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"Migrations of the Antarctic fish Pseudochaenichthys georgianus NORMAN, 1939
in the Atlantic sector of Antarctic". Gdynia 2012. Ryszard Traczyk
ABSTRACT
The material obtained between 1986 and 1991 has allowed to study the pattern of
geographical and depth distribution of icefish in the area of Scotia Arc Islands and on shelf of
South Georgia Island. In the Antarctic colder waters in the Palmer Archipelago there were
mostly young, small fishes. They have stomach filled well with krill so swim probably to match
with distribution of krill, which flow further to North and East with geostrophic currents.
Following after them, with stomach often full of them, they became large, so the numbers of
small individuals are disappearing farther from continent. Their gonad are more developed as
the temperature further from continent get warmer. Finally at the end of Scotia Arc Islands - at
South Orkney, Ps. georgianus are large with gonad ready to spam, and mostly concerning on
reproduction, they do not feeding, so they do not have reason to follow after krill driven to the
East to deep open Scotia Sea.
In vast Antarctic Zone, enlarged by ice cover species are distributed into age groups
separated geographically: first age group as it was not in sampled area, have to be assumed to
be near or under ice feeding on juvenile krill and fish; age 2 with 3 are in the South Shetlands;
age 4+ are accumulating at South Orkney. While on shelf of one, small sub-antarctic island the
group ages of Ps. georgianus are separated by time in a patterns of strong – week cohorts: age
group of 4 is separated from the 2th by low numbers fish in previous age group 3. This may
save them before feeding of own younger fish – common fish behavior among predators.
At South Orkney there are no small fishes. After hatching in cold winter waters as
coming summer warm them, they swim upstream (as young fish usually do) to colder water of
Palmer Archipelago to have more efficiency in food assimilation. This behavior save them
before to be driven to open pelagic water and ensure good reproductive success as old fish
drifting to warm water to gonad develop and to span.
At South Georgia similar pattern exist in the distribution of Ps. georgianus. Large
specimens were closer surface near Eastern shores of South Georgia where water is warmer and
sheltered before to be driven to Eastern large pelagic sea. In such conditions there were eddies
with larger krill aggregations (1). After spawn young postlarvae swim upstream to the West, to
the deep and cold waters. Span, feeding, diurnal and another migrations means changes of
environments and physiology - events that have appropriate marks in the otolith microstructure.
INTRODUCTION.
The results of ecological study create the movie which helps humans understand needs of
wild life. For Antarctic underwater life, it is difficult and very rare. We need information on
ecology. Change in life pattern can be explained by changes of environment. Now we face
global warming that has great impact on marine life at South Georgia and further to South. We
have already example influent of large change in environment on distribution and biology Ps.
georgianus from its cold conditions settlements in Scotia Sea Arc to warmer area South
~ 2 ~
Georgia. Data of all Antarctic expedition shown, that Pseudochaenichthys georgianus appear in
cold water on shelves Scotia Arc Island from Palmer Archipelago to South Orkney Island and
outside Antarctic border (in Aliasow classification) about half thousand km further on North
across open deep Scotia Sea in warmer Subantarctic South Georgia Island.
Additionally they have vertical feeding migrations from cold deep water to warmer
surface and came back following the krill and fish larvae diurnal migrations (2; 3; 4). Also
mature fish migrate inshore to warm surface water to spawn (5). So larvae after hatching have to
migrate to offshore to cold deep water across body transformation - process marked in otolith
microstructure as the other environment changes, e.g. from inside to outside of egg (6).
The biology and distributions of Ps. georgianus from South Georgia area were described
in papers by researches from British Antarctic Survey, Cambridge (7), but there is no the same
data for an Antarctic waters.
Older fish in this area migrate to shallow spawning waters warmer that are at North East
side of Island. After month, two later, depend on water temperature larva hatch in July and
August. They growth by the von Bertalanffy: Lt=66.11(1-e-0.283(t+0.008)
); L0=0.15 cm; r2
=0.98,
(’=3.09), which is as other icefish very high. Postlarvae, about 8 cm long migrate to deep
colder water out of shore. They also swim upstream to Western colder parts of South Georgia I,
were they growth faster. In first year, fish growth up to average 19 cm TL, in the second year -
33 cm. In the age group of III fish may have 44 cm TL length, have first spawn, and because of
that migrate to shallow and warmer water in the East. Next year, 4 year fish have 50 cm TL, and
as their maturity increase, more - above 50% fishes in population are ready to span, and then
they do not feed. Their length growth very slow – fish a year older reach only 53 cm TL.
As the Subarctic S. Georgia is bordered by deep ocean, migration of Ps. georgianus is
restricted to shelf this Island, probably because of warm temperature. Ps. georgianus unlike to
similar species of Ch. aceratus do not extend their appearance with Antarctic current to East,
and also not migrate to warmer surface water to which Ch. aceratus have better adaptation: e.g.:
they have lighter coloured in wave pattern skin of body to be invisible by avian and large sea
predators.
In Antarctic our species of South Orkneys hatch later in November – December and
during ontogenesis show similar temperature induced distributions that for fish depend on water
temperature means migrations to more familiar waters. Antarctic Ps. georgianus represents
growth to similar size and pattern, as in Subantarctic S. Georgia in that time, with Bertalanffy
equation: Lt=66.32*(1-e-0.261(t-0.009)
); r2
=0.99, ’=3.06, L0=0.15 cm. In summer in first year fish
have 17 cm, in second 31 cm. Age group of III have average length about 41.7 cm, age of IV
have 47.7 cm, age of V: 49.9 cm, VI have 51.3 cm TL, age of 0 (postlarvae): 5 cm (after
Ślósarczyk, pers. com.).
The smaller fish were distributed close to continent in cold water of Archipelago
Palmera, were they feeding the most on krill and follow after them to North East up to Orkney
Is. In this warmer region they do not feed but they spawn. The extreme unevenness in spatial
distribution of the species was there probably due to dense pre-spawning aggregations (8).
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Ps. georgianus in Antarctic waters feed more than at Subantarctic South Georgia. In the
South Georgia fish do not need as large energy and activity as in cold south and thus food as
well. Swimming upstream is usually performed by young fish, futures that play in both areas for
saving the species - not fast and long swimmers before spreading to unfamiliar for them large
open pelagic Scotia Sea. In another side unusual higher temperature at South Georgia because
of global warming may suppress feeding activity and in addition food concurrence with
Ch.aceratus may be empowered.
In area of South Orkney young larvae after the hatch in October oppose geotropic
currents carrying out into the open waters of the Scotia Sea by swimming up current to cold
waters of the Archipelago Palmera. They have higher growth in the colder water and in that area
the larval stages of the krill are concentrating in deep pelagic, which is ate by them together
with associated larvae. Pelagic postlarvae of Ps. georgianus are feed on krill (9). It is probably
true, as there were similar Western up currents directions in migration pattern at South Georgia.
In krill aggregations on East from South Georgia, there were fish larvae, e.g. of Ch. aceratus
but there were not larvae of Ps. georgianus. Larvae of Ps. georgianus were on the Western side
off the krill aggregations (10). Also in Eastern peripheries South Shetland Islands, in ecological
studies of the sea-ice zone near South Orkney in summer 1988/89, there were not larvae Ps.
georgianus among larvae such Ch. gunnari, Ch. aceratus found in krill aggregations. Larvae of
Ps. georgianus were reported only from Palmer Archipelago, and generally other larvae of
Channichthyidae were in greatest density on the West of South Shetland Is (11).
Opposing geotropic currents conditioning the reproductive success in Antarctic has the
similar course on the shelf of South Georgia. The most hatching larvae were fished in the
sheltered from current East-northern side of the Island. They hatch in shallow ~50m cold winter
water (12), so adapting to it at the beginning through the early developing in it. Consequently as
they growth, the water became warm as summer coming up, what driving them to sink deeper,
so must off shore, direct to familiar cold water this means upstream to West, where this species
was cached in larger amount. Inshore, in shallow water larvae of Ps. georgianus that hatched
there in winters, in summer were not cached, as e.g., Ch. aceratus (13; 14), and were not
numerous present on the East with krill as the other larvae, but outside of krill swarms and
farther to West (10). Postlarvae to get more effective utilization of the food they migrate into
deeper colder waters and upstream westerly to the Western side of Island where flow cold
current stimulating larger increases of the body; as 7 cm long postlarvae they have been better
prepared to swim in pelagic water (12) with their lighter colour of body (9), and well developed
fins. The perpendicular migrations of this species also reflect thermal preferences of young Ps.
georgianus to cold waters - in summer being on larger depths and of the older fish, shown
maturing depended extreme unevenness in spatial distribution (8) concentrating in shallow and
warmer waters for accelerating their maturation and the development to the spawn.
Observed decline of the sizes of body Ps. georgianus during many years observations of
the region of South Georgia, can result from the warming up of the climate to of whose Ps.
georgianus probably shows the slower growth and late maturation.
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MATERIALS AND METHODS.
Collecting sample.
Samples of Ps. georgianus were collected during one to three month long cruises in the
Antarctic Peninsula and the South Georgia area since 1976 up to 1991 in the almost every
summer. Collections were made by the international science teams of Fish Stock Assessment on
the research ships r/v "Professor Siedlecki", r/v "Professor Bogucki" and the trawlers m/t
"Gemini", "Sirius", "Taurus", "Carina", "Libra", and "Hill Cove".
The haul jobs were lead with the bottom trawl P-32 / 36 having the opening horizontal
17.5 m, perpendicular 4.5 m and a codend mesh of 80 mm with the insertion of the mesh side of
20 mm. The trawling was usually with the speed 3.5 knot during generally 0.5 hour of the time
of duration of the hall. Additionally in 11-12.02.1989, 10 pelagic net hauls were made on three
transects with pelagic net of WP 16/41 x4 with the insertion of mesh side 20 mm.
22.6
14.4
64 60 56 52 48 44 40 36
54
56
58
60
62
64
54
56
58
60
62
64
64 60 56 52 48 44 40 36
Feb.1979 – the R/V “Prof. Siedlecki” (N = 67)
Nov. 78 – Feb. 79 – the M/T “Sirius” (N = 30)
Ps. georgianus capture, [kg/h]
Mean water temperature [°C] at ~20 m in the summers
5.1
47.8
S o u t h
S h e t l a n d
South Orkney I.
South Georgia I.
Bransfield Strait
S c o t i a S e a
W e d d e l l S e a
Elephant I.
PalmerA.
shallow sea bed 0 - 500 m
2°
0°
>2° >2°
>2°
0°-2°
<0°
<0°
Fig. 1. Caches of Ps. georgianus during surveys on research vessel r/v “Prof. Siedlecki” and on travel m/t “Sirius” on the shelfs of South Shetland
and South Georgia Islands in summer 1978/79. The mean water temperatures at about 20 m in the summers were from oceanographic stations
from S. Shetland and S. Orkney of period 1927 to 1980 (15), that were extend to South Georgia area by hydrological study of period of 1955-
1975(16). Because Antarctic waters from region North of South Shetland Islands flow to North-East direction, the temperatures of upper waters
of South and West sides of South Georgia Island were similar to waters of South Shetland (0-2°C). Instead of that North and East sides of South
Georgia are in average warmer, above 2°C. Waters of Palmer Archipelago and South side S. Orkney are colder - below 0°C.
Investigations carried out on the shelves of South Georgia and South Shetland Islands in
depths 50 - 500 m at random chosen stations. At each station, total catch was estimated and its
fish subsamples were chosen. Fish were sorted by species with taxonomic key (17), with otolith
key (18) and key for larvae (19). Ps. georgianus like other industrial species were measured
following instructions of BIOMASS (20). Total length (cm), total weight (g), sex, maturity stage
in five-point scale and stomach contents in five-point scale for each individual of species
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sample were recorded, with otolith subtraction (21).
Density, biomass and length frequency presentations.
The mass of the each catch were linked to area and time trawled to estimate fish density.
From above data, biomass were estimated using swept area method, for all region, and in
account 3 depth stratum (50-150, 150-250, 250-500 m) of Everson’s bottom surfaces of fish
statistical squares (22; 23; 24). Catchability was set to 1. The larvae of Ps. georgianus, Ch.
aceratus, Ch. gunnari and other species in range of 3 - 7 cm total length were found in the
commercially travelled net. They could supply some information about postlarvae and juvenile
stages of fish.
Age determination.
The ages was determined on the base of the results obtained from daily increments count
in otoliths (6) and the otolith internal and external with body morphology analysis with taking
into account environmental and physiologic influences.
Fig. 2. EDTA removing CaCO3 left gaps in matrix of colagenlike fibres. Those gaps are arranged in a pattern of daily increments rings, that were
measured. Into ancles of those gaps cristals of aragonite are deposited to the size and form, which they were restricted by dimensions of gaps.
Aragonite from otolith, when were outside of gaps cristals in a long needles (picture on the right).
Central Primordium CP in 3 dimensions is a 0.064 mm diameter ball which describes no
movable larvae in egg. Larger otolith is two sides flattened and describe movable larvae in egg.
Otolith elliptic after 46 daily increments on long radius 0.098 mm having hatching mark
(increments 2 times wider 2,5·10-3
mm with less colagenlike fibres) describe about 1.5 cm
larvae which just get out from the egg, and free swimming, and having large changes in
physiologic and in the environment. Those great changes draw marks in otoliths of other
icefish.
Fig. 3. Hatching mark in otolith from the left: of Ps. georgianus at radius 0.09 mm with 46 daily incriments; of Ch. gunnari with 21 daily
incriments; of Ch. aceratus at radius 0.034 mm with 24 day incriments. On the rigcht gaps in otolith matrix protein from Ch. aceratus (25).
Flattened otolith having 0.15 mm Second Primordium SP (source of new radial
increments, that start to increase otolith length 2 times and perception of shorter frequencies) on
longer radius of 0.8 mm usually after 240 daily increments describe about 7 cm postlarvae of
age group 0, transforming, moving to deeper water and swimming 2 times faster. They were in
bottom catches in January. Otolith having longest radius more longer, of about 1.8 mm, and SP
0.01 mm 0.01 mm
0.01 mm
~ 6 ~
of 0.8 mm long describe about 20 cm young fish in I age group. Otolith with start of Third
Primordium at radius 2.1 mm, describe 36 cm TL fishes starting to mature gonads (6).
Fig. 4. Scheme of large marks of major events in the otolith microstructure describing A) LN- larval nucleus (in yellow), otolith of new hatched
1.5 cm larvae with yolc-sack; B) SP – otolith (in green colour) with start of second primordium indicate body tranformation of 7 cm postlarvae,
that change shalow inshore to deep cold waters; C) Larger part of SP construkt 2 times longer increase in otolith growth of 21 cm TL Ps.
georgianus that start gonad developing. On the right: SP on the otolith surface is indicated by incisions. The SEM photo showing growth of
otolith from second primordium.
Mainly for all samples, otolith weight frequency with linear relationship of their peaks to
age, were used to approximate age group for each specimen from its otolith weight (6; 26).
The new age group on the otolith weight axis is starting after larger break between
adjacent individuals ordered by weights otoliths. The wide width of above breaks resulting from
having an annual growth weight of the present generation over the nearest going to hatch
generation. That breaks for older fish between 5 and 6 age groups are diminishing because of
increased variation of otolith weight within age groups that cames from f.e. ratio of males to
females, since little differences in their sizes.
In spite of that variations, the external space prevail between new age group and previous
one started to growth a year earlier, because as otolith is growing during all fish life the year
difference of growth between age group is maintained.
Age composition was determined by taking into consideration mass measurements, using
the key: length-age (27), transforming length frequency of sample to length frequency of catch
with Gulland’s equation (27): NiPij; Ni – no. of fish in i-th length class in mass measurements,
Pij = nij/ni; where ni, nij – no. of fish in i-th length class and in j-th age group.
Lengths, age at maturity.
Icefish were considered to be sexually mature if the gonad was in maturity stage 3 to 5
(28). Lengths at which 50% of fish reach maturity, and spawn for the first time was established
from cumulating frequency of maturity equal and above stage 3, that were approximated by
sigmoid equation using Solver procedure.
Growth of the fish length by the von Bertalanffy equation.
Positions of peaks of otolith weight within each age group (seasonal variability) were
used to establish age in months (part of the year) from hatching to catch for each otolith, when
fitting Bertalanffy body growth equation.
