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1 mm Distribution of the Antarctic fish Pseudochaenichthys georgianus NORMAN, 1939 in the Atlantic sector of Antarctic. 1 mm 1 mm by Ryszard Traczyk 21.3 km/h ~0.9 km/h ~0.8 km/h

Otolith shape icefish

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17 minutes about changes of the otolith shape with the growth of the swimming speed and with the changes in the environment - sample to develop

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  • 1. 1 mm1 mmby Ryszard Traczyk21.3 km/h~0.9 km/h~0.8 km/h

2. Terrestrial observations of separate geographical and vertical living on different age groups andspecies of fish suggest that differences in otolith shape among them became from difference in theirenvironment conditions. (Extracted and enlarged otoliths are over or near the fish heads: Median or Transverse plane)S. Orkney SouthGeorgiaAntarctic Circumpolar0.1 km/hPs. georgianus Ch. aceratusWHITE-BLOODED: high Antarctic; ice pack zone; temperate80S 74, ~70S; 63, ~60S; 63, ~5730S; 52, ~45S 30S-100-200-300-400-500-600-700-800Current[m]S. japonicusChannichthyidaeMacrouridaeC. gunnariFisher, 1985; Kellermann, 1990; North 1990; Hecht, 198721 km/h1 km/h0.9 km/h 3. A S WAT E R T E M P E R AT U R E I S D R O P P I N GS. Orkney SouthGeorgiaAntarctic Circumpolar0.1 km/hPs. georgianus Ch. aceratusWHITE-BLOODED: high Antarctic; ice pack zone; temperate80S 74, ~70S; 63, ~60S; 63, ~5730S; 52, ~45S 30S-100-200-300-400-500-600-700-800Current[m]Decrease of Otolith Length, increase of Otolith HeightS. japonicusChannichthyidaeMacrouridaeC. gunnariincrease of Otolith LengthMand flattening (T plane)TMTMTTTTMMMMM21 km/h0.9 km/h1 km/h 4. Age groups of Ps. georgianus seperated in time scale confirm geographical divide ofotolith mass frequency as separate age groups on Antarctic islands: 2 aged at PamerArchipelago, in February, 3 aged at King George in March, and 4 aged and older fish atS. Orkney in December.4N4=102; MOr=0,0430,00051g;s=0,0026; TLr=47,1 cm; MCr=1113,7g;Gr=3,7; r=0,8N3=6; MOr=0,0320,00185g;s=0,0023; TLr=37 cm; MCr=453,3g;Gr=1,7; r=0,8N5=19; MOr=0,0520,00119g;s=0,0027; TLr=49,1 cm;MCr=1229,2g; Gr=3,4; r=0,57654321011109876543210Age groupNOW [g]South Orkney Is30.XII.1978N6=5; MOr=0,0670,00126 g;s=0,0014; TLr=50,8 cm;MCr=1471g; Gr=3,4; r=0,4N3=26;MOr=0,0340,00058g;s=0,0015;TLr=43,2cm;MCr=752,5g; Gr=2,3; r=1,7N4=9; MOr=0,0410,0011g;s=0,0017; TLr=47,6cm;MCr=1158,9g; Gr=4; r=2N5=3; MOr=0,0520,00233g;s=0,0021; TLr=52,3cm;MCr=1673,3g; Gr=3,3; r=2,787654321043210Age groupNOW [g]King George,25.III.1979N=39 N2=1; MO=0,021; TL=31cm;MC=210g; G=1; =2N3=62; MOr=0,0330,00045g;s=0,0018; TLr=42,1cm;MCr=740,6g; Gr=2,4; r=1,9N4=20; MOr=0,0410,00126g;s=0,0029; TLr=47,8cm;MCr=1224,8g; Gr=3,2; r=2,5N2=10; MOr=0,0230,00081g;s=0,0013; TLr=30,4cm; MCr=260g;Gr=1,5; r=1,2N6=1;MO=0,064;TL=51cm;MC=1750g;G=3; =376543210111098765432100.0100.0120.0140.0160.0180.0200.0220.0240.0260.0280.0300.0320.0340.0360.0380.0400.0420.0440.0460.0480.0500.0520.0540.0560.0580.0600.0620.0640.0660.0680.0700.072Age group0.074NOW [g]N5=4; MOr=0,0520,00151g;s=0,0015; TLr=50,8 cm;MCr=1548,8g; Gr=3,8; r=2Palmer A., Deception, Elephant Is:19-22.II.1979, N=97Traczyk, 2012 5. Age groups of Ps. georgianus seperated in time scale confirm geographical divide ofotolith mass frequency as separate age groups on Antarctic islands: 2 aged at PamerArchipelago, in February, 3 aged at King George in March, and 4 aged and older fish atS. Orkney in December.5N4=102; MOr=0,0430,00051g;s=0,0026; TLr=47,1 cm; MCr=1113,7g;Gr=3,7; r=0,8N3=6; MOr=0,0320,00185g;s=0,0023; TLr=37 cm; MCr=453,3g;Gr=1,7; r=0,8N5=19; MOr=0,0520,00119g;s=0,0027; TLr=49,1 cm;MCr=1229,2g; Gr=3,4; r=0,57654321011109876543210Age groupNOW [g]South Orkney Is30.XII.1978N6=5; MOr=0,0670,00126 g;s=0,0014; TLr=50,8 cm;MCr=1471g; Gr=3,4; r=0,4N3=26;MOr=0,0340,00058g;s=0,0015;TLr=43,2cm;MCr=752,5g; Gr=2,3; r=1,7N4=9; MOr=0,0410,0011g;s=0,0017; TLr=47,6cm;MCr=1158,9g; Gr=4; r=2N5=3; MOr=0,0520,00233g;s=0,0021; TLr=52,3cm;MCr=1673,3g; Gr=3,3; r=2,787654321043210Age groupNOW [g]King George,25.III.1979N=39 N2=1; MO=0,021; TL=31cm;MC=210g; G=1; =2N3=62; MOr=0,0330,00045g;s=0,0018; TLr=42,1cm;MCr=740,6g; Gr=2,4; r=1,9N4=20; MOr=0,0410,00126g;s=0,0029; TLr=47,8cm;MCr=1224,8g; Gr=3,2; r=2,5N2=10; MOr=0,0230,00081g;s=0,0013; TLr=30,4cm; MCr=260g;Gr=1,5; r=1,2N6=1;MO=0,064;TL=51cm;MC=1750g;G=3; =376543210111098765432100.0100.0120.0140.0160.0180.0200.0220.0240.0260.0280.0300.0320.0340.0360.0380.0400.0420.0440.0460.0480.0500.0520.0540.0560.0580.0600.0620.0640.0660.0680.0700.072Age group0.074NOW [g]N5=4; MOr=0,0520,00151g;s=0,0015; TLr=50,8 cm;MCr=1548,8g; Gr=3,8; r=2Palmer A., Deception, Elephant Is:19-22.II.1979, N=97 6. This same is for age groups in length frequency geographicallydivided on separated Antarctic Islands.30 N2520151050TL [cm]6543IIIIVVVIS. Orkney, 30.XII.1978; N = 142Theoretical; EmpiricalAge GroupN3=7;TL3=37,41,6 cm,N4=108;TL4=470,4 cm,s4=2,1N5=21;TL5=49,20,7s5=1,6 N6=6;TL6=511,32520151050NTL [cm]65432IIIIIIVVPalmer A., Deception, Elephant,18-22.II.1979, N=171;Theoretical; EmpiricalAge GroupN2=16; TL2=31,10,9 cm, s2=1,9N3=97;TL3=41,80,4s3=2N4=41;TL4=48,20,5s4=1,5N5=10;TL5=50,90,8s5=1,3N6=7;TL6=51,70,8s6=1,11614121086420NN4=34; TL4=48,50,5 cm,s4=1,4 N5=4;TL5=52,30,5 cm,s5=0,5TL [cm]2543IIIIIIVVPalmer A., Deception, King George,Elephant; 25.III.1979; N=145AgeGroupTheoretical; empiricalN2=24; TL2=30,80,7 cm, s2=1,7N3=83; TL3=420,5 cm,s3=2,425 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 7. This same is for age groups in length frequency geographically divided on separated AntarcticIslands.: 2 aged fish at Pamer A., in March, 3 aged fish dominated at Elephan in February, and 4aged and olders fish at South Orkney in December.30 N2520151050TL [cm]6543IIIIVVVIS. Orkney, 30.XII.1978; N = 142Theoretical; EmpiricalAge GroupN3=7;TL3=37,41,6 cm,N4=108;TL4=470,4 cm,s4=2,1N5=21;TL5=49,20,7s5=1,6 N6=6;TL6=511,32520151050NTL [cm]65432IIIIIIVVPalmer A., Deception, Elephant,18-22.II.1979, N=171;Theoretical; EmpiricalAge GroupN2=16; TL2=31,10,9 cm, s2=1,9N3=97;TL3=41,80,4s3=2N4=41;TL4=48,20,5s4=1,5N5=10;TL5=50,90,8s5=1,3N6=7;TL6=51,70,8s6=1,11614121086420NN4=34; TL4=48,50,5 cm,s4=1,4 N5=4;TL5=52,30,5 cm,s5=0,5TL [cm]2543IIIIIIVVPalmer A., Deception, King George,Elephant; 25.III.1979; N=145AgeGroupTheoretical; empiricalN2=24; TL2=30,80,7 cm, s2=1,7N3=83; TL3=420,5 cm,s3=2,425 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 8. Ps. georgianus from the younger age groups prefer the western part of the Atlantic Antarctic, andthe older age group East - North part. Age group is identified by otolith shape, which, indicate thedifferent habitats, development stages and strategies of life: type of swimming.8201816141210864221 %18151296310864220 %15105%6420King Edward I., N=7925 30 35 40 45 50 55TL [cm]Palmer Arch, N=5927.1%; TL=30.8cm; s=1.8 45.8%; TL=40cm; s=1.7 25.4%; TL=47cm; s=1.5 1.7%; TL=51.5cm; s=0.50TL [cm]12.3%; TL=30.6cm; s=1.660.7%; TL=41.7cm; s=2 20.9%;TL=47.9cm;s=1.44.3%; TL=50.4cm; s=0.81.8%; TL=51.1cm; s=0.2Deception I., N=1630%TL [cm]2.5%; TL=33.5cm; s=2.148.1%; TL=43.7cm; s=1.935.4%; TL=49.3cm;s=1.413.9%; TL=53.1cm; s=0.90TL [cm]S. Orkney, Western part, N=133 0.8%; TL=40.5cm;s=0.573.7%; TL=48cm;s=1.821.8%; TL=49.9cm; s=2.33.8%; TL=52.2cm;s=20%TL [cm]S. Orkney, Eastern part, N=1384.3%; TL=37cm; s=2.176.1%;TL=47.2cm; s=2.115.9%; TL=48.9cm; s=1.63.6%; TL=50.8cm;s=1.8 9. 9Ps. georgianus from the younger age groups prefer the western part of the Atlantic Antarctic, andthe older age group East - North part. Age group is identified by otolith shape, which, indicate thedifferent habitats, development stages and strategies of life: t y p e o f swimmi n g201816141210864221 %18151296310864220 %15105%6420King Edward I., N=7925 30 35 40 45 50 55TL [cm]Palmer Arch, N=5927.1%; TL=30.8cm; s=1.8 45.8%; TL=40cm; s=1.7 25.4%; TL=47cm; s=1.5 1.7%; TL=51.5cm; s=0.50TL [cm]12.3%; TL=30.6cm; s=1.660.7%; TL=41.7cm; s=2 20.9%;TL=47.9cm;s=1.44.3%; TL=50.4cm; s=0.81.8%; TL=51.1cm; s=0.2Deception I., N=1630%TL [cm]2.5%; TL=33.5cm; s=2.148.1%; TL=43.7cm; s=1.935.4%; TL=49.3cm;s=1.413.9%; TL=53.1cm; s=0.90TL [cm]S. Orkney, Western part, N=133 0.8%; TL=40.5cm;s=0.573.7%; TL=48cm;s=1.821.8%; TL=49.9cm; s=2.33.8%; TL=52.2cm;s=20%TL [cm]S. Orkney, Eastern part, N=1384.3%; TL=37cm; s=2.176.1%;TL=47.2cm; s=2.115.9%; TL=48.9cm; s=1.63.6%; TL=50.8cm;s=1.8Otolith, M-plane, after Hecht, 19870.4 km/h1.6 km/hand speed 10. Water temperature determine distribution of Antarctic fish. So older agegroups, fish 49 centimeters found at South Orkney Islands may havebeen resulted from lower temperature of waters below 0 up to minus 1Celsius degree. Younger age groups of 3 aged fish 43 cm found mostnumerous at Elephan have little warmer waters up to 0 degreescentigrade. The smallest fish 30 cm length of II age group appear atPalmer Archipelago have warm water of above 0 up to 1 Celsius degree.A, B, C - transects2Potential temperature [C] at 200 m110- 1Bransfield Strait- 1georgianus 34>3>4-2 -1 0 1 2 3 4 522.698.1South Orkney I.14.45.15658606264Feb.1979 the R/V Prof. Nov. 78 Feb. 79 the Ps. 64 60 56 52 48 44 B 11. Water temperature determine distribution of Antarctic fish.A, B, C - transects2Potential temperature [C] at 200 m110- 1Bransfield Strait- 1georgianus 34>3>4-2 -1 0 1 2 3 4 522.698.1South Orkney I.14.45.15658606264Feb.1979 the R/V Prof. Nov. 78 Feb. 79 the Ps. 64 60 56 52 48 44 B 12. A, B, C - transects2Potential temperature [C] at 200 m110- 1Bransfield Strait- 1georgianus 34>3>4-2 -1 0 1 2 3 4 522.698.1South Orkney I.14.45.15658606264Feb.1979 the R/V Prof. Nov. 78 Feb. 79 the Ps. 64 60 56 52 48 44 B 13. C. aceratus from larvae to fish of 2 year old, living in a warm pelagic water and preying krill,has weaker activity of AFGP proteins (equals to 0,21-0,41 C) than adult fish.0 year old feezingpoint drop by 0.21CTraczyk, 2013; Arkive nature.pl; Hureau, 19852 years old drop by0.41C of freezing point4.2 years drop by 0.57CBilyk 20110.45 km/h0.1 km/h1.8 km/h1.8 km/h 14. Adult fish at the age of 4.5 years, while descend to the colder depths of -1.47 Chave large activity of AFGP, that reduce freezing point of blood by 0.57C,0.45 km/h0.1 km/h1.8 km/hfood: fishC. aceratus 4 large swimming speed Arkive nature.pl; Hureau, 1985 15. that is accompanied with 4 times increase of their swimming possibility and diet change toeating fish. Young fish with delayed activation of AFGP protein can avoid cold water and swim to North, to waterArkive nature.pl; Hureau, 19850.45 km/h0.1 km/h1.8 km/hwithout ice.Protein activity of AFGP is suspended in icefish. It increases during adaptation of fishdevelopment stages to the transition from pelagic to bottom, or from warm to cold water. 