Characteristics of the Eggs and Larval Distribution and Transport Process in the Early Life Stage of the Chub Mackerel Scomber japonicus Near Korean Waters

666

Copyright © 2019 The Korean Society of Fisheries and Aquatic Science pISSN:0374-8111, eISSN:2287-8815

서 론

고등어

(Scomber japonicus)

(pelagic fish)

,

(Collette and Nauen, 1983; Scoles

et al., 1998).

는고등어는분포범위및회유경로

,

(Tsushima Current stock)

(Pacific stock)

나뉜다

(Shiraishi et al., 2008).

·

·

,

,

,

(Hiyama et al., 2002;

Yukami et al., 2009; Sassa and Tsukamoto, 2010; Lee et al.,

2016).

며

,

Izu Islands

(Murayama et al., 1995; Watanabe et al., 1999).

한국 연근해에 분포하는 고등어(Scomber japonicus) 난∙자치어의 분포특 성 및 초기 수송과정 연구

김소라·김중진*·William T. Stockhausen ¹ ·김창신·강수경·차형기·지환성 ² ·장서하 ³ ·백혜자 ³

국립수산과학원 연근해자원과, ¹미해양대기청 알래스카수산연구소, ²국립수산과학원 수산자원연구센터, ³부경대학교 자원생물학과

Characteristics of the Eggs and Larval Distribution and Transport

Process in the Early Life Stage of the Chub Mackerel Scomber japonicus Near Korean Waters

So Ra Kim, Jung Jin Kim*, William T. Stockhausen

¹

²

³

³

Costal Water Fisheries Resources Research Division, National Institute of Fisheries Science, Busan 46083, Korea

1Alaska Fisheries Science Center, NOAA/NMFS, 7600 Sand Point Way, NE, Seattle, WA 98115-6349, USA

2Fisheries Resources Research Center, National Institute of Fisheries Science, Tongyeong 53064, Korea

3Department of Marine Biology, Graduate School, Pukyong National University, Busan 48513, Korea

The horizontal distributions of eggs and larvae of chub mackerel Scomber japonicus were extensively surveyed in the vicinity of Korean waters between 31°75'N and 36°50'N during May and June in 2016 and 2017 (total of four sur- veys). We used a coupled bio-physical model (DisMELS) that combines an individual-based model (IBM) incorpo- rating vertical migration of larvae and temperature-dependent survival to understand transport processes in the early life stage. Using the distributions of eggs and larvae from surveys, the potential spawning grounds were estimated at the northwest and southeast of Jeju Island and the central East China Sea in May, and at the southwestern East Sea and southern West Sea in June by running the model backward in time. In forward experiments within 30 days from the backward results, most larvae were transported to both the Korean and Japanese sides of the East Sea through the Korea Strait. However, the larvae released in the central East China Sea were transported to the Japanese side only, while those released in the southern West Sea were retained within that region. The survival rates at 30 days after release based on the simulation incorporating temperature-dependent survival throughout May and June were 29.7%

in 2016 and 28.8% in 2017.

Key words: Scomber japonicus , Eggs and larvae, Spawning ground, Transport, Individual-based model (IBM)

*Corresponding author: Tel: +82. 51. 720. 2287 Fax: +82. 51. 720. 2277 E-mail address: [email protected]

This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial Licens (http://creativecommons.org/licenses/by-nc/3.0/) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

Received 25 November 2019; Revised 5 December 2019; Accepted 17 December 2019 저자 직위: 김소라(연구원), 김중진(연구사), William T. Stockhausen(연구원), 김창신(연구사), 강수경(연구관), 차형기(연구관), 지환성(연구사), 장서하(연 구원), 백헤자(교수)

https://doi.org/10.5657/KFAS.2019.0666

Korean J Fish Aquat Sci 52(6), 666-684, December 2019

고등어 난자치어 분포특성 및 수송과정

667

의산란시기는

3-6

,

4-5

(Cha et al., 2002),

15-23°C

(NIFS, 2017a).

0.9-1.3 mm

(Hwang et al., 2008; Kim et al., 2008; Jung et al., 2013; Lee et al., 2016).

25 m

,

50 m

(Jung et al., 2013).

19-24°C

41-56

(Hunter and Kimbrell, 1980; Hwang et al., 2008; Kim et al., 2008; Jung et al., 2013),

33-35 psu

90%

(Hwang et al., 2008).

