Otolith shape

Ryby antarktyczne

1 mm

Distribution of the Antarctic fish Pseudochaenichthys georgianus NORMAN, 1939 in the Atlantic sector of Antarctic. The Changes of otolith shape - a book of stories about the nature of the distribution of fish Ps. georgianus and not only for fish

1 mm

1 mmby Ryszard Traczyk

fast mackerel fromtemperate watersslow icefish from cold waterslarval otolithslow Macrouridefrom cold deep waters21.3 km/h~0.9 km/h~0.8 km/h

AntarcticaAntarctic Circumpolar

CDWEast Wind Drift

ABWABW

Cold water

HighsLows

Polar Fronticy windAntarctic Bottom WaterAntarctic Bottom WaterHighsWHITE-BLOODED: high Antarctic; ice pack zone; temperate 80S 74, ~70S; 63, ~60S; 63, ~5730S; 52, ~45S 30SPolar cellFerrel cell

S. OrkneySouthGeorgiaS.Shetland Is

-100-200-300-400-500-600-700-800Current[m]

S. japonicusChannichthyidae

Ch. aceratusPatagonia

Ps. georgianus

MacrouridaeM. holotrachys

C. gunnariDeep-WaterCircum-polardescentTerrestrial observations of separate geographical and vertical living on different age groups and species of fish suggest that differences in otolith shape among them became from difference in their environment conditions. (Extracted and enlarged otoliths are over or near the fish heads: Median or Transverse plane)

Fisher, 1985; Kellermann, 1990; North 1990; Hecht, 1987

21 km/h1 km/h0.9 km/h0.1 km/h Traczyk, 1992; 2012; 2013c; Grabowska, 2010; JACKSON, 1994


CDWEast Wind Drift

ABWABW

Cold water

HighsLows


S. OrkneySouthGeorgiaSouth Shetland Is

-100-200-300-400-500-600-700-800Current[m]

Decrease of Otolith Length, increase of Otolith Height

S. japonicusChannichthyidae


Ps. georgianus

MacrouridaeM. holotrachys

C. gunnariDeep-WaterCircum-polardescent

increase of Otolith LengthAS WATER TEMPERATURE IS DROPPING

MTransverseplaneMedianplaneand flattening (T plane)TMTMTTTTMMMMM

21 km/h0.9 km/h0.1 km/h1 km/h

4Age groups of Ps. georgianus seperated in time scale confirm geographical divide of otolith mass frequency as separate age groups on Antarctic islands: 2 aged at Pamer Archipelago, in February, 3 aged at King George in March, and 4 aged and older fish at S. Orkney in December. Palmer A., Deception, Elephant Is: 19-22.II.1979, N=97

Traczyk, 2012

5Age groups of Ps. georgianus seperated in time scale confirm geographical divide of otolith mass frequency as separate age groups on Antarctic islands: 2 aged at Pamer Archipelago, in February, 3 aged at King George in March, and 4 aged and older fish at S. Orkney in December. Palmer A., Deception, Elephant Is: 19-22.II.1979, N=97

23>4Palmer A.K. GeorgeS. Orkney

This same is for age groups in length frequency geographically divided on separated Antarctic Islands.

This same is for age groups in length frequency geographically divided on separated Antarctic Islands.: 2 aged fish at Pamer A., in March, 3 aged fish dominated at Elephan in February, and 4 aged and olders fish at South Orkney in December. 23>4Palmer A.K. GeorgeS. Orkney

8Geographical separation of age groups on shelf of different islands indicates that marine habitats of these areas have different properties.

9such that Ps. georgianus from the younger age groups prefer the western part of the Atlantic Antarctic, and the older age group East - North part. Age group is identified by otolith shape, which, indicate the different habitats, development stages and strategies of life: type of swimming.

West49 cm

43 North East

30

10such that Ps. georgianus from the younger age groups prefer the western part of the Atlantic Antarctic, and the older age group East - North part. Age group is identified by otolith shape, which, indicate the different habitats, development stages and strategies of life: type of swimming.

West49 cm

43 North East

30

Otolith, M-plane, after Hecht, 19870.4 km/h1.6 km/hand speed

Water temperature determine distribution of Antarctic fish? So older age groups fish 48 cm found at South Orkney Islands may have been resulted from lower temperature of waters below 0C up to - 1C. Younger age groups of 3 aged fish 43 cm found most numerous at Elephan have little warmer waters up to 0C. The smallest fish 30 cm length of 2 age group appear at Palmer Archipelago have warmer water of above 0C up to 1C.

Potential temperature [C] at 200 m>0C ~0C 0C ~0C 0C ~0C 34 at 20m)WSCWSCWSC

73

200

350

90

150

550Ps. georgianus show different geographic distributions in number of fish. At S. Georgia icefish were more numerous. This could be related to swimming posibility: at S. Georgia we can see strong turbulences and eddys, and also more krill for food .

23

14

krill density [g/m2] and cluster extending

AntarcticCircumpolarCurrent

48

Marschall, 1988; 2012; Sahrhage, 1988; Siegel, 1988

Closer to the continent catch drops 2 (on r/v Prof. Siedlecki).S. Georgia S. Orkney. K. George Palmer A. 10 kg/h20304050

47.8

22.6

3.2

14.4

5.1

98.1

South Orkney I.South Georgia I.Bransfield Str.S c o t i a S e aW e d d e l l S e a

60565248444036545658606264

Elephant I.Feb.1979 the R/V Prof. Siedlecki (N=67)

Nov. 78 Feb. 79 the M/T Sirius (N=30)

Ps. georgianus capture, [kg/h]545658606264

6460565248444036

0-500 m

48

23

14

K. GeorgePalmer A.AntarcticCircumpolarCurrent

Traczyk, 2012

47.8

22.6

3.2

14.4

5.1

98.1

South Orkney I.South Georgia I.Bransfield Str.S c o t i a S e aW e d d e l l S e a

60565248444036545658606264

Elephant I.Feb.1979 the R/V Prof. Siedlecki (N=67)

Nov. 78 Feb. 79 the M/T Sirius (N=30)

Ps. georgianus capture, [kg/h]545658606264

6460565248444036

AntarcticCircumpolarCurrent

48

23

14

0-500 m

K. GeorgePalmer A.

20>2>2>2>2>21C; 0,6C; 34 per 20 m)krill high density regions

Secondary Frontal Zone, SFZ

0,30,30,35

Krill 550g/m3South GeorgiaA large number of fish from old age groups of Ps. georgianus on the North East side of South Georgia Is., were coincided with the location of permanent turbulent flows causing local accumulation of large krill. This show that large fish of Ps. georgianus live in currents to prey krill. In 1978 to 1979 where krillo fagous of Ps. georgianus was being caught more (48kg / h), krill density was high (550g/m3). Such an arrangement is not accidental because is repeated year after year.

Witek, 1988; Sahrhage, 19881.6 km/h

51

42 41 40 39 38 37 36 35 34 43

43 42 41 40 39 38 37 36 35 345455

Shag Rocks

5554500m500m500m30'30'30'

150m 150m

30'30'30'w

48 kg fish/h

Antarctic Circumpolar Current

WSC-Weddella-Scotia Confluence (s > 34 per 20 m)krill high density regions

Secondary Frontal Zone, SFZ

0,30,30,35

Krill 550g/m3Georgia Pd.A large species, fish of Ps. georgianus have to be better prepared in balance perception to withstand currents when feeds on large krill being in stronger flows.