16. It seems to be the role, because also concentration of protein activity AFGPfor the high Antarctic fish depends on the stage of development of the fish -synchronizing with space system of water temperature and food type (Whrmann1996).-100-200-300-400_500E a s t Wi n d D r i f tlarvae, 0 yearPleurogramma antarcticumcm, SL 22 cmspawning adultsI c es h e l f0.150.1330.2190.1090.170.1390.1010.1940.188larvae, 1 yearSL 6 cmjuvenes, SL 10 cmT < -0,5 CIce ShelvesWaterT < -2,0 Cjuvenes, 2 yearAFGP (% wet weight)high molar m. AFGP (% wet weight)low molar m. AFGP (% wet weight)Wohrmann, 19960.3 km/h 17. Concentration, the activity of the AFGP protein in the white bloodedfish increasing in colder water in the direction to the pole, and withS. Orkney SouthGeorgiaC. gunnari: 0,98C-1,85CCh. wilsoni: 1,29C-2,23CAntarctic CircumpolarPs. georgianus: 1,03C-1,91CColdwaterdescentActivity AFGP [C]Blood feeezing: [ C]the depth., Bilyk, 2011.Ch. hamatus: 1,45C-2,44CWHITE-BLOODED: high Antarctic; ice pack zone; temperate80S 74, ~70S; 63, ~60S; 63, ~5730S; 52, ~45S 30S-100-200-300-400-500-600-700-800CurrentC. aceratus: 0,54C-1,47C[m] 18. Geographic age groups in otolith shape, we can find being inwell agreement with their food size of krill migrating, driftingand growing with currents to the East. II age group were catch atPalmer Archipelago among juvenile krill. Adults fish, age IV andabove were cought at S. Orkney Island among adults krill.5860Fish and environment data: Sahrhage, 1988, Traczyk, 2012; Sahrhage, 1988; Witek, 1988; lsarczyk, 1985; White, 199864 60 56 52 48 44Orkady Pd.(A- adult, S- small,J- juveniles) migrationin the spawning period62spring - summer extended byice edge to the S. Orkney Is.64KrillPs. georgianus catchFeb.1979, R/V Prof. Siedlecki (N=67)Nov 78 Feb 79 M/T Sirius (N=30)SWeddell-Scotia ConfluenceIce edge 1 - 8 Jan 1989Ice edge 8 - 15 Jan 1989geostrophic currentKrill migrationsJSS6062S350 g/m3S SJAAAAJAJS SAAA ASJ90 g/m3200g/m3150 g/m373 g/m36464 60 56 52 48 44 19. 586064 60 56 52 48 44Orkady Pd.(A- adult, S- small,J- juveniles) migrationin the spawning period62spring - summer extended byice edge to the S. Orkney Is.64KrillPs. georgianus catchFeb.1979, R/V Prof. Siedlecki (N=67)Nov 78 Feb 79 M/T Sirius (N=30)SWeddell-Scotia ConfluenceIce edge 1 - 8 Jan 1989Ice edge 8 - 15 Jan 1989geostrophic currentKrill migrationsSahrhage, 1988, Traczyk, 2012JSS6062S350 g/m3S SJAAAAJAJS SAAA ASJ90 g/m3200g/m3150 g/m373 g/m36464 60 56 52 48 44 20. Thus on the NorthEast of Shetland Islands large adults Ps. georgianus,of age IV (and above) from King George to South Orkney have towithstand stronger surface currents and whirls.In opposite SouthWest side of Shetlands larvaes and small fish haveprotective environment on their shelves extended by ice edge zone to theSouth Orkney.586064 60 56 52 48 44JAS SAAAS South Orkney I.Krill migrations(A- adult, S- small,62J- juveniles) migrationin the spawning periodspring - summer extended byice edge to the S. Orkney Is.64Ps. georgianus catch:Feb.1979, R/V Prof. Siedlecki (N=67)Nov 78 Feb 79 M/T Sirius (N=30)Weddell-Scotia ConfluenceIce edge 1 - 8 Jan 1989Ice edge 8 - 15 Jan 1989Surface currents 200 mA150 g/m373 g/m36062AS SAS S6490 g/m3SJJJJS200g/m3A64 60 56 52 48 44 21. 21kg/h10080604020023 kg/h7814 kg/h 3 kg/h827473414069676665565571Ice edge connecting K. George, Elephant with S. Orkney, were located over mountainsthat combine undersea above islands. Undersea seabed belt of mountains and ridgesplay role of channel for conducting sea currents which carries krill and fish larvaes 22. Anon. 199022kg/h100755025023 kg/h78SGI - 198914 kg/h 3 kg/h827473414069676665565571larvaehaul No 40 41 55 56 65 66 67 69 71 73 74 78 82 larvaehaul No 40 41 55 56 65 66 67 69 71 73 74 78 82coastal demersal bottom shelf, deep-waterC. aceratus 5 1 Cr. antarcticus 1 1 4 1C. rastrospinosus 5 2 1 Neopagetopsis sp. 1C. wilsoni 1 3 2 3 bathypelagicT. eulepidotus 1 N. ionach 2 1 1L. larseni 1 1 N. coatsi 2 1P. macropterus 2 pelagic, oceanicNotothenia sp. 1 Pl. antarcticum 1 4coastal pelagic E. carlsbergi 20C. gunnari 1 E. antarctica 48 35Pagetopsis sp. 5 2 3 1 G. opisthopterus 1bottom shelf, seamounts - ice edge: 26.XII.88 - 8.I. 89; 9.I - 13.I.89D. eleginoides 2 Seamounts and ice combine S. Orkney with Elephant I. 23. 403020100z71; z73; z7473 71SL, mm29 mm, I.;158 mg;712 szt/h,11 kg/h2015105%27 mm, XII;139 mg; 537szt/h, 7 kg/h%25 mm, XII;107 mg; 25,3ml/1000m3SL, mmb39; b459; 30; 55; 86 % 20; 46; 63 %3; 19; 52 %15; 41; 56 %%z56; z65; z6649 mm; 983 mg;1394szt/h; 137 kg/h6 11 16 21 26 31 36 41 46 51 56656655566769404182SL, mm93 857874606137 mm; 392mg; 774 szt/h;30 kg/hSL, mm3-364 kg/h; II-III 2009; SGI=7535; 35 %z7852 mm; 1171 mg;3339 szt/h; 391 kg/h252015105%z40; z41302010bongonetXII.1996, SGI2-0C 34 at 20m)Current flow rel. to 5MPa surface [Dyn m].25Area of high krill abundance.30Secondary Frontal Zone.35.40.30.2047.82322.62003.2350.2415014.4.35.28.265.198.173South Orkney I.South Georgia I.S c o t i a S e a.20.19W e d d e l l S e a545658606264Elephant I.WSCPs. georgianus capture, [kg/h]Feb.1979 the R/V Prof. Siedlecki (N=67)Nov. 78 Feb. 79 the M/T Sirius (N=30)545658606264.31.309014krill density [g/m2] and cluster extending64 60 56 52 48 44 40 3648This system together with ice edge zone inhabited by krill and fish larvaes havegreat potential for migration of fish even for shelf species. Such conditions areoften extend to South Georgia 27. 43 42 41 40 39 38 37 36 35 345330'5430'5530'5330'haul. 42 N=23x=49,36 11 16 21 26 31 36 41 46 51 565430'haul 45,50x=45,5N=736 11 16 21 26 31 36 41 46 51 5655haul 48, 49, 54, 5730'x=34,3N=65haul. 345 10 15 20 25 30 35 40 45 50 5536 34373938454651525455>1C; 0C; 1C; 0C; 34 per 20 m)krill high density regionsSecondary Frontal Zone, SFZ0,30,30,35Krill 550g/m3Witek, 1988; Sahrhage, 1988Ps. georgianus has krill as dominant food (Sarah Clarke, 2008; Chojnacki, 1987). 30. 2.5 2.25 2 2 2.251.750.51.51.25 10.250.750.751 1204060801001201401601802000 0.4 0.5 0.6 0.7 0.8 1 1.25 1.5 1.75 2 2.25 2.5 315001000500Krill biomass [t/nm2]mWitek, 1988; Sahrhage, 1988 31. 43 42 41 40 39 38 37 36 35 3430'5430'5530'3130'5455ShagRocks500m500m150m43 42 41 40 39 38 37 36 35 3430'30'Currents at surface at 5MPa [Dyn m]WSC-Weddella-Scotia Confluence (s > 34 per 20 m)krill high density regionsSecondary Frontal Zone, SFZ0,30,30,35Krill 550g/m3Witek, 1988; Sahrhage, 1988Ps. georgianus has krill as dominant food (Sarah Clarke, 2008; Chojnacki, 1987). 32. 32Surface current 200 m40W 38WSea currents pattern (Murphy, 2013) 33. 33Surface current 200 m40W 38WSea currents pattern (Murphy, 2013) 34. 34Surface current 200 m40W 38W 35. 0 10 20 30 40 50 60~49 cm TL, 1275 g. 105 fish1-10.II. 198945 cm TL, 808 g.46 cm TL, 870 g.501502505000 100 200 300 400 500 600 70049 cm TL, 1275 g. 1-10.II. 1989Fish/km245 cm TL, 808 g.46 cm TL, 870 g.5015025050025 %20151050TL, cm50-150 m: n=71, TL=48,9 cmS.Georgia Is., 1-10.II.1989151050%TL, cm150-250 m: n=348, TL=46,3 cmS.Georgia Is., 1-10.II.1989151050%S.Georgia Is., 1-10.II.1989TL, cm250-500 m: n=252, TL=44,6 cm15 20 25 30 35 40 45 50 55 60 36. 0 10 20 30 40 50 60~49 cm TL, 1275 g. 105 fish1-10.II. 198945 cm TL, 808 g.46 cm TL, 870 g.501502505000 100 200 300 400 500 600 70049 cm TL, 1275 g. 1-10.II. 1989Fish/km245 cm TL, 808 g.46 cm TL, 870 g.5015025050025 %20151050TL, cm50-150 m: n=71, TL=48,9 cmS.Georgia Is., 1-10.II.1989151050%TL, cm150-250 m: n=348, TL=46,3 cmS.Georgia Is., 1-10.II.1989151050%S.Georgia Is., 1-10.II.1989TL, cm250-500 m: n=252, TL=44,6 cm15 20 25 30 35 40 45 50 55 60 37. Surface current 200 m40W 38WOn the north eastern side of the island large fish supposed to be wellswimming, with using and opposing whirls (threatening to carry off from theshelf) could persist in them, and feed on accumulated in these whirls largespecimens of krill. Separated geographically age groups were differentiatedby the ability to swim in the currents and eddies needed to get food in them.This ability is managed by otolith shape. Why? 38. Surface current 200 m40W 38WSeparated geographically age groups were differentiated by the ability toswim in the currents and eddies needed to get food in them. This ability ismanaged by otolith shape. Why?swimming speed 39. 3921 2, ( ) R C Sv a h x 2SHydrodynamic resistance Ra,h is the smallest for flowing shape = 1 40. 4021 km/h21 km/h1 2, ( ) R C Sv a h x 2S0.001 km/hHydrodynamic resistance Ra,h is the smallest for flowing shape = 1thanks to have it, fish mastered ocean space.Success in obtaining food provides higher speed of swimming. 41. 4121 km/h21 km/h1 2, ( ) R C Sv a h x 2S0.001 km/hHydrodynamic resistance Ra,h is the smallest for flowing shape = 1thanks to have it, fish mastered ocean space.Success in obtaining food provides higher speed of swimming.This has also a reference to the otoliths provide in swimming:- balance and precision. 42. 42The flattened shape of the otoliths poses littleresistance in endolymph and increases:the perception of positions in fast swimming.21 km/h21 km/h1 2, ( ) R C Sv a h x 2S0.001 km/hHydrodynamic resistance Ra,h is the smallest for flowing shape = 1thanks to have it, fish mastered ocean space.Success in obtaining food provides higher speed of swimming.This has also a reference to the otoliths provide in swimming:- balance and precision.flatened otolith ofMackerelround otolith of medusabidirectionalotolith of squid 43. 