This means age groups of individuals were extended by adding or extracting to age group
part of the year derived from deviation of otolith weight from the mean otolith weight (or
replace independent variables, from age „t” on otolith weight by expression: „b[otolith
SP
~ 7 ~
weight]+a” in the growth equation, see below), f.e.: from August to January is 6 months + 1
year= 1.5 year for average otolith 0.017g in 1th age group. In a result otolith more heavy will
have large value of age data, f.e.: 1.6 year, otolith lighter will have less value of age: 1.4 year.
To age at length data the Bertalanffy equation was fitted modified by replacement of
unknown usually unreal parameter t0 to know real parameter L0 using following formula:
t0=ln(1-L0/L∞)/k, where L0=0.15 cm. The equation was fitted by minimizing the sum of square
differences between empirical data and Bertalanffy model:
Lt - L∞(1-e-K(b[otolith weight]+a - ln(1-L0/L∞)/k))]2
using solver in EXCEL (29;
30). Error was estimated by statistics of the regression between empirical data and model (31).
Length – weight and weight – weight relationships.
To study migrations by compare the size and development of fish body in the regions and
across a time, the pairs of the measurements - the length and the mass of the body were
described in their power relationship and the pairs of the body length and the otolith weight by
the von Bertalanffy and polynomial equations.
Ps. georgianus was divided on age groups by relationship between otolith weight and
body weight, Fig. 15, that arrange individuals into separated size age groups having difference
in a growth of a year.
Tab. 1. Biological materials of Ps. georgianus collected in the South Georgia I. region within 1976-1992.
season Name of vessel
Number of fish
season Name of vessel
Number of fish
Mass
measure
Detailed
Analysis
Age
estimate
Mass
measure
Detailed
Analysis
Age
estimate
1976/77 m/t “Gemini” 1072 350 1982/83 catches break
1977/78 r/v “Bogucki” 150 1983/84 m/t “Taurus” 1928 299 300
m/t “Sirius” 802 265 1984/85 m/t “Taurus” 495 166 161
m/t “Gemini” 1619 201 1985/86 m/t “Carina” 1176 500 500
m/t “Gemini” 4928 1409 147 1986/87 r/v “prof. Siedlecki” 811 1556 323
∑(4 vessels) 7499 1875 147 1987/88 r/v “prof. Siedlecki” 2996 306 712
1978/79 r/v “prof. Siedlecki” 2950 576 335 1988/89 r/v “prof. Siedlecki” 884 343 686
1979/80 lack of data 1989/90 m/t “Hill Cove” 850 508 1322
1980/81 m/t “Libra” 8517 900 1300 1990/91 m/t “Falklands Protector” 2097 350 588
1981/82 m/t “Neptun” 2724 800 300 1991/92 m/t “Falklands Protector” 1878 253 500
1976-92 ∑ 37877 8782 7174
Tab. 2. Biological materials of Ps. georgianus collected on r/v „prof. Siedlecki” in the Antarctic and Subantarctic waters.
REGION
FISH MEASUREMENTS IN 1978/79
ICHTIOLOGICAL OTOLITH
WEIGHTTotal length Detailed
Palmer A. 60 10 20
Deception I. 200 100 200
S. Shetland I. 164 136 272
King Edward I. 100 50 100
Elephant I. 11 11 22
S. Orkney I 343 343 276
Subantarctic S. Georgia I. 177 177 164
All 1055 827 1054
In data analysis and interpretation following additional data were included: length
frequencies of German expedition from fish base (1980) and CCAMLLR data base (2002-
~ 8 ~
2007), possibility presence of Ps. georgianus larvae 4-5 cm TL in Antarctic Zone, after the
memory of the personal comment of dr Wiesław Ślósarczyk (11).
Fish research were lead on r/v “Professor Siedlecki” with same others studies biological
and physical. They provided additional valuable information for ecological study and
comparison, such as distribution and biomass of krill from catch and hydroacoustic data and
fish biomass from hydroacoustic data. Also hydrological properties of water during cruise were
measured giving distribution of temperature and other water parameters. Apart that information
in this work available published temperature data were used to draw map of temperatures
comprised all regions of Ps. georgianus distribution: from Palmer Archipelago to South
Georgia, to show and study similarity and differences.
RESULTS
Differences between fish living in Subantarctic and Antarctica.
Otolith weight frequency at total length (TL). Ps. georgianus of South Georgia has a
heavier otoliths and larger TL than this species from the Antarctic, but increases their masses
are similar. The same distances between age group are well show in linear relationship between
otolith weight and body weight, arranging individuals into separated age groups, Fig. 15. This
linear relationship approximates the birth of in August to fish from South Georgia and in
October for Antarctic zone. Earlier birth of fish from warmer South Georgia may explain their
larger body size and greater weight of the otoliths in compare to fish from the Antarctic born
later.
Fig. 5. Age groups in the otolith mass frequency of Ps. georgianus of Subantarctic South Georgia I. and Antarctic Islands were approximated by
similar equations.
Separating indexes neighbouring peaks in the otolith frequency: I= ( n+1- n)/{(sn+1+sn)/2} are large then 2, and shows significant distances between
~ 9 ~
age groups in the otolith frequency. The distances were large showing that new age group are in a new year. From above equations mean birth
date was derived as in August for South Georgia in account of daily increment of otolith (6) and in October for Antarctic Zone. The lengths of fish
from Antarctic Zone in the appropriate groups of the otolith weight are larger but in the appropriate groups of the age are smaller than the lengths
of fish from the subantarctic South Georgia.
In the Antarctic, in the first age group there are found only males, and in the third age
group are smaller females.
Lengths, age at maturity. Icefish in Subantarctic zone mature for the first time in larger
sizes and older than in Antarctica.
Fig. 6. Length at which 50% of Ps. georgianus spawn for the first time, shown that in Subantarctic zone species mature in larger sizes and older.
1 – ♂♀ Immature
2 – ♂♀ Maturing virgin (
♂ developing) or resting
3 – ♂♀ Developing ( ♂
developed)
4 – ♂♀ Gravid ( ♂Ripe)
5 - ♂♀ Spent
Fig. 7. In Subantarctic Zone more fish were undevelop and smaller. Instead of this fish with developed gonads in Subantarctic, were bigger.
~ 10 ~
Fig. 7. The gonadal stage for
young is similar but older
fish in Subantarctic South
Georgia as there are warmer
have more developed gonad
than in Antarctic Zone.
Fig. 8. In Antarctic Zone undeveloped fish feed similar as in Subantarctic South Georgia, but mature fish in Antarctic Zone feed less and a lot of
them have empty stomach. Since the age group of 1 were not in studied areas in Antarctic Zone, undeveloped fish were larger and older there
than in South Georgia.
Fig. 9. In Antarctic Zone, more empty stomachs were in mature fish as the average degree fill of stomach was dropped. Fish in stage of gonad
development of 2 were most feeding. Both immature and mature fishes feed in slow degree, and a lot of them have empty stomachs.
~ 11 ~
Fig. 10. In Subantarctic South Georgia immature and developing fishes were feeding intense. Almost they have not empty stomachs. Opposite
to that, mature fish were not feeding and have a lot of numbers of empty stomaches.
Fig. 11. In Antarctic Zone fish were feeding more in Palmer Archipelago and less at Eastern islands of Scotia Sea Arc.
At South Orkney Ps. georgianus feed less and a lot of individuals have empty stomaches,
especially mature fish. A large number of empty stomachs is probably due to the lack of food
especially krill, which every year fishery removes in a large number from habitat of this species
(32).
Growth of the fish length by the von Bertalanffy equation. Growth curves of
Bertalanffy for fish from South Georgia and from Antarctic as was to be expected are similar.
Not much larger factor k for growth of fish from South Georgia apparent from adaptation to
climate change which result in the earlier development of the species in warmer South Georgia
giving larger body and a few months older age.
~ 12 ~
Fig. 12. Compare growth curves of Von Bertalanffy for Ps. georgianus, from Antarctic and Subantarctic Zones. Small marks are the estimated
age, and large marks are their averages.
Total Length and body weight to otolith weight relationships.
In Antarctic Zone fish weights growth faster for the given length, but their otolith weights
growth slower with TL than in Subantarctic South Georgia. The total length increments with
increasing otoliths weight in the region of South Georgia were smaller than in Antarctic zone.
Fig. 13. In Antarctic Zone fish weights growth faster for the given length, but otolith weights growth slower than in Subantarctic South Georgia.
In South Georgia body weight at given length was growth slower than today which have b=3.43-3.52(7).
~ 13 ~
Fig. 14. Depending of the body length on the otolith weight of fish
of Antarctic islands shelves can be described by a single formula.
Increases in length with increasing otolith weight of fish at the South
Georgia are slower.
This same is in a body weight that after with length relationship the regional difference
made even deeper. Increases in body length and weight with an increase in the otolith weight of
fish from the different islands in Antarctic zone are similar. Can be described by one single
model characterizing species for the zone of Antarctic.
Tab. 3. Mostly in average the males are stockier than females: bmale>bfemale. But regional in Antarctic Zone e.g. in Elephant I. only
large females are in sex relation. First line: BW=aTL
b
.
Region Males and females Females males
LT, cm b a R2 N LT,cm b A R2 N LT,cm b a R2 N
All: BW=aTLb 17÷56 3,144 0,0058 0,967 779 17÷56 3,0798 0,0074 0,965 393 19÷54 3,208 0,0045 0,969 386
Y=a×ebx 0,0881 16,554 0,969 0,0874 17,04 0,966 0,0889 16,044 0,972
S. Georgia.
17÷55
2,955 0,0111 0,983
176 17÷55
2,894 0,0138 0,982
82 19÷54
3,022 0,0087 0,985
94
Y=a×ebx 0,0872 15,864 0,99 0,0866 16,0092 0,989 0,0877 15,651 0,991
S. Orkney. 34÷56 3,141 0,006 0,74 342 34÷56 3,098 0,0071 0,741 188 37÷54 3.2085 0.004 0,736
154
0,068 44,191 0,73 0,0673 45,207 0,727 0,0689 42,222 0,722
S.Shetl+Palmer. 26÷53 3,475 0,0017 0,955 261 28÷53 3,439 0,0019 0,944 123 26÷52 3,4884 0,0016 0,961 138
0,0884 16,977 0,959 0,0856 19,237 0,946 0,0901 15,757 0,968
S.Shetland
27÷53
3,5796 0,0011 0,954 202 28÷53 3,68 0,0008 0,949 99 27÷52 3,519 0,0014 0,956 103
0,0896 16,201 0,955 0,0902 15,805 0,946 0,0892 16,467 0,959
K.George+Elephan 31÷53 3,6088 0,0010 0,934 67 40÷53 3,798 0,0005 0,894 34 31÷52 3,5505 0,0013 0,955 33
0,0853 19,403 0,941 0,0834 21,026 0,902 0,0864 18,558 0,96
Elephant 40÷52 3,2815 0,0037 0,965 17 40÷51 3,7294 0,0007 0,972 11 32÷52 3,1484 0,0063 0,974 6
0,0787 27,834 0,976 0,0829 23,06 0,975 0,077 30,237 0,979
K. George 31÷53 3,8665 0,0004 0,947 50 40÷53 3,992 0,0002 0,892 23 31÷52 3,859 0,0004 0,975 27
0,0902 15,167 0,946 0,0869 17,367 0,9 0,0928 13,696 0,971
Deception+Palmer 26÷52 3,4502 0,0018 0,957 194 28÷52 3,4074 0,0022 0,951 89 26÷52 3,4607 0,0017 0,959 105
0,0892 16,416 0,961 0,0863 18,727 0,950 0,091 15,156 0,967
Deception 27÷52 3,5844 0,0011 0,958 135 28÷50 3,6904 0,0007 0,963 65 27÷52 3,505 0,0015 0,954 70
0,0917 14,933 0,959 0,093 14.118 0,959 0,0906 15,595 0,958
Palmer A. 26÷41 3,1664 0,005 0,958 59 30÷52 2,932 0,0123 0,956 24 26÷50 3.295 0,0031 0,964 35
0,0834 20,231 0,964 0,0747 29,667 0,96 0,090 15,367 0,979
~ 14 ~
Fig. 15. In Antarctic Zone age group 2 were at Palmer A. and Deception Island. Similar was for age group of 3. Older fish were most numerous at
South Orkney Island. On linear regression of body weight and otolith weight age group are very well separated.
Tab. 4. Equations for deriving TL and BW from otolith weight (OW) in the range measured data.
Region LT, cm OW, [g] Equation R
2
N
Subantarctic
(S.Georgia)
17÷55 0.01241÷0.06837 LOW=57(1-e
-38.09(OW -0.0041)
); ’=5.09
BW= 29962.89OW-485.97 [g]
0.98
0.95
168
Antarctic.
1) + 2)
26÷56 0.02099÷0.07006 LOW=53(1-e
-60.96(OW+0.0082)
);’=5.23;
BW=1269.4ln(OW)+5100.2 [g]
0.99
0.67
394
1) S. Orkney. 34÷56 0.02982÷0.07006 LOW=53(1-e
-80(OW-0.02)
); ’=3.18 0.38 271
2) Shetland
+ Palmer. A,
26÷53 0.02099÷0.06368 LOW=53.1*(1-e
-82.66*(OW-0.012)
); ’=5.37
BW= 48779.05OW-843.49
0,71
0,78
330
King George,
Elephant I.
31÷53 0.02168÷0.05522 TL31÷53=-19737OW
2
+2165.1OW-7.3613
LOW=53.1(1-e-82(OW-0.0115))); ’=5.36
0.83
0.93
110
~ 15 ~
Elephant 40÷52 0.03279÷0.05359 TL40÷52=-41791OW
2
+4147.4OW-50.204
LOW=53.1(1-e-77.7(OW-0.011))); ’=5.34
0.92
0.83
12
King George 31÷53 0.02168÷0.05522 TL31÷53=-19307OW
2
+2120.2OW-6.2805
LOW=53.1(1-e-82(OW-0.0115))); ’=5.36
0.82
0.93
98
Deception+
Palmer A.
26÷52 0.02099÷0.06368 TL26÷52= 449368.57OW
3
-79837.57OW
2
+4726.81OW-42.58
LOW=53.1(1-e-78.71(OW-0.0134)); ’=5.35
0.85
0.73
220
Deception 30÷52 0.02378÷0.06368 TL30÷52=580724.44OW
3
-97145.53OW
2
+5451.66OW-52.17
LOW=53.1(1-e-78.71(OW-0.0134)); ’=5.35
0.82
0.7
200
Palmer A. 26÷41 0.02099÷0.03587 TL26÷41=-45760OW
2
+3549.2OW-27.677
LOW=53.1(1-e-56(OW-0.0085)); ’=5.16
0.86
0.87
20
The occurrence.
In summer 1978/79 the studied species was present in the cold Antarctic waters, uncovered by ice, in lower
density and with completely lack of small fish than in the warmer region of Subantarctic South Georgia I.
In winter the temperature surface of Antarctic waters fall below -5°C, while in S. Georgia is above, Fig. 17.
The summer’s temperatures of S. Georgia unlike the Antarctic were above 2°C, Fig. 1.
At S. Georgia there was 48 kg/h, consisted an average fish with TL = 40 cm in the range of
15-58 cm TL. Also, their food - krill was in high (223-550g/m3
) density of ~40 m swarms;
At South Orkney, there was 23 kg/h, cm TL in the range of 31 – 57 cm;
At Elephant I. 3 kg/h, cm TL in the range of 40 – 52 cm;
At King George I, 14 kg/h, cm TL in the range of 29 – 56 cm; Distance ~560 km
and in the Palmer Archipelago was 5 kg/h, cm TL in the range of 25 – 53 cm.
Fig. 16. Density differences between Antarctic and Subantarctic of Ps. georgianus and its main food – krill occurrence during expedition 1978/79
(33). Explosion mark – the only one place number 98 from sampling trawel stations of r/v “Prof. Siedlecki” at which larvae of Ps. georgianus
were reported during SIBEX (11). The highest krill density and biomass was at S. Georgia =1502 tons. At S. Orkney there was smallest krill density
but in larger swarms, about 120 m length. Krill density at S. Orkney may be despearsed by oscilation of ice border, Fig. 18, or low temperature,
Fig. 1.
At South Georgia during sampling in the summer
there are no any ice cover, instead of Antarctic
Zone, where ice close a large amounts of islands
shelf to sampling, Fig. 18. In addition sea ice have a
great amount of alga, copepods (Oithona spp) on
which krill feed. Up to 13 km to the South from the
edge of ice, there are high densities of the krill
(even 30000 specimens per m3
). As the krill preying
on algae, fish larvae and young fish prey on it as well. Krill by itself in clusters creates shelters and
environment for the development of fish larvae, physically and chemically changes properties of Antarctic
waters to more friendly for that(34; 35). Also sea ice offers a shelter from predators, and from fishing. In well
Fig. 17. Surface temperature in Antarctic Penninsular.