Currents at surface at 5MPa [Dyn m]

Ps. georgianus has krill as dominant food (Sarah Clarke, 2008; Chojnacki, 1987).1.6 km/h

43 42 41 40 39 38 37 36 35 34555430'30'30'

43 42 41 40 39 38 37 36 35 34

53555430'30'30'531C1C1C1C1C1C>1C; 0,6C; 1C; 0,6C; 1C; 0,6C; 1C; 0,6C; 1C; 0,6C; 1C; 0C; 0.6C; 0C; 1C; 0,6C; 1C; 0C; 0C; 0C; 1C; 0,6C; 1C; 0,6C; OL: OH=bOL+a above y=xproportionality of the otolith dimensions are constant

DorsalareaventralareaAnterior culliculumSimilar life stage confirms proportions of otolith, if this is same during life of fish. Large dorsal area indicated large vertical movements, has high growth rate.

Dorsal area

OH > OL: OH=bOL+a above y=xproportionality of the otolith dimensions are constant

DorsalareaventralareaAnterior culliculumFor 30 cm Ps. georgianus above 2 age group, high growth of Anterior culliculum give larger sensitivity of body balance in swimming forward need in surface currents.

How do we know that the shape of otolith indicates the speed of swimming? In comparison: faster species have them more flatter

S. japonicusPs. georgianusfaster species

Transverse planemedian planemedian planeTransverse plane1 mm

1 mmData for speed of swimming (bikowski, 2008, Fuiman, 2002)21.3 km/h~1,6 km/h

the shape of otolith is chageing among species of fish of different depth of livingChannichthyidaeCh. aceratusMacrouridaeM. holotrachysDeep water speciesshelves species

Transverse planemedian planeTransverse planemedian plane1 mm1 mmBody fish and otolith shape data (Hecht, 1987; Fischer, 1985; Grabowska, 2010; Traczyk, 1992)

1 mmS. japonicus, FL=39 cmMauretania 21.I.2011Haul 44. No 39 optical density annual incrementsLarge changes in the width increments of otolith mackerel on radii R3, R110246810NucleusBack edgedaily increments widthR3R11

otolith shape data (Traczyk, 2011)

1 mmS. japonicus, FL=39 cmMauretania 21.I.2011Haul 44. No 39 optical density annual incrementsOtolith mackerel on R3 has large width increments, on R11 small, so has larger length than high.0246810NucleusBack edgedaily increments widthR3R11

Extreme length of otolith compensated by reduction of it other sides because of

S. japonicusspawn I-V

0246810Direction of lack of otolith growth Fork Length FL; daily increments width from inner side

Otolith back radius (mm)high speed ofExtreme length of mackerel otoliths arise from high speed of swimming (show by torpedo shape of body) - from their incessant fast swimming in the pelagic ocean (to not to allow to fall, from their body heavier than water). Increase in the dorsal edge of mackerel otolith is reduced, because this fish has stability resulting from high inertia, or from high frictional force. swimming

Extreme length of otolith compensated by reduction of it other sides because of

Lack of the increments on the one side of the labyrinth is probably from a pressure of swimming speed removing otoliths substrates from that side to the other, where a large acceleration locally concentrate them thus determining the constant rapid increase of the otolith length and their local elongation at the expense of not growing of the otolith inner surface. In contrast Ps. georgianus is slower swimmer but migrates vertically with perfection ensured by larger high of the otoliths.

S. japonicusspawn I-V

0246810Direction of lack of otolith growth Fork Length FL; daily increments width from inner side

Otolith back radius (mm)swimminghigh speed of

S. japonicusPs. georgianus1 mmSpeed in swimming is the source of shape diversity Dorsal marginVentral marginnucleusrostrumAnnual incrementsOuter sideFrontal marginBack marginOuter dorsal sideconcave surface

Transverse planeTransverse planeone side incrementsDorsal marginrostrum1 mmnucleusChanges in the width increments on R3, R11

Dorsal marginventral marginback marginrostrum

Increments in all directions,around nucleusVentral marginnucleusNarrow annual incrementsand wide

S. japonicusDorsal marginVentral marginnucleusrostrumAnnual incrementsFrontal marginBack margin1 mm1 mmDorsal edge of mackerel otoliths have increments tightened up and the growth radius becomes 3.7 and 5 times smaller than the radii of growth to back and front edges. In Ps. georgianus otolith radii growth in opposite pattern, wide microincrements form dorsal radius of 1.8 and 1.5 times larger than the radii of the back and front edges.

Transverse plane

Ps. georgianusDorsal marginventral marginback marginrostrumIncrements in all directions,around nucleusTransverse planeone side incrementsChanges in the width increments on R3, R11

nucleus

Ventral marginnucleus

Outer sideOuter dorsal sideconcave surfacerostrumDorsal margin

Narrow annual incrementsand wide

S. japonicusDorsal marginVentral marginnucleusrostrumAnnual incrementsFrontal marginBack margin1 mm1 mmThe high otoliths of SGI icefish - as fishing floats inform about vertical stability needed for vertical migrations and for lifting with currents. In opposite to that long otoliths of mackerel are sensitive on changes during swimming in the horizontal direction. Information important in the fast swimming for far distances.

Transverse plane

Ps. georgianusDorsal marginventral marginback marginrostrumIncrements in all directions,around nucleusTransverse planeone side incrementsChanges in the width increments on R3, R11

nucleus

Ventral marginnucleus

Outer sideOuter dorsal sideconcave surfacerostrumDorsal margin

Narrow annual incrementsand wide

95 Macrourus carinatusSwimming depth is source of diversity in microstructure and shape of the otolithLength of otolith of fish Macrourus carinatus is large, more than two times than height: R3 >> R9. They are not very fast swimmers, so that is the impact of higher hydrostatic pressure of ~ 1200 m column water. Dorsal margin R9 against the pressure does not rise, but increases ventral R11 in the direction of pressure and most increase otolith length: R3, 7 (in the zero pressure gradient).

25y19cm0,000199 mmR11, 0,000463 mm0,0000887R3 0,00139 mmR10R12R9Dorsal marginVentral margin

=0,00139 mmR7R3

R9R11=0.000463 mmDorsal marginVentral margin1 mmBack marginfrontal margininner marginouter marginBack marginTransverse planemedian plane

Otolith shape data Grabowska, 2010

96Transverse section similar to Channichthyidae, but otolith length is large similar to length of otolith mackerel: ~2> otolith height: R3>>R9, M. carinatus live in deep waters 1200mDaily increments = 1 year increment/ 365Yearly increments along otolith inner radius of R10

19cm, 25 year oldMacrourus carinatusHigh depth of swimming is the source of shape diversity0,000199 mmR11, 0,000463 mm0,0000887R3 0,00139 mmR10R12R9Dorsal marginVentral margininner marginouter margin

97 M. carinatus

25y19cm0,000199 mmR11, 0,000463 mm0,0000887R3 0,00139 mmR10R12R9Dorsal marginVentral margin

=0,00139R7R3

R9R11=0.000463 DorsalVentral 1 mmBack marginfrontal marginInner sideouter sideBack margin

C. aceratus M. carinatus M. carinatus has longer OL than C. aceratus, but is not faster - it swim deeperdorsal edgedimensions of radii: small dorsal and large ventral for M. carinatus and vice wersa for Channichthyidae inversed proportions Ventral marginInner sideouter side

1 mmVentral marginBack margin1590 days R9=2.35 R9=0.82554 days45 cm SLOW= 0.0247 g OH = 3.44 mmR7R9R8R11R11R9R10R12

98

2 directional growth in otolith shape: determined by pulsed swimmingR9=0,5 mm/0,007 = 72 days = 2,4 monthsR9=1,21 mm/0,007 = 160 days = 5,3 monthsSquids swims slow with pulsation have twins hemispheres in otolith shape with the widest otolith increments 0,007 mm