43The flattened shape of the otoliths poses littleresistance in endolymph and increases:the perception of positions in fast swimming.21 km/h1 2, ( ) R C Sv a h x 2S21 km/h0.001 km/hHydrodynamic resistance Ra,h is the smallest for flowing shape = 1thanks to have it, fish mastered ocean space.Success in obtaining food provides higher speed of swimming.This has also a reference to the otoliths provide in swimming:- balance and precision.flatened otolith ofMackerelround otolith of medusabidirectionalotolith of squid 44. Otoliths have changes of shape from spherical to more elongated shapeduring the ontogenyThe surface of the otoliths is plastically formed by labyrinth and by measured thechanges of endolymphatic pressure induced by activity. Changes of the pressure in theendolymph arrange her ingredients from which at start of that changes in stationarylarvae they assembling into spherical otoliths.Swimming development in fish. asc, psc, lsc anterior, posterior, lateral semicircular canals, c cristae, l lagena, ml, ms,mu macula lagenae, sacculi, utriculi, s saccule, u utriculi, ed endolymphatic duct, c cochlea, bm basilarmembrane, pb papilla basilaris.muu44sensorialmicrovilliVestibular nervefiberslml21 cm TLcpsc asclscms smsdrift swimmingslow swimmingmsstationary fast swimming~0.01 km/h ~0.1 km/h ~0.3 km/h 45. as the speed of fish swimming increase4521 cm TL 46. 460.1 mmpostlarvaeotolithTraczyk, 2013 47. For Ps. georgianus the mark of that shift is as wide layer47Changing environment and physiology in hatching is aslarge as it is clearly marked in the microstructure.having more calcium that washed out separates thehatching nucleus from the rest of the otolith.LN, seperated larval microstruktureeasy disamble by EDTAfrom other parts of otolithTraczyk, 2013 48. Further flattening of otoliths runs from postlarvae to fish of I age group (2.8 to 3)by Second Primordium. Postlarvae swims faster than larvae as begins to migrate tothe waters further from shore and deeper, their otoliths become more flattered.Postlarvae drifting in coastal current accumulate on west side of island.1 0 . 30.6 0 .4 0.2 0 0.2 0.4 0.61.2 0 .8 0.4 0 0.4 0.8 1.2 1.62.81SP1 mmLN12.8AP 49. SEM2 nm platinum + palladiumSecond Primordium increase the flattening ofotolith and decrease its front profileSecond Primordium2.80.1 mm110. 3Traczyk, 2013 50. The increase in the otolith flattening by Second Primordium on medial cross-sectiongive the widest surface which in move of fish in dorsal directionincrease the perception of balance during vertical migration110.6 0 .4 0.2 0 0.2 0.4 0.61.2 0 .8 0.4 0 0.4 0.8 1.2 1.68. 23.0SP1 mmLN12.8 51. Inside egg inshore outshore deep water, below 200 m development0 1 2 3 4 5 6 7 8 9 10 mmAge group : 0 I II III IV V VI 52. OH= 0,8248OL1,3048R = 0,9116OH = 0,8064ORL + 1,9359R = 0,9382OH = 1,2623OLJ - 0,2123R = 0,9277876543210OL, mmOH, mmOH otolith height of adultsOLJ otolith lenght of juvenesORL otolith rostral length0 1 2 3 4 5 6 7TL = 4,2463e0,3834OHR = 0,9819TL = 3,3334e0,5477OLR = 0,96211,0356TL = 10,632R9R = 0,9805TL = 9,382ORL - 4,2732R = 0,95436050403020100OH height of adult otolithOLJ otolith length of juvenesOL otolith lengthORL otolith rostral lengthR9 dorsal radiusTL, cm0 1 2 3 4 5 6 7mmResults are in agegroupsOH > OL:OH=bOL+aabove y=xproportionality of the otolith dimensions are constantAnteriorculliculum 53. OH= 0,8248OL1,3048R = 0,9116OH = 0,8064ORL + 1,9359R = 0,9382OL, mmTL = 4,2463e0,3834OHR = 0,9819TL = 3,3334e0,5477OLR = 0,96211,0356OH > OL:OH=bOL+aabove y=xOH otolith height of adultsOLJ otolith lenght of juvenesORL otolith rostral lengthproportionality of the otolith dimensions are constantTL = 10,632R9R = 0,9805TL = 9,382ORL - 4,2732R = 0,9543876543216050403020100OH height of adult otolithOLJ otolith length of juvenesOL otolith lengthORL otolith rostral lengthR9 dorsal radiusAnteriorculliculumOH, mmTL, cm0 1 2 3 4 5 6 7mmResults are in agegroupsOH = 1,2623OLJ - 0,2123R = 0,927700 1 2 3 4 5 6 7 54. OH= 0,8248OL1,3048R = 0,9116OH = 0,8064ORL + 1,9359R = 0,9382OL, mmTL = 4,2463e0,3834OHR = 0,9819TL = 3,3334e0,5477OLR = 0,96211,0356OH > OL:OH=bOL+aabove y=xOH otolith height of adultsOLJ otolith lenght of juvenesORL otolith rostral lengthproportionality of the otolith dimensions are constantTL = 10,632R9R = 0,9805TL = 9,382ORL - 4,2732R = 0,9543876543216050403020100AnteriorculliculumOH, mmTL, cmOH height of adult otolithOLJ otolith length of juvenesOL otolith lengthORL otolith rostral lengthR9 dorsal radius0 1 2 3 4 5 6 7mmResults are in agegroupsOH = 1,2623OLJ - 0,2123R = 0,927700 1 2 3 4 5 6 7 55. Inside egg inshore outshore deep water, below 200 m development0 1 2 3 4 5 6 7 8 9 10 mmTraczyk, 2012; Traczyk, 2013Age group : 0 I II III IV V VI 56. 0 1 2 3 4 5 6 7 8 9 10 mmAge group : 0 I II III IV ~0.3 km/h ~ 0 . 9 km/h V VI 57. How do we know that the shape of otolith indicates the speed ofswimming? In comparison: faster species have them more flatterS. japonicusPs. georgianusfaster speciesTransverseplanemedian planemedian planeTransverseplane1 mm 1 mmData for speed of swimming (bikowski, 2008, Fuiman, 2002)21.3 km/h~1,6 km/h 58. The high otoliths inform about vertical stability needed for vertical migrations and for lifting with currents.Long otoliths of mackerel are sensitive on changes during swimming in the horizontal direction. Informationimportant in the fast swimming for far distances.Ps. georgianus1 mmrostrumTransverse planeone side incrementsChanges in thewidth incrementson R3, R11nucleus Y 0,000775 mmY 0,000276 mmnucleusOuter dorsal side 59. the shape of otolith is chageing among species of fish of differentdepth of livingCh. aceratusChannichthyidaeDeep water species M. holotrachysMacrouridaeshelves speciesTransverseplaneTransverseplane1 mm1 mmBody fish and otolith shape data (Hecht, 1987; Fischer, 1985; Grabowska, 2010; Traczyk, 1992) 60. the shape of otolith is chageing among species of fish of differentdepth of livingCh. aceratusDeep water species M. holotrachysMacrouridaeTransverseplaned e p t h1 mmBody fish and otolith shape data (Hecht, 1987; Fischer, 1985; Grabowska, 2010; Traczyk, 1992)Channichthyidaeshelves speciesTransverseplane1 mm_500 m_1200 m 61. Swimming depth is source of diversity in microstructure61and shape of the otolithTransverseplaneOtolith shape data Grabowska, 2010 62. 62C. aceratusproportions inversedDorsal margin R9 against the pressure doesnot rise, but increases ventral R11 in thedirection of pressure and most increase R3,7(pressure gradient = 0).M. carinatusdimensions ofradii: is small fordorsal part butlarge ventral forM. carinatus andvice wersa forChannichthyidae 63. Otolith shape differentiates pattern ofhigh energy swimming of mackerelScombrus japonicus ~21.3 km/hSquids 1-3 km/hpulsed swimming in squidsbikowski, 2008; Videler J.J., 1984Gosline, 1985 64. low energy swimming of icefish Channichthyidaeby using large pectoral finsfloating fish in the depths;The accuracy of vertical movements,are measured better by higher otolith.1 mmHigh, laterally flattened body having a great fins has about 20 times more resistance of the lateralthan the front and the current pressure on the concave side of curved body of flowing fishproduces a hydrodynamic force increasing speed of fish swimming forward. An asymmetricalshape with respect to the direction axis of swimming causes asymmetric flow, that createsdifferential pressure on opposite surfaces, and thus the driving force to forward.(Fuiman L, 2002; Anon, 2006)m i g r a t i o n 65. In lifting strategy and low-energy swimming of Ps. georgianus pectoralfins are moving their first rays as spars entailing the sheet of streamer.Forward with a minimum resistance of sheets fins flowing after trace ofthin first ray and back with a large opposition of all returning fin surface.Pectoral fins in the first phase of motion, horizontal spreading out to thefront and to the sides increase the horizontal plane of fish so keep, supportfish to float at required depth level.In the second phase the fins retractedhorizontally to the rear are pushing its allsurface on water and pushing fish forward.Also locomotor activity have a caudal finbut much smaller. Fin is bent on sidewayswith the body when fish is turning. Le Franois, 2014 66. Le Franois, 2014 67. Le Franois, 2014 68. Le Franois, 2014 69. Le Franois, 2014 70. Le Franois, 2014 71. Le Franois, 2014 72. Le Franois, 2014 73. low energy swimming of icefish Channichthyidaeby using large pectoral finsfloating fish in the depths;The accuracy of vertical movements,are measured better by higher otolith.1 mmHigh, laterally flattened body having a great fins has about 20 times more resistance of the lateralthan the front and the current pressure on the concave side of curved body of flowing fishproduces a hydrodynamic force increasing speed of fish swimming forward. An asymmetricalshape with respect to the direction axis of swimming causes asymmetric flow, that createsdifferential pressure on opposite surfaces, and thus the driving force to forward.(Fuiman L, 2002; Anon, 2006)m i g r a t i o n 74. 90L i f t i n g s t r a t e g ym i g r a t i o n 75. (abrowski, 2000)91The use of currents and the uplift force of fins.m i g r a t i o n(Le Franois, 2014; 2014a; PolarTrec, 2013; Detrich, 2012; (Walesby, 1982; Davison W., 1985; HARRISON, 1987; Twelves, 1972) Uve, 2008; Byrd, 2012) 76. High laterally flattened body takes over energy of sea current.An asymmetrical shape creates differential pressure onopposite surfaces, and thus the driving force to forward.V speed of the fish,Rh,c front resistRh,b side resist = 22 VFPFCFAEAnon, 2006Rh,c = 1,5 FAE aero-hydrodynamic force - the forceexerted on the body by the environment,which is the result of movement of the bodyrelative to the environment (gas or liquid).FC driving force, thrust (force of pressure induced by pressure of current exerted on the bodysurface area). Operates forward because of body shape an the resistance of the lateral is 20 timesgreater than the frontal; FP - drift force; viscosity force (friction at the surface of the body). 