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1 한수지 5(1), 65-76, 217 Original Article Korean J Fish Aquat Sci 5(1),65-76,217 생활사기반모델을이용한동해명태 (Gadus chalcogrammus) 의개체군평가 김규한 손명호 1 현상윤 * 부경대학교자원생물학과, 1 국립수산과학원제주수산연구소 A Life Stage-based Model for Assessing the Walleye Pollock Gadus chalcogrammus Population in the East Sea Kyuhan Kim, Myoung Ho Sohn 1 and Saang-Yoon Hyun* Department of Marine Biology, Pukyong National University, Busan 48513, Korea 1 Jeju Fisheries Research Institute, National Institute of Fisheries Science, Jeju 6368, Korea Since the late 199s, walleye pollock Gadus chalcogrammus fisheries in Korean waters have been considered collapsed. Although many fisheries scientists suspect that the collapse might have been triggered by overexploitation of juvenile pollock or environmental changes, such conjectures have been neither tested nor investigated, partially because of limited data on the population. There has been no survey of the population, and the ages of fish in fishery catch have rarely been identified. Instead, fishery catch data from included information about two life stages, juveniles and adults, and data on catch-per-unit-effort (CPUE) during and those on fish length and weight during had been sporadically collected from commercial fisheries. To test hypotheses about the collapse of the pollock fisheries, we used a statistical linear model with juvenile CPUE as the response variable, and abiotic (e.g., water temperatures) and biotic factors [e.g., adult pollock, flatfishes (Pleuronectidae sp.), and sandfish (Arctoscopus japonicus) CPUEs] as the explanatory variables. The model results indicated that depletion of the pollock population was associated with both biotic (adult pollock and flatfishes abundance) and abiotic factors (mid-water temperatures in February and October). We further interpreted the results from ecological and biological perspectives, suggesting possible mechanisms. Key words: Pollock population depletion, Generalized linear model, Catch-per-unit-effort, Fisheries selectivity 서론 (Gadus chalcogrammus) , (climate regime shift) (Zhang et al., 2; Zhang and Gong, 25; Kim et al., 27).., Nishimura (1969) 2-5 C 2 m (permanent thermocline) 5 C 2 m. Gong and Zhang (1986) Korean J Fish Aquat Sci 5(1) 65-76, February 217 This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial Licens ( which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited. Received 27 December 216; Revised 13 February 217; Accepted 14 February 217 *Corresponding author: Tel: Fax: address: shyun@pknu.ac.kr Copyright 217 The Korean Society of Fisheries and Aquatic Science 65 pissn: , eissn:

2 66 김규한ㆍ손명호ㆍ현상윤 Kooka et al. (1998) m 4-5 m. (ontogenetic vertical migration) (Yamamura et al., 22; Duffy-Anderson et al., 23; Smart et al., 213; Parker-Stetter et al., 215) (Kim, 1978), 1975 ( : ) ( ) (Fig. 1a). Kang et al. (213) % Longline CPUE (kg basket -1 ) Catch in MT ( 1 3 ) Danish-seine CPUE (kg haul -1 ) (b) (c) (a) Year Juvenile Adult Longline Gillnet Gillnet CPUE (kg sheet -1 ) Fig. 1. Data on fishery catches of adult and juvenile pollock Gadus chalcogrammus in the East Sea (a), and catch per unit effort (CPUE) collected from gillnet and longline fisheries (b), and Danish seine fishery (c). A sheet is a unit area of the gillnet whose width ranges from 45 m to 72 m and whose height ranges from 4 m to m. One basket in the longline contains about 3 hooks (recruitment to the adult population),,.,, ( ). Virtual Population Analysis (VPA) (Yabuki and Honda, 25; Mori et al., 212) (Shida et al., 27; Funamoto, 211; Funamoto et al., 213; Funamoto et al., 214). (i.e., ).,. (Generalized linear model:glm),.,, (Pleuronectidae sp.) (Arctoscopus japonicus),. 재료및방법 생물학적정보 ,,,.,,,. 12,45 5,699 (47%), 2,668 (22%), 1,632 (14%), 69 (6%) 1,356 (11%)., 39 1 ( , , 21) (Total