Data: Arkhipkin, 1996; Arkhipkin, 1999

0,1 mOtolith shape differentiates pattern of high energy swimming of mackerelwith body waves otolithScombrus japonicus ~21.3 km/h Squids 1-3 km/h

ejection of H2O download H2Otwo inhibitons of motion to even reverse by input propulsion and fin opposition shapeotolithpulsed swimming in squidsslower suction

rapid motionfunnel

bikowski, 2008; Videler J.J., 1984Gosline, 1985

wyrzut H2O pobr H2Olow energy swimming of icefish Channichthyidae using the pectoral fins

otolith

Large pectoral fins are floating fish in the depths; vertical movements, are measured more by deviation of otolith height from the vertical, that is, by higher otolith.High, laterally flattened body having a great fins has about 20 times more resistance of the lateral than the front and the current pressure on the concave side of curved body of flowing fish produces a hydrodynamic force increasing speed of fish swimming forward. An asymmetrical shape with respect to the direction axis of swimming causes asymmetric flow, that creates differential pressure on opposite surfaces, and thus the driving force to forward. axial musculature reduction1 mm

(Fuiman L, 2002; Anon, 2006)

101High body and wide, long fins reduce drift in currents. Fin fish and flat body during swimming provide hydrodynamic lifting force.

35 43%18,5 27,3%28 - 3123 - 24>100%29 - 318-10Ps. georgianusThe shape of the otoliths is plastically formed by distribution of endolymph pressure thank to that it provides for fish from otolith shape interpretation - the information about the speed and body movement and also provides information about sound and water vibrations causing vibration of endolymph and giving seeing yourself in surroundings. Similarly, by evolution the body shape is formed in order to obtain maximum speed to which the shape of otolith is adjusted. Thanks to this otolith shape and body shape are interrelated. Body shape of Ps. georgianus evolved as the shape of otoliths with respect to the same target is high, facilitating vertical migration and is laterally flattened facilitating swimming in currents with minimal energy consumption.

axial musculature reduction

102High body and wide, long fins reduce drift in currents. Fin fish and flat body during swimming provide hydrodynamic lifting force.

35 43%18,5 27,3%28 - 3123 - 24>100%29 - 318-10Ps. georgianus

axial musculature reductionBody shape as an indicator of the shape of otolith, because it results from the speed of swimming and life strategy and that all depends on body shape adapted to environmental conditions.

103

Lifting strategy, the use of currents and the uplift force of fins. spreading a large area of fins gives lifting and rapid flow with sea currents without the need for high energy consumption on the fins and body movements. From a large area it is also the possibility of a rapid rebound, catch fish for food and obtain success as a predator.Strategy of swimming: during the day fish focused at the bottom and at night migrates floating and falling vertically. Vertical migrations measure better the high of otolith, when radius R9>R3.

migration

Pectoral fins large, widely spread and billowing are floating body up and forward.

104

Lifting strategy, low-energy swimming

migration

Large, wide fins hover body, which is lighter due to the reduction of bone and axial muscleIcefish use of sea currents and hydrodynamic uplift force of fins. High laterally flattened body strongly reduces drift in side direction and takes over energy of sea current. Through this resistance frequency beats of swimming decrease and also needs for high energy and oxygen delivery.

(Le Franois, 2014; 2014a; PolarTrec, 2013; Detrich, 2012; Uve, 2008; Byrd, 2012)(Walesby, 1982; Davison W., 1985; HARRISON, 1987; Twelves, 1972)(abrowski, 2000)

In lifting strategy and low-energy swimming of Ps. georgianus pectoral fins are moving their first rays as spars entailing the sheet of streamer. Forward with a minimum resistance of sheets fins flowing after trace of thin 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 the front and to the sides increase the horizontal plane of fish so keep, support fish to float at required depth level.In the second phase the fins retracted horizontally to the rear are pushing its all surface on water and pushing fish forward. Also locomotor activity have a caudal fin but much smaller. Fin is bent on sideways with the body when fish is turning.

1 phase

1 phase

2 phase

1 phase2 phase

Le Franois, 2014

1 phase

1 phase

2 phase

2 phase

2 phase2 phase

1 phase

pelvic fins have static task and create the jets that smooth and accelerate the flow of water along the body.jetjet

Le Franois, 2014

1 phase

2 phase

1 phase

1 phase

2 phase

2 phase

Also between 1 and 2 dorsal fin there is jet that smooth and accelerate the flows along dorsal fin and bodyjetjetjetjetjetjetjet

Le Franois, 2014

Le Franois, 2014

Le Franois, 2014

Le Franois, 2014

Le Franois, 2014

reduction of the axial musculature of the body Le Franois, 2014

The water pressure creates hydrodynamic force acts on side of the flowing fish.

V speed of the fish, Rh,c front resistRh,b side resist = 22 V

FPFCFAERh,c = 1,5

FAE aero-hydrodynamic force - the force exerted on the body by the environment, which is the result of movement of the body relative to the environment (gas or liquid).FC driving force, thrust (force of pressure induced by pressure of current exerted on the body surface area). Operates forward because of body shape an the resistance of the lateral is 20 times greater than the frontal; FP - drift force; viscosity force (friction at the surface of the body).

Anon, 2006

An asymmetrical shape with respect to the axis of swimming direction causes asymmetric flow that creates differential pressure on opposite surfaces, and thus the driving force to forward.

V speed of the fish, Rh,c front resistRh,b side resist = 22 V

FPFCFAERh,c = 1,5

FAE aero-hydrodynamic force - the force exerted on the body by the environment, which is the result of movement of the body relative to the environment (gas or liquid).FC driving force, thrust (force of pressure induced by pressure of current exerted on the body surface area). Operates forward because of body shape an the resistance of the lateral is 20 times greater than the frontal; FP - drift force; viscosity force (friction at the surface of the body).

Anon, 2006

Factors increasing the hydrodynamic force acting on back of the body. Force: Fa,h = kv2; power: Pa,h = kv3; 2Vcurrent 4FAE

FAE aero hydrodynamic force increases on larger body of fish. Larger, stronger ones are occurring closer to the sea surface, where the currents are stronger with turbulences and eddys. Smaller fish so weaker live deeper where the currents are weaker and also in regions with weak currents, V

FPFC2FAERh,c = 1,5

Icefish have adaptation to cold water. One of them Ps. georgianus live and choose habitat of sea currents so to exist in it, it adopt the shape of the body, fins and otoliths in liftting strategy of low-energy swimming

Anon, 2006

132The smooth surface of the body increases the power of aero-hydrodynamic FA, H Channichthyidae have a smooth skin, without scales, allowing the feeling of each particle of the water flowing and gliding over the surface of the skin and react accordingly by deflection of the body, or by rearrangement positions of fins to reduce the resistance, to increase laminar flow and to eliminate turbulences. Lack of scales could be adopted as an adaptation of a low-energy swimming in cold strong currents, for which in a warm water there is high energy swimming. For example. Salmon, trout, or mackerel. We can find that the lack of scales for icefish is 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 to which oxygen diffuses and therefore scales do not interfere with the diffusion of oxygen through the skin. Certainly scaleless increase skin smoothness.

Jakubowski, 1971, 1982

The sensitivity and skin elasticity in the perception of the body bending

V

FPFCFAERh,c = 1,5

When the stream of water on the side of after current detach and move disordered (turbulent), this will reduce the hydrodynamic forces.The bending body must always be tailored to the nature of the currents. Too big bow causes break away water streams from the surface of the body, for small bow quite similar paths and velocity of water particles on both sides of the odd fins causing a lack of hydrodynamic forces.