77. Fac tor s inc reas ing the for ce (Fa ,h ) ac t ing on the body .Force: Fa,h = kv2; power: Pa,h = kv3; 2Vcurrent 4FAEGrowing of latteral surfaces of fishbodyLarger, stronger ones are occurringcloser to the sea surface, where thecurrents are stronger withturbulences and eddys. Smaller fishso weaker live deeper where thecurrents are weaker and also inregions with weak currents,VFP2FAEFCRh,c = 1,5Anon, 2006Icefish have adaptation to cold water. Oneof them Ps. georgianus live and choosehabitat of sea currents so to exist in it, itadopt the shape of the body, fins and otolithsin liftting strategy of low-energy swimming 78. The smooth surface of the body increases the power of aero-hydrodynamic FA, HChannichthyidae have a smooth skin, without scales, allowing the feeling of eachparticle of the water flowing and gliding over the surface of the skin and reactaccordingly by deflection of the body, or by rearrangement positions of fins toreduce the resistance, to increase laminar flow and to eliminate turbulences. Lackof scales could be adopted as an adaptation of a low-energy swimming in coldstrong currents, for which in a warm water there is high energy swimming. Forexample. Salmon, trout, or mackerel. We can find that the lack of scales for icefishis treat as an adaptation to increase the respiration of skin. Jakobowski however,argues that such a view is wrong, because the scales are below the epidermis towhich oxygen diffuses and therefore scales do not interfere with the diffusion ofoxygen through the skin. Certainly scaleless increase skin smoothness. Jakubowski, 1971, 198294 79. The sensitivity and skin elasticity in the perception of the body bendingVFPFCFAERh,c = 1,5The bending body must always be tailored to the nature of the currents. Too big bow causes breakaway water streams from the surface of the body, for small bow quite similar paths and velocityof water particles on both sides of the odd fins causing a lack of hydrodynamic forces.When the stream of water on the side of after current detach and move disordered (turbulent), thiswill reduce the hydrodynamic forces. 80. The fins increase the smoothness and flow velocity of after currentside of the bodyFAERh,c = 1,5VAfter the first front dorsal fin and before the second dorsal fin creates the nozzle that accelerateswater flow on after currant side of second fin and body.Body shape as an indicator of the shape of otolith, because it results from thespeed of swimming and life strategy and that all adapted to environmentalconditions. This could be show by compare of species. 81. Body shape as an indicator of the shape of otolith,because it results from the speed of swimming and lifestrategy and that all adapted to environmentalconditions.This could be show by compare of species. 82. Ps. georgianus has greater body height than the C. gunnari andC. aceratus body highest, large headand jaws definespredator creates anarrow, pelvic fins largeeffective for verticalmigration. Longer,unpaired fins increasebody side resistance .body less high, but fins:anal and dorsal longer ,smaller head largerpectoral fins so largerhorizontal migrationsBody less high, but verylarge anal fin, dorsal andpectoral, so the greatesthorizontal migrations.The smallest headreduces front resisting98Data: Fisher, 1985 when swimming. 83. Data: Fisher, 1985; Traczyk, 2013, Parkes, 1990y = 0.2251x - 0.0242R = 0.9798cmy = 0.3985x0.9976R = 0.99712018161412108642015-30% SL SL, cm0 10 20 30 40 50 604038363432302826242220181614121086420C. gunnariC. aceratusC. aceratusC. gunnariC. gunnariC. aceratusPs. georgianusSL, cmNPs. georgianus0 10 20 30 40 50 60Ps. georgianus has greater body height than the C. gunnari and C. aceratus also haveshorter dorsal and anal fins in favor of head size andthe decline of swimming opportunities. Otoliths of Ps. georgianus like its body are high.Shape of otoliths (determined by microstructure) can show the process and direction ofgrowth of the body, which is a response to factors of surrounding marine environment.Increase of otoliths taking place outside the cell in endolymph suffers from it the samefactors of the marine environment reaching endolymph through the bones of the body.Therefore, the body and otoliths become a models of the fish growth as reader of sufferedenvironmental influences - to which fish during growth is adapting the otolith shape bychanges of microstructure of the otoliths, that are treat as indicators of fish behavior. 84. Ps. georgianus has a smaller range of occurrence but higher vertical migration than C. aceratusC. aceratus have longer otoliths and has a greater range of occurrence than Ps. georgianus.100Ps. georgianus, otolith height OH> otolith length OL, TL body lengthC. aceratus, OH< OL, TLData: Hecht, 1978 85. Ps. georgianus has smaller range of occurrence than the C. gunnari101Ps. georgianus, otolith height OH> otolith length OL, TL body lengthC. gunnari: OH < OLData: Hecht, 1978 86. 3OH/OTflattening 9y = 0.1585x + 1.715387654321032.521.51mackerelSL, cm0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16OH otolith heightOT otolith thicky = 0.2521x + 5.2326y = 0.0339x + 1.089898765432102.521.51OH/OTScale for mackerelSL, cm0 10 20 30 40 50Data: Traczyk, 2012 87. 987654321OH/OTScale for mackerel1033OH/OTflattening 9y = 0.1585x + 1.715387654321032.521.51mackerelSL, cm0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16OH otolith heightOT otolith thicky = 0.2521x + 5.2326y = 0.0339x + 1.089802.521.51SL, cm0 10 20 30 40 50 88. S.Georgia I.Shag RockS.Sandwich I.S. Orkney I.Elephan I.K.George I.DeceptionPalmer A.Known catch location of fish:Ps. georgianus: 0-475 m; 53S-66SS. Sandwich I. Germany 1975/76; 1980/81Kerguelen I. Australia 2003/04BallenyKerguelen I.Bouvet I.Heard I.Ch. aceratus: 5-770 m; 53S-65SCh. gunnari : 0-700 m; 48S-66SUnusual catch location of Ps. georgianusBalleny I. Russia 2004/05 89. S.Georgia I.Shag RockS.Sandwich I.S. Orkney I.Elephan I.K.George I.DeceptionPalmer A.Known catch location of fish:Ps. georgianus: 0-475 m; 53S-66SS. Sandwich I. Germany 1975/76; 1980/81Kerguelen I. Australia 2003/04BallenyKerguelen I.Bouvet I.Heard I.Ch. aceratus: 5-770 m; 53S-65SCh. gunnari : 0-700 m; 48S-66SUnusual catch location of Ps. georgianusBalleny I. Russia 2004/05? 90. S.Georgia I.Shag RockS.Sandwich I.S. Orkney I.Elephan I.K.George I.DeceptionPalmer A.Known catch location of fish:Ps. georgianus: 0-475 m; 53S-66SS. Sandwich I. Germany 1975/76; 1980/81Kerguelen I. Australia 2003/04BallenyKerguelen I.Bouvet I.Heard I.Ch. aceratus: 5-770 m; 53S-65SCh. gunnari : 0-700 m; 48S-66SUnusual catch location of Ps. georgianusBalleny I. Russia 2004/05? 91. )) .2y ( A sin(i91x constTiiiR9=2,35 mm1590daysCh. aceratus, 45 cm SLS. Georgia , 29.III.1979hol 136, No 75OW=0,0247 gOH=3,44mmSP APC. aceratus body less high, but fins are longer.Additional centers, AP are also available in otoliths ofC. aceratus.. They give however a lower elongationthan the radius R9 of Ps. georgianus. Dorsal edge forotolith of older fish of Ps. georgianus grows morestrongly than in otoliths of C. aceratus..Otoliths of greater length than height indicate agreater range and speed of swimming. This confirmsthe elongated shape of the body with less weight Asotoliths of C. aceratus are not high, so height of theirbody is reduced. Data: Traczyk, 1992; 2014 92. Realization various opportunities of swimming arising from variousconstructions of body that are adapted to the best use of differenthabitats of environment allows the perception of this swimming byotolith recording it with appropriate shape.Traczyk, 2013; Traczyk, 1992; Fischer, 1985Otoliths of C. aceratus are longer than hights.In otoliths of C. aceratus proportion: lengthwith respect to height is reversed. Ps.georgianus are smaller, TLy = 1.0484x1.1338R = 0.9877OH, mmy = 0.8064x + 1.9359R = 0.9382y = 0.9743x0.9426R = 0.991876543210OH Ps. georgianusORL Ps. georgianusOH C. aceratusOL, mm0 1 2 3 4 5 6 7TL, cmy = 6.6517e0.9547xR = 0.9889y = 6.7436e0.4989xR = 0.9909y = 7.3835e0.424xR = 0.9718706050403020100~OLPs. georgianusOH Ps. georgianusOL Ps. georgianusR9 Ps. georgianusOH C. aceratusOL C. aceratusR9 C. aceratusmmC. aceratus0 1 2 3 4 5 6 7 93. S.Georgia I.Shag RockS.Sandwich I.S. Orkney I.Elephan I.K.George I.DeceptionPalmer A.Known catch location of fish:Ps. georgianus: 0-475 m; 53S-66SS. Sandwich I. Germany 1975/76; 1980/81Kerguelen I. Australia 2003/04BallenyKerguelen I.Bouvet I.Heard I.Ch. aceratus: 5-770 m; 53S-65SCh. gunnari : 0-700 m; 48S-66SUnusual catch location of Ps. georgianusBalleny I. Russia 2004/05 94. S.Georgia I.Shag RockS.Sandwich I.S. Orkney I.Elephan I.K.George I.DeceptionPalmer A.Known catch location of fish:Ps. georgianus: 0-475 m; 53S-66SS. Sandwich I. Germany 1975/76; 1980/81Kerguelen I. Australia 2003/04BallenyKerguelen I.Bouvet I.Heard I.Ch. aceratus: 5-770 m; 53S-65SCh. gunnari : 0-700 m; 48S-66SUnusual catch location of Ps. georgianusBalleny I. Russia 2004/05? 95. S.Georgia I.Shag RockS.Sandwich I.S. Orkney I.Elephan I.K.George I.DeceptionPalmer A.Known catch location of fish:Ps. georgianus: 0-475 m; 53S-66SS. Sandwich I. Germany 1975/76; 1980/81Kerguelen I. Australia 2003/04BallenyKerguelen I.Bouvet I.Heard I.Ch. aceratus: 5-770 m; 53S-65SCh. gunnari : 0-700 m; 48S-66SUnusual catch location of Ps. georgianusBalleny I. Russia 2004/05? 96. Otoliths Height is less than Otolith Length, indicating a wider occurrence andgreater speed of swimming. It confirms the elongated body with a lower height.Otoliths C. gunnari nearly square, two times smaller than otoliths Ps. georgianus.TL, cmy = 9.5723x1.2236180R = 0.98431,52,5y = 8.6308x1.2495R = 0.9848y = 21.245x1.2292R = 0.98686050403020100OH Ps. georgianusOL Ps. georgianusR9 Ps. georgianusOH C. gunnariOL C. gunnariR9 C. gunnari0 1 2 3 4 5 6 7mmOL > OH1,07 : 16,415,3TL, cm2427,83445 5035y = 0.92x1.0199R = 0.9979y = 0.4834x1.0103R = 0.985943.532.521.510.50mmOL, otolith length [mm]0 0.5 1 1.5 2 2.5 3 3.5 4Data: Hecht, 1978;Traczyk, 2013; 2014 97. R9=0,046 mm, 48 daysR9=2,35mm1590daysThe otoliths shape of larvae is similar to an oval onmedian plane and flattened on the transverse planeto reduce resistance. The biggest flattened otolithhas C. gunnari so it swims the fastest andfarthest. Older fish swim faster, so flatteningof its otoliths increases.C. aceratus, 45 cm SLS. Georgia , 29.III.1979hol. 136, s. 75OW=0,0247 g. OH=3,44 mmAPSPData: Traczyk, 20130.1 mmC. gunnari - otolith most flattened 98. R9=0,046 mm, 48 daysR9=2,35mm1590daysThe otoliths shape of larvae is similar to an oval onmedian plane and flattened on the transverse planeto reduce resistance. The biggest flattened otolithhas C. gunnari so it swims the fastest andfarthest. Older fish swim faster, so flatteningof its otoliths increases.C. aceratus, 45 cm SLS. Georgia , 29.III.1979hol. 136, s. 75OW=0,0247 g. OH=3,44 mmAPSPData: Traczyk, 20130.1 mmC. gunnari - otolith most flattened 99. S.Georgia I.Shag RockS.Sandwich I.S. Orkney I.Elephan I.K.George I.DeceptionPalmer A.Known catch location of fish:Ps. georgianus: 0-475 m; 53S-66SS. Sandwich I. Germany 1975/76; 1980/81Kerguelen I. Australia 2003/04BallenyKerguelen I.Bouvet I.Heard I.Ch. aceratus: 5-770 m; 53S-65SCh. gunnari : 0-700 m; 48S-66SUnusual catch location of Ps. georgianusBalleny I. Russia 2004/05? 