3 생활사기반동해명태개체군평가 67 length: TL) (Fork length: FL) , 1989, cm. (survey). (Fig. 2). (Table 1)., Kim and Huh (1978) ( 27 cm) 27 cm., (i.e., ) ( 27 cm) (> 27 cm) 9:1, 199 (i.e., ) 3:1., ( ). Table 1. Annual juvenile and adult pollock Gadus chalcogrammus catches by four major pollock fisheries in Korean waters Year Adult catch by fishery (MT) Juvenile catch by fishery (MT) Danish Gillnet Longline Trawl Total Danish Gillnet Longline Trawl Total ,264 1, ,369 38,69 11,592 5, , ,175 52, ,4 69, ,721 8, ,349 64, ,714 92, ,11 1, ,444 52, ,322 88, ,291 7,78 1,533 1,68 11,582 5, ,463 34, ,464 13,916 4,863 5,698 26,941 33, ,922 65, ,79 18,89 2,992 5,219 46,81 47, ,296 89, ,826 19,5 1,738 4,779 37,843 47, ,88 94, ,4 12,78 7,624 4,177 28,585 31, ,75 55, ,435 18,395 12,11 3,458 39,398 37, ,792 65, ,534 22,892 5,817 9,96 46,23 22, ,67 37, ,182 21,454 7,411 8,36 46,47 19, ,554 31, ,674 5,953 5,219 3,945 19,791 8, ,825 13, ,382 5,753 2,457 1,649 13,241 1, ,32 2, ,11 6,41 2,939 2,162 15,612 6,4 4 1,236 7, ,154 4,85 1,867 1,496 9,62 13, ,34 15, ,664 1,137 1,951 2,533 8,285 6, ,295 1, ,71 1,612 1, ,962 3, ,584 4, ,649 2,422 2,446 1,558 8,75 2, ,53 7, ,94 2,131 3, ,2 1, ,839 3, ,635 2, ,723 1, , ,76 1, ,332 1, ,79 3, , ,19 6,

68 김규한ㆍ손명호ㆍ현상윤 CPUE 자료 1963 27,, (catch-per-uniteffort: CPUE) (Fig. 1b, 1c and Fig. 3)., CPUE.,,, CPUE CPUE. (i.e., ) (1-3 ) (1-3 ). CPUE CPUE CPUE.,., CPUE, CPUE CPUE. 반응변수 27 cm,.

4 68 김규한ㆍ손명호ㆍ현상윤 CPUE 자료 ,, (catch-per-uniteffort: CPUE) (Fig. 1b, 1c and Fig. 3)., CPUE.,,, CPUE CPUE. (i.e., ) (1-3 ) (1-3 ). CPUE CPUE CPUE.,., CPUE, CPUE CPUE. 반응변수 27 cm,., CPUE 2 (recruitment to the fishery). CPUE, CPUE., ( cm) Park et al. (1978) von Bertalanffy [L t =65.4 (1 exp.223 (t+.249) )]. t, L t t (cm) von Bertanalffy FL TL FL. FL-TL, FL= TL (r 2 =.98, n=8189) FL (a) 39 Frequency 3 (b) (c) Total length (cm) Fig. 2. Length-frequency distribution of walleye pollock Gadus chalcogrammus caught by gillnet (a), longline (b), and Danish seine (c) fisheries Fig. 3. Blocks [approximately 3 3 (nautical mile) 2 ] in dark gray where the CPUE data were collected. Those encompassed by the solid and broken lines denote major locations where gillnet and longline fisheries, and Danish seine fishery were operated, respectively

5 생활사기반동해명태개체군평가 69 (13-25 cm) von Bertalanffy.7-2. CPUE CPUE CPUE (r>.8, P-value<.5) CPUE CPUE , (1-3 ) 17-2 cm (i.e., cm) 1 (Fig. 2c), t CPUE 1-3 t 2, 6-8 t 1. CPUE. 설명변수 (abiotic) (Fig. 3). (Kim and Yoo, 27), 7. (KODC) (KODC, 211) Feb Apr Depth (m) Jun Oct Aug Dec Temperature ( ) Fig. 4. Bimonthly vertical temperature profiles of Block 7 during Each solid line indicates an annual temperature profile of every other month (i.e., February, April, June, August, October, and December).