The fins increase the smoothness and flow velocity of after current side of the bodyFAE

VRh,c = 1,5

After the first front dorsal fin and before the second dorsal fin creates the nozzle that accelerates water flow on after currant side of second fin and body.

vvv

Body shape as an indicator of the shape of otolith, because it results from the speed of swimming and life strategy and that all depends on body shape adapted to environmental conditions. This could be show by compare of species.

13535,7 - 40%38 - 4037 - 3923 - 2628,5 - 31%37 - 4135 - 3925 - 2860%35 43%18,5 27,3%28 - 3123 - 24>100%14,3 20%13,8 -16,6%29 - 31Larger lateral surface of the body increases the FAE strength: 9-108-107-8body highest, large head and jaws defines predator creates an arrow, pelvic fins large effective for vertical migration. Longer, unpaired fins increase body side resistance .

body less high, but fins: anal and dorsal longer , smaller head larger pectoral fins so larger horizontal migrations

Body less high, but very large anal fin, dorsal and pectoral, so the greatest horizontal migrations. The smallest head reduces front resisting when swimming.Ps. georgianusC. aceratusC. gunnarifactors modeling the shape of otoliths.

Data: Fisher, 1985

13635,7 - 40%38 - 4037 - 3923 - 2628,5 - 31%37 - 4135 - 3925 - 2860%35 43%18,5 27,3%28 - 3123 - 24>100%14,3 20%13,8 -16,6%29 - 31Environmental requirements with respect to efficiency of swimming9-108-107-8High body helps in swimming using shelf currents and countercurrents and vertical migration. Big mouth helps predation.

Intermediate species is not as high as Ps. georgianus so has greater diffusion and slim as C. gunnari, so has less diffusion than it

The most slender body sacrifice the species for predators, but it gives little front resistance, with the big fins giving the greatest diffusion.Ps. georgianusC. aceratusC. gunnarifactors modeling the shape of otoliths.

Data: Fisher, 1985

Shape of otoliths (determined by microstructure) can show the process and direction of growth 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 same factors 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 suffered environmental influences - to which fish during growth is adapting the otolith shape by changes of microstructure of the otoliths, that are treat as indicators of fish behavior.Ps. georgianus has greater body height and shorter lengths than the C. gunnari and C. aceratus also have shorter dorsal and anal fins in favor of head size and the decline of swimming opportunities. Otoliths of Ps. georgianus like its body are high.

Data: Fisher, 1985; Traczyk, 2013, Parkes, 1990

Realization various opportunities of swimming arising from various constructions of body that are adapted to the best use of different habitats of environment allows the perception of this swimming by otolith recording it with appropriate shape.Otoliths of Ps. georgianus and C. aceratus as species are similar, have same shapes similar (~OL), but instead of that have important differences. In otoliths of C. aceratus increments are narrower and proportion: length with respect to height is reversed. Ps. georgianus are smaller, TLPs. georgianus

Traczyk, 2013; Traczyk, 1992; Fischer, 1985

139

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.Ps. georgianus, otolith height OH> otolith length OL, TL body lengthC. aceratus, OH< OL, TL

Data: Hecht, 1978

x = 0,0024 mm;

R9=0,046 mm 48 daysR9=0,82 mm, 554 daysR9=2,35 mm1590 daysCh. aceratus, 45 cm SLS. Georgia , 29.III.1979hol 136, No 75OW=0,0247 gOH=3,44 mmSP APAdditional centers, AP are also available in otoliths of C. aceratus.. They give however a lower elongation than the radius R9 of Ps. georgianus. Dorsal edge for otolith of older fish of Ps. georgianus grows more strongly than in otoliths of C. aceratus..Otoliths of greater length than height indicate a greater range and speed of swimming. This confirms the elongated shape of the body with less weight and with longer dorsal and anal fins by about 10 rays. As otoliths of C. aceratus are not high, so height of their body is reduced.

Data: Traczyk, 1992; 2014

141Ps. georgianus has smaller range of occurrence than the C. gunnariC. gunnari: OH < OLPs. georgianus, otolith height OH> otolith length OL, TL body length

Data: Hecht, 1978

142

Bars on the sides of the body camouflage the fish swimming near the surface C. gunnariPs. georgianusOtoliths C. gunnari are longer than height, indicating a wider occurrence and greater speed of swimming. It confirms the elongated body with a lower height. Otoliths C. gunnari nearly square, two times smaller than otoliths Ps. georgianus.

Data: Hecht, 1978;Traczyk, 2013; 2014

otoliths C. gunnari have more circadian microincrements this suggests a wide geographic distribution in which the length of the day changes.

Data: Traczyk, 2013; 2013

0,1 mm

0,1 mm2,8 : 12,3 : 11,96 : 1C. gunnarii 6,5 cm SLC. aceratus 7,6 cm SLPs. georgianus 8,2 cm TL

C. gunnari6, 3 cm TL

R9=0,046 mm 48 daysR9=0,82 mm, 554 daysR9=2,35 mm1590 daysC. aceratus, 45 cm SLS. Georgia , 29.III.1979hol. 136, s . 75OW=0,0247 gOH==3,44 mmAPSP The otoliths shape of larvae is similar to an oval on median plane and flattened on the transverse plane to reduce resistance. The biggest flattened otolith has C. gunnari so it swims the fastest and farthest. Older fish swim faster, so flattening of its otoliths increases.

Data: Traczyk, 2013

0,1 mm

0,1 mm2,8 : 12,3 : 11,96 : 1C. gunnarii 6,5 cm SLC. aceratus 7,6 cm SLPs. georgianus 8,2 cm TL

C. gunnari6, 3 cm TL

R9=0,046 mm 48 daysR9=0,82 mm, 554 daysR9=2,35 mm1590 daysC. aceratus, 45 cm SLS. Georgia , 29.III.1979hol. 136, s . 75OW=0,0247 gOH==3,44 mmAPSP Larvae otoliths shape is similar to an oval on median plane and flattened on the transverse plane to reduce resistance. The biggest flattened otolith has C. gunnari so it swims the fastest and farthest. Older fish swim faster, so flattening of its otoliths increases.

Median planeTransverse planeTransverse planeData: Traczyk, 2012

1 mm0,1 mm

0,1 mm2,8 : 12,3 : 11,96 : 11,6 : 1C. gunnarii 6,5 cm SLC. aceratus 7,6 cm SLPs. georgianus 8,2 cm TLAlliroteuthis antarcticus ~15 cm ML

9 : 10,1 mm

Larger flattened otolith indicates a faster swimmerS. japonicus (15) 39 cm FL

Data: Traczyk, 2012

147

1 mm0,1 mm

0,1 mmS. japonicus (15) 39 cm FL2,8 : 12,3 : 11,96 : 11,6 : 1C. gunnarii 6,5 cm SLC. aceratus 7,6 cm SLPs. georgianus 8,2 cm TLAlliroteuthis antarcticus ~15 cm ML

9 : 10,1 mm

Scotia ArcWeddell- -ScotiaConfluenceShag Rock

S.Georgia I.

S.Sandwich I.

S. Orkney I.Elephan I.K.George I.DeceptionPalmer A.BallenyKerguelen I.

Bouvet I.

Heard I.

Ch. aceratus: 5-770 m; 53S-65S

Ch. gunnari : 0-700 m; 48S-66SUnusual catch location of Ps. georgianus Balleny I. Russia 2004/05

S. Sandwich I. Germany 1975/76; 1980/81

Kerguelen I. Australia 2003/04Known catch location of fish:

Ps. georgianus: 0-475 m; 53S-66SRange of occurrenceChannichthyidae and mackerel otoliths have large differences confirmed by their complete separation in occurence. Mackerel is not an antarctic fish

Data: Fischer, 1985; CCAMLR, 2012; Traczyk, 2013


S.Georgia I.

