~. i. , Inlernalional Council ror C.H. 1992/G:22 lhe Exploralion Demersal Fish Cammillee o~ lhe Sea A comparison or Lhe growLh raLe i~ walleye pollock (Theragra chalcogramma) ~aken rrom Lhe open waLers or Lhe Bering Sea and Lhose rrom ~he open waLers or Lhe OkhoLsk Sea in 1991. Kowalewska-Pahlke Sea Fisheries Inslilule 81-332 Gdynia, Koll~laja 1 POLAND t~gdalena • A B S T R A C T The growlh o~ walleye pollock (Theragra chalcogramma) rrom lhe inlernalional walers o~ lhe Bering and Okholsk Sea was examined in 1991. Hean lenglh al age o~ pol lock was dislincly higher in lhe Bering Sea lhal in lhe Okholsk Sea. A comparison o~ lhe von Berlalanrry's growlh curves indicaled dirrerences in growlh belween lhe lwo regions. These dir~erences are expressed by lhe L-inr and K paramelers, ror bolh males and remales. Con~idence limils ror lhese lwo paramelers are dirrerenl and do nol overlap. The lesl o~ lhe hypolheses concerning lhe dirrerences belween maens lenglh al age o~ pol lock rrom lhe -inlernalional walers or lhe Bering and Okholsk Sea also indicaled slgniricanl dirrerences belween lhese lwo regions. • I N T ROD U C T ION Pollock (Theragra chalcogramma) is a species dislribuled over wide area or lhe Norlh Paciric. conlinenlal slope rrom lhe Sea or Il Japan occurs lhrough on lhe lhe shelr Okholsk a and Sea, Bering Sea and Gulr or Alaska,lo Vancouver. Il has been belived unlil recenlly lhal lhe species lives ne ar lhe boltom it however, lhal subslanlial pollack slocks occur also in turned deep out, walers, ., . ~he especially in or/~his stocks 45 M. ~ons. Bering Sea, leading ~he species in ~he and a pelagic mode or es~imated North Paciric are catches in reached 1989 living. 6 259 a~ The about ~hous ~ons ex~rac~ed rrom I CYearbook 1990). ~he abou~ Bering Sea and recent years species ~his in and large eas~ern populations are less visible. been:~he gene~ic Okhotsk ~he Sea. In greatest single- pollock be~ween North ~he Paciric Genetic dirrerences regional al~hough ~he in object or dirrerences or isola~ed. . were world rishery. ~he parts ~hous ~ons ~hous ~ons 1 800 species has exploi~ation Rela~ively wes~ern or wh ich 3 713 ou~ show rrom ~ha~ wi~hin ~hese dirrerences in grow~h ~he ~hese areas are ra~e. age • composition. meristi~ and morphometric rea~ures occur ~here. biology and stocks or pol lock in Al~hough ~he rela~ively weIl known. Sea. The objective or composition or the waters or growth between ~Iaterials in 1991,by the pol lock stock exploi~ed Okho~sk ~he to were rrom ~he otoliths were removed rrom 3001 Bering rish were measured and otoli~hs ror pol lock In and were removed rrom age a split visible accepted according reccomenda~ions o~ Sea in Gdynia. In or ~he on ~he and pollock December. 1801 Hay- and international 35 646 individuals. between reading. increments Okho~sk January specimens interpretation of growth ~he in individuals. Fork length or rish was measured with served Insti~ute Sea, Okhotsk Sea, between September ~o age on Polish rishing vessels Sea Fisheries ~he or collec~ed waters or the Bering and interna~ional otoliths and international ~he Sea and show dirrerences in August. measurements were made or 39 389 collected in leng~h Okhotsk 11 E T H·· 0 D ~he s~udy scien~ists ~he presen~ ~he two areas. in~Jinational wa~ers waters or Bering Sea are pollock rrom is AND ror abou~ paper ~hese tlATERIAL is known ~his Bering and ~he operating in lit~le ~he sexes. The The me~hod and the otolith were Special Group on Ageing rJethodology or Walleye Pollock. held in Gdynia in 1990 [Report 2 • , ~rom ~he chalcogramma)~ Areas o~ 1991 are presen~ed in open ~he wa~ers according ~he on leng~hs i~~ ~o basis (Theragra or pol lock ~he conduc~ed Bering and by Polish ~he Okho~sk Sea in was used in ~or calcula~ion years subsequen~ or or grow~h are li~e eqa~ion: o~ L~ = L inr ( * 1 - e -K. ~ ( * - ~o » ~ where: L~ - leng~h or ~ish a~ parame~ers calcula~ed Linr K. - ~ mean grow~h ~he age on ~~ basis or empirical ~he da~a: asymp~o~ie leng~h~ index), (ca~abolism coerricien~ de~ermines shape ~he or curve~ LO ~heore~ieal age~ The parameLers or aL whieh ~his calcula~ed In were ~his in~rodueed~ assuming.~ha~ ~he calculaLed me~hod~ addi~ion, in leng~h o~ leng~h would have a o~ on Lhe basis or mean lengLhs by abundaneei weigh~ed ~he ~ish equaLion described by Allen (1966). • Pol lock in Fig.1. Ber~alanr~y~s me~hod de~ermina~ed • ~ishing opera~ions ~or ~lee~ The von Walleye 1990]. deep-sea ra~ej Me~hology o~ Workshop on Ageing ~he ~ish ~heore~ieal by o~ 0 cm. meLhod ~he a~ a poin~ given O~ age, is O~ age 0 is elose ~ish a~ are coerrieien~s ~o 0 em. In order be~ween ~o L inr and K. or examined. An analysis or me an leng~hs ~he signi~ica~ce popula~ions ~rom ~hese ~wo pol lock ~or parame~ers de~ermine or ~ish carried ouL ~or bo~h where: x n nn mean in ~he ~he age-groups areas. The rollowing leng~h. number in sampie, 3 ~he di~:ferences· equa~ion ~he di~~erences or limi~s conridence Ber~alanr~y's signi:ficance or same ~he von areas. or pol lock s~a~isLies beLween was was used: were also . ". S2 - variance. n For larger'samples (n 1 > 30 and n 2 > 30). .. re~~ered Lo anormal disLribuLion (2). and s~a~is~ical small ~or values were samples LO L-SLudenL disLribuLion (L). c 0 M POS I T ION L ENG T 11 In Lhe internaLional lengLhs wa~ers the o~ Bering Sea pol lock 34-62 cm. mean being 50.2 cm (Fig.2.3). The main o~ the sLock (over 75 %) consisLed o~ rish below 41 cm and cm di~~erence over 55 between mean lengLh o~ with ~ish lengLhs barely cons~iLuted males and 3.6 was ~emales par~ 48-53 o~ had %. small o~ cm') The (0.5 4t cm). In Lhe inLernaLional waLers lengLhs o~,11-62 Lhe o~ cm. mean being 40.9 cm. The main (60 %) consisLed or rish Sea par~ pollock or di~~erence be~ween mean had sLock ~he lengLhs or 39-43 cm; rish below 33 wi~h over 47 cm consLiLued only 4.5 %. The o~ OkhoLsk and leng~h males and remales was 1.3 cm. Mean lengLhs pollock by sex in sLudy areas in o~ 1991 are Lhe ~ollowing: Bering Sea Okhot.sk Sea males 49.9 c. 40 . .1 c. f"e.ales 50.4 c. 41.4 c. .. & f 50.2 c. 4t ~ ~ 40.9 c. Comparing lengLh disLributions o~ pollack ~rom Lhe Lwo areas one may nOLice LhaL Lhe curves have a dicisievly single-peak disLribuLion and are similar. ~lean greater by over 9 cm AGE The lengLhs or pol lock ~han in ~rom 01' ~he Lhe Bering Sea were the Okhotsk Sea. C 0 M POS I T ION age considerably in composi~ion ~he two areas. 4 L ~ish ~rom exploi~ed st.ocks dirrered In waters or ~he in~erna~ional ~he were present in the stock, with groups represented by only ~he single Bering Sea 4- to 23-year-olds oldest specimens. 13-year old pol lock (year class or 1978); population occuring in ~he 12 and 11 % or In ~he ~his youngest age numerous ~Ios~ were 22 % or the ~hey cons~i~ued area (Fig.4,5) and rish'aged 12 and 14 population, respectively.