Fish:
: 15-58 cm TL,
South Georgia I.
23 kg/h
S. Orkney I.
31-57, cm TL3 kg/h, 40-52, cm TL
14 kg/h
29-56, cm TL
King George I.
Palmer A.
5 kg/h, 25-53: cm TL
14
23
48
kg
/h
Krillmax550g/m3
Krill, 73 g/m3
Krill, 150 g/m3
Krill,
200
g/m3
Krill, 350 g/m3
Elephant I.
Krill, 90 g/m3
~ 16 ~
conditions for fishing, catch of this species in Antarctic zone was high, over 2000 tonnes, usually in the
Northern and Western parts of Islands, Fig. 18, sheltered and upstream of currents – like at South Georgia I.,
Fig. 16, that usually were(7), and in the middle stratum zone: 150-250 m (7).
Fig. 18. Localization of catch of Ps. georgianus on Island’s shelfs (<500m) in the Scotia Sea during sampling in 1978/1979 with the fish larvaes
found in krill catch on open deep pelagic waters (1000-4350m) near the Northern ice edge during expedition on r/v “prof. Siedlecki” in the
summer 1988/89(36). Arrows – fishing catch of krill that had fish larvaes: Ch. aceratus (40,66); Ch. gunnarii (41); Chionodraco rastrospinosus
(41,66,78); Chaenodraco wilsoni (69,73,74); Dissostichus eleginoides (73); Pleurogramma antarcticum (40,74); Gryodraco antarcticus
(40,73,74,82); Neopagetopsis sp. (55); Neopagetopsis ionach (56,71,73); Trematomus eulepidotus (65); Notolepis coatsi (67,71); Pagetopsis sp.
(69,78,82); Notothenia sp. (69); Notothenia larseni (73;74); Pagetopsis macropterus (73); Electrona carlsbergi (78).
Length, weight and growth.
The length frequency show that smaller range of the lengths of fish in Antarctic Zone mean lack of all
fish from first length group at age. This show that Ps. georgianus is distributed in separated age groups - not
stay together. The age youngest group probably are swimming under ice and feeding on krill and fish larvae.
kg/h
3 kg/h14 kg/h
78
82
74
73 71
40
41
69 67
66 65
5655
~ 17 ~
Fig. 19. Length frequency distribution of Ps. georgianus cached on r/v „Prof. Siedlecki” in 1978/79 showing lack of all small fish 14-
24 cm TL from first age at length groups in Antarctic Zone. Also that group of fish were not present in German data from fishing
off S. Orkney I. in 1980 (Fishbase data (37)). Only 3 specimens this group was cached in Antarctic Zone by Germany from 2002 to
2007 and few by Spain in 1991 (8) in the summers. Larvae of 4.5 cm TL were assumed as probably present, but as an invaluable
there were never accounted by fisheries in that time (personal comments).
The length distributions shown that length groups at age, in the Antarctic Zone are smaller than their
appropriate groups in the South Georgia. That change is expected as at South Georgia is warmer water
allowing taking span earlier and start growth earlier than in Antarctic waters.
The age structures shown the age groups are separated geographically in the vast region of Antarctic
Zone. First age group is not in sampled area, age 2 with 3 are in South Shetlands I., age 4+ cumulate at South
Orkney I, Fig. 20. While in a small one island of Subantarctic Zone, age groups are time separated in patterns
of strong – week cohorts: strong group of age 4 is separated from the strong age group 2 by low numbers of
previous age of 3, Fig. 20. Offspring of the age group of 4 this study dominate too, in summer 1981/82 (38).
Also offspring of numerous age group of 2 dominate in summer 1983/84 as age group of 3. This event is
normal for South Georgia I.
Fig. 20. Age structure showing lack of 1th age group in Antarctic Zone in which species distribute age groups separated geographically: 1 in not
sampled area, age 2 with 3 in Shetlands, age 4+ cumulate in South Orkney I. While in Subantarctic ages are time separated in patterns of strong
– week cohorts: group of age 4 is separated from the 2th by low numbers of previous age of 3.
Strong age groups by their offspring determine the next strong age groups in the future, so the events
of high level of species biomass. The secondary offspring, offspring 1984/85 of above mentioned numerous 2
age groups this study, dominate in two summers 1986/87; 87/88, as an age group 2 and 3. Their offspring of
1986/87 (third level offspring of age group of 2 from 1978/79, Fig. 20) were dominating even in 3 summers
from 1989/90 to 1991/92, as age group 1, 2, 3 (38). This is may be confirmed in the BAS research: fourth level
offspring - offspring 1992/93 seems to dominate in summer 1993/94 as an age group of 4. After two offspring
(6 level offspring) seems to dominate in summer 2099/00 as age group 3. New offspring 2000/01 (7 level) is
numerous in summer 2001/02 as an age group one and seems to dominate in summers 2002/03 and 2003/04
in age group of 2 and 3. Their offspring 2004/05 (eight level from span of 4 aged specimens) is numerous in
summer 2005/06 as an age group of 1 (39). From group of ancestors in age of 4, being in age group 2 in
1978/79 the nine level offspring this year 2012 is in age group 3.
The concentrates of the large fish of Ps. georgianus in the region of South Orkney Island.
In Antarctic Zone the numerous small fish of Ps. georgianus stepped out near continent in the cold
waters in the Palmer Archipelago. Farther to the West from the continent in the direction of South Orkney the
number of small fish got smaller. However the part of large fish increased. They stepped out in the largest
condensations in the warmest region, most farthest from the continent at South Orkney I.
~ 18 ~
Fig. 21. The large fish of Ps. georgianus are in the region of South Orkney I.
Those above regional divide of distribution of Ps. georgianus on different size groups may
have been in relation with biology and migrations of krill to which this species as to its dominant food
(7) have strong dependence (32). In the summer near continent at Palmer Archipelago were Ps.
georgianus was small, there were reported small and juvenile krill (40). Gravid and spawning adult’s
krill during summer active migrate into off shore along the continental slope and in oceanic waters.
Those waters along Antarctic Peninsula are influenced by the West Wind Drift. This current is very
~ 19 ~
strong and in summer flows closer to the Peninsula and even from coastal waters would carry
juvenile and subadult krill to the North – East, resulting in high concentrations this species there –
between South Orkney and South Georgia – the most bigger and larger in South Ocean- where ice
cover is very large too (1). As adult krill actively migrate offshore for spawning as larger and even
small Ps. georgianus as semipelagic species may follow their main food: juvenile Ps. georgianus are
vulnerable to exploitation as bycatch in the krill fishery (9; 41).
Fig. 22. Spatial separation in the maturity stages of krill. Adults occur along the continental slope. Nearer to the coast subadult krill dominate while
the juvenile stages are confined to coastal shelf waters, where young and small of Ps. georgianus were catch. Similar pattern of krill distribution is
proposed for sea ice zone between Elephant Island and South Orkney, from ecological expedition result: small juveniles near ice, larger adults
farther from ice edge.
Both krill and associated fish larvae and juvenes may drift in a safe environment of krill clumps
and clusters in the vicinity of shelter and source of food provided by sea ice, which join Antarctic
Peninsula Islands with South Orkney. Ice cover is physical obstacle for sea currents, slowing their
velocity and directions and creating new ones opposite to the main giving possibility to return. Also
that places have preferable conditions for aggregating of krill (1). Near sea ice edge that is nurseling
and save environment, the small krill and fish larvae were reported (36), Fig. 18. So ice edge may play
similar function as coastal shore and be useful for deep and shallow water species by joining those
environments. In this way sea ice not only join island shores and support to many species the 2 ways
of migration between them (by create altercurrents), but by itself containing green algae enlarge food
source and nurseling home for krill and fish larvae. This may explaining the great increase of all krill
stages during spring after retreat of sea ice conceal them in winter. This may explain unexpected high
aggregations of Ps. georgianus around South Orkney I. Also this may explain possible migrations this
species from S. Orkney to S. Georgia.
Krillmax550g/m3
Krill, 73
g/m3Krill,
200
g/m3
Krill, 350 g/m3
Krill, 90 g/m3
Krill, 150 g/m3
~ 20 ~
0
2
4
6
8
10
12
16 26 36 46 56
N=500
Near the ice edge
trawls: 40, 55, 56, 67, 71.
0
2
4
6
8
10
12
N=300
Far to the ice edge
trawls: 65, 55, 66, 78
0
2
4
6
8
10
12
N=600
Near the ice edge
Bongo :24, 27, 31, 39, 45, 52.
N=600
Far to the ice edge
Bongo: 25, 33, 37, 46, 50, 53
0
2
4
6
8
10
12
TL, mm
%
%
%
%
0
10
20
30
40
50
1 2 3 4 5 6 7
25 38 47 33 40 48
_
TL, mm
0
10
20
30
40
1 2 3 4 5 6 7
25 36 48 33 45 49 46
_
TL, mm
0
10
20
30
40
1 2 3 4 5 6 7
29 40 49 35 42
_
TL, mm
0
10
20
30
40
50
60
1 2 3 4 5 6 7
3 0 4 4 5 0 4 7 4 7
_
TL, mm
100
20
40
60
80
kg/h0
100
200
300
400
500
37
62
Near
iceedge
Far from
ice edge
80
218kg/h
39
47mm
1636
3690No/h
kg/h
8
16
24
32
40
·10No/h
0
2.2ºC
2.6ºC
0
10
20
30
40
50
60
Near
ice edge Far from
ice edge
44ml;32mmTL
53ml;35mmTL
ml/1000m3
Fig. 23. Krill length frequency and
maturity stage composition and
abundance for trawls and bongo
sampling in the ice edge zone between
Elephant I. and South Orkney I
obtained in the ecological expeditions
on board R/V “Professor Siedlecki”,
BIOMAS IV, December 1988.
~ 21 ~
Fig. 24. Large cover of Scotia Sea by sea ice sometime reaching South Georgia may play transition bridge that expand migration possibility: reduce
velocity and power of currents, creating altercurrents giving returning ways, support shelter and food, change water chemically and physically to
more familiar for larvae and young. Ice edge is placed often along water mass confluence and it like islands create perturbations in laminar flow,
that cumulate plankton and krill in places that catch of Ps. georgianus were successful.
Ice cover sometime can reach South Georgia in the North with its large treasure, source of
food for fish: krill in swarms can stretch under sea ice to the about 13 km from the edge. There krill is
preying ice algae on the bottom planted.
Ice cover brings together 2 environments: sheltering shallow coast with deep pelagic water
reduce distance between stages inside the species. For example under sea ice juvenile forms of the
krill are prey for a long time in a way to reaching the adult stage. In this way krill support food for fish
larvae and for young and adults fish too.
Ice cover give conditions to great arise of krill numbers. Now we know that krill is preying algae
from the bottom of ice like the mower, as it has a lot of them there. Layer of algae placed on bottom of
ice is present on very extensive areas. They contain more organic carbon than the whole pole of
water under ice. When the algae rise e.g. in spring, the bottom of pack ice is the big source of energy
– base for biomass escalation.
It was reported that in winter after spawning krill adults leave the oceanic regions and migrate
into neritic areas where larger food supply than in oceanic waters are. Instead adult krill was in
greater number near the islands in coastal shelf waters: in zones protected from winds and currents,
it was also present near the ice edge, which may play here similar sheltering function and large food
supply too.
Movements of adult krill into coastal waters were explained by the impact of East Wind Drift,
greater during winter and carrying krill and associated fish larvae to South and East. This passive drift
may affect on larvae of Ps. georgianus.
~ 22 ~
Fig. 25. Distribution of Ps. georgianus in relationship with krill high abundance in the Scotia Sea. Krill derived from the Weddell Sea Gyre and from
the Bransfield Strait entrained by currents expand in high abundance on large area of Scotia Sea up to South Georgia. Krill congregate in the centers
of local current of meanders and eddies, that usually were placed on Northern sides of Islands. In that places Ps. georgianus appeared numerous
too. At South Georgia the currents have higher velocity than in Antarctic Zone and could faster cumulate the krill, and to high density. It was true
for summer 1978/79. Biomass and densities of krill swarms and were very high in the current eddies at North East South Georgia, where Ps
georgianus was cached in large quantities too. At South Shetland the currents were slower, and eddy was smaller, so krill swarms had the smallest
biomass and swarm lengths. Ps. georgianus were there in lower numbers. At South Orkney krill was in the largest swarms, that cam probably from
low probability for their breaking: current velocity was smaller, that expressed by the low density of krill swarms. Ps. georgianus were often in large
quantities. Vicinity of Weddell Gyre supply krill inflow here. Palmer Archipelago is at beginning and outside of the border of WSC that flow from
South Shetland to South Orkney and South Georgia. At S. Orkney low density of krill is probably from horizontal oscillation of ice edge.
If Ps. georgianus are not migrate even in one way from South Orkney to South Georgia on the
base of feeding on krill and fish located in the sea ice zone, bonding above Islands, the South
Georgia population this species should have larger differences from fish off South Orkney.
Additionally Ps. georgianus was found at South Sandwich Is, probably expanding there from South
Orkney in a results of drifting with currents and krill, and moving with sea ice and with Secondary
Frontal Zone.
Their proposed migration from S. Orkney – to S. Georgia seems to be more probable after
taking into account, that the water temperature at the depth of 200 meter, where this species larger
occurrence has, were similar at a large distance between above mentioned Islands, and so by this
connect them. Ps georgianus during vertical migrations may cross the layers of water with
temperature change of 1-2 degrees.
Krill, 73
g/m3
Krillmax550g/m3
Krill,
200
g/m3Krill, 350 g/m3
Krill, 90 g/m3
Krill, 150 g/m3
~ 23 ~
BransfieldStrait
- 1
- 1
0
1
1
2
3
4
>3
>4
1
1
-2 0-1 1 2 3 4 5
47.8
22.6
98.1
South Orkney I.
14.4
5.1
64 60 56 52 48 44 40 36
54
56
58
60
62
64
54
56
58
60
62
64
64 60 56 52 48 44 40 36
B
A
C
Feb.1979 – the R/V “Prof. Siedlecki” (N = 67)
Nov. 78 – Feb. 79 – the M/T “Sirius” (N = 30)
Ps. georgianus capture, [kg/h]
Potential temperature [°C] at 200 m
A, B, C - transects
Fig. 26. Catches of Ps. georgianus. Potential temperature in the range of 0-1°C at 200 m(15; 16), join underwater Antarctic Scotia Arc Islands with
Subantarctic South Georgia, especially when ice cover were existed too. Transect A in March 1977, B in February 1977, C in April 1977 (33).
Fig. 27. Temperatures at transect B in the region of Palmer Archipelago, February 21-22, 1977(33). The coldest region: Fig. 30.
~ 24 ~
Fig. 28. Temperatures at transect A from the South Orkney Islands to the northeast across the Weddell – Scotia Confluence, March 4-6, 1977(33).
So near S. Orkney there were temperature lower 0°C and near South Georgia greater 2°C, Fig. 30, Fig. 26.
Fig. 29. Temperatures at transect C from North-East of South Georgia in April 14-15, 1977 (33) - in the region of appearance of Ps. georgianus.
Warmer side of Island because upper water have above 2°C, Fig. 30. Temperature at depth of 200 m was lower than 1°C as in South Shetland, Fig.
26.
~ 25 ~
- 1
1
22.6
14.4
5.1
64 60 56 52 48 44 40 36
54
56
58
60
62
64
54
56
58
60
62
64
64 60 56 52 48 44 40 36
B
A
Nov. 78 – Feb. 79 – the M/T “Sirius” (N = 30)
2°
South Orkney I.
1°
0°
0°
-1°
3°
4°
5°
6°7°
-2 0-1 1 2 3 4 5 6 7 8
Potential temperature [°C] at 20 m
Ps. georgianus capture, [kg/h]
Feb.1979 – the R/V “Prof. Siedlecki” (N = 67)
47.8
A, B, C - transects
C
Fig. 30. Catches of Ps. georgianus and the temperatures at its habitat arranging warm North – East islands sides and cold at opposite sides.