100. S.Georgia I.Shag RockS.Sandwich I.S. Orkney I.Elephan I.K.George I.DeceptionPalmer A.Known catch location of fish:Ps. georgianus: 0-475 m; 53S-66SS. Sandwich I. Germany 1975/76; 1980/81Kerguelen I. Australia 2003/04BallenyKerguelen I.Bouvet I.Heard I.Ch. aceratus: 5-770 m; 53S-65SCh. gunnari : 0-700 m; 48S-66SUnusual catch location of Ps. georgianusBalleny I. Russia 2004/05? 101. The shape of the otoliths Channichthyidae:Ps. georgianus, C. aceratus and C. gunnari are similardue to the similar strategy of swimming they have asimilar body shape. However that species have a littledifferent otolith shape and body what indicate 102. Why are there swimming differences, where are the causes and how it is go?C. gunnari:Together or parallel with their evolution. Bilyk, 2011; Whrmann, 1996; Clarke A., 1996; Cheng, 1999; Chen, 250)Ch. wilsoni: 1,29C 0,98C-2,23CAntarctic CircumpolarPs. georgianus: 1,03C-1,91CColdwaterdescentCh. hamatus: 1,45C-2,44CWHITE-BLOODED: high Antarctic; ice pack zone; temperate80S 74, ~70S; 63, ~60S; 63, ~5730S; 52, ~45S 30SActivity AFGP [C]Blood feeezing: [ C]S. Orkney SouthGeorgia-100-200-300-400-500-600-700-800CurrentC. aceratus: 0,54C-1,47C[m]increase in activity and production of antifreeze proteins-1,85Cincrease in activity and production of AFGPBilyk 2011 103. All species of Channichthyidae have lost hemoglobin(that reduced oxygen transport) Jakubowski, 1971; Near, 2010; Everson, 1977; Fisher, 1985S. Orkney SouthGeorgiaAntarctic CircumpolarCh. aceratusC. gunnariWHITE-BLOODED: high Antarctic; ice pack zone; temperate80S 74, ~70S; 63, ~60S; 63, ~5730S; 52, ~45S 30S-100-200-300-400-500-600-700-800Current[m]Ch. hamatus:Nototheniidae; 0,4; 2,5P. antarcticum[mln./mm3]; [g/100 ml]Nototheniidae; 0,8, 8,3S. japonicus: 4, 18Channichthyidae:0; 0Ch. esox: 0; 0L. squamifronsPs. georgianus0; 0Bathydraconidae: 0,2; 0,8Macrouridae; 0,99, 3,90; 00; 0 0; 0 104. but each species has a different compensation of reduced oxygen transport to fulfill strategyof swimming speed to different environment(what differentiates the shape of the body and the otoliths. Icefish have different lifestrategies and occupy different habitats, La Mesa M., 2004)At first larger body, so they have reduced heat loss due to the lower surface. Bergmansrule of energy benefits.412Mass = 1 Mass=8Area = 28 Area = 112Area/mass = 28 Area/mass = 14additional oxygen from skin breathing of large head for the heart (Detrich, 2012).Channichthyidae achieve larger body size.248Jakubowski, 1977; Wells, 1985; Kock, 1991; Johnston, 1983eggs, mm Larvae, mm Adult, cm incr, cm/yWhite-blooded Channichthyidae 5 4-17 43 6-10Red-blooded Nototheniidae 36Red-blooded Bathydraconidae 26Red-blooded Harpagiferidae 15 105. Bottom lifestyle of Channichthyidae. Chaenocephalus aceratus has the body weakwith a reduction in the axial muscles, which is probably due to the large volume of blood, up to9% of body weight usually poorly vascularised.Jakubowski, 1977; Wells, 1985 106. White-blooded also have a reduction in ossification as a result of displacement of benthic fishfauna from shelf by glaciers to greater depth or to the pelagic. It enforced a reduction in bodyweight, because the white-blooded does not have a swim bladder.abrowski, 2000; Byrd, 2012Walesby, 1982; Jakubowski, 1977 107. Icefish larvae. Bone mass replaced by cartilage give large head.Additional oxygen from skin breathing for the heart (Detrich, 2012; Kils, 2008). 108. Pelagic life of Channichthyidae. Lack of myoglobin, which enhances oxygen diffusion by 600%should limited locomotion activity of muscles becase lack of oxygen. It is not for C. gunnari, whichincreases oxygenation by larger capillaries and large spaces supplying capillaries with blood.Walesby, 1982; Twelves, 1972C. gunnari 109. It is agree with Allen's Rule for energy benefits of having a more slender (less resistance)in the warm waters.The activity of alkaline phosphatase determines the size of the muscle vasculature. Reis, 1970.Greatest is in C. gunnari hence this species has m o r e c a p i l l a r i e s .1s [enzyme units/g wet muscleh]species / muscle type pectoral fin oxidative glycolyticC. gunnari, n=7, 8, 8 1185,5101,4 770,062,5 400,043,4Notothenia rossii, n=8, 9.698,1118,5483,271,0310,037,6Gadus morhua, n=10405,837,5353,565,1100,319,4Its s l ende r body shape increases heat loss, by Allens rule.166Mass = 8Area = 192+12+32=236Area/mass = 236/8=29,5Mass=8Area = 96+48+16=160Area/mass = 160/8=2022412 Ps. georgianus C. gunnari 110. O2, o-oxidative fibres, slow reaction -actATP: 2-5lowermore and larger capillaries and large spaces with bloodN. rossii C. gunnari340 155Gglycolytic fibres transport more efficient( large)N. rossii 75reduces distance and amount diffusion O2320 C. gunnari small, many capillaries, lipids, glycogen, SDHenzymeWalesby, 1982 111. Low-energy swimming is on the pectoral fins only? No!ChannichthyidaeC. gunnariNototheniidae AntarcticN. rossiimyomer muscule pectoral fin muscleHigh energy swimming by body wavesGadidaePollachius virensWhitefibers,fastreactionoxi 1oxi 2Oxi 1Oxi 2 mosaiclateral linechannelswimming on the pectoral fins savesmore energy (than by body waving),because the muscles of that fins areslow oxidative fibers adapted tocontinuous low-intensity movements(consuming less energy), ensuring thelong-term swimming at low speed.C. gunnari has also oxidative andglycolytic fibers in axial muscles.mosaic ofwhite andred fibersred fibers oxidative slow reactionoxi 1 & oxi 2SalmonidaeOncorhynchus mykissAltringham, Ellerby, 1999; Davison i Macdonald 1985, Harrison i in. 1987, Walesby 1982 112. Semipelagic lifestyle of Channichthyidae.Ps. georgianus has a shape more compact than C. gunnari andin a result (by no change in body weight) it reduces the ratio ofits body surface to the body weight, hence from that reducingloss of heat through the smaller surface of the body - whichis important in cold water.It is agree with Allen's Rule for energy benefits of having a more stocky body shape incold water.1616Mass = 8Area = 192+12+32=236Area/mass = 236/8=29,5Mass=8Fisher, 1985Area = 96+48+16=160Area/mass = 160/8=2022412 Ps. georgianus C. gunnari 113. Hofinger, 2010; Kils,1 230008 114. 0 100 200 300 400 500 600 700 800 9001-10.II. 198937,7 cm TL, 296,667 g.39,2 cmTL, 315g.47,3 cm TL, 665 g.ryb/km250150250500Different flattening and proportions of otoliths indicating differences in swimmingeven among icefish could be confirmed in their occurence and interspeciec ratio.54Biomass distribution of Ps. georgianus & noteson domination main species in a square, andcases domination Ch. aceratus over Ps. georgianus552 3(Traczyk, 2012a, 2012b; Sosiski, 1989; 1989a; Traczyk, 2013; Traczyk, 2012; Traczyk, 2013a924 5 6 935796585960 6197 65 64 63 62 99depth 500 mSampling period: February 110, 1989.Biomass of Ps. georgianus [tons]43 41 39 37 35105ShagRocks545543 41 39 37 35N. gibberifrons N. gibberifrons> Ch. aceratusCh. gunnarii> Ch. aceratusN. marmorata> Ch. aceratus54565591103 10450 100 200 500 1000 1500 2000 3500 115. 0 100 200 300 400 500 600 700 800 9001-10.II. 198937,7 cm TL, 296,667 g.39,2 cmTL, 315g.47,3 cm TL, 665 g.ryb/km250150250500Different flattening and proportions of otoliths indicating differences in swimmingeven among icefish could be confirmed in their occurence and interspeciec ratio.54Biomass distribution of Ps. georgianus & noteson domination main species in a square, andcases domination Ch. aceratus over Ps. georgianus552 3924 5 6 935796585960 6197 65 64 63 62 99depth 500 mSampling period: February 110, 1989.Biomass of Ps. georgianus [tons]43 41 39 37 35105ShagRocks545543 41 39 37 35N. gibberifrons N. gibberifrons> Ch. aceratusCh. gunnarii> Ch. aceratusN. marmorata> Ch. aceratus54565591103 10450 100 200 500 1000 1500 2000 3500 116. 43 42 41 40 39 38 37 36 35 34>0C; 1C; 0C; Ch. aceratusN. gibberifrons> Ch. aceratusCh. gunnarii> Ch. aceratusCh. aceratus> N. gibberifronsCh. gunnarii> Ch. aceratusN. gibberifrons> Ch. aceratusCh. gunnariiN. gibberifrons> Ch. aceratus54565591103 104Ch. gunnarii> Ch. aceratusN. gibberifrons> Ch. aceratusCh. gunnarii> Ch. aceratus2 35455545543 41 39 37 3543 41 39 37 35 124. 41 39 37 3517 18 19 20 21 22141D. mawsoni4 5 6 7 8 9 10 1112 131415 162324 25depth 500 mShag RockSampling period: January, 1992.Competitor C. aceratus displaces Ps. georgianus from 8 statistical squares. ACE = 8> 7 = SGI262743ShagRocksCh. gunnarii> Ch. aceratus Ch. gunnarii> Ch. aceratusN. gibberifrons> Ch. aceratusCh. gunnariiCh. gunnariiN. gibberifrons> Ch. aceratusN. marmorata> Ch. aceratusP. guntheriP. guntheri> Ch. aceratus D. mawsoni50 100 200 500 1000 1500 2000 2500545543 41 39 37 355455Biomass distribution of Ps. georgianus & noteson domination main species in a square, andcases domination Ch. aceratus over Ps. georgianusBiomass of Ps. georgianus [tons] 125. 41 39 37 3517 18 19 20 21 22142D. mawsoni4 5 6 7 8 9 10 1112 131415 162324 25depth 500 mSampling period: January, 1992.Competitor C. aceratus displaces Ps. georgianus from 8 statistical squares. ACE = 8> 7 = SGI262743ShagRocksCh. gunnarii> Ch. aceratus Ch. gunnarii> Ch. aceratusN. gibberifrons> Ch. aceratusCh. gunnariiCh. gunnariiN. gibberifrons> Ch. aceratusN. marmorata> Ch. aceratusP. guntheriP. guntheri> Ch. aceratus D. mawsoni50 100 200 500 1000 1500 2000 2500545543 41 39 37 355455Biomass distribution of Ps. georgianus & noteson domination main species in a square, andcases domination Ch. aceratus over Ps. georgianusBiomass of Ps. georgianus [tons]Parkes, 1990; SARAH CLARKE, 2008; Traczyk, 2012; Traczyk, 2012; Traczyk, 2013 126. 