· waters interna~ional 18-year-olds were present. Young classes or 1986 and 1985) tt ~he and or rish Okhotsk ~he aged 5 and Sea 6 predominatedi~hey consti~ued 1- years ~o (year 26 and 42 % or the population, respectively (Fig.4,5). Mean age or pol lock in the areas was ~wo ~he Bering Sea 6.3 "10.7 6.1 11.6 6.2 The considerable remales rrom ~he Okho1.sk Sea 12.3 aales :feaales rollowing: dirrerence Hering Sea is a in mean resul~ or age ~he between large males abundance and or remales rrom the 1983 and 1984 year classes. .' RAT E GR01'fTH The resul~s or analyses or mean lengths or age-groups exhibit considerable dirrerences pollock inhabiting the ~wo be~ween areas (Table 1). Mean rrom the international waters or age-groups between 5 and 15 years ~he rish ~"1.n leng~hs Bering Sea were ~han ~hose ~he individual stocks or larger in rish rrom ~he or pol lock in all Okhotsk Sea. Growth rate curves calculated on mean lengths in subsequent years lire point CFig.6,7). to The considerable calcula~ed dirrerences values or parameters have the following form: 5 ~he von between the ~wo Ber~alanrry's or areas equa~ion .-------------------- ---- Okhotsk Sea Hering Sea males .. 1'e.ales ..ales & l' 1'e.ales .. & l' 51.2 54.3 52.6 44.5 48.4 ~ 0.3293 0.2814 0.3031 0.47i2 0.3659 0.4357 1. 0 0.0096 0.0062 0.0083 -0.1237 -0.2051 -0.i393 LinC 45.6 The growth or pol lock rrom the Bering Sea is slower than that in rish rrom the Okhotsk Sea. Approximately at age or 9 years, length or about 50 cm, growth is arrested and in later increments are insigniricant. In t.he Okhotsk Sea at years the a length growth or pollock in t.he rirst years or lire is raster, but at the age or about 7 years it reaches an almost asymptotic lengt.h and its rurther growth • . is arrested. Conridence limits ror the Linr and K parameters or t.he von Bertalanrry's equation are dirrerent ror rish rrom these two areas (Fig.8,9). Only ror remales, t.he conridence limits or parameter K overlap ror the range or 0.28 1.0 0.30. The t.ested dirrerence bet.ween mean lengths or rish in the same ;i' age-group turned out 1.0 be signiricant ror the populations studied at probability p < < and showed probability p remales 0.001, with the exception or age-group 15, ror 0.01. mean Testing the t.hat ror males age-gr.oup 5 and lengths dirrerences remales ror separately are age-group ror which males insigniricant 12. This may be explained by the large dirrerence in abundance or tested rish between t.he Bering and Okhotsk Sea. DIS C U S S ION A comparison or length and age st.ructure or the international waters or the Bering and pol lock Okhotsk distinct. dirferences between t.hese two stocks. Genetic, and meristic studies (Iwata., 1975 Bakkala et al, stocks Sea in reveals morphometric 1983) suggested that the stocks of the Bering and Okhot.sk Sea do not mingle. However, within these stocks there 6 is a number of isolated spawning • populaLions. , In Lhe Bering Sea, despiLe expending in research potenLial, Lhe· sLrucLure clearly derined <AFSC proc. conLroversy surrounds Lhe or Rep. origin Lhe recenL years a pollock stock was 91-06,' 1991). or Lhe The pol lock large noL greatesL sLock rrom Lhe internaLional waters, where currenL invesLigaLion are cenLred. LengLh and age composiLion, growth raLe or pol lock rrom waters dirrer considerably pollock rrom Lhose or Lhe inLernaLional inhabiLing Lhe shelves or Lhe Bering Sea <Dawson 