6 7 김규한ㆍ손명호ㆍ현상윤 15 m -15 m 5 m (Fig. 4). (multicollinearity)..7 (P-value<.5).,, CPUE. CPUE CPUE CPUE (Spawning Stock Biomass: SSB). CPUE. CPUE CPUE CPUE. 시차적용 (Winter and Swartzman, 26). CPUE 1 2 CPUE 1 2 (timelag). CPUE Response variable Juvenile CPUE (Jan~Mar in year t) Juvenile CPUE (Jun~Aug in year t) Age-2 in year t Age-1 in year t Candidate explanatory variables Temperatures (year t-2) Adult CPUEs (Oct~Mar in year t-2) Flatfishes CPUE (Jan~Mar in year t) Sandfish CPUE (Jan~Mar in year t) Temperatures (year t-1) Adult CPUEs (Oct~Mar in year t-1) Flatfishes CPUE (Jun~Aug in year t) Sandfish CPUE (Jun~Aug in year t) Fig. 5. Diagram depicting all the variables considered for the linear model. In the candidate explanatory variables, the numbers subtracted from t (e.g., t-1 and t-2, where t denotes the year index), indicate time delays by which CPUE of juveniles at age-1 and age- 2 were used as that of juveniles at age (Fig. 5). 일반화선형모델 (Generalized linear model: GLM). Y=X β+ε Y (n 1) log ( CPUE t ). n t CPUE (skewness). X (n p) (design matrix) (interaction term). p β. ε. (analysis of deviance: AN- ODEV) (Hyun et al., 214). (D) (saturated model) (proposed model) (likelihood ratio) 2 (McCullagh and Nelder, 1989; Dobson and Barnett, 28)., D = 2 (l saturated l proposed ) = 2 (l proposed l saturated ) l., D (null model). (nested model) ( D). 2 ( df) (McCullagh and Nelder, 1989; Dobson and Barnett, 28): D~ 2 df. 모델진단.. (Cook s distance), C i (Cook and Weisberg, 1982) C i =(b b (i) ) T (Χ T X)(b b (i) )/ (p MSE). i C i 1 b, b (i)

7 생활사기반동해명태개체군평가 71 i, MSE T. (externally studentized residual) d i. d i =e i /( MSE (i) (1 h ii )) e i i, MSE (i) i h ii (hat matrix) i. (n p 1) t 1.5 d i >t(1 /2; n p 1) (Kutner et al., 25). n, p.. (ridge trace) (mean-centering). (Variance Inflaction Factor: VIF). (ridge estimator), ^ β( )=X T X+ I) -1 X T Y (biasing parameter), ^β( ) (Hoerl and Kennard, 197)., ^β( ). 가입과정분석 SSB. CPUE., 1 6 CPUE CPUE. (i.e., CPUE),. 결과 모델의선택과진단 Table 2 1+Temp1.15_Feb+Long+Temp15_Oct+Flat+Long Temp15_Oct. Shapiro-Wilk (Shapiro and Wilk, 1965; R Core Team, 213) (Pvalue=.35) (Fig. 6), Breusch-Pagan (Breusch and Pagan, 1979; R Core Team, 213; Hothorn et al., 215) (P-value=.55) Sample quantiles Theoretical quantiles Fig. 6. Quantile-quantile plot of residuals for the final model. Table 2. Model selection summary. Temp 1.15_Feb, Long, Temp 15_Oct, and Flat denote the average of water temperatures at the depths of 1 and 15 m in Feburary, longline CPUE, that of water temperatures at the depth of 15 m in October, and flatfishes CPUE, respectively Model Linear form D df ΔD Δdf P-value AIC 1 Null NA NA NA Temp 1.15_Feb Temp 1.15_Feb + Long Temp 1.15_Feb + Long + Temp 15_Oct Temp 1.15_Feb + Long + Temp 15_Oct + Flat Temp 1.15_Feb + Long + Temp 15_Oct + Flat + Long Temp 15_Oct