143Phylogeny statocyst and otolith - the relationship with themovement, that determine changes in their shape.Statocysts containing otoliths in the evolution of animal play one of the most importanttasks: they give the opportunity to gain space, a new environment, food, escape frompredators, thus ensure for species the survival and success. Without them, need to abovethe precise registration of changes in body position in the animal kingdom (with theexception of volatile insects) would be impossible. Otoliths serve not only maintain abalance, but also participate in the perception of sounds.Like the change of otolith shape sign up the development stages of the fish: from theball in a stationary fish embryo to elongated in swimming adult fish, it can be tracedin the interspecies changes of statoliths, the evolution of capabilities and speed ofswimming of animals on successive organization level of the animal kingdom. Fromthe simplest forms detection of balance changes in metazoan to a specialized organ ofbalance and hearing in bony fish in the process of concurring a new environment. Free-livinganimals having statoliths that during ontogeny transform to sedentary life stylewith a radial symmetry were lost statoliths.lacks a centralized brainLost statocystsSea cucumberbipinnaria larva brachiolaria larva 5 statocystsJura., 1983havestatocysts bisymmetry 127. Development swimming,moving and flying possibilitywith statocysts in animals:free living radialthe bilateralsymmetry.The otolith shape andmicrostructure evolved to bethe best in serving theperception of oscillationscaring information on bodymove and sounds. All movinganimals largest and small,high and low organized haveit in water and also this heavywatered organ was taken onland and in the air, excludingonly Pterygota because thoseones do not have neededappropriate strong osseousbase to carry it. And it is soconcerning that sessile,parasites and simpler withoutnerve system organisms donot maintain it.asc, psc, lsc anterior, posterior,lateral semicircular canals, c cristae, l lagena, ml, ms, mu macula lagenae, sacculi, utriculi,s saccule, u utriculi, ed endolymphatic duct, co cochlea, bm basilarmembrane, pb papilla basilaris144 128. 145Several statocysts at the edges giving orientation in space;Radial symmetry of free-floating organisms have spherical statolithsEvery step forward to improve the possibility of moving to extend migrating isdisplayed in change of statolith shape. If animal swim faster enough it has appropriatelarge deviation of otolith shape from a ball. In slow moving, the changes work onbody shape and statolith localizations! Animal only floating are radiallysymmetrical with same statoliths in the edges around of body.The statocysts in the margin around umbrella of the medusae of hydrozoans: e- the ectoderm, g-gonad,j- the absorptive lacuna digesting, k- the radial canal of the absorptive lacuna digesting, m-mesoglea,n- endoderm, s- statocyst, t- the stomatic bell, - pendentive.The cut across statocyst of the medusae: l-statolith,z- the sensorial cell, r- tentacle.emnsgtk ztrzlJura., 1983 129. One statocyst at peak, the perception of sound vibration. Bilateralsymmetry, actively swimming organisms: statolith sphericalIn this evolution step animal are with one statolith in the topside of body, to get similarinformation from all directions.More swimming animals (ctenophora) are changing body to bilateral symmetry andstatolith localization in a front to the head, or to head part as a very importantperception organ for swimming after a food or escaping from predators. In head partthat informations stymulate the development of the brain for their interpretations.In annelid one statocyst are embedded in the brain in a cephalad section or in the frontof the body, and are only in living free animals, having birateral symmetry.Statocyst of ctenophore comb jellies.Statocystof Ctenophorestatolithctenophoretentacletile Statolithcilianervous Statocystsystema mouth trochoforaanusa mouthJura., 1983 comb jellies - ctenophore 130. Statocysts at two sides of the brain or at larger distance, butinnervated by the cerebral ganglia; The bilateral symmetry,statoliths elongated.In a further evolution of the bilateral symmetry in the expansion of the newenvironment molluscs and arthropods form a pair of statocyst which are most optimallylocalised on both sides of the brain. These animals winning land and air environment.Statolith starts the change of spherical shape already in Molluscs in cephalopods.They are no longer balls like. Their shape and growth rate varies according to theneeds of the increasing swimming speed, as well as to changes in the depth zones.Changes in the shape and microstructure of cephalopods otoliths was also found to berelated to ontogenesis, the change in the living from epipelagic larvae to adult life,swimming deep into the meso and bathypelagic waters.cecumgonadInner shellInk bagventricle mantle cavity esophagus cerebralgratergrater and beak of tonguestatocyst funnel jawgillsstomach Gili heartJura., 1983 131. As squid swim faster their larvae otoliths originally spherical (N nucleus) in the elderlyhave become more elongated and acquired the shape of the dual spherical waves.148Alliroteuthisantarcticus 100mGaliteuthis glacialis200mArkhipkin, 1996; JACKSON, 1994The sphericalnucleus (N)andelongatedpost-larvalgrowth zone(PZ) of squidotoliths. 132. 5 statocyst in place of 2 in bilaterian animals. Deuterostomes repeat thelarvae have statocysts Lost statocystsd b149beginning of the development of the sense of balance.In the new stage of evolution in which into place of the blastopore of the embryoan anus was re-formed, the vulnerable period of embryogenesis is extended. Hence thefirst Deuterostomes - echinoderms develop in a safe aquatic environment, which is atthe bottom, and leading sedentary lives. Accordingly to that, the bilateral symmetry oflarvae was dislodged during development to the mature adults by the radiant symmetry.The abandonment of bilateral symmetry related to free life style causes the loss ofstatocyst. Statocysts are localized in old numerous mode, only if they have abilateral symmetry as a form more useful in the free mode of life.c bJura., 1983Sea cucumber: a- tentakles, b- cloaca, c- ambulacral feet, d papillae, e mouth, f throat, g- circular canal, h- esophagus,i - water system, j- stomach, k-Water lungs, l intestine, m gonads, n - duct stone, o - about oesophageal ring.kjihgfemlnSquirts (sea cucumbers), drilling and pelagic animals,has five statocyst located around the oesophageal ringoa cbipinnaria larva brachiolaria larva 133. 5 statocyst in place of 2 in bilaterian animals. Deuterostomes repeat thelarvae have statocysts Lost statocystsd b150beginning of the development of the sense of balance.In the new stage of evolution in which into place of the blastopore of the embryoan anus was re-formed, the vulnerable period of embryogenesis is extended. Hence thefirst Deuterostomes - echinoderms develop in a safe aquatic environment, which is atthe bottom, and leading sedentary lives. Accordingly to that, the bilateral symmetry oflarvae was dislodged during development to the mature adults by the radiant symmetry.The abandonment of bilateral symmetry related to free life style causes the loss ofstatocyst. Statocysts are localized in old numerous mode, only if they have a bilateralsymmetry as a form more useful in the free mode of life.c bJura., 1983Sea cucumber: a- tentakles, b- cloaca, c- ambulacral feet, d papillae, e mouth, f throat, g- circular canal, h- esophagus,i - water system, j- stomach, k-Water lungs, l intestine, m gonads, n - duct stone, o - about oesophageal ring.kjihgfemlnSquirts (sea cucumbers), drilling and pelagic animals,has five statocyst located around the oesophageal ringoa cbipinnaria larva brachiolaria larva 134. 5 statocyst in place of 2 in bilaterian animals. Deuterostomes repeat thed b151beginning of the development of the sense of balance.In the new stage of evolution in which into place of the blastopore of the embryoan anus was re-formed, the vulnerable period of embryogenesis is extended. Hence thefirst Deuterostomes - echinoderms develop in a safe aquatic environment, which is atthe bottom, and leading sedentary lives. Accordingly to that, the bilateral symmetry oflarvae was dislodged during development to the mature adults by the radiant symmetry.Statocysts are localized in old numerous mode, only if they have a bilateralsymmetry as a form more useful in the free mode of life.sea cucumbersc bJura., 1983Sea cucumber: a- tentakles, b- cloaca, c- ambulacral feet, d papillae, e mouth, f throat, g- circular canal, h- esophagus,i - water system, j- stomach, k-Water lungs, l intestine, m gonads, n - duct stone, o - about oesophageal ring.kjihgfemlnSquirts (sea cucumbers), drilling and pelagic animals,has five statocyst located around the oesophageal ringoa c 135. 1 central instead of 2 statocyst in bilaterians. Deuterostomes repeatedthe centralization of the sense of balance.The last line Deuterostomes of chordates, the tunicates confirm the centralization ofsense of balance to 1 statocyst in the vicinity of nerve ganglia. Wherein in some ofthem, the central statocyst probably corresponds to the central cavity of a brain as inother chordates. Appendicularia - Larvaceans lead free lifestyle, or settled, are withoutexcretory system and have opened the circulatory system. They are considered, asneotenic larvae. Those organisms have lost during evolution the mature stage butgained the ability to sexual reproduction before the full diversity of the body.In other sedentary tunicates, sea squirt they have statocysts only in their larvae whichare pelagic organisms. In the next organism in Thaliacea statocysts occurs in caseswhere there is free-living larval stage.Larvaceans and Thaliacea, order Doliolida, Doliolium denticulatum: a- statocyst, b- cloaca, c- ice, d heart,e mouth, f throat, g- stomach, h- notochord, i - cerebral ganglia j- gill crevice k- endostyl.aaebcd fdgghijksea squirt larvaDolioliumdenticulatumJura., 1983 136. 