S.Sandwich I.


Bouvet I.

Heard I.





Ps. georgianus: 0-475 m; 53S-66S


S.Georgia I.

S.Sandwich I.


Bouvet I.

Heard I.






Ps. georgianus

?

Shelves of Islandsotolith little flattenedsmall range of occurence


S.Georgia I.

S.Sandwich I.


Bouvet I.

Heard I.






Ps. georgianus

?

Shelves of Islands


S.Georgia I.

S.Sandwich I.


Bouvet I.

Heard I.






C. aceratus

?

Shelves of Islandsotolith more flattenedaverage range of occurence


S.Georgia I.

S.Sandwich I.


Bouvet I.

Heard I.






C. aceratus

?

Shelves of Islands


S.Georgia I.

S.Sandwich I.


Bouvet I.

Heard I.






C. gunnari

?

on shelves of Islandsotolith most flattenedthe greatest range of occurrence


S.Georgia I.

S.Sandwich I.


Bouvet I.

Heard I.






C. gunnari

?

on shelves of Islands


S.Georgia I.

S.Sandwich I.


Bouvet I.

Heard I.

C. aceratus: 5-770 m; 53S-65S

C. gunnari : 0-700 m; 48S-66SUnusual catch location of Ps. georgianus Balleny I. Russia 2004/05




Ps. georgianus

?

Shelves of IslandsUnusual catch?

?


S.Georgia I.

S.Sandwich I.


Bouvet I.

Heard I.

C. aceratus: 5-770 m; 53S-65S

C. gunnari : 0-700 m; 48S-66SUnusual catch location of Ps. georgianus Balleny I. Russia 2004/05




Ps. georgianus

?

Shelves of IslandsUnusual catch?

East Wind DriftAntarctic Circumpolar CurrentPolar Front

East Wind DriftAntarctic Circumpolar CurrentPolar FrontChannichthyidae concentrated in eddies and swim in current greatest in a World facilitating migration - they have running in large and small back and forth branches. In winter, the ice cover with reach under icy living world distributes the larvae of fish from shelf to the open ocean, connecting habitat between islands. Wide near and under-ice distribution of krill secures there the food to near-shore larvae lives in that caring under ice world on the open ocean. Channichthyidae have different flattening and proportions of otoliths that indicate different swimming capabilities specialized to cold sea currents in different environments.

The fish larvae drift for food (Hecq, 2007)

In winter, the ice cover with reach under icy living world distributes the larvae of fish from shelf to the open ocean, connecting habitat between islands. In the 80s can reached South Georgia.


East Wind DriftICE

Data: Sahrhage, 1988; Murphy, 2013; Bargelloni L., 2000; Kaufmann R.S., 1995; Eicken, 1992; Vincent, 1988


krillkrill

Wide near and under-ice distribution of krill secures there the food to near-shore larvae lives in that caring under ice world on the open ocean.

East Wind Drift

Data: Sahrhage, 1988; Murphy, 2013; Bargelloni L., 2000; Kaufmann R.S., 1995; Eicken, 1992; Vincent, 1988

161At the surface, where the currents are strong, there were large fish. In the surface waters there were found only 3 postlarvae of Ps. georgianus. Rest, about 100 postlarvae were deeper, where water currents are weaker.

Data: Traczyk, 2012, 2013

Juvenile Ps. georgianus also occurred in shallow pelagic waters, but in the case of running isobath of 150 m and in the northeastern part of the island, more sculptured and sheltered from the wind and West current. In places where there were greater depths juvenile observed deeper.In the surface water, in summer warmer by about 2 C, instead that the fish may have a higher rate of digestion and growth, they were not colonized by larvaes and juveniles, but only by a few large fish, better than larvaes swimers .

Often in the layer of water limited by 150 m isobath juvenile fish did not occur with adults. Hence it can be assumed that, as in the Antarctic zone, here juvenile fish inhabit cooler, deeper water, with weak currents. Older fish inhabit warmer shallower waters, with strong current.Postlarvae of Ps. georgianus do not occur at the surface, they have high activity of antifreeze proteins. Post-larva of C. aceratus do so, because it has low activity of antifreeze proteins AFGP and from that could swim in warmer water.C. aceratus Ps. georgianusAll larvaeNo / 1000 m3

Data: Traczyk, 2013; Traczyk, 2012; North, 1991; Bilyk, 2011

Ps. georgianusC. aceratus

C. gunnariThe shape of the otoliths Channichthyidae: Ps. georgianus, C. aceratus and C. gunnari are similar due to the similar strategy of swimming they have a similar body shape. However that species have a little different otolith shape and body what indicate

C. gunnari: 0,98C-1,85CAntarctica

Ch. wilsoni: 1,29C-2,23CAntarctic Circumpolar

CDW

Ps. georgianus: 1,03C-1,91CEast Wind Drift

ABWABW

Cold water descent

HighsLows

Polar Fronticy windAntarctic Bottom WaterCircum-polarDeep WaterAntarctic Bottom WaterHighsActivity AFGP [C]Blood feeezing: [C]change tolerance of water temperature and swimming speed (pelagic life to bottom; temperate to high Antarctic) in a result of different content of AFGP. Why there are swimming differences, where are the causes and how it is go?Ch. hamatus: 1,45C-2,44CWHITE-BLOODED: high Antarctic; ice pack zone; temperate 80S 74, ~70S; 63, ~60S; 63, ~5730S; 52, ~45S 30SPolar cellFerrel cell


-100-200-300-400-500-600-700-800CurrentC. aceratus: 0,54C-1,47C[m]

Bilyk 2011

C. gunnari: 0,98CAntarctica

Ch. wilsoni: 1,29C-2,23CAntarctic Circumpolar

CDW

Ps. georgianus: 1,03C-1,91CEast Wind Drift

ABWABW

Cold water descent

HighsLows

Polar Fronticy windAntarctic Bottom WaterCircum-PolarAntarctic Bottom WaterHighsActivity AFGP [C]Blood feeezing: [C]Ch. hamatus: 1,45C-2,44CWHITE-BLOODED: high Antarctic; ice pack zone; temperate 80S 74, ~70S; 63, ~60S; 63, ~5730S; 52, ~45S 30SPolar cellFerrel cell


-100-200-300-400-500-600-700-800CurrentC. aceratus: 0,54C-1,47C[m]

increase in activity and production of antifreeze proteinsDeep-Water-1,85Cincrease in activity and production of AFGPWhy are there swimming differences, where are the causes and how it is go?

Bilyk 2011Together or parallel with their evolution. Bilyk, 2011; Whrmann, 1996; Clarke A., 1996; Cheng, 1999; Chen, 250)


CDWEast Wind Drift

ABWABW

Cold water

HighsLows



-100-200-300-400-500-600-700-800Current[m]Deep-Water

Ch. hamatus: Nototheniidae; 0,4; 2,5

P. antarcticum[mln./mm3]; [g/100 ml]Pa. georgianusNototheniidae; 0,8, 8,3

S. japonicus: 4, 18Channichthyidae: 0; 0

Ch. esox: 0; 0L. squamifrons


Ps. georgianus0; 0Bathydraconidae: 0,2; 0,8

Macrouridae; 0,99, 3,9M. holotrachys

C. gunnari0; 00; 0Circum-polar0; 0descentAll species of Channichthyidae have lost hemoglobin that reduced oxygen transport

Jakubowski, 1971; Near, 2010; Everson, 1977; Fisher, 1985


CDWEast Wind Drift

ABWABW

Cold water

HighsLows



-100-200-300-400-500-600-700-800Current[m]

reduction in the number of and contentDeep-Water



S. japonicusChannichthyidae: 0; 0





C. gunnari0; 00; 0Circum-polar0; 0descent

reduction of red cells and hem4; 1.8BUT INSTEAD OF THE MAIN TREND (WITH LOWERING WATER TEMPERATURE)LOW TEMPERATURE

hemred celsJakubowski, 1971; Everson, 1977


CDWEast Wind Drift

ABWABW

Cold water

HighsLows



-100-200-300-400-500-600-700-800Current[m]Deep-Water



S. japonicus: 4, 18Channichthyidae: 0; 0





C. gunnari0; 00; 0Circum-polar0; 0descentthey spread on all Antarctica from temperate to high Antarctic waters, from surface across pelagic to bottom deep waters.