8 72 김규한ㆍ손명호ㆍ현상윤. (Fig. 7a). 5 (numerical instability). Fig. 7b. (Table 3). Standardized coefficient (a) (b) Temp 1.15_Feb Long Temp 15_Oct Flat Long Temp 15_Oct Temp 1.15_Feb Long Temp 15_Oct Flat Long Temp 15_Oct Biasing parameter (θ) Fig. 7. Ridge trace plots describing trends of the estimated coefficients as a function of biasing parameters θ before (a) and after (b) mean-centering. 모델결과 (final model) (Table 3b).., (Afshartous and Preston, 211). 2 1 m 15 m., m CPUE (interaction effect) (Fig. 8). 어획의영향 3 CPUE (r>.7) 4 (r>.9) (Table 4). CPUE CPUE 1 3,, 3 4. Fig. 9 Table 3. Estimates of coefficients (Coef) of explanatory variables and their standard errors (SE) in the best model before mean-centering (a) and those after mean-centering (b). Temp 1.15_Feb, Long, Temp 15_Oct, and Flat denote the average of water temperatures at the depths of 1 and 15 m in Feburary, longline CPUE, the average of water temperatures at the depth of 15 m in October, and flatfishes CPUE, respectively Explanatory variable (a) Before mean-centering (b) After mean-centering Coef SE t P-value Coef SE t P-value Intercept Temp 1.15_Feb Long Temp 15_Oct Flat Long Temp15_Oct

9 생활사기반동해명태개체군평가 73 6 Environmental effect Fishing effect 5 Temp15_Oct ( ) Juvenile CPUE (log of Kg/haul) Long (kg/basket) Temp 15_Oct ( ) Fig. 8. Three-dimensional plot of the interaction term, Long and Temp 15_Oct. Those two variables associated with the interaction term were mean-centered (e.g., Long c =Long-Long and Temp 15_Oct c =Temp 15_Oct-Temp 15_Oct, where the subscript c denotes the mean-centering procedure; the means of the two variables, Long and Temp 15_Oct are 9.9 kg/basket and 4.91 C, respectively).. 고찰,. (Montgomery et al., 215)., Temp1.15_Feb (+) Flat (+) Age- Age-1, 2 Adult Long ( ) Fig. 9. Summary for interpreting study results. The dashed arrows show pathways of significant explanatory variables and those in parentheses indicate a direction of effect (i.e., -, + and denote negative, positive and interaction effects, respectively). Areas separated by gray represent a major source of mortality on each life stage of walleye pollock Gadus chalcogrammus; environmental effect on those in white; fishing effect on those in gray m. 15 m 15 m (Temp 1.15_Feb Temp 15_Oct)., 15 m m. Table 4. Time-lagged correlations between catch and CPUE of juvenile, and those of adult. Those in parentheses denote a type of fishery used for data collection, and values followed by an asterisk denote significant difference (P-value<.5) Data Shift value in year Juvenile catch Adult catch.576*.688*.836*.939*.867*.716* Juvenile CPUE (Danish seine) Adult CPUE (gillnet) *.787*.672*.4.32 Adult CPUE (longline) *.796*.671*

10 74 김규한ㆍ손명호ㆍ현상윤. (1) Temp 1.15_Feb: 2 1 m 15 m Cushing (1969). 1 m 15 m. (spring bloom) (Yamada et al., 24),., Shida et al. (27) 2 1-2, Funamoto et al. (214) m, 2., Shida et al. (27) 2. (2) Flat: (3) Long Temp 15_Oct: CPUE 1 15 m., 1 15 m.,..,. 1 (fall bloom) (Heintz and Vollenweider, 21)., (1) Temp 1.15_Feb.,.. Francis and Bailey (1983) Olla and Davis (199).,, m. (uncertainty).. CPUE (survey), CPUE 25. (variance). Park et al. (1978) von Bertalanffy (deterministic model),.. Kooka (212) von Bertalanffy Park et al. (1978), Park et al. (1978)., Kooka (212), Park et al. (1978)