Potential temperature of 2°C and above in upper waters, 20 m(15; 16), were only in North-East South Georgia. But temperatures below 2°C to 0°C
propagate with currents from South Shetland in North East directions to the South Georgia. The colder region is Palmer Archipelago with mean
temperature below 0°C. Also those cold temperatures were partially flow from southern Weddell Sea to the South Orkney and Elephant. This may
be explain why larvae of Ps. georgianus were not in upper 50 meters in warm sides of Island as its swim to colder waters. Transect A in March 1977,
B in February 1977, C in April 1977 (33).
~ 26 ~
Distribution of Ps. georgianus on the shelf of S. Georgia I.
In the summers Ps. georgianus was cached from postlarvae to adult, inside the belt of 10 km around
South Georgia in the depth range from 50 to 500 m. Juvenes was in the pelagic and bottom hauls around the
whole Island: about 2 times more often on the side North and East (24 positive recruitments) than on South,
West (12 recruitments positive). The adult stepped out 2 times more often on North Eastern side of island (249
positive recruitments) than on the West-South (117 positive recruitments). The North-East side of the island,
has the more varied bank, possesses numerous gulfs and is sheltered to the acting of the Western Current. In
that a condition meanders and eddies where formed there, that mechanically concentrates plankton and krill
(33).
Fig. 31. Catches of Ps. georgianus on the shelf of S. Georgia I.
In the season 1988/89 Ps. georgianus predominated on the North Eastern shelf S. Georgia. Ch. gunnari
and Ch. aceratus predominated in the remaining part of the shelf.
43° 41° 39° 37° 35°
54°
55°
2 3
4 5 6 93
5796
58
59
60 61
97 65 64 63 62 99
92
105
Shag
Rocks
54°
55°
43° 41° 39° 37° 35°
N. gibberifrons N. gibberifrons
> Ch. aceratus
Ch. gunnarii
> Ch. aceratus
N. marmorata
> Ch. aceratus
54
5655
91
103 104
50 100 500200 1000 1500 2000 3500
depth 500 m
Biomass distribution of Ps. georgianus & notes
on domination main species in a square, and
cases domination Ch. aceratus over Ps. georgianus
Biomass of Ps. georgianus [tons]
Sampling period: February 1–10, 1989.
Fig. 32. Domination of Ps. georgianus in 5 squares over Ch.aceratus that dominate in 7 remain in biomass of S. Georgia in February 1989 yr.
Next season in 1989 / 90, there were more Ps. georgianus when where more of Ch aceratus.
~ 27 ~
4 5 6 93
5796
58
59
60 61
97 65 64 63 62 99
92
105
Shag
Rocks
D. mawsoni
N. marmorata
> Ch. aceratus
N. gibberifrons
>Ch. aceratus
Ch.gunnarii
>Ch. aceratus
Ch.aceratus
>N. gibberifrons
Ch.gunnarii
> Ch. aceratus
N. gibberifrons
>Ch. aceratus
Ch. gunnarii
N. gibberifrons
>Ch. aceratus
54
5655
91
103 104
Ch.gunnarii
>Ch. aceratus
N. gibberifrons
>Ch. aceratus
Ch. gunnarii
> Ch. aceratus
2 3
54°
55°
54°
55°
43° 41° 39° 37° 35°
43° 41° 39° 37° 35°
50 100 500200 1000 1500 2000 2500
Biomass distribution of Ps. georgianus & notes
on domination main species in a square, and
cases domination Ch. aceratus over Ps. georgianus
depth 500 m
Biomass of Ps. georgianus [tons]
Sampling period: January, 1990.
Fig. 33. Domination of Ps. georgianus in 7 squares over Ch.aceratus that dominate in 10 remain in biomass of S. Georgia in January 1990yr.
The similar distribution of Ps. georgianus is well-kept from the previous period of investigations in
season 1991/92. Ps. georgianus predominated on the western colder shelf of S. Georgia and interior north-east
part, while Blackfin icefish hold the superiority of the numerousness on the eastern shelf of the Island.
4 5 6 7 8 9 10 11
12 13
14
15 16
17 18 19 20 21 22
23
24 25
26
27
Shag
Rocks
D. mawsoni
Ch. gunnarii
> Ch. aceratus Ch. gunnarii
> Ch. aceratus
N. gibberifrons
> Ch. aceratus
Ch. gunnarii
Ch. gunnarii
N. gibberifrons
> Ch. aceratus
N. marmorata
> Ch. aceratus
P. guntheri
P. guntheri
> Ch. aceratus D. mawsoni
50 100 500200 1000 1500 2000 2500
depth 500 m
43° 41° 39° 37° 35°
54°
55°
43° 41° 39° 37° 35°
54°
55°
Biomass distribution of Ps. georgianus & notes
on domination main species in a square, and
cases domination Ch. aceratus over Ps. georgianus
Biomass of Ps. georgianus [tons]
Sampling period: January, 1992.
Fig. 34. Domination of Ps. georgianus in 7 squares over Ch.aceratus that dominate in 9 remain in biomass of S. Georgia in January 1992yr.
West of S. Georgia in the region of Shag Rocks both species Ps. georgianus and Ch. aceratus replaces
one to another completely. This is the Patagonian toothfish region - the larger than them consumer of fish. In
summers 1989-1992, in the region Shag Rocks the occurrence of Ps. georgianus and Ch. aceratus accompanied
the lack of one inversely.
Vertical appearance.
Data put together show that Ps. georgianus most numerous was in the layer between 150 - 250 m, but
larger fish were above in warmer water. Below that layer, deeper smaller fish were found. Larger biomass in the
layer between 150 and 250 were reported in overall catch of Ps. georgianus. This result with the similarity this
layer of water in all regions of Ps. georgianus range appearance, suggest that this species have large possibility
~ 28 ~
for horizontal migrations, that in same period. The confirmation of this is dis- and appearing of Ps. georgianus
in the Shag Rocks migrate from South Georgia I.
Fig. 35. Vertical distribution of postlarvae.
Fig. 36. Vertical distribution Ps. georgianus from S. Georgia I in 1988/89 (red) and 1989/90 (green).
Changes of body size.
During period of 14 summers from 1976 to 1992 the length of species was changed from 46 cm to 39
cm, and parallel their age was dropped from age of 4.4 to age of 3.2 of year. This change was operated on larger
individuals in old age groups from age of 4 to reported 6. Unlike the older fish, the bodies of younger ones were
enlarged. Exploitation may have influence on this when fishing older individuals congregate in summer for
spawning, that later causing freeing more space for younger fast growth individuals. But in opposite side human
eliminated their natural predators: f.e. seals and whales and overall press on species may be at similar level.
Another future that may influence on changes of length of fish body is an increase of water temperature that
reported as warm after cold period around South Georgia (16) or in account of global warming from
anthropogenic influences. Also changes in ice cover, changes in geotropic currents may influence directly or
indirectly across their food (f.e. biomass of krill) on changes in biology of Ps. georgianus. Enlarge set of
naturally influencing futures may determine vertical and horizontal migration of Ps. georgianus – described as
semipelagic species, previously, before anthropogenic period reported as benthic species.
Fig. 37. Change of averages of body length for Ps. georgianus during 14 summers off S. Georgia I.
~ 29 ~
Fig. 38. Change of averages of body length at age of Ps. georgianus during 14 summers off S. Georgia I.
~ 30 ~
Discussion.
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.
Fig. 39. Development swimming, moving and flying possibility with statocysts in animal word. 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 – basilar membrane, pb – papilla basilaris.
~ 31 ~
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 because of, f.e.: Doppler Effect is different between forward
and afterward and backward of moving directions and modifies width of otolith increments to
compensate those differences in perception of move and sound. It compensate by arranging
dipoles of otolith substrates on liquid crystallise surface of otolith in order of picks of high and
lower pressure of oscillations.
Fig. 40. Ernst Chladni (1756-1827) settled small
particles in the net and different figures using the
sounds.
Hans Jenny registered that low frequencies
produced the simple circle described rings,
and number of circles placed around one
circle centrically grows up at higher
frequencies. The same tone always
creates the same shape. Jenny was
convinced that biological evolution was a
result of vibrations. Different frequencies
influence genes, cells and various
structures in the body. He also suggested
that through the study of the ear and
larynx we would be able to come to a
deeper understanding of the ultimate
cause of vibrations (42).
Fig. 41. Assembling the particles in the nods of the acoustic wave.
So, because of above influence environment and moving on otolith growth, the
migrations nature of Ps. georgianus is put simply in its otoliths microstructure and on otolith
external morphology and of course in body shape.
Fig. 42. The pattern of organic surface over polycrystal
could be determined by environment and physiology Fig. 43. Before the otolith's polycrystalline is, its surface in liquid crystal phase (without
hydrogen bonds) could be modified on same stages.
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 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
~ 32 ~
spicule forming, which percept 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.
Fig. 44. Spicules in the gaps between sponge cells.
Also in the gaps is mesohyl: mostly collagen
polymerizes into spongin - collagen fibers (43).
Fig. 45. Gaps in real collagen fibres – aragonite crystallizing in the corners.
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, Fig. 47, or with one in the topside of body Fig. 46, to get similar information
from all directions.
The cut across stocyst of the
medusae: l- statolith, z- the sensorial
cell, r- tentacle.
Fig. 46. Statocyst of ctenophore –
comb jellies: up and
down.
Fig. 47. 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,
ż- pendentive.
More swimming are changing body to bilateral symmetry and statolith localization in a
front to the head, or to head part as a very important perception organ for swimming after a food
or escaping from predators. Statolith starts the change of spherical shape already in Molluscs in
cephalopods, Fig. 48, but large changes have fish vertebrate.
~ 33 ~
Fig. 48. Spherical nucleus (N) and not centrically growth of postlarvae zones of squid otolith. In bottom squid otolith of Alliroteuthis antarcticus,
scale 100µm and Galiteuthis glacialis, scale 200µm. The changes of shape and microstructure of squid otoliths were related with the
ontogenesis, with the change of life from the epipelagic larva to adult swimming deeply in meson and bathypelagic
The shape change is under motion regulation working during species specific
ontogenesis, Fig. 48, and as above in phylogenesis. Taking into account larger species
difference to expose Ps. georgianus swimming strategy, and so the possibility for migration in
its environment condition, it could be see that Ps. georgianus unlike scombrus do not have body
for long and fast swimming in torpedo shape to cut and across the currents. Ps. georgianus is a
fish that rather flying with currents then swimming actively. It has body in shape well
manoeuvrable in currents, that ensure more static in lower own driving velocities, but with
greater possibility to use current force for swimming. Its wide and large fins provide additional
static for turning, and in appropriate angles to currents, like sails can drive them in almost every
direction, as fast as current is and with fast-turning.
Otolith shape of Ps. georgianus show this same as body, the order and relationship with
fish motion. They are not narrow, disc like as otolith of scombrus; they are softly rounded to
diminish any turbulence, what is important in use the driving force from environment: such as
water currents. Height of otolith large than length show that important is perception of stability
on sides during vertical migrations and during swimming, among currents, when currents press
on their body sides.
~ 34 ~
Fig. 49. Scombrus japonicus is in the faster’s fish and swimming
with use power of his body.
Fig. 50. Ps. georgianus like sailor swim with use the power of stream on its
fins.
Fig. 51. Postlarvae have fins even larger and have better possibility to
heaven.
Fig. 52. Havening larvae.
Fig. 53. Otolith of S. japonicus is flat long and short show adaption of perception in high velocity of
swimming.
Fig. 54. Otolith of Ps. georgianus is
thick.
The catch data with current map show, that Ps. georgianus was in current eddies that
congregate the krill – its main food. Its swimming strategy is very useful to follow after a krill
driven physically by currents.
On tendency that as scombrus is faster, as its otoliths are very flat and longer then
higher, in account that otolith of Ch. aceratus - Antarctic white blood fish are longer then
higher it could be concluded that the last fish are more pelagic swimmers than Ps. georgianus.
True Ch. aceratus has wider range of settlings and swim where Ps. georgianus not: to surface
water into 0-50 m depth. This is confirmed by body colour. Ch. aceratus has light colour and at
~ 35 ~
surface is invisible for predators. Ps. georgianus is darker and this shows its better possibility to
hide under ice, where water conditions are darker.
Fig. 55. Ps. georgianus has height of otolith OH > length OL.
Fig. 56. Ch. aceratus, has height of otolith OH< length OL – from this relation similar to otolith S. japonicus is that: Ch. aceratus swim further
than Ps. georgianus.
Fig. 57. Ch. aceratus has body color lighter then Ps. georgianus so more invisible at the surface.
~ 36 ~
As otolith percept vibrations from swimming, also depend on environment conditions, it
during ontogenetic development of Ps. georgianus record all large changes of environments, so
its migrations.
First migration is migration from internal fluids of egg outside to water, that undergo
about 1.5 cm larvae, with otolith mark as above described, Fig. 3.
The second migration is migration of about 7 cm postlarvae from shallow to deep waters,
marked in otolith by SP – second primordium. SP double the length of otolith – the resonator,
and from that perception of higher frequencies of oscillations. The larvae and juvenes percept
low frequencies, older higher.
Seasonal catching of Antarctic fish reported agglomerate of Ps. georgianus in shallow
water for spawning. That swimming to shallow water for spawning could be indicated by
appearance of next additional primordia in otolith microstructures. The spawning of Ps.
georgianus is long time period occur in shallow warm water, that usually cause increasing of
growth in this case an additional centre, AP – large marks in otolith microstructure –as
spawning means change environment on longer time. Several organisms during reproduction
and several fish during spawn and during migration to spawn do not feed. A lot of stomachs are
empty among larger, older fish with large numbers of mature Ps. georgianus that directly may
be suppressed by higher temperature of shallow spawning water. Large change in environment
and in physiology can cause large change in otolith growth.
Ps. georgianus every day change environment conditions on a short time in diurnal
migrations after food in which fish cross the layers of water having different temperature and
pressure that attracts short changes in metabolism and activity. That short changes may explain
small marks of daily increment of otolith. Ps. georgianus following after krill percept the
vibrations they emitted. The vibrations are propagating from all water animals with species
specific frequency (vibrations from swimming 1000-100Hz, pressure: ~10Nm2
) and could be
percept from tens and more meters. In aggregation they interfere and carry the information
about that and attract predators adapted to percept it. With change of depth the oscillation
pressure and other characteristic are changing as well (velocity, amplitude), and operates on
labyrinth and otolith surface and modifies it when surface of otolith growth in a state of liquid
crystal. Sounds have important influence on modifying its perception. Oscillations propagate in
water could describe different animals and their different activity: swimming, feeding
aggregation, spend. Also it is depend on seasons. Otolith microstructure of Ps. georgianus
shown a lot of periodic increments, and from this it say species swim and migrate with large
changes of environment.
So Ps. georgianus have large possibility and already have same migrations reported.
Conclusions
Since we do not have direct observations, in account of above work we could not throw
aside the possibility migration of semipelagic fish of Ps. georgianus after krill and under the
ice between shelves of two islands S. Orkney and S. Georgia in the depth 150-250 m having
similar temperatures in all area extended from Palmer Archipelago through South Orkney and
South Sandwich to South Georgia – just exact covering species range limits.
Above is in opposite to opinion that Ps. georgianus off South Georgia and off South
Orkney are isolated populations.
~ 37 ~
It is most likely that the isolation of island populations takes place currently in connection
with global warming, leading to the loss of ice cover and in connection with krill overfishing
leading to the disappearance of their large cluster enabling the migration of this species sailing
in the depths. To resolve this subject more studies needed.
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%20klinicznych_2.pdf, 2012.