1988,1990). • As regards Lhe OkhoLsk Sea ,Lhere is no agreemenLs eiLher as Lhe number or pollock stocks inhabiLing disLinguishes six sLocks, while Fadiejew Lhis (1987) sea. IwaLa jusL as with respecL Lo the Bering Sea, there is (1975) disLinguishes spawning stocks only in Lhe norLhern parL or Lhe sea. In no Lhis rive case, certainty which stock rish taken ror examination came. Bakkala (1983) Lo rrom presents great suscepLibility or spawning stocks to environmenLal changes Lhe size or the spawning stock. Such siLuaLion place in the internaLional waLers or Lhe Bering had probably Sea, where very intensive exploitation or the pollock stock distinct its growLh rate occured <Janusz 1991), mosLly as a and taken due to changes in result or a decrease in reeding competition. The dirrerences in the strucLure and growth or pol lock rrom the inLernaLional waters or Lhe Bering and OkhoLsk Sea, presented in this ~~ paper, cannoL b~ treated as the only' deLerminanLs or independenL IL may be said, however, thaL there are signiricant dirrerences sLocks. in Lhe growth or pol lock beLween these two areas expressed by parameters Linr and K or Lhe von BerLalanrry's equaLion, both ror males and remales. This is conrirmed also by signiricance tesLs between mean lengLhs or rish in thc same age-group. R E F E REN CES Allen K.P.; 1966. A method or 7 ritLing growtll curves or the von BerLalanrry Lype observed vol.23. no 2. 163-179. , daLa. j. Fish. Res. Board. Can .• Disscusion Summaries rrom Lhe AFSC PROCESSED REPORT 91-06.; 1991. InLernaLional Workshop on Bering Sea Pollock SLock Assessment. February 4-8. 1991. Bakkala R.• T. Maeda .• G. McFarlane.; 1983. DistribuLion and sLock sLrucLure or pol lock (Theragra chalcogramma) in Lhe NorLh Paciric Ocean. Unpubl. rep ..• 23p. ~orLh~esL and Alaska Fi~heries CenLer.Natl. Mar. Fish. Serv .• NOAA.2725 Montlake Blvd. E.• Seattle. WA 98112. (Document submiLLed to Lhe InLernational NorLh Paciric Fisheries Commision in OCLober 1983) Dawson P.; 1988. Walleye Pollack Stock SLrucLure ImplicaLions rrom Age ComposiLion. Length-aL-Age. and MorphomeLric DaLa rrom Lhe CenLral and Eastern Bering Sea. Proceedings or the InLernational Symposium on Lhe Biology and ManagemenL or Walleye Pollock. Anchorage. Alaska USA.p. 605-642 • Dawson P.; 1990. InrormaLion on Lhe SLock SLrucLure or Bering Sea Pollack. CompilaLion or Papers PresenLed aL Lhe InLernaLional Symposium on Bering Sea Fisheries. Khabarovsk. USSR. 1990. p.65-73 Fadiejew N.• tl.G. Suczkova.; 1987. Raspriedielenie nagulnogo minLaja na sjewierie OChoLskogo Morja.-Populacjonnaja sLrukLura. dinamika czisljennosLi i sledowia minLaja. Sbornik naucznych Lrudow TINRO. WladywosLOk. 1987 p.69 ;IwaLaM.; 1975. PopulaLion idenLiricaLion ör walleye pol lock Theragra chalcogramma (PALLAS). in Lhe vicinity or japan. Memo. Fac. Fish. Hokkaido Univ. 22: 193-258 janusz j.; 1991. Impact or PopulaLion DensiLy on GrowLh RaLe or Walleye Pol lock in Lhe AleuLian Basin. 'BulleLin or Lhe Sea Fisheries IriSLiLute. No 1-2 (123-124): 22-30. Gdynia 1991 ReporL From The Workshop On Ageing MeLhology or Walleye Pol lock (Theragra chalcogramma). Held at the Sea Fisheries InsLiLuLe. Gdynia. 