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First Reliable Record of the Dusky Shark Carcharhinus obscurus (Lesueur, 1818) (Carcharhinidae, Pisces) from Korea

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696

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

서 론

흉상어목

(Carcharhiniformes)

흉상어과

(Carcharhinidae)

류는전계적으로

12

60

(White et al., 2019),

우리나라에

5

12

종이알려져있다

(MABIK, 2018).

흉상어과어류의주요 분류형질로주둥이

,

이빨

,

눈의모양

,

지느러미의크기와위치 등이알려져있다

(Compagno, 1984).

흉상어과어류에속하는 흉상어속

(Carcharhinus)

어류에는 전세계적으로

36

(White et al., 2019),

우리나라에

6

[

무태상어

(Carcharhinus brachy- urus),

흰뺨상어

(Carcharhinus dussumieri),

남방상어

(Carcha- rhinus limbatus),

흑상어

(

가칭

) (Carcharhinus obscurus),

상어

(Carcharhinus plumbeus),

검은꼬리상어

(Carcharhinus sorrah)]

보고되어 있다

(MABIK, 2018).

흉상어속 어류는 주둥이가둥글고눈이원형에가까우며

,

2

등지느러미가 작고꼬리자루에융기선이없는특징을가진다

(White et al.,

2019).

연구는우리나라에출현기록만있을아직표본에

근거하여보고된없는

Carcharhinus obscurus 5

개체를대상 으로상세한형태분자특징을제공하고자한다

.

재료 및 방법

연구에서사용된흉상어속어류

5

개체는

2010

9

경상 남도남해군

, 2013

10

강원도삼척시

, 2014

10

전라남 신안군

, 2015

1

제주특별자치도제주시

, 2019

9

산광역시에서채집되었다

(Fig. 1).

계수계측은

Compagno (1984), Voigt and Weber (2011)

방법을따랐으며

,

줄자

vernia caliper

이용하여

0.1 mm

단위까지측정하였고

, 2

(PKU 10234, PKU 11402)

경우꼬리지느러미앞까지 추골수를

X-ray (Sehwa Medical System SMS CM, Gwangju, Korea)

계수하였다

.

부위의 명칭은

Compagno (1984), Ebert et al. (2013), Choi (2016)

따랐다

.

표본은실험실로 반하여사진촬영측정을마친

15% formalin

30

일간

한국산 흉상어과(Carcharhinidae) 어류 1미기록종, Carcharhinus ob- scurus (Lesueur, 1818)

이우준·김윤지

1

·나정현

1

·김진구*

부경대학교 자원생물학과, 1오송첨단의료산업진흥재단 신약개발지원센터

First Reliable Record of the Dusky Shark Carcharhinus obscurus (Lesueur, 1818) (Carcharhinidae, Pisces) from Korea

Woo Jun Lee, Yoonji Kim

1

, Jung-Hyun Na

1

and Jin-Koo Kim*

Department of Marine Biology, Pukyong National University, Busan 48513, Korea

1 New Drug Development Center, Osong Medical Innovation Foundation (Kbio health), Chungbuk 28160, Korea

Five specimens (839 - 1280 mm TL, Total length) of the dusky shark Carcharhinus obscurus , belonging to the fam- ily Carcharhinidae, were collected from the waters off Jejudo Island, Sinan, Namhae, Busan and Gangneung, Korea, during 2010-2019. Carcharhinus obscurus was similar to Carcharhinus brachyurus but was distinguished by the interdorsal ridge (present in C. obscurus vs. absent in C. brachyurus ) and the shape of upper jaw teeth (broad in C.

obscurus vs. narrow in C. brachyurus ). In addition, 479 base-pair sequences in the mitochondrial DNA cytochrome c oxidase subunit I (COI) of our two specimens corresponded to those of C. obscurus (genetic distance, 0.000-0.003), but clearly distinguished from those of the Korean C. brachyurus (genetic distance, 0.03). We adopted the Korean name “Heuk-sang-eo” for Carcharhinus obscurus , after Kim and Ryu (2017).

Key words: New record, Shark, Carcharhinidae, Carcharhinus obscurus , Korea

*Corresponding author: Tel: +82. 51. 629. 5927 Fax: +82. 51. 629. 5931 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 29 October 2019; Revised 18 November 2019; Accepted 27 November 2019 저자 직위: 이우준(대학원생), 김윤지(연구원), 나정현(연구원), 김진구(교수) https://doi.org/10.5657/KFAS.2019.0696

Korean J Fish Aquat Sci 52(6), 696-701, December 2019

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한국산 흉상어과 어류 1미기록종, 흑상어 Carcharhinus obscurus

697

정시킨

,

최종적으로

70% ethyl alcohol

담아보존하였다

.

측정이끝난표본은부경대학교

(Pukyoung National Univer- sity, PKU)

등록

,

보관하였다

.

분자분석을 위해

2

개체

(PKU 10234, PKU 17878)

체측 근육에서

Chelex 100 resin (Bio-rad, USA)

사용하여

ge- nomic DNA

추출하였다

.