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Migration ps.georgianus

  • 1. ~ 1 ~ "Migrations of the Antarctic fish Pseudochaenichthys georgianus NORMAN, 1939 in the Atlantic sector of Antarctic". Gdynia 2012. Ryszard Traczyk ABSTRACT The material obtained between 1986 and 1991 has allowed to study the pattern of geographical and depth distribution of icefish in the area of Scotia Arc Islands and on shelf of South Georgia Island. In the Antarctic colder waters in the Palmer Archipelago there were mostly young, small fishes. They have stomach filled well with krill so swim probably to match with distribution of krill, which flow further to North and East with geostrophic currents. Following after them, with stomach often full of them, they became large, so the numbers of small individuals are disappearing farther from continent. Their gonad are more developed as the temperature further from continent get warmer. Finally at the end of Scotia Arc Islands - at South Orkney, Ps. georgianus are large with gonad ready to spam, and mostly concerning on reproduction, they do not feeding, so they do not have reason to follow after krill driven to the East to deep open Scotia Sea. In vast Antarctic Zone, enlarged by ice cover species are distributed into age groups separated geographically: first age group as it was not in sampled area, have to be assumed to be near or under ice feeding on juvenile krill and fish; age 2 with 3 are in the South Shetlands; age 4+ are accumulating at South Orkney. While on shelf of one, small sub-antarctic island the group ages of Ps. georgianus are separated by time in a patterns of strong – week cohorts: age group of 4 is separated from the 2th by low numbers fish in previous age group 3. This may save them before feeding of own younger fish – common fish behavior among predators. At South Orkney there are no small fishes. After hatching in cold winter waters as coming summer warm them, they swim upstream (as young fish usually do) to colder water of Palmer Archipelago to have more efficiency in food assimilation. This behavior save them before to be driven to open pelagic water and ensure good reproductive success as old fish drifting to warm water to gonad develop and to span. At South Georgia similar pattern exist in the distribution of Ps. georgianus. Large specimens were closer surface near Eastern shores of South Georgia where water is warmer and sheltered before to be driven to Eastern large pelagic sea. In such conditions there were eddies with larger krill aggregations (1). After spawn young postlarvae swim upstream to the West, to the deep and cold waters. Span, feeding, diurnal and another migrations means changes of environments and physiology - events that have appropriate marks in the otolith microstructure. INTRODUCTION. The results of ecological study create the movie which helps humans understand needs of wild life. For Antarctic underwater life, it is difficult and very rare. We need information on ecology. Change in life pattern can be explained by changes of environment. Now we face global warming that has great impact on marine life at South Georgia and further to South. We have already example influent of large change in environment on distribution and biology Ps. georgianus from its cold conditions settlements in Scotia Sea Arc to warmer area South
  • 2. ~ 2 ~ Georgia. Data of all Antarctic expedition shown, that Pseudochaenichthys georgianus appear in cold water on shelves Scotia Arc Island from Palmer Archipelago to South Orkney Island and outside Antarctic border (in Aliasow classification) about half thousand km further on North across open deep Scotia Sea in warmer Subantarctic South Georgia Island. Additionally they have vertical feeding migrations from cold deep water to warmer surface and came back following the krill and fish larvae diurnal migrations (2; 3; 4). Also mature fish migrate inshore to warm surface water to spawn (5). So larvae after hatching have to migrate to offshore to cold deep water across body transformation - process marked in otolith microstructure as the other environment changes, e.g. from inside to outside of egg (6). The biology and distributions of Ps. georgianus from South Georgia area were described in papers by researches from British Antarctic Survey, Cambridge (7), but there is no the same data for an Antarctic waters. Older fish in this area migrate to shallow spawning waters warmer that are at North East side of Island. After month, two later, depend on water temperature larva hatch in July and August. They growth by the von Bertalanffy: Lt=66.11(1-e-0.283(t+0.008) ); L0=0.15 cm; r2 =0.98, (’=3.09), which is as other icefish very high. Postlarvae, about 8 cm long migrate to deep colder water out of shore. They also swim upstream to Western colder parts of South Georgia I, were they growth faster. In first year, fish growth up to average 19 cm TL, in the second year - 33 cm. In the age group of III fish may have 44 cm TL length, have first spawn, and because of that migrate to shallow and warmer water in the East. Next year, 4 year fish have 50 cm TL, and as their maturity increase, more - above 50% fishes in population are ready to span, and then they do not feed. Their length growth very slow – fish a year older reach only 53 cm TL. As the Subarctic S. Georgia is bordered by deep ocean, migration of Ps. georgianus is restricted to shelf this Island, probably because of warm temperature. Ps. georgianus unlike to similar species of Ch. aceratus do not extend their appearance with Antarctic current to East, and also not migrate to warmer surface water to which Ch. aceratus have better adaptation: e.g.: they have lighter coloured in wave pattern skin of body to be invisible by avian and large sea predators. In Antarctic our species of South Orkneys hatch later in November – December and during ontogenesis show similar temperature induced distributions that for fish depend on water temperature means migrations to more familiar waters. Antarctic Ps. georgianus represents growth to similar size and pattern, as in Subantarctic S. Georgia in that time, with Bertalanffy equation: Lt=66.32*(1-e-0.261(t-0.009) ); r2 =0.99, ’=3.06, L0=0.15 cm. In summer in first year fish have 17 cm, in second 31 cm. Age group of III have average length about 41.7 cm, age of IV have 47.7 cm, age of V: 49.9 cm, VI have 51.3 cm TL, age of 0 (postlarvae): 5 cm (after Ślósarczyk, pers. com.). The smaller fish were distributed close to continent in cold water of Archipelago Palmera, were they feeding the most on krill and follow after them to North East up to Orkney Is. In this warmer region they do not feed but they spawn. The extreme unevenness in spatial distribution of the species was there probably due to dense pre-spawning aggregations (8).
  • 3. ~ 3 ~ Ps. georgianus in Antarctic waters feed more than at Subantarctic South Georgia. In the South Georgia fish do not need as large energy and activity as in cold south and thus food as well. Swimming upstream is usually performed by young fish, futures that play in both areas for saving the species - not fast and long swimmers before spreading to unfamiliar for them large open pelagic Scotia Sea. In another side unusual higher temperature at South Georgia because of global warming may suppress feeding activity and in addition food concurrence with Ch.aceratus may be empowered. In area of South Orkney young larvae after the hatch in October oppose geotropic currents carrying out into the open waters of the Scotia Sea by swimming up current to cold waters of the Archipelago Palmera. They have higher growth in the colder water and in that area the larval stages of the krill are concentrating in deep pelagic, which is ate by them together with associated larvae. Pelagic postlarvae of Ps. georgianus are feed on krill (9). It is probably true, as there were similar Western up currents directions in migration pattern at South Georgia. In krill aggregations on East from South Georgia, there were fish larvae, e.g. of Ch. aceratus but there were not larvae of Ps. georgianus. Larvae of Ps. georgianus were on the Western side off the krill aggregations (10). Also in Eastern peripheries South Shetland Islands, in ecological studies of the sea-ice zone near South Orkney in summer 1988/89, there were not larvae Ps. georgianus among larvae such Ch. gunnari, Ch. aceratus found in krill aggregations. Larvae of Ps. georgianus were reported only from Palmer Archipelago, and generally other larvae of Channichthyidae were in greatest density on the West of South Shetland Is (11). Opposing geotropic currents conditioning the reproductive success in Antarctic has the similar course on the shelf of South Georgia. The most hatching larvae were fished in the sheltered from current East-northern side of the Island. They hatch in shallow ~50m cold winter water (12), so adapting to it at the beginning through the early developing in it. Consequently as they growth, the water became warm as summer coming up, what driving them to sink deeper, so must off shore, direct to familiar cold water this means upstream to West, where this species was cached in larger amount. Inshore, in shallow water larvae of Ps. georgianus that hatched there in winters, in summer were not cached, as e.g., Ch. aceratus (13; 14), and were not numerous present on the East with krill as the other larvae, but outside of krill swarms and farther to West (10). Postlarvae to get more effective utilization of the food they migrate into deeper colder waters and upstream westerly to the Western side of Island where flow cold current stimulating larger increases of the body; as 7 cm long postlarvae they have been better prepared to swim in pelagic water (12) with their lighter colour of body (9), and well developed fins. The perpendicular migrations of this species also reflect thermal preferences of young Ps. georgianus to cold waters - in summer being on larger depths and of the older fish, shown maturing depended extreme unevenness in spatial distribution (8) concentrating in shallow and warmer waters for accelerating their maturation and the development to the spawn. Observed decline of the sizes of body Ps. georgianus during many years observations of the region of South Georgia, can result from the warming up of the climate to of whose Ps. georgianus probably shows the slower growth and late maturation.
  • 4. ~ 4 ~ MATERIALS AND METHODS. Collecting sample. Samples of Ps. georgianus were collected during one to three month long cruises in the Antarctic Peninsula and the South Georgia area since 1976 up to 1991 in the almost every summer. Collections were made by the international science teams of Fish Stock Assessment on the research ships r/v "Professor Siedlecki", r/v "Professor Bogucki" and the trawlers m/t "Gemini", "Sirius", "Taurus", "Carina", "Libra", and "Hill Cove". The haul jobs were lead with the bottom trawl P-32 / 36 having the opening horizontal 17.5 m, perpendicular 4.5 m and a codend mesh of 80 mm with the insertion of the mesh side of 20 mm. The trawling was usually with the speed 3.5 knot during generally 0.5 hour of the time of duration of the hall. Additionally in 11-12.02.1989, 10 pelagic net hauls were made on three transects with pelagic net of WP 16/41 x4 with the insertion of mesh side 20 mm. 22.6 14.4 64 60 56 52 48 44 40 36 54 56 58 60 62 64 54 56 58 60 62 64 64 60 56 52 48 44 40 36 Feb.1979 – the R/V “Prof. Siedlecki” (N = 67) Nov. 78 – Feb. 79 – the M/T “Sirius” (N = 30) Ps. georgianus capture, [kg/h] Mean water temperature [°C] at ~20 m in the summers 5.1 47.8 S o u t h S h e t l a n d South Orkney I. South Georgia I. Bransfield Strait S c o t i a S e a W e d d e l l S e a Elephant I. PalmerA. shallow sea bed 0 - 500 m 2° 0° >2° >2° >2° 0°-2° <0° <0° Fig. 1. Caches of Ps. georgianus during surveys on research vessel r/v “Prof. Siedlecki” and on travel m/t “Sirius” on the shelfs of South Shetland and South Georgia Islands in summer 1978/79. The mean water temperatures at about 20 m in the summers were from oceanographic stations from S. Shetland and S. Orkney of period 1927 to 1980 (15), that were extend to South Georgia area by hydrological study of period of 1955- 1975(16). Because Antarctic waters from region North of South Shetland Islands flow to North-East direction, the temperatures of upper waters of South and West sides of South Georgia Island were similar to waters of South Shetland (0-2°C). Instead of that North and East sides of South Georgia are in average warmer, above 2°C. Waters of Palmer Archipelago and South side S. Orkney are colder - below 0°C. Investigations carried out on the shelves of South Georgia and South Shetland Islands in depths 50 - 500 m at random chosen stations. At each station, total catch was estimated and its fish subsamples were chosen. Fish were sorted by species with taxonomic key (17), with otolith key (18) and key for larvae (19). Ps. georgianus like other industrial species were measured following instructions of BIOMASS (20). Total length (cm), total weight (g), sex, maturity stage in five-point scale and stomach contents in five-point scale for each individual of species
  • 5. ~ 5 ~ sample were recorded, with otolith subtraction (21). Density, biomass and length frequency presentations. The mass of the each catch were linked to area and time trawled to estimate fish density. From above data, biomass were estimated using swept area method, for all region, and in account 3 depth stratum (50-150, 150-250, 250-500 m) of Everson’s bottom surfaces of fish statistical squares (22; 23; 24). Catchability was set to 1. The larvae of Ps. georgianus, Ch. aceratus, Ch. gunnari and other species in range of 3 - 7 cm total length were found in the commercially travelled net. They could supply some information about postlarvae and juvenile stages of fish. Age determination. The ages was determined on the base of the results obtained from daily increments count in otoliths (6) and the otolith internal and external with body morphology analysis with taking into account environmental and physiologic influences. Fig. 2. EDTA removing CaCO3 left gaps in matrix of colagenlike fibres. Those gaps are arranged in a pattern of daily increments rings, that were measured. Into ancles of those gaps cristals of aragonite are deposited to the size and form, which they were restricted by dimensions of gaps. Aragonite from otolith, when were outside of gaps cristals in a long needles (picture on the right). Central Primordium CP in 3 dimensions is a 0.064 mm diameter ball which describes no movable larvae in egg. Larger otolith is two sides flattened and describe movable larvae in egg. Otolith elliptic after 46 daily increments on long radius 0.098 mm having hatching mark (increments 2 times wider 2,5·10-3 mm with less colagenlike fibres) describe about 1.5 cm larvae which just get out from the egg, and free swimming, and having large changes in physiologic and in the environment. Those great changes draw marks in otoliths of other icefish. Fig. 3. Hatching mark in otolith from the left: of Ps. georgianus at radius 0.09 mm with 46 daily incriments; of Ch. gunnari with 21 daily incriments; of Ch. aceratus at radius 0.034 mm with 24 day incriments. On the rigcht gaps in otolith matrix protein from Ch. aceratus (25). Flattened otolith having 0.15 mm Second Primordium SP (source of new radial increments, that start to increase otolith length 2 times and perception of shorter frequencies) on longer radius of 0.8 mm usually after 240 daily increments describe about 7 cm postlarvae of age group 0, transforming, moving to deeper water and swimming 2 times faster. They were in bottom catches in January. Otolith having longest radius more longer, of about 1.8 mm, and SP 0.01 mm 0.01 mm 0.01 mm
  • 6. ~ 6 ~ of 0.8 mm long describe about 20 cm young fish in I age group. Otolith with start of Third Primordium at radius 2.1 mm, describe 36 cm TL fishes starting to mature gonads (6). Fig. 4. Scheme of large marks of major events in the otolith microstructure describing A) LN- larval nucleus (in yellow), otolith of new hatched 1.5 cm larvae with yolc-sack; B) SP – otolith (in green colour) with start of second primordium indicate body tranformation of 7 cm postlarvae, that change shalow inshore to deep cold waters; C) Larger part of SP construkt 2 times longer increase in otolith growth of 21 cm TL Ps. georgianus that start gonad developing. On the right: SP on the otolith surface is indicated by incisions. The SEM photo showing growth of otolith from second primordium. Mainly for all samples, otolith weight frequency with linear relationship of their peaks to age, were used to approximate age group for each specimen from its otolith weight (6; 26). The new age group on the otolith weight axis is starting after larger break between adjacent individuals ordered by weights otoliths. The wide width of above breaks resulting from having an annual growth weight of the present generation over the nearest going to hatch generation. That breaks for older fish between 5 and 6 age groups are diminishing because of increased variation of otolith weight within age groups that cames from f.e. ratio of males to females, since little differences in their sizes. In spite of that variations, the external space prevail between new age group and previous one started to growth a year earlier, because as otolith is growing during all fish life the year difference of growth between age group is maintained. Age composition was determined by taking into consideration mass measurements, using the key: length-age (27), transforming length frequency of sample to length frequency of catch with Gulland’s equation (27): NiPij; Ni – no. of fish in i-th length class in mass measurements, Pij = nij/ni; where ni, nij – no. of fish in i-th length class and in j-th age group. Lengths, age at maturity. Icefish were considered to be sexually mature if the gonad was in maturity stage 3 to 5 (28). Lengths at which 50% of fish reach maturity, and spawn for the first time was established from cumulating frequency of maturity equal and above stage 3, that were approximated by sigmoid equation using Solver procedure. Growth of the fish length by the von Bertalanffy equation. Positions of peaks of otolith weight within each age group (seasonal variability) were used to establish age in months (part of the year) from hatching to catch for each otolith, when fitting Bertalanffy body growth equation. This means age groups of individuals were extended by adding or extracting to age group part of the year derived from deviation of otolith weight from the mean otolith weight (or replace independent variables, from age „t” on otolith weight by expression: „b[otolith SP
  • 7. ~ 7 ~ weight]+a” in the growth equation, see below), f.e.: from August to January is 6 months + 1 year= 1.5 year for average otolith 0.017g in 1th age group. In a result otolith more heavy will have large value of age data, f.e.: 1.6 year, otolith lighter will have less value of age: 1.4 year. To age at length data the Bertalanffy equation was fitted modified by replacement of unknown usually unreal parameter t0 to know real parameter L0 using following formula: t0=ln(1-L0/L∞)/k, where L0=0.15 cm. The equation was fitted by minimizing the sum of square differences between empirical data and Bertalanffy model: Lt - L∞(1-e-K(b[otolith weight]+a - ln(1-L0/L∞)/k))]2 using solver in EXCEL (29; 30). Error was estimated by statistics of the regression between empirical data and model (31). Length – weight and weight – weight relationships. To study migrations by compare the size and development of fish body in the regions and across a time, the pairs of the measurements - the length and the mass of the body were described in their power relationship and the pairs of the body length and the otolith weight by the von Bertalanffy and polynomial equations. Ps. georgianus was divided on age groups by relationship between otolith weight and body weight, Fig. 15, that arrange individuals into separated size age groups having difference in a growth of a year. Tab. 1. Biological materials of Ps. georgianus collected in the South Georgia I. region within 1976-1992. season Name of vessel Number of fish season Name of vessel Number of fish Mass measure Detailed Analysis Age estimate Mass measure Detailed Analysis Age estimate 1976/77 m/t “Gemini” 1072 350 1982/83 catches break 1977/78 r/v “Bogucki” 150 1983/84 m/t “Taurus” 1928 299 300 m/t “Sirius” 802 265 1984/85 m/t “Taurus” 495 166 161 m/t “Gemini” 1619 201 1985/86 m/t “Carina” 1176 500 500 m/t “Gemini” 4928 1409 147 1986/87 r/v “prof. Siedlecki” 811 1556 323 ∑(4 vessels) 7499 1875 147 1987/88 r/v “prof. Siedlecki” 2996 306 712 1978/79 r/v “prof. Siedlecki” 2950 576 335 1988/89 r/v “prof. Siedlecki” 884 343 686 1979/80 lack of data 1989/90 m/t “Hill Cove” 850 508 1322 1980/81 m/t “Libra” 8517 900 1300 1990/91 m/t “Falklands Protector” 2097 350 588 1981/82 m/t “Neptun” 2724 800 300 1991/92 m/t “Falklands Protector” 1878 253 500 1976-92 ∑ 37877 8782 7174 Tab. 2. Biological materials of Ps. georgianus collected on r/v „prof. Siedlecki” in the Antarctic and Subantarctic waters. REGION FISH MEASUREMENTS IN 1978/79 ICHTIOLOGICAL OTOLITH WEIGHTTotal length Detailed Palmer A. 60 10 20 Deception I. 200 100 200 S. Shetland I. 164 136 272 King Edward I. 100 50 100 Elephant I. 11 11 22 S. Orkney I 343 343 276 Subantarctic S. Georgia I. 177 177 164 All 1055 827 1054 In data analysis and interpretation following additional data were included: length frequencies of German expedition from fish base (1980) and CCAMLLR data base (2002-
  • 8. ~ 8 ~ 2007), possibility presence of Ps. georgianus larvae 4-5 cm TL in Antarctic Zone, after the memory of the personal comment of dr Wiesław Ślósarczyk (11). Fish research were lead on r/v “Professor Siedlecki” with same others studies biological and physical. They provided additional valuable information for ecological study and comparison, such as distribution and biomass of krill from catch and hydroacoustic data and fish biomass from hydroacoustic data. Also hydrological properties of water during cruise were measured giving distribution of temperature and other water parameters. Apart that information in this work available published temperature data were used to draw map of temperatures comprised all regions of Ps. georgianus distribution: from Palmer Archipelago to South Georgia, to show and study similarity and differences. RESULTS Differences between fish living in Subantarctic and Antarctica. Otolith weight frequency at total length (TL). Ps. georgianus of South Georgia has a heavier otoliths and larger TL than this species from the Antarctic, but increases their masses are similar. The same distances between age group are well show in linear relationship between otolith weight and body weight, arranging individuals into separated age groups, Fig. 15. This linear relationship approximates the birth of in August to fish from South Georgia and in October for Antarctic zone. Earlier birth of fish from warmer South Georgia may explain their larger body size and greater weight of the otoliths in compare to fish from the Antarctic born later. Fig. 5. Age groups in the otolith mass frequency of Ps. georgianus of Subantarctic South Georgia I. and Antarctic Islands were approximated by similar equations. Separating indexes neighbouring peaks in the otolith frequency: I= ( n+1- n)/{(sn+1+sn)/2} are large then 2, and shows significant distances between
  • 9. ~ 9 ~ age groups in the otolith frequency. The distances were large showing that new age group are in a new year. From above equations mean birth date was derived as in August for South Georgia in account of daily increment of otolith (6) and in October for Antarctic Zone. The lengths of fish from Antarctic Zone in the appropriate groups of the otolith weight are larger but in the appropriate groups of the age are smaller than the lengths of fish from the subantarctic South Georgia. In the Antarctic, in the first age group there are found only males, and in the third age group are smaller females. Lengths, age at maturity. Icefish in Subantarctic zone mature for the first time in larger sizes and older than in Antarctica. Fig. 6. Length at which 50% of Ps. georgianus spawn for the first time, shown that in Subantarctic zone species mature in larger sizes and older. 1 – ♂♀ Immature 2 – ♂♀ Maturing virgin ( ♂ developing) or resting 3 – ♂♀ Developing ( ♂ developed) 4 – ♂♀ Gravid ( ♂Ripe) 5 - ♂♀ Spent Fig. 7. In Subantarctic Zone more fish were undevelop and smaller. Instead of this fish with developed gonads in Subantarctic, were bigger.