Jakubowski, 1971; Fisher, 1985

descent of cold waterHSSWHigh Salinity Shelf Water

0.96C34,84 Antarctic Surface34,18 -100-200-300-400-500-600-1.91CRemoving fish from the shelf by glaciers

Increase O2 pressure in the blood and tissues

increase saturation O2 in hemoglobin

drop in temperature when pO2 = const.

increasing the affinity of hemoglobin for O2 impedes, even impairs putting into tissues;Blood viscosity increase when the drop in temperature100% O2Dyfuzja70-90% O2

CDW

Channichthyidae

diffusion

Water70-90% O2diffusion

lowers blood viscosity, and this > flow, transport of O2[m]

?Not ! Bottom fishFish migrationFish migrationreduction of red blood cells-1.91Cchanges withchanges withchanges withFish migration

migration to bottomThe links in which loss of red cells and heme content determine a variety of strategies of low-energy swimming and thus different otolith shapes

Jakubowski, 1971; Near, 2010; Everson, 1977; Fischer, 1985Kunzmann, 1991; Rakusa - Suszczewski, 1989; White, 1977

descent of cold waterHSSWHigh Salinity Shelf Water

0.96C34,84 Antarctic Surface34,18 -100-200-300-400-500-600-1.91CRemoving fish from the shelf by glaciers

Increase O2 pressure in the blood and tissues

increase saturation O2 in hemoglobin

drop in temperature when pO2 = const.

increasing the affinity of hemoglobin for O2 impedes, even impairs putting into tissues;Blood viscosity increase when the drop in temperature100% O2Dyfuzja70-90% O2

CDW

Channichthyidae

diffusion

Water70-90% O2Oxygen diffusion

lowers blood viscosity, and this > flow, transport of O2[m]

?Not ! Bottom fishFish migrationFish migrationreduction of red blood cells-1.91Cchanges withchanges withchanges withFish migration

Reduction in the number of red cells and heme reduces transport, storage of oxygen to the muscles of the body and thus their reduction and develop swimming strategy with low energy in all Antarctic species.

migration to bottom

At first white-blooded in relation to red-blooded have a larger size so they have reduced heat loss due to the lower surface. Bergmans rule of energy benefits.Channichthyidae achieve larger body size.eggs, mmLarvae, mmAdult, cmincr, cm/yWhite-blooded Channichthyidae 54-1743 6-10 Red-blooded Nototheniidae36Red-blooded Bathydraconidae26Red-blooded Harpagiferidae15

To fulfill strategy of swimming speed to different environment each species has a different compensation of reduced oxygen transport what differentiates their ability to swim and thus differentiates the shape of the body and the otoliths. Icefish have different life strategies and occupy different habitats (La Mesa M., 2004)

412

248Mass = 1Mass=8Area = 28Area = 112Area/mass = 28Area/mass = 14

Jakubowski, 1977; Wells, 1985; Kock, 1991; Johnston, 1983additional oxygen from skin breathing of large head for the heart (Detrich, 2012).

sink of cold icy wind from the poleIce formation

warm water from the equator

salty water Bottom lifestyle of Channichthyidae. Chaenocephalus aceratus has the body weak with a reduction in the axial muscles, which is probably due to the large volume of blood, up to 9% of body weight usually poorly vascularised.

Jakubowski, 1977; Wells, 1985

sink of cold icy wind from the poleIce formation


salty water White-blooded also have a reduction in ossification as a result of displacement of benthic fish fauna from shelf by glaciers to greater depth or to the pelagic. It enforced a reduction in body weight, because the white-blooded does not have a swim bladder.

Bone mass is replaced by cartilage by inhibition of skeletal development in the early stages of ossification of cartilage skeleton. Reduction of fish bone were observed in several ways: replacing the entire bone to cartilage, ossification of the surface (eg. Ethmoid region), the separation of the bones by large areas of cartilage (such as in the case of the brain), reducing the dimensions of the bones and even the lack of it. Reductions ossification observed in white-blooded (large in C. wilsoni) cause large osteological variation, the asymmetry of the bones even. The reduction process is still in progress, and skeletal elements undergo constant metamorphosis.

Walesby, 1982; Jakubowski, 1977matured form similar to the larvalabrowski, 2000; Byrd, 2012

Icefishhuk. Additional oxygen from skin breathing of large head for the heart (Detrich, 2012; Kils, 2008).

icy wind from the poleIce formation


salty water

Additionally C. gunnari have no reduction an axial muscles, between vertebrae, its streamlined body shape indicates a pelagic life. Dense vascularization in skeletal muscle reduces the distance of oxygen diffusion - increases oxygen transport. A similar body shape and pelagic lifestyle has species Champsocephalus esox living in warmer waters outside the Antarctic near Falkland Islands and Patagonia. Its gills are not as specialized as in other species of the family.

sink of cold

Walesby, 1982; Twelves, 1972Pelagic 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, which increases oxygenation by larger capillaries and large spaces supplying capillaries with blood.

Pelagic lifestage expect for C. gunnari to be a good swimmer with low supplay in energy.

The activity of alkaline phosphatase determines the size of the muscle vasculature. Reis, 1970.Greatest is in C. gunnari hence this species has most capillaries. s [enzyme units/g wet muscleh]species / muscle typepectoral finoxidativeglycolyticC. gunnari, n=7, 8, 81185,5101,4770,062,5400,043,4Notothenia rossii, n=8, 9.Gadus morhua, n=10698,1118,5405,837,5483,271,0353,565,1310,037,6100,319,4

Its slender body shape increases heat loss, by Allens rule. 1661Mass = 8Area = 192+12+32=236Area/mass = 236/8=29,5Mass=8Area = 96+48+16=160Area/mass = 160/8=2022412

Ps. georgianusC. gunnari

It is agree with Allen's Rule for energy benefits of having a more slender (less resistance) in the warm waters.