1 central instead of 2 statocyst in bilaterians. Deuterostomes repeatedthe centralization of the sense of balance.The last line Deuterostomes of chordates, the tunicates confirm the centralization ofsense of balance to 1 statocyst in the vicinity of nerve ganglia. Wherein in some ofthem, the central statocyst probably corresponds to the central cavity of a brain as inother chordates. Appendicularia - Larvaceans lead free lifestyle, or settled, are withoutexcretory system and have opened the circulatory system. They are considered, asneotenic larvae. Those organisms have lost during evolution the mature stage butgained the ability to sexual reproduction before the full diversity of the body.In other sedentary tunicates, sea squirt they have statocysts only in their larvae whichare pelagic organisms. In the next organism in Thaliacea statocysts occurs in caseswhere there is free-living larval stage.Larvaceans and Thaliacea, order Doliolida, Doliolium denticulatum: a- statocyst, b- cloaca, c- ice, d heart,e mouth, f throat, g- stomach, h- notochord, i - cerebral ganglia j- gill crevice k- endostyl.aaebcd fdgghijksea squirt larvaDolioliumdenticulatumJura., 1983 137. 2 statocysts, each developing a labyrinth in bilaterians. Deuterostomes154reach near the brain as the best location for statocysts.In vertebrates, each of a pair of statocyst develops in the membranous labyrinthcontaining statoliths adjacent on both sides of the brain. This structure has threechannels for decomposing the measure of the movement into 3 components. Theydevelop sequentially starting from one channel only in the first aquatic vertebratescharacterized by achieving high speed: in accrania bottom living hagfish. Although notyet have a dorsal fin needed to sedate of faster movement, they have 1 channel, thatincreases the measuring precision a more important component of their swimming.Large changes of spherical shape of otoliths have fish vertebrate. In state of no movable, or slowmovable the sphere is the best to percept the vibrations carrying the information on body changes and on sounds inenvironment from all directions. If fish velocity is large the signals from space are different between forward and afterwardand backward. It is compensate by changethe sphere to the elongate shape of otolith.Longer radius of otolith that isin motion percept the samesignal in similar period as radiiof otolith sphere no moving(Radtke 1985) 138. No significant changes in labyrinth of terrestrial vertebratesThe development of sound perception by labyrinth.Labyrinth evolved among aquatic vertebrates as a whole has moved further by them inspace acquiring on the land and in the air.In these new environments, fish labyrinth still provides for the birds and land animalsthe balance without major changes in its structure and operation.Also reading sounds of terrestrial vertebrates despite excelling in a result of extension of Ladena intocochlea, but it proceeds as in fish in the aquatic environment in the labyrinth as a result of the vibrationtransfer of endolymph on cilia of innervated hair cell of fish lagena elongated to the twisted cochlea ofmammals.Low sounds in lagena high in cochlea (Roy, 1994; umech.mit.edu, 2013; Inoue, 2013)In the organisms evolution, the process of development of statocysts containingstatoliths combines them into a common direction for getting a better perception of theirposition during faster swimming or moving, by measure vibration and in addition byinterpretation of their acoustic parts for the knowledge of environment and to generatethem in order to communicate within and between species. 139. cited works and the materials usedAllan, D., 2014. Blackfin icefish (Chaenocephalus aceratus) image. www: Arkive.org, natureepl.com.Altringham, J. D. D. J. E., 1999. Fish swimming: patterns in muscle function.. GB: The Journal of Experimental Biology 202,15633973403, The Comp. of Biol. Limited..Anon., 1., 1983. Encyklopedia Fizyki Wspczesnej. W-wa: PWN.Anon., 1990. State of fish population around the South Shetland Islands (Antarctic).. Hel: Workshop, PL..Anon, 2006. Podstawy mechaniki pynw biofizyka ukadu krenia. U: Wykad 30 X 2006.Arkhipkin, A., 1996. Age and growth of planktonic squids Cranchia scabra and Liocranchia reinhardti (Cephalopoda,Cranchiidae) in epipelagic waters of the central-east Atlantic. In: U.K.: Journal of Plankton Research, pp. l675-1683,.Arkhipkin, A. H. B., 1999. Statolith shape and microstructure as indicators of ontogenetic shifts in the squid Gonatus fabricii(Oegopsida, Gonatidae) from the Norwegian Sea. In: U.K.: Polar Biol, pp. 1-10.BARGELLONI L., Z. L. D. N. L. G. P. T., 2000. Molecular zoogeography of Antarctic euphausiids and notothenioids: fromspecies phylogenies to intraspecific patterns of genetic variation. UK: BAS, Antarctic Science 12 (3): 259-268.BAS, 2014. Impacts of Southern Ocean warming on marine connectivity. http: isow.bangor.ac.uk/rationale.php.en?subid=0.Bilyk, K., 2011. The influence of environmental temperature on the thermal tolerance of Antarctic notothenioid fishes.. UC:PhD dissertation University of Illinois at Urbana-Champaign.Byrd, D., 2012. Kristin OBrien: Antarctic icefishes have translucent bodies and blood.. U: http://earthsky.org/biodiversity.Image Credits to Kristin-OBrien..CCAMLR, 2012. CCAMLR_2012_Statistical_Bulletin_Volume_24_(Database_Version).mdb. Hobart:https://www.ccamlr.org/.CCAMLR, H., 2011. CCAMLR Statistical Bulletin, Vol. 24 (20022011). [Online]Available at: http://www.ccamlr.org/en/document/publications/ccamlr-statistical-bulletin-vol-24-2002%E2%80%932011Cheng, C. L. C., 1999. Evolution of an antifreeze glycoprotein.. U: Mac.Mag.Nature Vol. 401: 443.Chen, W., 250. Strange Things of the South. U: Temple, R..Chojnacki, J. M. P., 1987. Odywianie si i pokarm antarktycznych ryb biaokrwistych (Chaenocephalus aceratus,Pseudochaenichthys georgianus oraz Champsocephalus gunnari).. Gdynia: Studia Materiay. MIR/A/28..Clarke A., I. A. J., 1996. Evolution and adaptive radiation of Antarctic fishes. UK: Elsevier Science Ltd., TREE vol. 11. no.5.DAMERAU M., M. M. S. W. H. R., 2014. Population divergences despite long pelagic larval stages: lessons from crocodileicefishes (Channichthyidae). NY: John Wiley & Sons Ltd. Molecular Ecology 23, 284299.Davison W., J. A. M., 1985. A histochemical study of the swimming musculature of Antarctic fish. NZ: New Zealand Journalof Zoology, 12:4, 473-483,. 140. 157cited works and the materials usedDetrich, B. C. C. a. A. D., 2012. The Birth and Death of Genes. NY: Dnatube.com. Scientific video site. Film of HowardHughes Medical Institute; https://www.youtube.com/watch?v=WpIe8_pUSu4.Eicken, H., 1992. The role of sea ice in structuring Antarctic ecosystems. G: Springer-Verlag. Polar Biol. 12: 3-13..Everson, I., 1981. Antarctic fish age determination methods. Rome: BIOMASS handbook no. 8.Everson, I., 2014. Young blackfin icefish (Chaenocephalus aceratus). www: Arkive.org. Photoshot.com. .Everson, I. T. E., 1977. The Evolution of the Haemoglobinless Condition in the Antarctic Channichthyidae.. Washington:Proc Third SCAR Symp Antarct Biol.Fischer, W. J. H., 1985. FAO Species Identification Sheets For Fishery Purposes. Southern Ocean.. Rome: CCAMLR.Fuiman, L. D. R. W. T., 2002. Behavior of midwater fishes under the Antarctic ice: observations by a predator. USA:Springer-Verlag. Marine Biology 140: 815822.Gosline, J. M. M. E. D., 1985. Jet-propelled Swimming in Squids. NY: Sci. Amer. 256: 96-103.Grabowska, B. R. T., 2010. OKRELANIE WIEKU RYB FALKLANDZKICH Z OTOLITW DLA PICIU GATUNKW:1. SALILOTY AUSTRALIS, 2. GENYPTERUSA BLACODES,3. MACROURUSA CARINATUS, 4.COELORHYNCHUSA FASCIATUS, 5. MACROURUSA HOLOTRACHYS. Gdynia: MIR, raport.HARRISON, P. N. J. M. I. A. J., 1987. Gross morphology, fibre composition and mechanical properties of pectoral finmuscles in the Antarctic teleost, Notothenia neglecta. Nybelin. In Proceedings of the Fifth Congress on EuropeanIchthyology, (ed. S. O. Kullander & B. Fernholm), pp. 459-465.. Stockholm: Swedish Museum of Natural History.Hecht, T., 1978. A descriptive systematic study of the otoliths of theneopterygean marine fishes of South Africa.. In:Transactions of the Royal Society of South Africa. s.l.:s.n., pp. 191-197.Hecht, T., 1987. A Guide to the Otoliths of Southern Ocean Fishes. 1 ed. S. Afr.: T. Nav. Antarkt., Deel 17.Hecq, J.-H. F. V., 2007. Status, control and role of the pelagic diversity of the austral ocean (PELAGANT). Part 2: Globalchange, Ecosystems and Biodiversity. Belgium: Belgian Sci. Policy, SPSD II, D/2007/1191/48.Hofinger, E., 2010. photo Pseudochaenichthys georgianus. Fishbase:http://www.fishbase.se/Photos/ThumbnailsSummary.php?Genus=Pseudochaenichthys&Species=georgianus.Inoue, M. M. T. Y. O., 2013. The role of ear stone size in hair cell acoustic sensory transduction. Nagoya: Scientific reports.JACKSON, G. C. L., 1994. Statolith microstructure of seven species of Antarctic squid captured in Prydz Bay, Antarctica..In: Perth: Antarctic Science, pp. 195-200.Jakubowski, M., 1960b. The structure and vascularization of the skin of the eel (Anguilla anguilla L.) and viviparous blenny(Zoarces viviparous L.).. Cracov: Ibid., 3: 1-22;.Jakubowski, M., 1971. Biaokrwisto i inne osobliwoci ichtiofauny Antarktyki.. W-wa: Przeg. Zol., XV, 3.. 141. 158cited works and the materials usedJakubowski, M., 1982. Dimensions of Respiratory Surfaces of the Gills and Skin in the Antarctic White-Blooded Fish,Chaenocephalus aceratus Lonnberg (Chaenichthyidae).. Leipzing 96 /1: Z.mikrosk.-anat. Forsch..Johnston, A. N. F. G. Z. R. E. W. P. H. a. B. T., 1983. Morphometric and ultrastructural features of the ventricularmyocardium of the haemoglobin-less icefish Chaenocephalus aceratus.. U: Camp. Biachem. Physial. Vol. 76A, No.3,pp. 475-480.Jura, C., 1983. Bezkrgowce. W-wa: PWN.Kaufmann R.S., S. J. K. B. R. G. R. R. B. R. K., 1995. Effects of seasonal pack ice on the distribution of macrozooplanktonand micronekton in the northwestern Weddell Sea. USA: Springer-Verlag. Marine Biology 124:387-397..Kellermann, A. (., 1990. Identification key and catalogue of larval Antarctic fishes.. Berlin: Polarforsch. 