  • 10. ~ 10 ~ Fig. 7. The gonadal stage for young is similar but older fish in Subantarctic South Georgia as there are warmer have more developed gonad than in Antarctic Zone. Fig. 8. In Antarctic Zone undeveloped fish feed similar as in Subantarctic South Georgia, but mature fish in Antarctic Zone feed less and a lot of them have empty stomach. Since the age group of 1 were not in studied areas in Antarctic Zone, undeveloped fish were larger and older there than in South Georgia. Fig. 9. In Antarctic Zone, more empty stomachs were in mature fish as the average degree fill of stomach was dropped. Fish in stage of gonad development of 2 were most feeding. Both immature and mature fishes feed in slow degree, and a lot of them have empty stomachs.
  • 11. ~ 11 ~ Fig. 10. In Subantarctic South Georgia immature and developing fishes were feeding intense. Almost they have not empty stomachs. Opposite to that, mature fish were not feeding and have a lot of numbers of empty stomaches. Fig. 11. In Antarctic Zone fish were feeding more in Palmer Archipelago and less at Eastern islands of Scotia Sea Arc. At South Orkney Ps. georgianus feed less and a lot of individuals have empty stomaches, especially mature fish. A large number of empty stomachs is probably due to the lack of food especially krill, which every year fishery removes in a large number from habitat of this species (32). Growth of the fish length by the von Bertalanffy equation. Growth curves of Bertalanffy for fish from South Georgia and from Antarctic as was to be expected are similar. Not much larger factor k for growth of fish from South Georgia apparent from adaptation to climate change which result in the earlier development of the species in warmer South Georgia giving larger body and a few months older age.
  • 12. ~ 12 ~ Fig. 12. Compare growth curves of Von Bertalanffy for Ps. georgianus, from Antarctic and Subantarctic Zones. Small marks are the estimated age, and large marks are their averages. Total Length and body weight to otolith weight relationships. In Antarctic Zone fish weights growth faster for the given length, but their otolith weights growth slower with TL than in Subantarctic South Georgia. The total length increments with increasing otoliths weight in the region of South Georgia were smaller than in Antarctic zone. Fig. 13. In Antarctic Zone fish weights growth faster for the given length, but otolith weights growth slower than in Subantarctic South Georgia. In South Georgia body weight at given length was growth slower than today which have b=3.43-3.52(7).
  • 13. ~ 13 ~ Fig. 14. Depending of the body length on the otolith weight of fish of Antarctic islands shelves can be described by a single formula. Increases in length with increasing otolith weight of fish at the South Georgia are slower. This same is in a body weight that after with length relationship the regional difference made even deeper. Increases in body length and weight with an increase in the otolith weight of fish from the different islands in Antarctic zone are similar. Can be described by one single model characterizing species for the zone of Antarctic. Tab. 3. Mostly in average the males are stockier than females: bmale>bfemale. But regional in Antarctic Zone e.g. in Elephant I. only large females are in sex relation. First line: BW=aTL b . Region Males and females Females males LT, cm b a R2 N LT,cm b A R2 N LT,cm b a R2 N All: BW=aTLb 17÷56 3,144 0,0058 0,967 779 17÷56 3,0798 0,0074 0,965 393 19÷54 3,208 0,0045 0,969 386 Y=a×ebx 0,0881 16,554 0,969 0,0874 17,04 0,966 0,0889 16,044 0,972 S. Georgia. 17÷55 2,955 0,0111 0,983 176 17÷55 2,894 0,0138 0,982 82 19÷54 3,022 0,0087 0,985 94 Y=a×ebx 0,0872 15,864 0,99 0,0866 16,0092 0,989 0,0877 15,651 0,991 S. Orkney. 34÷56 3,141 0,006 0,74 342 34÷56 3,098 0,0071 0,741 188 37÷54 3.2085 0.004 0,736 154 0,068 44,191 0,73 0,0673 45,207 0,727 0,0689 42,222 0,722 S.Shetl+Palmer. 26÷53 3,475 0,0017 0,955 261 28÷53 3,439 0,0019 0,944 123 26÷52 3,4884 0,0016 0,961 138 0,0884 16,977 0,959 0,0856 19,237 0,946 0,0901 15,757 0,968 S.Shetland 27÷53 3,5796 0,0011 0,954 202 28÷53 3,68 0,0008 0,949 99 27÷52 3,519 0,0014 0,956 103 0,0896 16,201 0,955 0,0902 15,805 0,946 0,0892 16,467 0,959 K.George+Elephan 31÷53 3,6088 0,0010 0,934 67 40÷53 3,798 0,0005 0,894 34 31÷52 3,5505 0,0013 0,955 33 0,0853 19,403 0,941 0,0834 21,026 0,902 0,0864 18,558 0,96 Elephant 40÷52 3,2815 0,0037 0,965 17 40÷51 3,7294 0,0007 0,972 11 32÷52 3,1484 0,0063 0,974 6 0,0787 27,834 0,976 0,0829 23,06 0,975 0,077 30,237 0,979 K. George 31÷53 3,8665 0,0004 0,947 50 40÷53 3,992 0,0002 0,892 23 31÷52 3,859 0,0004 0,975 27 0,0902 15,167 0,946 0,0869 17,367 0,9 0,0928 13,696 0,971 Deception+Palmer 26÷52 3,4502 0,0018 0,957 194 28÷52 3,4074 0,0022 0,951 89 26÷52 3,4607 0,0017 0,959 105 0,0892 16,416 0,961 0,0863 18,727 0,950 0,091 15,156 0,967 Deception 27÷52 3,5844 0,0011 0,958 135 28÷50 3,6904 0,0007 0,963 65 27÷52 3,505 0,0015 0,954 70 0,0917 14,933 0,959 0,093 14.118 0,959 0,0906 15,595 0,958 Palmer A. 26÷41 3,1664 0,005 0,958 59 30÷52 2,932 0,0123 0,956 24 26÷50 3.295 0,0031 0,964 35 0,0834 20,231 0,964 0,0747 29,667 0,96 0,090 15,367 0,979
  • 14. ~ 14 ~ Fig. 15. In Antarctic Zone age group 2 were at Palmer A. and Deception Island. Similar was for age group of 3. Older fish were most numerous at South Orkney Island. On linear regression of body weight and otolith weight age group are very well separated. Tab. 4. Equations for deriving TL and BW from otolith weight (OW) in the range measured data. Region LT, cm OW, [g] Equation R 2 N Subantarctic (S.Georgia) 17÷55 0.01241÷0.06837 LOW=57(1-e -38.09(OW -0.0041) ); ’=5.09 BW= 29962.89OW-485.97 [g] 0.98 0.95 168 Antarctic. 1) + 2) 26÷56 0.02099÷0.07006 LOW=53(1-e -60.96(OW+0.0082) );’=5.23; BW=1269.4ln(OW)+5100.2 [g] 0.99 0.67 394 1) S. Orkney. 34÷56 0.02982÷0.07006 LOW=53(1-e -80(OW-0.02) ); ’=3.18 0.38 271 2) Shetland + Palmer. A, 26÷53 0.02099÷0.06368 LOW=53.1*(1-e -82.66*(OW-0.012) ); ’=5.37 BW= 48779.05OW-843.49 0,71 0,78 330 King George, Elephant I. 31÷53 0.02168÷0.05522 TL31÷53=-19737OW 2 +2165.1OW-7.3613 LOW=53.1(1-e-82(OW-0.0115))); ’=5.36 0.83 0.93 110
  • 15. ~ 15 ~ Elephant 40÷52 0.03279÷0.05359 TL40÷52=-41791OW 2 +4147.4OW-50.204 LOW=53.1(1-e-77.7(OW-0.011))); ’=5.34 0.92 0.83 12 King George 31÷53 0.02168÷0.05522 TL31÷53=-19307OW 2 +2120.2OW-6.2805 LOW=53.1(1-e-82(OW-0.0115))); ’=5.36 0.82 0.93 98 Deception+ Palmer A. 26÷52 0.02099÷0.06368 TL26÷52= 449368.57OW 3 -79837.57OW 2 +4726.81OW-42.58 LOW=53.1(1-e-78.71(OW-0.0134)); ’=5.35 0.85 0.73 220 Deception 30÷52 0.02378÷0.06368 TL30÷52=580724.44OW 3 -97145.53OW 2 +5451.66OW-52.17 LOW=53.1(1-e-78.71(OW-0.0134)); ’=5.35 0.82 0.7 200 Palmer A. 26÷41 0.02099÷0.03587 TL26÷41=-45760OW 2 +3549.2OW-27.677 LOW=53.1(1-e-56(OW-0.0085)); ’=5.16 0.86 0.87 20 The occurrence. In summer 1978/79 the studied species was present in the cold Antarctic waters, uncovered by ice, in lower density and with completely lack of small fish than in the warmer region of Subantarctic South Georgia I. In winter the temperature surface of Antarctic waters fall below -5°C, while in S. Georgia is above, Fig. 17. The summer’s temperatures of S. Georgia unlike the Antarctic were above 2°C, Fig. 1. At S. Georgia there was 48 kg/h, consisted an average fish with TL = 40 cm in the range of 15-58 cm TL. Also, their food - krill was in high (223-550g/m3 ) density of ~40 m swarms; At South Orkney, there was 23 kg/h, cm TL in the range of 31 – 57 cm; At Elephant I. 3 kg/h, cm TL in the range of 40 – 52 cm; At King George I, 14 kg/h, cm TL in the range of 29 – 56 cm; Distance ~560 km and in the Palmer Archipelago was 5 kg/h, cm TL in the range of 25 – 53 cm. Fig. 16. Density differences between Antarctic and Subantarctic of Ps. georgianus and its main food – krill occurrence during expedition 1978/79 (33). Explosion mark – the only one place number 98 from sampling trawel stations of r/v “Prof. Siedlecki” at which larvae of Ps. georgianus were reported during SIBEX (11). The highest krill density and biomass was at S. Georgia =1502 tons. At S. Orkney there was smallest krill density but in larger swarms, about 120 m length. Krill density at S. Orkney may be despearsed by oscilation of ice border, Fig. 18, or low temperature, Fig. 1. At South Georgia during sampling in the summer there are no any ice cover, instead of Antarctic Zone, where ice close a large amounts of islands shelf to sampling, Fig. 18. In addition sea ice have a great amount of alga, copepods (Oithona spp) on which krill feed. Up to 13 km to the South from the edge of ice, there are high densities of the krill (even 30000 specimens per m3 ). As the krill preying on algae, fish larvae and young fish prey on it as well. Krill by itself in clusters creates shelters and environment for the development of fish larvae, physically and chemically changes properties of Antarctic waters to more friendly for that(34; 35). Also sea ice offers a shelter from predators, and from fishing. In well Fig. 17. Surface temperature in Antarctic Penninsular. Fish: : 15-58 cm TL, South Georgia I. 23 kg/h S. Orkney I. 31-57, cm TL3 kg/h, 40-52, cm TL 14 kg/h 29-56, cm TL King George I. Palmer A. 5 kg/h, 25-53: cm TL 14 23 48 kg /h Krillmax550g/m3 Krill, 73 g/m3 Krill, 150 g/m3 Krill, 200 g/m3 Krill, 350 g/m3 Elephant I. Krill, 90 g/m3
  • 16. ~ 16 ~ conditions for fishing, catch of this species in Antarctic zone was high, over 2000 tonnes, usually in the Northern and Western parts of Islands, Fig. 18, sheltered and upstream of currents – like at South Georgia I., Fig. 16, that usually were(7), and in the middle stratum zone: 150-250 m (7). Fig. 18. Localization of catch of Ps. georgianus on Island’s shelfs (<500m) in the Scotia Sea during sampling in 1978/1979 with the fish larvaes found in krill catch on open deep pelagic waters (1000-4350m) near the Northern ice edge during expedition on r/v “prof. Siedlecki” in the summer 1988/89(36). Arrows – fishing catch of krill that had fish larvaes: Ch. aceratus (40,66); Ch. gunnarii (41); Chionodraco rastrospinosus (41,66,78); Chaenodraco wilsoni (69,73,74); Dissostichus eleginoides (73); Pleurogramma antarcticum (40,74); Gryodraco antarcticus (40,73,74,82); Neopagetopsis sp. (55); Neopagetopsis ionach (56,71,73); Trematomus eulepidotus (65); Notolepis coatsi (67,71); Pagetopsis sp. (69,78,82); Notothenia sp. (69); Notothenia larseni (73;74); Pagetopsis macropterus (73); Electrona carlsbergi (78). Length, weight and growth. The length frequency show that smaller range of the lengths of fish in Antarctic Zone mean lack of all fish from first length group at age. This show that Ps. georgianus is distributed in separated age groups - not stay together. The age youngest group probably are swimming under ice and feeding on krill and fish larvae. kg/h 3 kg/h14 kg/h 78 82 74 73 71 40 41 69 67 66 65 5655
  • 17. ~ 17 ~ Fig. 19. Length frequency distribution of Ps. georgianus cached on r/v „Prof. Siedlecki” in 1978/79 showing lack of all small fish 14- 24 cm TL from first age at length groups in Antarctic Zone. Also that group of fish were not present in German data from fishing off S. Orkney I. in 1980 (Fishbase data (37)). Only 3 specimens this group was cached in Antarctic Zone by Germany from 2002 to 2007 and few by Spain in 1991 (8) in the summers. Larvae of 4.5 cm TL were assumed as probably present, but as an invaluable there were never accounted by fisheries in that time (personal comments). The length distributions shown that length groups at age, in the Antarctic Zone are smaller than their appropriate groups in the South Georgia. That change is expected as at South Georgia is warmer water allowing taking span earlier and start growth earlier than in Antarctic waters. The age structures shown the age groups are separated geographically in the vast region of Antarctic Zone. First age group is not in sampled area, age 2 with 3 are in South Shetlands I., age 4+ cumulate at South Orkney I, Fig. 20. While in a small one island of Subantarctic Zone, age groups are time separated in patterns of strong – week cohorts: strong group of age 4 is separated from the strong age group 2 by low numbers of previous age of 3, Fig. 20. Offspring of the age group of 4 this study dominate too, in summer 1981/82 (38). Also offspring of numerous age group of 2 dominate in summer 1983/84 as age group of 3. This event is normal for South Georgia I. Fig. 20. Age structure showing lack of 1th age group in Antarctic Zone in which species distribute age groups separated geographically: 1 in not sampled area, age 2 with 3 in Shetlands, age 4+ cumulate in South Orkney I. While in Subantarctic ages are time separated in patterns of strong – week cohorts: group of age 4 is separated from the 2th by low numbers of previous age of 3. Strong age groups by their offspring determine the next strong age groups in the future, so the events of high level of species biomass. The secondary offspring, offspring 1984/85 of above mentioned numerous 2 age groups this study, dominate in two summers 1986/87; 87/88, as an age group 2 and 3. Their offspring of 1986/87 (third level offspring of age group of 2 from 1978/79, Fig. 20) were dominating even in 3 summers from 1989/90 to 1991/92, as age group 1, 2, 3 (38). This is may be confirmed in the BAS research: fourth level offspring - offspring 1992/93 seems to dominate in summer 1993/94 as an age group of 4. After two offspring (6 level offspring) seems to dominate in summer 2099/00 as age group 3. New offspring 2000/01 (7 level) is numerous in summer 2001/02 as an age group one and seems to dominate in summers 2002/03 and 2003/04 in age group of 2 and 3. Their offspring 2004/05 (eight level from span of 4 aged specimens) is numerous in summer 2005/06 as an age group of 1 (39). From group of ancestors in age of 4, being in age group 2 in 1978/79 the nine level offspring this year 2012 is in age group 3. The concentrates of the large fish of Ps. georgianus in the region of South Orkney Island. In Antarctic Zone the numerous small fish of Ps. georgianus stepped out near continent in the cold waters in the Palmer Archipelago. Farther to the West from the continent in the direction of South Orkney the number of small fish got smaller. However the part of large fish increased. They stepped out in the largest condensations in the warmest region, most farthest from the continent at South Orkney I.