Low-energy swimming is on the pectoral fins only?High energy swimming by body waves ChannichthyidaeNototheniidae AntarcticGadidae C. gunnariN. rossiiPollachius virensOncorhynchus mykissglycolyticoxidative

glycolyticoxidativeoxidativemyomer musculesmall diameter fibres(without enzymes)lateral line channel skinoxi 1oxi 2skin

oxidativeskinOxi 2mosaicoxi 1 & oxi 2White fibers, fast reactionmosaic of white and red fibersred fibers oxidative slow reaction Altringham, Ellerby, 1999; Davison i Macdonald 1985, Harrison i in. 1987, Walesby 1982swimming on the pectoral fins saves more energy (than by body waving), because the muscles of that fins are slow oxidative fibers adapted to continuous low-intensity movements (consuming less energy), ensuring the long-term swimming at low speed. C. gunnari has also oxidative and glycolytic fibers in axial muscles.Oxi 1pectoral fin muscle glycolytic

Low-energy swimming is on the pectoral fins only?High energy swimming by body waves ChannichthyidaeNototheniidae AntarcticC. gunnariN. rossiiglycolyticoxidative

glycolyticoxidativeoxidativemyomer musculesmall diameter fibres(without enzymes)lateral line channel skinoxi 1oxi 2skin

oxidativeskinOxi 2mosaicoxi 1 & oxi 2mosaic of white and red fibersred fibers oxidative slow reaction Altringham, Ellerby, 1999; Davison i Macdonald 1985, Harrison i in. 1987, Walesby 1982Antarctic Nototheniidae with pattern of low energy swimming has also mosaic of white and red fibers as living in currents high energy swimming fish of SalmonidaeOxi 1pectoral fin muscle

SalmonidaeOncorhynchus mykiss

O2, o-oxidative fibres, slow reaction -actATP: 2-5lower C. gunnariN. rossii

N. rossii

75340155

320C. gunnariHistochemical properties similar; lack of myoglobin compensatehigh vascularization, thicker capillaries and numerousbays reduces distance and amount diffusion O2 transport more efficientGglycolytic fibres ( large) small, many capillaries, lipids, glycogen, SDHenzyme

Walesby, 1982

1661Mass = 8Area = 192+12+32=236Area/mass = 236/8=29,5Mass=8Area = 96+48+16=160Area/mass = 160/8=20Ps. georgianus has a shape more compact than C. gunnari and in a result (by no change in body weight) it reduces the ratio of its body surface to the body weight, hence from that reducing loss of heat through the smaller surface of the body - which is important in cold water.It is agree with Allen's Rule for energy benefits of having a more stocky body shape in cold water.22412

Ps. georgianusC. gunnariSemipelagic lifestyle of Channichthyidae.

Fisher, 1985

182

A more massive Ps. georgianus is less susceptible to chilling of the body. Large individuals need relatively less food, having a slower metabolism. Increasing the size of an individual (without changing its shape) need increase of the strength in muscle by fourfold - muscles must carry and move 8 higher body weight.Additionally Ps. georgianus have in the myocardium hem that accumulate oxygen to the heart muscle so in this case fish has more energy to transport oxygen in enlarged body.

Hofinger, 2010; Kils, 2008

density of C. aceratus in 2 depth zone was greater than for Ps. georgianusPs. georgianus occurred on the northern and eastern shelf of Georgia I., while C. aceratus on the west side.Different flattening and proportions of otoliths indicating differences in swimming even among icefish could be confirmed in their occurence and interspeciec ratio.

(Traczyk, 2012a, 2012b; Sosiski, 1989; 1989a; Traczyk, 2013; Traczyk, 2012; Traczyk, 2013a

density of C. aceratus in 2 depth zone was greater than for Ps. georgianusPs. georgianus occurred on the northern and eastern shelf of Georgia I., while C. aceratus on the west side.Different flattening and proportions of otoliths indicating differences in swimming even among icefish could be confirmed in their occurence and interspeciec ratio.

47.8

43 42 41 40 39 38 37 36 35 34555430'30'30'

43 42 41 40 39 38 37 36 35 34Vertical temperature profile.

53555430'30'30'531C1C1C1C1C1C>1C; 0,6C; 1C; 0C; 1C; 0C; 1C; 0C; 1C; 0C; 3C; 1C; 1C; 2C; 2C; 4C; 1C; 6 = SGIThe dominant age group I of Ps. georgianus whose otoliths shape is more oval, is spreading on the shelf away from the island and to the west to cooler water. It migrates even west into the peripheral habitat at rocks of Shag Rock. A small number of large Ps. georgianus relieve their main area of focus north - eastern shelf and it is now colonized by C. aceratus - in the absence of its predator Ps. georgianus. Pelagic C. gunnari inhabits the western colder part of the shelf.

192Competitor C. aceratus displaces Ps. georgianus from 12 statistical squares. ACE = 12> 6 = SGIThe dominant age group I of Ps. georgianus whose otoliths shape is more oval, is spreading on the shelf away from the island and to the west to cooler water. It migrates even west into the peripheral habitat at rocks of Shag Rock. A small number of large Ps. georgianus relieve their main area of focus north - eastern shelf and it is now colonized by C. aceratus - in the absence of its predator Ps. georgianus. Pelagic C. gunnari inhabits the western colder part of the shelf.

Ps. georgianusC. aceratusC. aceratusC. aceratus

193Dominant in 1989-1990 age group of I of Ps. georgianus, now dominates as 3 age group of mature fish, in the Northeast, in a warmer area of larger current turbulences accumulating krill. This season Ps. georgianus is large and numerous, so C. aceratus under her pressure is maintained on a small area of cool water and more threatened remowing with current beyond shelf.Competitor C. aceratus displaces Ps. georgianus from 8 statistical squares. ACE = 8> 7 = SGI

194Dominant in 1989-1990 age group of I of Ps. georgianus, now dominates as 3 age group of mature fish, in the Northeast, in a warmer area of larger current turbulences accumulating krill. This season Ps. georgianus is large and numerous, so C. aceratus under her pressure is maintained on a small area of cool water and more threatened remowing with current beyond shelf.Competitor C. aceratus displaces Ps. georgianus from 8 statistical squares. ACE = 8> 7 = SGI

C. aceratusC. aceratus

Parkes, 1990; SARAH CLARKE, 2008; Traczyk, 2012; Traczyk, 2012; Traczyk, 2013

Larvae of Ps. georgianus occurred frequently in winter in the northern side of South Georgia, where there were large numbers of larvae of C. gunnari, and in the summer on the west, cold side of the island, where there were large numbers of larvae of C. aceratus. After North, 1989195High depths of the ocean surrounding South Georgia Island and the strong West current flowing around determines Ps. georgianus, rather gliding on pectoral broad fins than actively opposing sea currents, to focus in the limits of the shelf of the island. Within these limits there are all ages of Ps. georgianus: larvaes in summer inside a belt of 10 km, 5 km in winter (after North, 1989). Post-larval forms, accumulate in the coastal zone for searching small food. Stomachs of 7 cm Ps. georgianus contained 3 cm larvae of C. aceratus and C. gunnari. Ps. georgianus could prey fish of up to 89% of the length of their body. Larvae of Ps. georgianus migrate vertically for feed on C. aceratus and other larvae - they swim as their flattened otoliths show, must in currents with enough speed to cach a fish.

C. gunnari C. aceratus Ps. georgianus all larvaeThe influence of various factors on focusing of fish larvae on shelf of South Georgia in the seasons 1980/81, 83/84, 86/87, 87/88. After North, 1989

1.0--0.9--0.8--0.7--0.6--0.5--0.4--0.3--0.2--0.1--0.0--All factorsdistance from the coastdepthDay night, run offTime of dayseasonarea

South Georgia I., summary

The larvae of Ps. georgianus migrate to western waters, cooler, in which oxygen absorption through the skin is higher.

SARAH CLARKE, 2008

On the small shelf of Shag Rock, large fish Ps. georgianus and C. aceratus compete strongly for food, which is krill. In summers between1989 - 1992, at Shag Rocks occurrence of Ps. georgianus, was accompanied by lack of C. aceratus and vice versa.

mass, average lengthNumber of larvaewintersummeraveragePs. georgianusC. aceratus

Depth zone

NumbersStat.squareS. GeorgiaS. GeorgiaS. G.S. G.

Horizontal fish migrations. Large adult of Ps. georgianus, occurred in large numbers in the north, and in summer also in colder western side of the island. They also have vertical food migrations and show concentrations: fish larger - shallower in vortex warmer waters; smaller - deeper in colder waters. Horizontal in Antarctic zone: colder Palmer has a smaller fish, warm Orkney, bigger.

