinity gradient

37. Scheme of aquatic fauna pattern change in water bodies with different salinities(by: Remane, 1934; Khlebovich, 1962; Kinne, 1971; with additions and corrections)1 – freshwater, 2 – brackish-water, 3 – marine, 4 – hyperhaline and ultrahaline species

38. Changing of the species number following salinity gradient in the Baltic Sea

39. Number of fishes, free-living invertebrates and plants without micro-Metazoa, Protozoa and Bacteria in the Baltic Sea

40. Decreasing of marine species biodiversity following decreasing of the Baltic Sea salinity

41. Scheme of halopathy of main types of aquatic fauna of Azov and Black Seas basin (by: Mordukhai-Boltovskoi, 1953)Abscissa – salinity, ‰; ordinate axis – number of species

42. Scheme of aquatic fauna pattern change in the Caspian and Aral seas(by: Zenkevich, 1977; Andreeva, Andreev, 2001 with additions and corrections)1 – freshwater, 2 – brackish-water, 3 – marine species

43. Percentage of different types of osmoconformers and osmoregulators in the World Ocean and fully saline seas as Barents Sea, Sea of Japan, etc. A1 – Stenohaline marine hydrobionts (osmoconformers I) – 30% A2 – Marine hydrobionts (osmoconformers II) - 25% A3 – Euryhaline marine hydrobionts (osmoconformers III) – 15% B1 – Widely euryhaline marine hydrobionts (confohyperosmotics I) – 5% B2 – Brackish water hydrobionts of marine origin (confohyperosmotics II) – 3% C1 – Freshwater hydrobionts (hyperosmotics I) – 0% C2 – Brackish water hydrobionts of freshwater origin (hyperosmotics II or secondary confohyperosmotics) – 1% D1 – Some Caspian Brackish water hydrobionts (amphiosmotics I) – 0% D2 – Some euryhaline Australian hydrobionts of freshwater origin (amphiosmotics II) – 0% D3 – Euryhaline hydrobionts of freshwater origin (amphiosmotics III) – 1% D4 – Widely euryhaline hydrobionts of freshwater origin (amphiosmotics IV) – 0% E– Euryhaline marine hydrobionts of freshwater origin (hypoosmotics) – 20%

44. Percentage of different types of osmoconformers and osmoregulators in the World Ocean and fully saline seas as Barents Sea, Sea of Japan, etc.

45. Percentage of different types of osmoconformers and osmoregulators in brackish water seas as Black Sea, Sea of Azov, etc. A1 – Stenohaline marine hydrobionts (osmoconformers I) – 3% A2 – Marine hydrobionts (osmoconformers II) - 5% A3 – Euryhaline marine hydrobionts (osmoconformers III) – 10% B1 – Widely euryhaline marine hydrobionts (confohyperosmotics I) – 10% B2 – Brackish water hydrobionts of marine origin (confohyperosmotics II) – 15% C1 – Freshwater hydrobionts (hyperosmotics I) – 5% C2 – Brackish water hydrobionts of freshwater origin (hyperosmotics II or secondary confohyperosmotics) – 10% D1 – Some Caspian Brackish water hydrobionts (amphiosmotics I) – 5% D2 – Some euryhaline Australian hydrobionts of freshwater origin (amphiosmotics II) – 0% D3 – Euryhaline hydrobionts of freshwater origin (amphiosmotics III) – 5% D4 – Widely euryhaline hydrobionts of freshwater origin (amphiosmotics IV) – 2% E– Euryhaline marine hydrobionts of freshwater origin (hypoosmotics) – 30%

46. Percentage of different types of osmoconformers and osmoregulators in brackish water seas as Black Sea, Sea of Azov, etc.

47. Percentage of different types of osmoconformers and osmoregulators in freshwater lakes as Ladoga, Onega, etc. A1 – Stenohaline marine hydrobionts (osmoconformers I) – 0% A2 – Marine hydrobionts (osmoconformers II) - 0% A3 – Euryhaline marine hydrobionts (osmoconformers III) –0% B1 – Widely euryhaline marine hydrobionts (confohyperosmotics I) – 0% B2 – Brackish water hydrobionts of marine origin (confohyperosmotics II) – 0% C1 – Freshwater hydrobionts (hyperosmotics I) – 98% C2 – Brackish water hydrobionts of freshwater origin (hyperosmotics II or secondary confohyperosmotics) – 1% D1 – Some Caspian Brackish water hydrobionts (amphiosmotics I) – 1% D2 – Some euryhaline Australian hydrobionts of freshwater origin (amphiosmotics II) – 0% D3 – Euryhaline hydrobionts of freshwater origin (amphiosmotics III) – 0% D4 – Widely euryhaline hydrobionts of freshwater origin (amphiosmotics IV) – 0% E– Euryhaline marine hydrobionts of freshwater origin (hypoosmotics) – 0%

48. Percentage of different types of osmoconformers and osmoregulators in freshwater lakes as Ladoga, Onega, etc.

49. Percentage of different types of osmoconformers and osmoregulatorsin the Baltic Sea A1 – Stenohaline marine hydrobionts (osmoconformers I) – 0% A2 – Marine hydrobionts (osmoconformers II) - 0% A3 – Euryhaline marine hydrobionts (osmoconformers III) – 5% B1  – Widely euryhaline marine hydrobionts (confohyperosmotics I) – 15% B2  – Brackish water hydrobionts of marine origin (confohyperosmotics II) – 24% C1 – Freshwater hydrobionts (hyperosmotics I) – 14% C2 – Brackish water hydrobionts of freshwater origin (hyperosmotics II or secondary confohyperosmotics) – 9% D1 – Some Caspian Brackish water hydrobionts (amphiosmotics I) – 9% D2 – Some euryhaline Australian hydrobionts of freshwater origin (amphiosmotics II) – 0% D3 – Euryhaline hydrobionts of freshwater origin (amphiosmotics III) – 10% D4 – Widely euryhaline hydrobionts of freshwater origin (amphiosmotics IV) – 0% E– Euryhaline marine hydrobionts of freshwater origin (hypoosmotics) – 14%

50. Percentage of different types of osmoconformers and osmoregulators in the Baltic Sea