  • 18. ~ 18 ~ Fig. 21. The large fish of Ps. georgianus are in the region of South Orkney I. Those above regional divide of distribution of Ps. georgianus on different size groups may have been in relation with biology and migrations of krill to which this species as to its dominant food (7) have strong dependence (32). In the summer near continent at Palmer Archipelago were Ps. georgianus was small, there were reported small and juvenile krill (40). Gravid and spawning adult’s krill during summer active migrate into off shore along the continental slope and in oceanic waters. Those waters along Antarctic Peninsula are influenced by the West Wind Drift. This current is very
  • 19. ~ 19 ~ strong and in summer flows closer to the Peninsula and even from coastal waters would carry juvenile and subadult krill to the North – East, resulting in high concentrations this species there – between South Orkney and South Georgia – the most bigger and larger in South Ocean- where ice cover is very large too (1). As adult krill actively migrate offshore for spawning as larger and even small Ps. georgianus as semipelagic species may follow their main food: juvenile Ps. georgianus are vulnerable to exploitation as bycatch in the krill fishery (9; 41). Fig. 22. Spatial separation in the maturity stages of krill. Adults occur along the continental slope. Nearer to the coast subadult krill dominate while the juvenile stages are confined to coastal shelf waters, where young and small of Ps. georgianus were catch. Similar pattern of krill distribution is proposed for sea ice zone between Elephant Island and South Orkney, from ecological expedition result: small juveniles near ice, larger adults farther from ice edge. Both krill and associated fish larvae and juvenes may drift in a safe environment of krill clumps and clusters in the vicinity of shelter and source of food provided by sea ice, which join Antarctic Peninsula Islands with South Orkney. Ice cover is physical obstacle for sea currents, slowing their velocity and directions and creating new ones opposite to the main giving possibility to return. Also that places have preferable conditions for aggregating of krill (1). Near sea ice edge that is nurseling and save environment, the small krill and fish larvae were reported (36), Fig. 18. So ice edge may play similar function as coastal shore and be useful for deep and shallow water species by joining those environments. In this way sea ice not only join island shores and support to many species the 2 ways of migration between them (by create altercurrents), but by itself containing green algae enlarge food source and nurseling home for krill and fish larvae. This may explaining the great increase of all krill stages during spring after retreat of sea ice conceal them in winter. This may explain unexpected high aggregations of Ps. georgianus around South Orkney I. Also this may explain possible migrations this species from S. Orkney to S. Georgia. Krillmax550g/m3 Krill, 73 g/m3Krill, 200 g/m3 Krill, 350 g/m3 Krill, 90 g/m3 Krill, 150 g/m3
  • 20. ~ 20 ~ 0 2 4 6 8 10 12 16 26 36 46 56 N=500 Near the ice edge trawls: 40, 55, 56, 67, 71. 0 2 4 6 8 10 12 N=300 Far to the ice edge trawls: 65, 55, 66, 78 0 2 4 6 8 10 12 N=600 Near the ice edge Bongo :24, 27, 31, 39, 45, 52. N=600 Far to the ice edge Bongo: 25, 33, 37, 46, 50, 53 0 2 4 6 8 10 12 TL, mm % % % % 0 10 20 30 40 50 1 2 3 4 5 6 7 25 38 47 33 40 48 _ TL, mm 0 10 20 30 40 1 2 3 4 5 6 7 25 36 48 33 45 49 46 _ TL, mm 0 10 20 30 40 1 2 3 4 5 6 7 29 40 49 35 42 _ TL, mm 0 10 20 30 40 50 60 1 2 3 4 5 6 7 3 0 4 4 5 0 4 7 4 7 _ TL, mm 100 20 40 60 80 kg/h0 100 200 300 400 500 37 62 Near iceedge Far from ice edge 80 218kg/h 39 47mm 1636 3690No/h kg/h 8 16 24 32 40 ·10No/h 0 2.2ºC 2.6ºC 0 10 20 30 40 50 60 Near ice edge Far from ice edge 44ml;32mmTL 53ml;35mmTL ml/1000m3 Fig. 23. Krill length frequency and maturity stage composition and abundance for trawls and bongo sampling in the ice edge zone between Elephant I. and South Orkney I obtained in the ecological expeditions on board R/V “Professor Siedlecki”, BIOMAS IV, December 1988.
  • 21. ~ 21 ~ Fig. 24. Large cover of Scotia Sea by sea ice sometime reaching South Georgia may play transition bridge that expand migration possibility: reduce velocity and power of currents, creating altercurrents giving returning ways, support shelter and food, change water chemically and physically to more familiar for larvae and young. Ice edge is placed often along water mass confluence and it like islands create perturbations in laminar flow, that cumulate plankton and krill in places that catch of Ps. georgianus were successful. Ice cover sometime can reach South Georgia in the North with its large treasure, source of food for fish: krill in swarms can stretch under sea ice to the about 13 km from the edge. There krill is preying ice algae on the bottom planted. Ice cover brings together 2 environments: sheltering shallow coast with deep pelagic water reduce distance between stages inside the species. For example under sea ice juvenile forms of the krill are prey for a long time in a way to reaching the adult stage. In this way krill support food for fish larvae and for young and adults fish too. Ice cover give conditions to great arise of krill numbers. Now we know that krill is preying algae from the bottom of ice like the mower, as it has a lot of them there. Layer of algae placed on bottom of ice is present on very extensive areas. They contain more organic carbon than the whole pole of water under ice. When the algae rise e.g. in spring, the bottom of pack ice is the big source of energy – base for biomass escalation. It was reported that in winter after spawning krill adults leave the oceanic regions and migrate into neritic areas where larger food supply than in oceanic waters are. Instead adult krill was in greater number near the islands in coastal shelf waters: in zones protected from winds and currents, it was also present near the ice edge, which may play here similar sheltering function and large food supply too. Movements of adult krill into coastal waters were explained by the impact of East Wind Drift, greater during winter and carrying krill and associated fish larvae to South and East. This passive drift may affect on larvae of Ps. georgianus.
  • 22. ~ 22 ~ Fig. 25. Distribution of Ps. georgianus in relationship with krill high abundance in the Scotia Sea. Krill derived from the Weddell Sea Gyre and from the Bransfield Strait entrained by currents expand in high abundance on large area of Scotia Sea up to South Georgia. Krill congregate in the centers of local current of meanders and eddies, that usually were placed on Northern sides of Islands. In that places Ps. georgianus appeared numerous too. At South Georgia the currents have higher velocity than in Antarctic Zone and could faster cumulate the krill, and to high density. It was true for summer 1978/79. Biomass and densities of krill swarms and were very high in the current eddies at North East South Georgia, where Ps georgianus was cached in large quantities too. At South Shetland the currents were slower, and eddy was smaller, so krill swarms had the smallest biomass and swarm lengths. Ps. georgianus were there in lower numbers. At South Orkney krill was in the largest swarms, that cam probably from low probability for their breaking: current velocity was smaller, that expressed by the low density of krill swarms. Ps. georgianus were often in large quantities. Vicinity of Weddell Gyre supply krill inflow here. Palmer Archipelago is at beginning and outside of the border of WSC that flow from South Shetland to South Orkney and South Georgia. At S. Orkney low density of krill is probably from horizontal oscillation of ice edge. If Ps. georgianus are not migrate even in one way from South Orkney to South Georgia on the base of feeding on krill and fish located in the sea ice zone, bonding above Islands, the South Georgia population this species should have larger differences from fish off South Orkney. Additionally Ps. georgianus was found at South Sandwich Is, probably expanding there from South Orkney in a results of drifting with currents and krill, and moving with sea ice and with Secondary Frontal Zone. Their proposed migration from S. Orkney – to S. Georgia seems to be more probable after taking into account, that the water temperature at the depth of 200 meter, where this species larger occurrence has, were similar at a large distance between above mentioned Islands, and so by this connect them. Ps georgianus during vertical migrations may cross the layers of water with temperature change of 1-2 degrees. Krill, 73 g/m3 Krillmax550g/m3 Krill, 200 g/m3Krill, 350 g/m3 Krill, 90 g/m3 Krill, 150 g/m3
  • 23. ~ 23 ~ BransfieldStrait - 1 - 1 0 1 1 2 3 4 >3 >4 1 1 -2 0-1 1 2 3 4 5 47.8 22.6 98.1 South Orkney I. 14.4 5.1 64 60 56 52 48 44 40 36 54 56 58 60 62 64 54 56 58 60 62 64 64 60 56 52 48 44 40 36 B A C Feb.1979 – the R/V “Prof. Siedlecki” (N = 67) Nov. 78 – Feb. 79 – the M/T “Sirius” (N = 30) Ps. georgianus capture, [kg/h] Potential temperature [°C] at 200 m A, B, C - transects Fig. 26. Catches of Ps. georgianus. Potential temperature in the range of 0-1°C at 200 m(15; 16), join underwater Antarctic Scotia Arc Islands with Subantarctic South Georgia, especially when ice cover were existed too. Transect A in March 1977, B in February 1977, C in April 1977 (33). Fig. 27. Temperatures at transect B in the region of Palmer Archipelago, February 21-22, 1977(33). The coldest region: Fig. 30.
  • 24. ~ 24 ~ Fig. 28. Temperatures at transect A from the South Orkney Islands to the northeast across the Weddell – Scotia Confluence, March 4-6, 1977(33). So near S. Orkney there were temperature lower 0°C and near South Georgia greater 2°C, Fig. 30, Fig. 26. Fig. 29. Temperatures at transect C from North-East of South Georgia in April 14-15, 1977 (33) - in the region of appearance of Ps. georgianus. Warmer side of Island because upper water have above 2°C, Fig. 30. Temperature at depth of 200 m was lower than 1°C as in South Shetland, Fig. 26.
  • 25. ~ 25 ~ - 1 1 22.6 14.4 5.1 64 60 56 52 48 44 40 36 54 56 58 60 62 64 54 56 58 60 62 64 64 60 56 52 48 44 40 36 B A Nov. 78 – Feb. 79 – the M/T “Sirius” (N = 30) 2° South Orkney I. 1° 0° 0° -1° 3° 4° 5° 6°7° -2 0-1 1 2 3 4 5 6 7 8 Potential temperature [°C] at 20 m Ps. georgianus capture, [kg/h] Feb.1979 – the R/V “Prof. Siedlecki” (N = 67) 47.8 A, B, C - transects C Fig. 30. Catches of Ps. georgianus and the temperatures at its habitat arranging warm North – East islands sides and cold at opposite sides. Potential temperature of 2°C and above in upper waters, 20 m(15; 16), were only in North-East South Georgia. But temperatures below 2°C to 0°C propagate with currents from South Shetland in North East directions to the South Georgia. The colder region is Palmer Archipelago with mean temperature below 0°C. Also those cold temperatures were partially flow from southern Weddell Sea to the South Orkney and Elephant. This may be explain why larvae of Ps. georgianus were not in upper 50 meters in warm sides of Island as its swim to colder waters. Transect A in March 1977, B in February 1977, C in April 1977 (33).
  • 26. ~ 26 ~ Distribution of Ps. georgianus on the shelf of S. Georgia I. In the summers Ps. georgianus was cached from postlarvae to adult, inside the belt of 10 km around South Georgia in the depth range from 50 to 500 m. Juvenes was in the pelagic and bottom hauls around the whole Island: about 2 times more often on the side North and East (24 positive recruitments) than on South, West (12 recruitments positive). The adult stepped out 2 times more often on North Eastern side of island (249 positive recruitments) than on the West-South (117 positive recruitments). The North-East side of the island, has the more varied bank, possesses numerous gulfs and is sheltered to the acting of the Western Current. In that a condition meanders and eddies where formed there, that mechanically concentrates plankton and krill (33). Fig. 31. Catches of Ps. georgianus on the shelf of S. Georgia I. In the season 1988/89 Ps. georgianus predominated on the North Eastern shelf S. Georgia. Ch. gunnari and Ch. aceratus predominated in the remaining part of the shelf. 43° 41° 39° 37° 35° 54° 55° 2 3 4 5 6 93 5796 58 59 60 61 97 65 64 63 62 99 92 105 Shag Rocks 54° 55° 43° 41° 39° 37° 35° N. gibberifrons N. gibberifrons > Ch. aceratus Ch. gunnarii > Ch. aceratus N. marmorata > Ch. aceratus 54 5655 91 103 104 50 100 500200 1000 1500 2000 3500 depth 500 m Biomass distribution of Ps. georgianus & notes on domination main species in a square, and cases domination Ch. aceratus over Ps. georgianus Biomass of Ps. georgianus [tons] Sampling period: February 1–10, 1989. Fig. 32. Domination of Ps. georgianus in 5 squares over Ch.aceratus that dominate in 7 remain in biomass of S. Georgia in February 1989 yr. Next season in 1989 / 90, there were more Ps. georgianus when where more of Ch aceratus.