Lack of Ps. georgianusLack of C. aceratusLack of C. aceratusTraczyk, 2012; Traczyk, 2013; 2013North 1990

197Phylogeny statocyst and otolith - the relationship with the movement, that determine changes in their shape.Statocysts containing otoliths in the evolution of animal play one of the most important tasks: they give the opportunity to gain space, a new environment, food, escape from predators, thus ensure for species the survival and success. Without them, need to above the precise registration of changes in body position in the animal kingdom (with the exception of volatile insects) would be impossible. Otoliths serve not only maintain a balance, but also participate in the perception of sounds. Like the change of otolith shape sign up the development stages of the fish: from the ball in a stationary fish embryo to elongated in swimming adult fish, it can be traced in the interspecies changes of statoliths, the evolution of capabilities and speed of swimming of animals on successive organization level of the animal kingdom. From the simplest forms detection of balance changes in metazoan to a specialized organ of balance and hearing in bony fish in the process of concurring a new environment. Free-living animals having statoliths that during ontogeny transform to sedentary life style with a radial symmetry were lost statoliths.

lacks a centralized brainLost statocyst

brachiolaria larva

bipinnaria larva

5 statocystsSea cucumberJura., 1983

198Development swimming, moving and flying possibility with statocysts in animals:free livingradial the bilateral symmetry.The otolith shape and microstructure evolved to be the best in serving the perception of oscillations caring information on body move and sounds. All moving animals largest and small, high and low organized have it in water and also this heavy watered organ was taken on land and in the air, excluding only Pterygota because those ones do not have needed appropriate strong osseous base to carry it. And it is so concerning that sessile, parasites and simpler without nerve system organisms do not 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 basilar membrane, pb papilla basilaris

At all, the otolith means the possibility to swim and migrate, go to the new space to extent the border of species settlings. All evolution based on motion for challenging space and environment show it. The medusa free living hydrozoans have statoliths which lack of them in sessile sponge. But even sessile sponge in a free space of their bodies that are filled with collagens, the spicule forming, which percept the turns of their body - showing build up of the origin of otolith microstructure and its idea. In this same way in the gaps of otolith organic net, the aragonite crystals growth.

Spicules in the gaps between sponge cells. Also in the gaps is mesohyl: mostly collagen polymerizes into spongin - collagen fibersGaps in real collagen fibres aragonite crystallizing in the corners.

Anon., 1983Jura., 1983

200Several statocysts at the edges giving orientation in space;Radial symmetry of free-floating organisms have spherical statoliths Every step forward to improve the possibility of moving to extend migrating is displayed in change of statolith shape. If animal swim faster enough it has appropriate large deviation of otolith shape from a ball. In slow moving, the changes work on body shape and statolith localizations. Animal only floating are radially symmetrical 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.

m e n s g t z k t

r z l

Jura., 1983

One statocyst at peak, the perception of sound vibration. Bilateral symmetry, actively swimming organisms: statolith sphericalIn this evolution step animal are with one statolith in the topside of body, to get similar information from all directions. More swimming animals are changing body to bilateral symmetry and statolith localization in a front to the head (ctenophora), or to head part as a very important perception organ for swimming after a food or escaping from predators. In head part that 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 front of the body, and are only in living free animals, having birateral symmetry.

Statocystof Ctenophore statolithStatocyst of ctenophore comb jellies.

ctenophoretentacleStatolith tile cilia a mouth

trochoforaStatocyst nervous system anus a mouth

Jura., 1983

Statocysts at two sides of the brain or at larger distance, but innervated by the cerebral ganglia; The bilateral symmetry, statoliths elongated.In a further evolution of the bilateral symmetry in the expansion of the new environment molluscs and arthropods form a pair of statocyst which are most optimally localised 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 the needs 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 be related to ontogenesis, the change in the living from epipelagic larvae to adult life, swimming deep into the meso and bathypelagic waters.

gonad cecum Inner shell ventricle mantle cavityesophagus cerebral grater and beak of tongue grater jawfunnel statocyst gills Ink bag Gili heart stomach

Jura., 1983

203As squid swim faster their larvae otoliths originally spherical (N nucleus) in the elderly have become more elongated and acquired the shape of the dual spherical waves.

Alliroteuthis antarcticus 100m Galiteuthis glacialis 200mThe spherical nucleus (N) and elongated post-larval growth zone (PZ) of squid otoliths.

Arkhipkin, 1996; JACKSON, 1994

2045 statocyst in place of 2 in bilaterian animals. Deuterostomes repeat the beginning of the development of the sense of balance.In the new stage of evolution in which into place of the blastopore of the embryo an anus was re-formed, the vulnerable period of embryogenesis is extended. Hence the first Deuterostomes - echinoderms develop in a safe aquatic environment, which is at the bottom, and leading sedentary lives. Accordingly to that, the bilateral symmetry of larvae 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 of statocyst. Statocysts are localized in old numerous mode, only if they have a bilateral symmetry as a form more useful in the free mode of life.

c b Sea 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.e f g i h j k m l n Squirts (sea cucumbers), drilling and pelagic animals, has five statocyst located around the oesophageal ringo a c d b

Jura., 1983

1 central instead of 2 statocyst in bilaterians. Deuterostomes repeated the centralization of the sense of balance.The last line Deuterostomes of chordates, the tunicates confirm the centralization of sense of balance to 1 statocyst in the vicinity of nerve ganglia. Wherein in some of them, the central statocyst probably corresponds to the central cavity of a brain as in other chordates. Appendicularia - Larvaceans lead free lifestyle, or settled, are without excretory system and have opened the circulatory system. They are considered, as neotenic larvae. Those organisms have lost during evolution the mature stage but gained 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 which are pelagic organisms. In the next organism in Thaliacea statocysts occurs in cases where 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.a a e b c d f d g g h i j k sea squirt larva Doliolium denticulatum

Jura., 1983

2062 statocysts, each developing a labyrinth in bilaterians. Deuterostomes reach near the brain as the best location for statocysts.In vertebrates, each of a pair of statocyst develops in the membranous labyrinth containing statoliths adjacent on both sides of the brain. This structure has three channels for decomposing the measure of the movement into 3 components. They develop sequentially starting from one channel only in the first aquatic vertebrates characterized by achieving high speed: in accrania bottom living hagfish. Although not yet have a dorsal fin needed to sedate of faster movement, they have 1 channel, that increases 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 slow movable the sphere is the best to percept the vibrations carrying the information on body changes and on sounds in environment from all directions. If fish velocity is large the signals from space are different between forward and afterward and backward. It is compensate by change the sphere to the elongate shape of otolith. Longer radius of otolith that is in motion percept the same signal in similar period as radii of otolith sphere no moving

(Radtke 1985)

No significant changes in labyrinth of terrestrial vertebrates The development of sound perception by labyrinth.Labyrinth evolved among aquatic vertebrates as a whole has moved further by them in space acquiring on the land and in the air.In these new environments, fish labyrinth still provides for the birds and land animals the balance without major changes in its structure and operation. Also reading sounds of terrestrial vertebrates despite excelling in a result of extension of Ladena into cochlea, but it proceeds as in fish in the aquatic environment in the labyrinth as a result of the vibration transfer of endolymph on cilia of innervated hair cell of fish lagena elongated to the twisted cochlea of mammals.In the organisms evolution, the process of development of statocysts containing statoliths combines them into a common direction for getting a better perception of their position during faster swimming or moving, by measure vibration and in addition by interpretation of their acoustic parts for the knowledge of environment and to generate them in order to communicate within and between species.

Low sounds in lagena high in cochlea (Roy, 1994; umech.mit.edu, 2013; Inoue, 2013)

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