미토콘드리아

DNA cytochrome c oxidase subunit I

VF2 (5´-TCAACCAACCACAAAGA- CATTGGCAC-3´)

FishR2 (5´-ACTTCAGGGTGAC- CGAAGAATCAGAA-3´) universal primer

증폭하였다

(Ivanova et al., 2007).

이후

,

염기서열은

BioEdit ver. 7 (Hall, 1999)

Clustal W (Thompson et al., 1994)

사용하여정렬하 였으며

,

근린결합수

(neighbor-joining tree)

Mega 7 (Kumar et al., 2016)

Kimura two-parameter model (Kimura, 1980)

이용하여작성하였다

.

근린결합수의신뢰도는

bootstrap

법으로

1,000

수행하여구하였다

.

결 과

Carcharhinus obscurus (Lesueur, 1818) (Korean name: Heuk-sang-eo) (Fig. 2A)

Squalus obscurus Lesueur, 1818: 223 (type locality: North America).

Carcharhinus obscurus: Garrick, 1982: 120; Compagno, 1984: 489; Masuda et al., 1984: 6; Voigt and Weber 2011: 84;

Ebert et al., 2013: 472; Aonuma et al., 2013: 176; Kim and Ryu, 2017: 43; MABIK, 2018: 22.

관찰표본

PKU 4211, 1

개체

,

전장

839 mm,

경상남도남해군상주면

, 2010

9

5

; PKU 10234, 1

개체

,

전장

1088 mm,

강원도 삼척시

, 2013

10

26

,

정치망

; PKU 11402, 1

개체

,

전장

1135 mm,

전라남도신안군흑산면

, 2014

10

1

,

안강망

; PKU 53073, 1

개체

,

전장

1280 mm,

제주특별자치도제주시 추자면

, 2015

1

5

,

선망

; PKU 17878, 1

개체

,

전장

1020 mm,

부산광역시해운대구

, 2019

9

26

,

자망

.

비교표본

무태상어

Carcharhinus brachyurus PKU 10860, 1

개체전장

720 mm (Fig. 2B),

경상남도거제시

,

선망

.

기재

몸은길고방추형이다

.

주둥이는둥글고넙적하다

.

콧구멍은 주둥이와입의사이중간에위치하며

,

콧구멍의전방콧구멍덮 개는작다

.

눈은원형에가깝고아래쪽에흰색순막이있다

.

아치형으로길다

.

위턱의이빨은넙적하며약간구부러져 가장자리가톱니모양을하고있는반면

,

아래턱의이빨은 Fig. 1. Map showing the sampling sites of Carcharhinus obscurus

(●).

Fig. 2. Carcharhinus obscurus, PKU 17878, female, 1020 mm TL, Busan, Korea (A), and Carcharhinus brachyurus, PKU 10860, fe- male, 720 mm TL, Geoje, Korea (B). TL, total length.

(A)

(B)

(A)

(B)

(A)

(B)

Carcharhinus obscurus (MN723160)

PKU 17878

Carcharhinus obscurus (MG811817)

SF 512

Carcharhinus obscurus (MN723159)

PKU 10234

Carcharhinus brachyurus (KX379525)

PKU 10860

Squalus suckleyi (KX379519)

PKU 10244 99

91

0.020

(3)

이우준

김윤지

나정현

김진구

698

늘고길며가장자리가미세한톱니모양을보인다

(Fig. 3A).

수공은흔적적이다

.

새열은

5

쌍이줄지어있고

,

모두비슷한 기이다

.

가슴지느러미기점은

4

번째새열에위치하며모서리가 둥근삼각형으로길고크다

(

전장의

12.5-13.9%).

1

등지느러 미는가슴지느러미기저의말단보다뒤쪽에위치하며모서리가 둥근삼각형이다

.

배지느러미는몸의중앙보다뒤쪽에위치하 크기가작다

(

전장의

5.2-5.6%).

2

등지느러미는꼬리자루 가깝게위치하며매우작다

(

전장의

2.0-2.7%).

뒷지느러미 배지느러미와꼬리지느러미사이중간에위치한다

.

등지느 러미앞에는가시가없다

.

등지느러미사이에융기선이있다

(Fig. 4A).

꼬리자루의측면에융기선이없다

.

꼬리지느러미 엽은하엽에비해길며꼬리지느러미말단각이있고말단은 족하다

.

꼬리지느러미앞까지척추골수는

93-94

개이다

(Fig. 5).

몸은전체적으로방패비늘로덮여있다

. 체색

신선할

,

머리와몸의등쪽은짙은회색또는회갈색을

,

측면 옅은회색또는옅은회갈색을

,

배쪽은흰색을띤다

.

가슴지 느러미

,

1

등지느러미

,

2

등지느러미

,

배지느러미

,

꼬리지느 러미는회색회갈색을띠며뒷지느러미는흰색을띤다

.

포르

말린고정후에도신선할때의체색이유지되었다

. 분자분석

연구에서흉상어속

2

개체

(PKU 10234, PKU 17878)

미토콘드리아

DNA cytochrome c oxidase subunit I (COI)

기서열

479 bp

NCBI

등록하였다

(

등록번호

, MN723159- MN723160).