(total length, TL)

2.5-3.2 mm

(Kim et al., 2008;

Park et al., 2008).

7-8

10-11

(Choi, 2003).

고등어난과자어등의초기생활사단계는해양환경변화에 취약하기때문에매우높은사망률을나타내며

,

(Kim et al., 1999;

Sassa and Tsukamoto, 2010).

,

∙

(Sassa and

Konishi, 2006).

거스를수있는유영능력이미약하기때문에주로해류의흐름 에의해그분포가결정된다

.

,

(Kim et al., 2011).

(Tsushima Warm Current)

(Kuroshio)

,

,

(Isobe et al., 2002; Lee et al., 2016).

(Yellow Sea Warm Current)

,

(Jeju Warm Current)

.

,

.

·

.

·

·

(individual based models, IBM)

,

·

,

.

,

어유생의분포특성재현과가입변동성연구에활용될수있는 가능성을제시한바

(Kim et al., 2015)

.

,

(population connectiv- ity)

(Stockhausen et al., 2019a; Stockhausen et al., 2019b; Gibson et al., 2019).

최근동중국해남부해역에서의대규모난

·

(Sassa and Konishi, 2006; Sassa et al., 2008; Sassa and Tsukamoto, 2010).

·

(Lee et al., 2006; Lee et al., 2016).

, Lee et al. (2016)

,

0 2 4 6

Southern East Sea

Southern East Sea

South Sea

South Sea

Northern East China Sea

Northern East China SeaSouthern East SeaSouth SeaNorthern East China Sea

122°

38°

36°

34°

32°

30° 124° 126° 128° 130° 132°

122°

38°

36°

34°

32°

30° 124° 126° 128° 130° 132°

자치어분포특성및수송과정을이해하기에는채집해역이제 한적이었고

,

·

.

,

(

,

)

.

,

·

2016

2017

(5-6

)

,

∙

(Fig. 1),

(

,

)

.

3

,

,

.

재료 및 방법

연근해 고등어 난∙자치어 조사

국립수산과학원 자원조사전용선인 탐구

20

(890

), 21

(990

)

2016

5, 6

(

130, 248

), 2017

5, 6

(

115

)

(

)

,

·

4

(Fig. 1, Table 1).

고등어의난과자

∙

(Ø80 cm,

335 μm

500 μm)

(Hydro Bios co., Altenholz, Germany)

.

2 knot

2016

년조사는난과자

∙

0-100 m

15

, 2017

분포를고려하여수심약

25 m

15

하였다

(Table 1).

하기위해난자치어조사시매정점에서

CTD (conductivity- temperature-depth; Sea-bird 911plus, Sea-Bird Electoronics Inc, Washington, USA)

100 m

.

난과 자어의 샘플처리 및 동정

채집된시료는즉시

10%

의형태를고정하였고

, 2-4

94% ethyl alcohol

.

Kim et al. (2011)

,

0.001 mm

(body length, BL)

.

5-6 mm

(total length)

(Kim et al., 2008; Park et al., 2015)

,

5 mm

(BL<5 mm)

‘

(small larva)’, 5 mm

(BL≥5 mm)

‘

(large larva)’

(Fig. 2).

,

,

,

,

색소포의 유무등형태적인특징을고려한

1

(sorting)

을실시하였고

,

(NIFS, 2017b).

total DNA (de- oxyribonucleic acid)

accuPrep genomic DNA extraction kit (Bioneer, Daejeon, Korea)

10% QIAquick gel extraction kit (Qiagen, Hilden, Germany)

, uni- versal primer

mtDNA COI, 12s rRNA, 16S rRNA

control region

PCR (polymerase chain re-

action)

.

열은

NCBI (National center for biotechnology information)

blast

.

난과 자어의 분포밀도 및 해양환경과의 관계

조사해역에서의난과자어의분포밀도는경사채집을실시한

2016

(ind./10 m

)

,

2017

(ind./1000 m

)

.

(Kim et al., 2005),

.

고등어난은주로표층

25 m

으며

(Jung et al., 2013),

·

수심

100 m

0-50 m

포한다

(Watanabe, 1970; Morser and Smith, 1993).