67:1-136., .Kils, U., 2008. Photos in film "Antarctic Fish". www: https://www.youtube.com/watch?v=VbqHQss1SSg;http://en.wikipedia.org/wiki/User:Kils.Kock, K.-H. a. A. K., 1991. Reproduction in Antarctic notothenioid fish: a review.. Hobart: Ant. Sci., 3 (2): 125-150..Kunzmann, A., 1991. Blood Physiology and Ecological Consequences in Weddell Sea Fishes (Antarctica). Germany: Ber.Polarforsch. 91, ISSN 01 76 - 5027.La Mesa M., E. J. T. V. M., 2004. The role of notothenioid fish in the food web of the Ross Sea shelf waters: a review. USA:Springer-Verlag. Polar Biol 27: 321338.La Mesa, M. A. D. F. C. J. K. K., 2009. AGE AND GROWTH OF SPINY ICEFISH (CHAENODRACO WILSONI REGAN,1914) OFF JOINVILLEDURVILLE ISLANDS (ANTARCTIC PENINSULA). In: Hobart: CCAMLR Science, pp.115-130.LA MESA, M. J. A. E. L. M. V., 2004. Age and growth of Scotia Sea icefish, Chaenocephalus aceratus, from the SouthShetland Islands, Roma: Ant.Sci. Ltd. .La Mesa, M. M. V., 2001. Review. Age and growth of high Antarctic notothenioid fish.. In: U.K.: Antarcfic Science, pp. 227-235.Le Franois, N., 2014. C aceratus, le Grande Gueule ou Blackfin icefish. www:https://www.youtube.com/watch?v=EBiLWBpUTcY.Le Franois, N., 2014. P georgianus, the South Georgia icefish.. www: YouTube.https://www.youtube.com/channel/UCJLVlLySAkZC9SdmrqESp_g.Marschall, H.-P., 1988. The overwintering strategy of Antarctic krill under the pack ice of the Weddell Sea. 2 ed. UK: PolarBiology.Marschall, P. U. K., 2012. Antarctic krill Euphausia superba in ice cave. www.zuckerspeicher.de: Ecoscope.com. 142. 159cited works and the materials usedMaslennikov, V. E. S., 1988. Patterns of fluctuations in the hydrological Conditions of the Antarctic and their effect on thedistribution of Antarctic krill.. In: Antarctic Ocean and Resources Variability. Berlin Heidelberg: Springer-Verlag.Molina, R. a. F. L., 1989. Hidrografia en la region canaria. Campana "Canarias I". Bol. Inst. Esp. Oceanogr., 5(2), pp. 71-86.Murphy, e. a., 2013. Comparison of the structure and function of Southern Ocean regional ecosystems: The AntarcticPeninsula and South Georgia. UK,: Journal of Marine Systems 109110, 2242, www.elsevier.com/locate/jmarsys..Near, T. S. P. a. H. W. D., 2010. A Genomic Fossil Reveals Key Steps in Hemoglobin Loss by the Antarctic Icefishes.. U:Mol. Biol. Evol. 23(11):20082016..North, A., 1988. Distribution of fish larvae at South Georgia: horizontal, vertical and temporal distribution and early lifehistory relevant to monitoring year-class strength and recruitment.. In: Hobart: SC-CAMLR.North, A., 1990. Ecological studies of Antarctic fish with emphasis on early development of inshore stages at South Georgia..Cambridge: Br.Antarct.Surv. NERC.North, A., 1991. Review of the early life history of Antarctic Notothenioid fish. In: Biology of Antarctic Fish . U.K.:Springer-Verlag.North, A. W., 1990. Ecological studies of antarctic fish with emphasis on early development of inshore stages at SouthGeorgia.. Cambridge, U.K.: Ph. D. Thesis, Br. Antarct. Sur., Natural Environment Research Council,.Parkes, G. I. E. J. A. Z. C. J. S. R. T., 1990. Report of the UK/Polish fish stock assessment survey around South Georgia inJanuary 1990. In: London: Imp.Coll. of Sci. & Techn., p. 20.PolarTrec, 2013. Icefish in action. www: https://www.youtube.com/watch?v=VbqHQss1SSg.Radtke, R. T. T. J. B., 1985. Growth resolution of antarctic fish. In: s.l.:Ant.Jour., pp. 157-159.Rakusa - Suszczewski, S., 1989. W Antarktyce. W-wa: KAW.Reis, D. W. G. F., 1970. The relationship of blood flow to myoglobin, capillary density, and twitch characteristics in red andwhite skeletal muscle in cat.. GB: J. Physiol., 210, pp. 121-135; http://jp.physoc.org/content/210/1/121.full.pdf+html.Robinson, A., 2014. Underwater Photos: How Bass Eat. www: Outdoorlife.http://www.outdoorlife.com/photos/gallery/fishing/freshwater/largemouth-bass/2012/06/underwater-photos-how-bass-eat/?image=2.Roy, D., 1994. Development of hearing in vertebrates with special reference to anuran acoustic communication.. Shillong: J.Biosci..Sahrhage, D., 1988. Antarctic Ocean and Resources Variability.. In: Berlin Heidelberg: Springer-Verlag.SARAH CLARKE, W. D. R. M. A. C. a. M. B., 2008. Biology and distribution of South Georgia icefish (Pseudochaenichthysgeorgianus) around South Georgia and Shag Rocks. 20 ed. UK: Antarctic Science Ltd.. 143. 160cited works and the materials usedSiegel, V., 1988. A Concept of Seasonal Variation of Krill (Euphausia superba) Distribution and Abundance West of theAntarctic Peninsula.. In: Antarctic Ocean and Resources Variability. Berlin: Springer-Verlag.Sievers, H. W. N. J., 1988. Upper ocean Characteristics in Drake Passage and Adjoining Areas of the Southern Ocean, 39W-95W. In: Antarctic Ocean and Resources Variability. Berlin Heidelberg: Springer-Verlag.Sosiski, J. W. . A. K. S. W. K. S. R. T. S. L., 1989. Ocena stanu ywych zasobw na owiskach eksploatowanych przezpolskie rybowstwo dalekomorskie. In: Gdynia: MIR.Sosiski, J. Z. C. A. K. K. S. J. S. R. T. S. W. R. Z., 1989. Biologiczno-rybackie badania ywych zasobw Antarktyki wsezonie 1988/1989. In: Gdynia: MIR.lsarczyk, W. Z. C., 1985. Postlarval and juvenile fish (Pisces, Perciformes and Myctophiformes) in the Antarctic Peninsularegion the Antarctic Peninsula region during BIOMASS-SIBEX 1983/1984.. 1-2 ed. W-wa: Pol. Polar Res..Traczyk, R., 1992. AGE AND GROWTH OF THE ANTARCTIC FISH Chaenocephalus aceratus based on OTOLITHweight, microstructure and TL frequency; some relations with Pseudochaenichthys georgianus.. [Online].Traczyk, R., 1993. The occurrence of krill Euphausia superba in the floating ice edge zone and some its biological data..[Online].Traczyk, R., 2010. OKRELANIE WIEKU Z OTOLITW FALKLANDZKIEJ MIRUNY MACRURONUSAMAGELLANICUS I FALKLANDZKIEGO MORSZCZUKA MERLUCCIUSA POLYLEPIS. ZAPIS CYFROWYOTOLITW. Gdynia: MIR, Sprawozdanie z pracy.Traczyk, R., 2011. WIEK I WZROST MAKRELI KOLIAS (SCOMBER JAPONICUS) Z WD RODKOWOWSCHODNIEGO ATLANTYKU (MAURETANIA) I POUDNIOWO WSCHODNIEGO PACYFIKU (CHILE),Gdynia: MIR, raport.Traczyk, R., 2012a. Migrations of Antarctic fish Pseudochaenichthys georgianus Norman, 1939 in the Scotia Sea. Hobart:WG-FSA-12/68 Rev. 1, WG-FSA-13..Traczyk, R., 2012b. Ps. georgianus data base: materia_segr_bez_formu.xlsx. Gdask: https://drive.google.com/file/d/0B-QIBKvRn8b1cGdYaXlPY2FfZ2c/view?usp=sharing.Traczyk, R., 2012. Migrations of Antarctic fish Pseudochaenichthys georgianus Norman, 1939 in the Scotia Sea.. Hobart:CCAMLR. WG-FSA-12/68 Rev. 1.Traczyk, R., 2013a. fish data: materiay_wyniki_.docx. Gdask: https://drive.google.com/file/d/0B-QIBKvRn8b1THRhVEt4T3hiREk/view?usp=sharing; http://www.slideshare.net/ryszardtraczyk/materiay-wyniki.Traczyk, R., 2013b. Measurement and analysis of optical density profiles of otolith from Ps. georgianus and C. gunnari..www: http://www.slideshare.net/ryszardtraczyk/analysis-of-theopticaldensityprofileofotolithoficefish. 144. cited works and the materials usedTraczyk, R., 2013c. Zastosowanie analizy mikroprzyrostw i morfologii otolitw georgianki (Pseudochaenichthys georgianus161Norman, 1939) z rejonu Georgii Poudniowej (Antarktyka) do okrelenia wieku, wzrostu oraz waniejszych okreswrozwoju ryb. Krakw: Ryszard Traczyk In Altum, ISBN: 978-83-62841-10-3;http://www.slideshare.net/ryszardtraczyk/psgeorgianus-druk.Traczyk, R., 2013. Economic competition for high profits from Antarctic living resources in protection area and Mercurycontaminants of fish from outside.. Gdynia: unpublish. http://www.slideshare.net/ryszardtraczyk/antarctica-fish-mercury.Traczyk, R., 2014. Wspomnienia z antarktycznych ekspedycji Morskiego Instytutu Rybackiego.. Krakw: InAltum, ISBN:978-83-62841-20-2, str.: 183.Twelves, E., 1972. Blood volume of two Antarctic fishes. UK: Br. Antarct. Surv. Bull., No. 31, p. 85-92.umech.mit.edu, 2013. Introduction to Cochlear Micromechanics. Eng: umech.mit.Van Cise, A., 2009. AMLR 2008/2009 FIELD SEASON REPORT.. NY: NOAA-TM-NMFS-SWFSC-445.Vincent, W., 1988. Microbial ecosystems of Antarctica.. Great Britain: Press Syndicate of the University of Cambridge..VlDELER J. J., F. H., 1984. FAST CONTINUOUS SWIMMING OF TWO PELAGIC PREDATORS, SAITHE(POLLACHIUS VIRENS) AND MACKEREL (SCOMBER SCOMBRUS): A KINEMATIC ANALYSIS. GB: J. exp.Biol. 109, 209-22.Walesby, N. C. N. I. J., 1982. Metabolic differentiation of muscle fibres from a haemoglobinless (Champsocephalus gunnariLonnberg) and a red-blooded (Notothenia rossii Fisher) Antarctic fish.. GB: Br. Antarct. Surv. Bull., No 51. p.201-214..Wells, R. G. S. L. A. B. a. G. C. G., 1985. Ecological and Behavioural Correlates of Intracellular Buffering Capacity in theMuscles of Antarctic Fishes. New Zealand: unpubl. Dep. of Zoology, Un. of Auckland.Wsawski, J., 2011. ywy wiat lodu (2). Sopot: CESSS, IO PAN. [email protected], M., 1977. Ecological adaptations by Antarctic poikilotherms to the Polar Marine Environment.. Washington: ProcThird SCAR Symp Antarct Biol..White, M. K.-H. K. V. S., 1998. Larval and juvenile fish collected during December 1996 in the Elephant Island region. InKattner G.: The expedition Antarktis XIV/2 of rv "Polarstern" in 1996/97.. Berlin: Ber. Polarforsch. 274, ISSN 0176-5027 .Witek, Z. J. K. A. G., 1988. Formation of Antarctic Krill Concentrations in Relation to Hydrodynamic Processes and SocialBehaviour. In: Antarctic Ocean and Resources Variability. Berlin: Springer-Verlag.Whrmann, A., 1996. Antifreeze glycopeptides and peptides in Antarctic fish specles.. UK: Mar. Ecol. Prog.Ser., Vol. 130:47-59. 145. 162cited works and the materials usedabrowski, M., 2000. The osteology and ossification variability of the skull of antarctic white-blooded fish Chaenodracowilsoni Regan, 1914 (Channichthyidae, Notothenioidei). W-wa: Acta Ichthyol. Piscat. 30 (2): 111-126.bikowski, D., 2008. Szybko pywania ryb. [Online]Available at: http://www.fishing.pl/desde-agua/curiosidades/szybkosc-plywania-ryb