11 생활사기반동해명태개체군평가 75,.,.,, CPUE.,,. 사사,,,,,,,. (215 ). References Afshartous D and Preston RA Key results of interaction models with centering. J Stat Educ 19, Breusch TS and Pagan AR A simple test for heteroscedasticity and random coefficient variation. Econometrica 47, Cook RD and Weisberg S Residuals and influence in regression. Chapman and Hall, London, U.K., Cushing D The regularity of the spawning season of some fishes. ICES J Mar Sci 33, icesjms/ Dobson AJ and Barnett A. 28. An introduction to generalized linear models. CRC press, Boca Raton, U.S.A., Duffy-Anderson J, Ciannelli L, Honkalehto T, Bailey KM, Sogard S, Springer A and Buckley T. 23. Distribution of age-1 and age-2 walleye pollock in the Gulf of Alaska and eastern Bering Sea: sources of variation and implications for higher trophic levels. In: The Big Fish Bang: Proceedings of the 26th Annual Larval Fish Conference. Browman HI and Skiftesvik AB, eds. Institute of Marine Research, Bergen, Francis RC and Bailey KM Factors Affecting Recruitment of Selected Gadoids in the Northeast Pacific and East Bering Sea in: From Year to Year, Wooster WS, ed. A Washington Sea Grant Publication, Seattle, U.S.A., Funamoto T Causes of walleye pollock (Theragra chalcogramma) recruitment decline in the northern Sea of Japan: implications for stock management. Fish Oceanogr 2, Funamoto T, Yamamura O, Kono T, Hamatsu T and Nishimura A Abiotic and biotic factors affecting recruitment variability of walleye pollock (Theragra chalcogramma) off the Pacific coast of Hokkaido, Japan. Fish Oceanogr 22, Funamoto T, Yamamura O, Shida O, Itaya K, Mori K, Hiyama Y and Sakurai Y Comparison of factors affecting recruitment variability of walleye pollock Theragra chalcogramma in the Pacific Ocean and the Sea of Japan off northern Japan. Fish Sci 8, s z. Gong Y and Zhang C The walleye Pollock (Theragra chalcogramma) stock in Korean waters. Int North Pac Fish Comm 45, Heintz RA and Vollenweider JJ. 21. Influence of size on the sources of energy consumed by overwintering walleye pollock (Theragra chalcogramma). J Exp Mar Biol Ecol 393, Hoerl AE and Kennard RW Ridge regression: Biased estimation for nonorthogonal problems. Technometrics 12, Hothorn T, Zeileis A, Farebrother R, Cummins C, Millo G and Mitchell D lmtest: Testing linear regression models. R package. Hyun S-Y, Cadrin SX and Roman S Fixed and mixed effect models for fishery data on depth distribution of Georges Bank yellowtail flounder. Fish Res 157, dx.doi.org/1.116/j.fishres Kang S, Park JH and Kim S Size-class estimation of the number of walleye pollock Theragra chalcogramma caught in the southwestern East Sea during the 197s-199s. Korean J Fish Aquat Sci 46, kfas Kendall A, Incze L, Ortner P, Cummings S and Brown P The vertical distribution of eggs and larvae of walleye pollock, Theragra chalcogramma, in Shelikof Strait, Gulf of Alaska. Oceanograph Lit Rev 5, 399. Kim H and Yoo S. 27. Relationship between phytoplankton bloom and wind stress in the sub-polar frontal area of the Japan/East Sea. J Mar Syst 67, org/1.116/j.jmarsys Kim S, Zhang C, Kim J, Oh J, Kang S and Lee JB. 27. Climate variability and its effects on major fisheries in Korea. Ocean Sci J 42, Kim WS and Huh S Meristic and morphometric observations on nogari and Alaska Pollack. J Oceanol Soc 13, Kim YM Stock Assessment of small size fishes of Alaska Pollack, Theregra chalcogramma (PALLAS), in the Eastern Sea of Korea. Bull Fish Res Dev Agency 19, KODC (Korea Oceanographic Data Center) NFRDI (National Fisheries Research and Development Institute) Serial Oceanographic Observation[Internet]. Retrieved from