1990 Yearbook or SLaLis~ics - FAO or UN, 1990. vol.68 8 • Table 1. Mean length at age and test for hypotheses concerning the differences between mean lengths of walleye pollock from the international waters of the Bering and Okhotsk Sea in 1991 MLES :-------------------------------------------------------------------------------------: Bering Sea Okhotsk Sea :--------------:------------:---------------------------: age : .ean : length t variance: .ean : length : H.o. variance: : :i: 6: 7: • N.a. 40.13 43.47 45.83 8: 47.'14 9: 10: 11: 12: 13: 14: 15: 49.10 49.69 49.52 49.41 49.48 49.83 49.45 2.29 4.10 3.84 3.31 4.51 3.03 4.39 3.97 2.93 3.21 2.75 23 83 120 129 115 100 144 162 352 166 20 : : : : : : : : : : : t I • IBB 38.84 41.18 43.10 44.35 45.00 45.45 46.21 45.67 45.08 46.35 45.45 2.48 2.77 2.45 2.05 5.09 4.05 4.27 2.89 3.08 5.69 2.98 183 67 46 22 20 19 18 24 26 11 • : p<O.OOl : p<O.OOl p<O.OOl p<O.OOl p<O.OI p<0.05 p<0.05 p<O.OOl p<O.OOl p<O.Q01: :-------------------------------------------------------------------------------------: FEMLES :-------------------------------------------------------------------------------------; Bering Sea Okhotsk Sea :---------------------------:---------------------------; .ean ·• length age variance: aean •• length N.o. I I N.o. variance: t I . ' _________ : ___________________________ ' _______________- - - - - - - - - - - - ' - - -______ 1 ______ - - - ; :> : 11 37 97 179 149 120 106 96 124 244 119 40.19 43.29 46.28 49.36 51.20 52.27 52.29 51.99 52.41 52.56 54.38 6: 7: 8: 9: 10 : I • 12 : 13: 14: 15 I 2.64 5.13 5.25 6.42 6.49 7.82 7.79 4.78 4.93 3.88 2.73 8 39.35 42.33 44.35 45.70 46.86 48.48 49.14 50.53 49.76 50.15 50.60 2.97 3.24 3.76 5.13 7.15 5.54 7.17 p<O.OOl 13.45 1.24 p<O.OOl 6.57 • p<O.OOl 15.84 : p<O.OI 173 315 132 50 35 31 21 15 17 27 15 p(O.Ol p<O.OOl p<O.OOI p<O.OOl p<O.OOl p<O.OOl I I I I I I I I I I I :-------------------------------------------------------------------------------------, I'IALES Ir FElIALES :---------------------------------------------------------------------------: Bering Sea Okhotsk Sea :---------------------------:---------------------------: age .ean : length I I variance: .ean : length H.o. variance: H.o. I :---------'---------------------------'---------------------------:---------' 5 6 7 3 9 10 11 40.12 43.37 46.10 48.70 50.17 51.04 50.63 60 180 299 278 235 206 240 2.41 4.61 4.74 5.48 6.62 7.12 7.59 : 39.08 41.91 43.93 45.05 46.14 47.29 47.75 361 498 199 96 57 51 40 2.78 3.38 3.66 4.11 7.17 7.15 7.94 : : : : : : : p}O.OOl p}O.OOl p}O.OOI p}0.OO1 p}0.QQ1 p}O.OOI p}O.QOl 12 50.53 28b 5.90 : 47.88 33 13.5b : p>O.OOl 13 : H: 50.68 50.97 596 285 5.83 : 5.30 : 47.02 48.28 41 53 10.12 : p}O.OOI 9.75 : p}O.OOl 15 : 50.8b 28 7.b9 : 48.42 2b lb.86 : p>O.Ol ;---------------------------------------------------------------------------. 9 ~; 11,0 0 100:' i / I I i I .. / f:OON i .. ' i \BERING \ rv/r':~E o . KIlO T S K I» S E A I \ ij "i I \ " I I '\ ./ ,/ • / ./ . .. ,..... ./ . /~.~.;~:-...\' ~ ;." .:. ..:.::.::::.,. \ " P .' i . • .. .t! -.-.-.. {) " " .......... ,............ ..... ~'-'-'-'-' ...... .... -,,_.-. .... ..... .,,/ -"-~ ._.- .,- / ;/// ./ "", /' ! )-........ \ .I ....."._._. • ~ " .. ;" .-._._i , .,.... ", I 'J i f A / . /". ./ i.. ,/_ _-J.._ _-J.._ _- - J '--'-'-'-~~-L.--"':'<:-..L. l -_ _. 1 -_ _...L-_ _. l -_ _-1-_ _...J-_ _...J Fig. 1. Sarnpling areas of walleye pollock futhe international waters of the Hering and Okhotsk Sea ". in 1991. 4ff' Length distribution wal.leye pollack 1991 16..,.