·

, CTD

25

Table 1. Summary of surveys for collecting egg and larvae of Scomber japonicus during 2016-2017

Year Date Research

vessel Sampling type

(Bongo net) Sampling

stations Positive stations (Ratio of positive stations, %)

Egg Larvae

2016 May 12-31 Tamgu20 Oblique tow (0-100 m) 130 8 (6.2) 14 (10.8)

2016 June 07-24 Tamgu20, 21 Oblique tow (0-100 m) 248 7 (2.8) 39 (15.7)

2017 May 16-28 Tamgu21 Surface tow (0-30 m) 115 17 (14.8) 17 (14.8)

2017 June 16-29 Tamgu20 Surface tow (0-30 m) 115 15 (13.0) 7 (6.1)

m

.

고등어초기생활사단계에서서식적합환경을추정하기위해 각정점의수심

25 m

Quotient

(Lluch-Belda et al., 1991; Ibaibarriaga et al., 2007; Ale- many et al., 2010; Sassa and Tsukamoto, 2010)

.

Q

.

Q

= A

/∑

A

N

/∑

N

여기서

A

(

) i

(

1°C,

0.25 psu

)

(ind./1000 m

), N

i

. Q

1

(Q

>1),

. 자어 체장 분포 및 해역별 체장 비교

2016-2017

5

6

(

)

.

(

)

,

(˚Ν)

(31˚75’-33˚50’Ν),

(33˚75’-34˚75’Ν),

(35˚-36˚50’Ν)

,

(Fig. 1). 5

6

T-test

(P<0.05)

,

별로채집된자어의체장차이를분석하기위해

one-way ANO-

VA test

(P<0.05)

. 해수유동모델

해양순환모델인

ROMS (regional ocean modeling system;

Shchepetkin and Mcwilliams, 2005)

하였다

. ROMS

,

,

기본방정식을계산하는

3

.

(orthogonal-curvilinear coordinate)

,

Arakawa-C

.

S-

(Stretched terrain-following coordinate)

.

18.5-48.5˚N,

117.5-155.5˚E

.

334×332

,

0.1˚×0.1˚

.

20

(level)

.

1

(Sungkyunkwan University’s 1 min digital bathymetric data;

Choi et al., 2002)

ETOPO5 (5’

)

,

5 m

.

영역의남쪽과동쪽은개방경계

(open boundaries)

고

, HYCOM

,

,

,

.

ECMWF (European center for medium range weather forecasting)

를 사용했다

(ECMWF, 2019).

30

spin-up

.

ROMS

,

(

,

Stockhausen and Hermann, 2007); BL, body length.

염분

,

)

(Kim et al., 2014; Kim et al., 2015),

2016-2017

,

,

. 개체기반모델(Individual based model, IBM)

본 연구에서는 미해양대기청

(National oceanic and atmo- spheric administration, NOAA)

DisMELS (dispersal model for early life stages; Stockhausen

and Hermann, 2007)

부터고등어의산란장을역추정하였으며

,

. DisMELS

(

·

,

,

)

생물학적모델과해양순환모형

ROMS

-

물리접합모델

(coupled bio-physical model)

(Kim et al., 2015; Stockhausen et al., 2019a; Stockhausen et al., 2019b; Gibson et al., 2019).

본개체기반모델에서는고등어의초기생활사단계

(life stage)

,

,

3

,

(stage duration)

2

, 10

, 20

(Hunter and Kimbrell, 1980; Hwang et al., 2008; Park et al., 2008; Kim et al., 2008;

Jung et al., 2013)(Fig. 2).

0-25 m,

(

,

)

0-50 m

(Kramer, 1960;

Watanabe, 1970; Moser and Smith, 1993; Tsukamoto et al., 2001, Jung et al., 2013).

(vertical swim- ming speed)

0 m/s

,

(

,

)

0.01 m/s, 0.01 m/s

(Visser, 1997; Stockhausen et al., 2019b).

random-walk process

·

,

(Stockhausen and Her- mann, 2007).

(

)

(advection)

(diffusion)

.

산란장 추정과 초기 수송과정 실험

연근해고등어의잠재적인산란장을추정하기위해난과자어 가채집된정점에서각개체

(

)

,

,

2

, 10

, 20

backward

(Fig.

9, Fig.10).

backward

(

)

2016-2017

5

1

15

, 6

1

15

,

30

일동안난과자어의수송과정을실험하였다

(Fig. 11, Fig. 12).