12 76 김규한ㆍ손명호ㆍ현상윤 page?id=eng_obs_2_1 on March 15, 216. Kooka K Life-history traits of walleye pollock, Theragra chalcogramma, in the northeastern Japan Sea during early to mid 199s. Fish Res 113, fishres Kooka K, Takatsu T, Kamei Y, Nakatani T and Takahashi T Vertical distribution and prey of Walleye Pollock in the Northern Japan sea. Fish Sci 64, Kutner MH, Nachtsheim CJ, Neter J and Li W. 25. Applied linear statistical models. McGraw-Hill Irwin, New York, U.S.A., McCullagh P and Nelder JA Generalized linear models. CRC press. Boca Raton, U.S.A.. Montgomery DC, Peck EA and Vining GG Introduction to linear regression analysis. John Wiley and Sons, Hoboken, U.S.A., Mori K, Funamoto T, Yamashita Y and Chimura M Stock assessment and evaluation for Japanese Pacific stock of walleye pollock (fiscal year 211) In: Marine Fisheries Stock Assessment and Evaluation for Japanese Waters (fiscal year 211/212). Fisheries Agency and Fisheries Research Agency of Japan, Tokyo, Japan, Nishimura S The Zoogeographocal aspects of the Japan sea -Part V. Publ Seto Mar Biol Lab 17, Olla BL and Davis MW Effects of physical factors on the vertical distribution of larval walleye pollock Theragra chalcogramma under controlled laboratory conditions. Mar Ecol Prog Ser 63, meps6315. Park B, Hue J and Kim H Age and growth of Alaska pollack, Theragra Chalcogramma, in the Eastern Sea of Korea. Bull Fish Res Dev Agency 2, Parker-Stetter SL, Horne JK, Urmy SS, Heintz RA, Eisner LB and Farley EV Vertical Distribution of Age- Walleye Pollock during Late Summer: Environment or Ontogeny?. Mar Coast Fish 7, Shapiro SS and Wilk MB An analysis of variance test for normality (complete samples). Biometrika 52, Shida O, Hamatsu T, Nishimura A, Suzaki A, Yamamoto J, Miyashita K and Sakurai Y. 27. Interannual fluctuations in recruitment of walleye pollock in the Oyashio region related to environmental changes. Deep-sea Res Pt II 54, Smart T, Siddon E and Duffy-Anderson J Vertical distributions of the early life stages of walleye pollock (Theragra chalcogramma) in the Southeastern Bering Sea. Deep-sea Res Pt II 94, dsr R Core Team R: A language and environment for statistical computing. Foundation for Statistical Computing, Vienna, Austria. Winter AG and Swartzman GL. 26. Interannual changes in distribution of age- walleye pollock near the Pribilof Islands, Alaska, with reference to the prediction of pollock year-class strength. ICES J Mar Sci 63, Yabuki K and Honda S. 25. Stock assessment of Japanese Pacific Population of walleye pollock in 23. In: Assessments of fishery stocks in the Japanese Waters. Fisheries Agency and Fisheries Research Agency of Japan, Tokyo, Japan, Yamada K, Ishizaka J, Yoo S, Kim H and Chiba S. 24. Seasonal and interannual variability of sea surface chlorophyll a concentration in the Japan/East Sea (JES). Prog Oceanogr 61, Yamamura O, Honda S, Shida O and Hamatsu T. 22. Diets of walleye pollock Theragra chalcogramma in the Doto area, northern Japan: ontogenetic and seasonal variations. Mar Ecol Prog Ser 238, meps Zhang CI and Gong Y. 25. Effect of ocean climate changes on the Korean stock of Pacific saury, Cololabis saira (BREVOORT). J Oceanogr 61, org/1.17/s Zhang CI, Lee JB, Kim S and Oh J. 2. Climatic regime shifts and their impacts on marine ecosystem and fisheries resources in Korean waters. Prog Oceanogr 47,

수산과학회지46-4.indd

수산과학회지46-4.indd 한수지 46(4), 445-453, 213 Original Article Kor J Fish Aquat Sci 46(4),445-453,213 197-199 년대동해에서어획된명태 (Theragra chalcogramma) 의체장에따른체급별어획마릿수추정 강수경ㆍ박정호 1 ㆍ김수암 2 * 국립수산과학원자원관리과, 1 국립수산과학원동해수산연구소, 2 부경대학교자원생물학과