--------------------~ 14 12 • - 10 c ~ L. 8 (D n.. 6 4 2 o ' 11 14 17 20 23 26 29 32 35 38 41 44 47 50 53 56 59 62 Length (ern) I-+Fig. c.' 2~ Bering Sea -B- Okhock Sea Length distribution of walleye pollock caught in the international waters of the Bering and Okhotsk Sea in 1991 (males and females). 11 - ,..--------------- - -- --~~ Length distribution a) walleye pollack 1991 males 25.----------------------, 20 -c: 15 (iJ m\ Cl o m I L r ~ ? b Cl a.. 10 p 5 \ q c;j o ~ O • I t • i. 11 14 17 20 23 26 29 32 35 38 41 44 47 50 53 56 59 62 Length (ern) I-+b) Bering Ses • -e- Okhotsk Sea Length distribution walleye pollack 1991 femaJes 25.----------------------.., 20 ~ --c: 15 1\ rp\ •o ! ci i i o r: ':1 I!I \ \ C!.l \\ l!J\ / i oL - . , 11 14 17 20 23 26 29 32 35 38 41 44 47 50 53 56 59 62 Length (ern) I-+?ig. 3. Length in the Bering See distributic~ of walleye pollack internatic~al and Okhotsk Sea a) males b) -=- Cxhctsk Ses females - 12 i~ caugh~ waters of the Bering 1991 Age distribution walleye pollack 1991 400 1 350 85 300 86 Q)250 78 ,. E-2OQ 0 L- a... 150 100 J 50 , 0 I I • , I I I I I I I 1 2 3 4 5 6 7 8 910111213141516171 81 ~ Aga I0 8erin~.Sea _ Okhock Sea • Fig. 4. Age distribution of walleye pollock caught in the international waters of the Bering and Okhotsk Sea in 1991 - 13 (males and females) Age distribution a) walleye pollock 1991 males , 350 300 86 85 1 250 .; ~200 E o 0:. 150 • 78 I I 87 100 50 o. . 84ß3...~1 88 . 12345678 I 77 73 75 ~ ... ~ . 7q1 I- grJ32~ eil9 r, ~~ 91011121314151617181~O Aga ID Bering Ses ~ Okhotsk Ses I Age distribution b) walleye pollock 1991 femaIes 85 IDEering See ~ Fig. 5. Age distributic~ Okhctsk See I of walleye pollack caught in the international waters of the Bering and Okhotsk Sea in 1991 a) males b) females 14 - • ". ' . • walleye pollack 1991 males &females 55,...------------------------. 50 45 • ..-.40 E ~35 .c ~ "&30 c 0)' --l 25 20 15 10 .l.-...y----r----r--,--,---r--,---r---r---r---r--,---r---r---r---r-~~~_r__l 1 234 5 6 7 8 91011121314151617181920 Aga I 0 8.S. v.8art - • 8.S. 0.8. v.8art - 0.8. Fig. 6. Growth curves of walleye pollock caught in the international waters of the Bering and Okhotsk Sea in 1991 (males and females) - 15 ..---------- - walleye pollack 1991 a) females. 55...--------::-:::=:::;::;:::E:;=~:=T=1 I + + 50 45 -.40 E ~35 ..c "&30 c:I) -J 25 20 15 10 ...L...r--r----r-~.....___.__.__-r-_,_-r---r-..__~r__r___T__,..__..___r_____r_l 123 4 5 6 7891011121314151617181920 Aga 1- 8. S. v.Bert b) - + 8. S. O. S. v.Bert O. S. 0 walleye pollack 1991 males 55,....----------------------.. + 50 o 45 + o _40 E " ~35 ..r:: 0,30 c: Cl -J 25 20 15 10 ...L...r-r"---r-~.....__.___.___,__,_.,..__..__..__,..__,r__r_.,.__..___r___.___..__J 123 4 5 6 7891011121314151617181920 . Aga' 1- 8, S. v,Bert + - B. S. . O. S. v.Bert 0 O. S. Fig. 7. Growth curves of walleye pollack caught in the international waters of the Bering and Okhotsk Sea in 1991 a) females b) males 16 • Contidence limits tor L-int walleye pollack 1991 56 ~ 54 I ~ 52 50 r IBerlng I· I Sec I 48 46 44 I CM-otsk Sea I 42 males females m&f males femaJes m&f Fig. 8. Confidence limits for L-inf of walleye pol lock caught in the international waters of the Bering and Okhotsk Sea in 1991 Confidence limits for K walleye pollack 1991 0.6 0.55 10000tsk Sec 0.5 I 0.45 I 0.4 I Berfng Sec 0.35 0.3 0.25 I ~ males females Fig. 9. Confidence limits m&f fo~ males femaJes m&f K of walleye pollock caught in the international waters of the Bering and Okhotsk Sea in 1991 17