  • 27. ~ 27 ~ 4 5 6 93 5796 58 59 60 61 97 65 64 63 62 99 92 105 Shag Rocks D. mawsoni N. marmorata > Ch. aceratus N. gibberifrons >Ch. aceratus Ch.gunnarii >Ch. aceratus Ch.aceratus >N. gibberifrons Ch.gunnarii > Ch. aceratus N. gibberifrons >Ch. aceratus Ch. gunnarii N. gibberifrons >Ch. aceratus 54 5655 91 103 104 Ch.gunnarii >Ch. aceratus N. gibberifrons >Ch. aceratus Ch. gunnarii > Ch. aceratus 2 3 54° 55° 54° 55° 43° 41° 39° 37° 35° 43° 41° 39° 37° 35° 50 100 500200 1000 1500 2000 2500 Biomass distribution of Ps. georgianus & notes on domination main species in a square, and cases domination Ch. aceratus over Ps. georgianus depth 500 m Biomass of Ps. georgianus [tons] Sampling period: January, 1990. Fig. 33. Domination of Ps. georgianus in 7 squares over Ch.aceratus that dominate in 10 remain in biomass of S. Georgia in January 1990yr. The similar distribution of Ps. georgianus is well-kept from the previous period of investigations in season 1991/92. Ps. georgianus predominated on the western colder shelf of S. Georgia and interior north-east part, while Blackfin icefish hold the superiority of the numerousness on the eastern shelf of the Island. 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 Shag Rocks D. mawsoni Ch. gunnarii > Ch. aceratus Ch. gunnarii > Ch. aceratus N. gibberifrons > Ch. aceratus Ch. gunnarii Ch. gunnarii N. gibberifrons > Ch. aceratus N. marmorata > Ch. aceratus P. guntheri P. guntheri > Ch. aceratus D. mawsoni 50 100 500200 1000 1500 2000 2500 depth 500 m 43° 41° 39° 37° 35° 54° 55° 43° 41° 39° 37° 35° 54° 55° Biomass distribution of Ps. georgianus & notes on domination main species in a square, and cases domination Ch. aceratus over Ps. georgianus Biomass of Ps. georgianus [tons] Sampling period: January, 1992. Fig. 34. Domination of Ps. georgianus in 7 squares over Ch.aceratus that dominate in 9 remain in biomass of S. Georgia in January 1992yr. West of S. Georgia in the region of Shag Rocks both species Ps. georgianus and Ch. aceratus replaces one to another completely. This is the Patagonian toothfish region - the larger than them consumer of fish. In summers 1989-1992, in the region Shag Rocks the occurrence of Ps. georgianus and Ch. aceratus accompanied the lack of one inversely. Vertical appearance. Data put together show that Ps. georgianus most numerous was in the layer between 150 - 250 m, but larger fish were above in warmer water. Below that layer, deeper smaller fish were found. Larger biomass in the layer between 150 and 250 were reported in overall catch of Ps. georgianus. This result with the similarity this layer of water in all regions of Ps. georgianus range appearance, suggest that this species have large possibility
  • 28. ~ 28 ~ for horizontal migrations, that in same period. The confirmation of this is dis- and appearing of Ps. georgianus in the Shag Rocks migrate from South Georgia I. Fig. 35. Vertical distribution of postlarvae. Fig. 36. Vertical distribution Ps. georgianus from S. Georgia I in 1988/89 (red) and 1989/90 (green). Changes of body size. During period of 14 summers from 1976 to 1992 the length of species was changed from 46 cm to 39 cm, and parallel their age was dropped from age of 4.4 to age of 3.2 of year. This change was operated on larger individuals in old age groups from age of 4 to reported 6. Unlike the older fish, the bodies of younger ones were enlarged. Exploitation may have influence on this when fishing older individuals congregate in summer for spawning, that later causing freeing more space for younger fast growth individuals. But in opposite side human eliminated their natural predators: f.e. seals and whales and overall press on species may be at similar level. Another future that may influence on changes of length of fish body is an increase of water temperature that reported as warm after cold period around South Georgia (16) or in account of global warming from anthropogenic influences. Also changes in ice cover, changes in geotropic currents may influence directly or indirectly across their food (f.e. biomass of krill) on changes in biology of Ps. georgianus. Enlarge set of naturally influencing futures may determine vertical and horizontal migration of Ps. georgianus – described as semipelagic species, previously, before anthropogenic period reported as benthic species. Fig. 37. Change of averages of body length for Ps. georgianus during 14 summers off S. Georgia I.
  • 29. ~ 29 ~ Fig. 38. Change of averages of body length at age of Ps. georgianus during 14 summers off S. Georgia I.
  • 30. ~ 30 ~ Discussion. 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. Fig. 39. Development swimming, moving and flying possibility with statocysts in animal word. 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 – basilar membrane, pb – papilla basilaris.
  • 31. ~ 31 ~ 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 because of, f.e.: Doppler Effect is different between forward and afterward and backward of moving directions and modifies width of otolith increments to compensate those differences in perception of move and sound. It compensate by arranging dipoles of otolith substrates on liquid crystallise surface of otolith in order of picks of high and lower pressure of oscillations. Fig. 40. Ernst Chladni (1756-1827) settled small particles in the net and different figures using the sounds. Hans Jenny registered that low frequencies produced the simple circle described rings, and number of circles placed around one circle centrically grows up at higher frequencies. The same tone always creates the same shape. Jenny was convinced that biological evolution was a result of vibrations. Different frequencies influence genes, cells and various structures in the body. He also suggested that through the study of the ear and larynx we would be able to come to a deeper understanding of the ultimate cause of vibrations (42). Fig. 41. Assembling the particles in the nods of the acoustic wave. So, because of above influence environment and moving on otolith growth, the migrations nature of Ps. georgianus is put simply in its otoliths microstructure and on otolith external morphology and of course in body shape. Fig. 42. The pattern of organic surface over polycrystal could be determined by environment and physiology Fig. 43. Before the otolith's polycrystalline is, its surface in liquid crystal phase (without hydrogen bonds) could be modified on same stages. 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 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
  • 32. ~ 32 ~ spicule forming, which percept 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. Fig. 44. Spicules in the gaps between sponge cells. Also in the gaps is mesohyl: mostly collagen polymerizes into spongin - collagen fibers (43). Fig. 45. Gaps in real collagen fibres – aragonite crystallizing in the corners. 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, Fig. 47, or with one in the topside of body Fig. 46, to get similar information from all directions. The cut across stocyst of the medusae: l- statolith, z- the sensorial cell, r- tentacle. Fig. 46. Statocyst of ctenophore – comb jellies: up and down. Fig. 47. 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, ż- pendentive. More swimming are changing body to bilateral symmetry and statolith localization in a front to the head, or to head part as a very important perception organ for swimming after a food or escaping from predators. Statolith starts the change of spherical shape already in Molluscs in cephalopods, Fig. 48, but large changes have fish vertebrate.
  • 33. ~ 33 ~ Fig. 48. Spherical nucleus (N) and not centrically growth of postlarvae zones of squid otolith. In bottom squid otolith of Alliroteuthis antarcticus, scale 100µm and Galiteuthis glacialis, scale 200µm. The changes of shape and microstructure of squid otoliths were related with the ontogenesis, with the change of life from the epipelagic larva to adult swimming deeply in meson and bathypelagic The shape change is under motion regulation working during species specific ontogenesis, Fig. 48, and as above in phylogenesis. Taking into account larger species difference to expose Ps. georgianus swimming strategy, and so the possibility for migration in its environment condition, it could be see that Ps. georgianus unlike scombrus do not have body for long and fast swimming in torpedo shape to cut and across the currents. Ps. georgianus is a fish that rather flying with currents then swimming actively. It has body in shape well manoeuvrable in currents, that ensure more static in lower own driving velocities, but with greater possibility to use current force for swimming. Its wide and large fins provide additional static for turning, and in appropriate angles to currents, like sails can drive them in almost every direction, as fast as current is and with fast-turning. Otolith shape of Ps. georgianus show this same as body, the order and relationship with fish motion. They are not narrow, disc like as otolith of scombrus; they are softly rounded to diminish any turbulence, what is important in use the driving force from environment: such as water currents. Height of otolith large than length show that important is perception of stability on sides during vertical migrations and during swimming, among currents, when currents press on their body sides.
  • 34. ~ 34 ~ Fig. 49. Scombrus japonicus is in the faster’s fish and swimming with use power of his body. Fig. 50. Ps. georgianus like sailor swim with use the power of stream on its fins. Fig. 51. Postlarvae have fins even larger and have better possibility to heaven. Fig. 52. Havening larvae. Fig. 53. Otolith of S. japonicus is flat long and short show adaption of perception in high velocity of swimming. Fig. 54. Otolith of Ps. georgianus is thick. The catch data with current map show, that Ps. georgianus was in current eddies that congregate the krill – its main food. Its swimming strategy is very useful to follow after a krill driven physically by currents. On tendency that as scombrus is faster, as its otoliths are very flat and longer then higher, in account that otolith of Ch. aceratus - Antarctic white blood fish are longer then higher it could be concluded that the last fish are more pelagic swimmers than Ps. georgianus. True Ch. aceratus has wider range of settlings and swim where Ps. georgianus not: to surface water into 0-50 m depth. This is confirmed by body colour. Ch. aceratus has light colour and at
  • 35. ~ 35 ~ surface is invisible for predators. Ps. georgianus is darker and this shows its better possibility to hide under ice, where water conditions are darker. Fig. 55. Ps. georgianus has height of otolith OH > length OL. Fig. 56. Ch. aceratus, has height of otolith OH< length OL – from this relation similar to otolith S. japonicus is that: Ch. aceratus swim further than Ps. georgianus. Fig. 57. Ch. aceratus has body color lighter then Ps. georgianus so more invisible at the surface.
  • 36. ~ 36 ~ As otolith percept vibrations from swimming, also depend on environment conditions, it during ontogenetic development of Ps. georgianus record all large changes of environments, so its migrations. First migration is migration from internal fluids of egg outside to water, that undergo about 1.5 cm larvae, with otolith mark as above described, Fig. 3. The second migration is migration of about 7 cm postlarvae from shallow to deep waters, marked in otolith by SP – second primordium. SP double the length of otolith – the resonator, and from that perception of higher frequencies of oscillations. The larvae and juvenes percept low frequencies, older higher. Seasonal catching of Antarctic fish reported agglomerate of Ps. georgianus in shallow water for spawning. That swimming to shallow water for spawning could be indicated by appearance of next additional primordia in otolith microstructures. The spawning of Ps. georgianus is long time period occur in shallow warm water, that usually cause increasing of growth in this case an additional centre, AP – large marks in otolith microstructure –as spawning means change environment on longer time. Several organisms during reproduction and several fish during spawn and during migration to spawn do not feed. A lot of stomachs are empty among larger, older fish with large numbers of mature Ps. georgianus that directly may be suppressed by higher temperature of shallow spawning water. Large change in environment and in physiology can cause large change in otolith growth. Ps. georgianus every day change environment conditions on a short time in diurnal migrations after food in which fish cross the layers of water having different temperature and pressure that attracts short changes in metabolism and activity. That short changes may explain small marks of daily increment of otolith. Ps. georgianus following after krill percept the vibrations they emitted. The vibrations are propagating from all water animals with species specific frequency (vibrations from swimming 1000-100Hz, pressure: ~10Nm2 ) and could be percept from tens and more meters. In aggregation they interfere and carry the information about that and attract predators adapted to percept it. With change of depth the oscillation pressure and other characteristic are changing as well (velocity, amplitude), and operates on labyrinth and otolith surface and modifies it when surface of otolith growth in a state of liquid crystal. Sounds have important influence on modifying its perception. Oscillations propagate in water could describe different animals and their different activity: swimming, feeding aggregation, spend. Also it is depend on seasons. Otolith microstructure of Ps. georgianus shown a lot of periodic increments, and from this it say species swim and migrate with large changes of environment. So Ps. georgianus have large possibility and already have same migrations reported. Conclusions Since we do not have direct observations, in account of above work we could not throw aside the possibility migration of semipelagic fish of Ps. georgianus after krill and under the ice between shelves of two islands S. Orkney and S. Georgia in the depth 150-250 m having similar temperatures in all area extended from Palmer Archipelago through South Orkney and South Sandwich to South Georgia – just exact covering species range limits. Above is in opposite to opinion that Ps. georgianus off South Georgia and off South Orkney are isolated populations.
  • 37. ~ 37 ~ It is most likely that the isolation of island populations takes place currently in connection with global warming, leading to the loss of ice cover and in connection with krill overfishing leading to the disappearance of their large cluster enabling the migration of this species sailing in the depths. To resolve this subject more studies needed. Bibliography 1. Sahrhage, D. Antarctic Ocean and Resources Variability. Berlin Heidelberg : Springer-Verlag, 1988. 2. Eastman, J.T. The nature of the diversity of Antarctic fishes. UK : Polar Biology, 2005. pp. 93-107. Vol. 28. 28. 3. LINKOWSKI, T.B., J.M. REMBISZEWSKI. Ichthyological observations off the South Georgia coasts. W-wa : Pol.Arch.Hydrobiologii, 1978. pp. 697-704. Vol. 25. 4. Skóra, K. Wstępne wyniki z badań ichtiologicznych prowadzonych na r/v „Profesor Siedlecki” w czasie IV Morskiej Ekspedycji Antarktycznej. Gdynia : MIR, 1979. 5. Kock, K.-H. Antarctic fish and fisheries. Cambridge : Cambridge University Press, 1992. p. 359. 6. Traczyk, R. ANALYSIS OF THE MICROINCREMENTS RHYTHMIC STRUCTURES IN OTOLITH AND OTOLITH MORPHOMETRY FOR THE AGE, GROWTH AND IMPORTANT LIFE PERIODS DETERMINATION OF THE ANTARCTIC FISH Pseudochaenichthys georgianus NORMAN, 1939 OFF SOUTH GEORGIA AREA. http://georgianka.strefa.pl/pseudoch.html. [Online] INTERIA.PL S.A., 1990. 7. SARAH CLARKE, WILLIAM D.K. REID, MARTIN A. COLLINS* and MARK BELCHIER. Biology and distribution of South Georgia icefish (Pseudochaenichthys georgianus) around South Georgia and Shag Rocks. 20. UK : Antarctic Science Ltd., 2008. pp. 343–353. Vol. 4. 8. Jones, C.D., K.-H. Kock, E. Balguerias. CHANGES IN BIOMASS OF EIGHT SPECIES OF FINFISH AROUND THE SOUTH ORKNEY ISLANDS (SUBAREA 48.2) FROM THREE BOTTOM TRAWL SURVEYS. Hobart : CCAMLR Science, 2000. pp. 53-74. Vol. 7. 9. Rembiszewski, J. M., M. Krzeptowski, T.B. Linkowski,. Fishes (Pisces) as by-catch in fisheries of krill Euphausia superba Dana (Euphausiacae, Crustacea). Szczecin : Pol. Arch. Hydrobiol., 1978. pp. 677-695. Vol. 25. 10. Ślósarczyk, W. Preliminary estimation of abundance of juvenile Nototheniidae and Channichthyidae within krill swarms East of South Georgia I. 1. Szczecin : Acta Icht. et Piscatoria, 1983. Vol. XIII. 11. Ślósarczyk, W., Z. Cielniaszek. Postlarval and juvenile fish (Pisces, Perciformes and Myctophiformes) in the Antarctic Peninsula region the Antarctic Peninsula region, 1983/1984. 1-2. W-wa : Pol. Polar Res., 1985. pp. 159-165. Vol. 6. 12. Chojnacki, J., P. Palczewski. Age and growth of Pseudochaenichthys georgianus Norman 1937 (family Chaenichthyidae) of the South Georgia Region. W-wa : Pol. Polar Res., 1978, Vols. 2(3-4) , pp. 45-5. 13. Burchett, M.S. Abundance of the nearshore fish population at South Georgia (Antarctica) sampled by trammel net. Cambridge : Br. Antarct. Surv. Bull., 1983. pp. 39-43. Vol. 61. 14. North, A.W. Ecological studies of Antarctic fish with emphasis on early development of inshore stages at South Georgia. Cambridge : Br.Antarct.Surv. NERC, 1990. pp. 83-94. Vol. 62.
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