또한고등어난과자어단계의적정서식수온을바탕으로초기 수송과정에서의생존및가입과정을모의하기위해

2016

2017

30

.

5

6

5

1

6

30

10

,

30

.

한난의부화

,

(Hunter and

Kimbrell, 1980)

Quotient

13-23°C

,

어난개체들은사망한것으로간주하였다

.

결 과

고등어 난의 분자동정 결과

2016

423

166

(39.2%)

.

5

186

146

(78.4%)

,

,

11

. 6

237

20

(8.4%)

,

,

,

37

. 2017

1,267

662

(52.2%)

.

5

423

221

(52.2%)

,

,

,

10

. 6

844

441

(52.2%)

,

,

,

20

.

고등어 난의 분포특성

2016-2017

5-6

4

·

4,853

. 2016

5

25 m

13.3-17.5°C

.

현정점에서의난밀도는

3-210 ind./10 m

(Fig. 3).

2016

6

15.9-22.4°C

,

6-90 ind./10 m

.

.

한편

, 2017

5

12.2-22.2°C

,

1-133 ind./1000 m

. 2017

6

15.7-19.1°C

,

1-5095 ind./1000 m

.

5

25 m

12.2-22.2°C

,

,

.

6

15.0-22.8°C

,

밀도분포특성을보였다

. 고등어 자어의 월별 분포특성

2016-2017

5-6

4

339

. 2016

5

,

25 m

15.5-20.0°C

-

,

.

2-13 ind./10 m

(Fig. 4).

,

-

. 2016

6

,

9.4-

22.9°C

,

부해역에서채집되었다

.

1-61 ind./10 m

5

.

,

. 2017

5

15.2-22.2°C

주로출현하였다

.

0.4-8 ind./1000 m

.

,

. 2017

6

,

,

14.5-21.2°C

범위였다

.

1-11 ind./1000 m

.

,

.

전체적으로

5

12.7-22.2°C

-

,

.

,

-

. 6

13.2-22.9°C

,

.

,

.

고등어 자어의 월별 체장 변화

2016-2017

사하기위하여월별로채집된자어의체장빈도분포표를나타 내었다

(Fig. 5). 2016

5

2.99- 9.78 mm,

5.97 mm (SD=1.57 mm)

, 4-6 mm

. 2016

6

1.92-10.23 mm,

4.59 mm (SD=1.82 mm)

,

(Fig. 5).

2016

5

2016

6

된자어의평균체장보다약

1.38 mm

(T-test, P<0.05). 2017

5

2.35-12.13 mm,

5.97 mm (SD=2.12 mm)

4-6 mm

높은빈도를나타냈다

. 2017

6

는

2.04-10.08 mm,

4.06 mm (SD=2.56 mm)

,

작은크기에서높은빈도를나타냈다

. 2017

5

된자어의평균체장이

6

1.91 mm

(T-test, P<0.05). 2016-2017

5

6

, 5

6

(

2-3 mm)

.

자어의 출현 해역에 따른 체장 특성

위도에따른출현해역별로채집된자어의체장특성을비교하 기위해조사해역을동해남부

,

,

,

(Fig. 6).

2016-2017

5

위는

2.54-6.54 mm,

4.35 mm (SD=1.05)

,

6.47-12.13 mm,

8.69 mm (SD=2.12)

로나타났다

(Fig. 6).

2.35-

11.17 mm,

6.14 mm (SD=1.68)

(Fig. 6). 5

이를보였고

(One-way ANOVA, P<0.05),

을보였다

. 2016-2017

6

어의체장범위는

1.92-8.85 mm,

4.11 mm (SD=1.57)

,

2.56-10.23 mm,

5.94 mm (SD=1.78)

(Fig. 6).

1.97-6.53 mm,

3.31 mm (SD=1.19)

나타났다

(Fig. 6). 6

평균체장이통계적으로유의한차이를나타냈으며

(One-way ANOVA, P<0.05),

,

.

초기생활사 단계에서의 서식지 환경특성

조사해역에서난과자어가서식하기적합한환경특성을추정 하기위하여

Quotient

(Fig. 7, Fig. 8).

(2016-2017

5

, 6

)

7.4-24.0°C

, 15-16°C

40%

(Fig. 7). Quotient

18-20°C

Q

1

.

18-22°C

Q

1

.

,

13-14°C, 18-22°C

Q

1