흉상어속

2

개체의염기서열은호주서부에서 집된

C. obscurus (MG 811817)

99.7-100%

유전적으로일치 하였으며

,

한국산무태상어

(C. brachyurus, PKU 10860)

와는 Fig. 3. Teeth of Carcharhinus obscurus, PKU 10234 (A) and Car-

charhinus brachyurus, PKU 10860 (B).

(A)

(B)

(A)

(B)

(A)

(B)

Carcharhinus obscurus (MN723160)

PKU 17878

Carcharhinus obscurus (MG811817)

SF 512

Carcharhinus obscurus (MN723159)

PKU 10234

Carcharhinus brachyurus (KX379525)

PKU 10860

Squalus suckleyi (KX379519)

PKU 10244 99

91

0.020

Fig. 4. Photos showing interdorsal ridge of Carcharhinus obscurus, PKU 10234 (A) and Carcharhinus brachyurus, PKU 10860 (B).

(B)

(A)

(B)

(A)

(B)

Carcharhinus obscurus (MN723160)

PKU 17878

Carcharhinus obscurus (MG811817)

SF 512

Carcharhinus obscurus (MN723159)

PKU 10234

Carcharhinus brachyurus (KX379525)

PKU 10860

Squalus suckleyi (KX379519)

PKU 10244 99

91

0.020

Fig. 5. X-ray photo showing vertebrae of Carcharhinus obscurus, PKU 10234.

(4)

한국산 흉상어과 어류 1미기록종, 흑상어 Carcharhinus obscurus

699

Table 1. Comparison of the morphological characters of Carcharhinus obscurus and Carcharhinus brachyurus

C. obscurus C. brachyurus

Present study Garrick, 1982 Present study Garrick, 1982

Number of specimens 5 11 1 10

Total length (mm) 839-1280 812-3200 720 660-2725

In % of total length

Snout tip to eye 8.1-9.7 5.9-8.1 9.3 5.9-8.2

mouth 7.2-7.9 5.9-8.4 7.3 5.7-8.4

1st gill-opening 18.5-20.4 17.9-20.3 19.7 17.0-19.1

3rd gill-opening 20.6-22.6 20.5-22.3 22.4 19.5-24.4

5th gill-opening 22.2-24.3 22.3-24.0 24.2 21.5-24.1

pectoral origin 21.2-22.4 20.9-22.7 24.5 21.7-25.2

pelvic origin 48.2-51.8 48.9-53.6 52.5 49.0-55.2

1st dorsal origin 31.0-32.8 29.3-32.9 32.2 31.2-34.0

2nd dorsal origin 62.1-65.0 61.6-65.8 63.9 62.1-67.3

anal fin origin 62.3-65.0 61.1-65.3 63.9 59.9-67.3

upper caudal origin 73.2-76.2 72.9-75.8 73.6 72.4-76.2

lower caudal origin 72.8-74.9 71.8-75.0 73.6 71.4-76.4

Mouth width 7.8-8.9 7.1-10.1 8.0 6.7-8.8

length 3.5-5.1 4.0-4.6 4.2 4.0-5.2

Labial furrow lengths upper 0.3-1.0 0.3-0.9 0.6 0.4-0.7

lower 0.3-0.7 0.4-0.9 0.5 0.3-0.7

Gill opening lengths 1st 2.4-4.1 2.1-3.6 3.1 2.4-3.5

3rd 3.2-4.3 2.7-4.0 3.2 2.5-4.1

5th 2.3-3.3 2.1-2.7 2.7 2.0-2.7

Eye horizontal diameter 1.7-2.2 1.0-2.1 2.2 1.1-2.2

1st dorsal fin length of base 8.6-9.2 8.9-11.7 9.1 8.9-10.8

height 8.1-9.7 6.0-9.7 6.9 6.8-9.7

2nd dorsal fin length of base 2.8-3.4 2.5-4.2 4.7 3.1-4.3

height 2.0-2.7 1.8-2.3 2.1 2.0-2.5

Anal fin length of base 3.6-3.9 3.5-4.6 4.3 3.8-5.4

height 2.7-3.7 2.4-3.8 3.5 2.9-3.7

Pectoral fin length of base 5.2-5.8 5.4-7.0 5.9 5.4-6.8

length distal margin 11.5-13.9 10.3-18.8 12.9 12.0-18.0

Pelvic fin length of base 4.2-4.8 4.2-5.8 4.6 4.2-6.2

length distal margin 5.2-5.6 4.3-6.3 5.3 4.7-6.0

Caudal fin length of upper lobe 25.5-27.8 25.8-27.5 25.0 25.3-27.8

length of lower lobe 10.6-11.7 10.8-12.4 9.9 10.4-13.4

Trunk at pectoral origin width 11.1-12.8 10.7-14.4 11.8 10.3-13.2

height 10.8-14.1 9.6-14.7 11.7 9.5-13.6

Vertebrae precaudal 93-94 (two specimens) 89-97 - 98-110

3%

차이를보였고

,

외집단으로사용된한국산곱상어

(Squalus

suckleyi)

와는

24.9-25.2%

차이로명확히구분되었다

(Fig. 6). 분포

한국전해역

(

연구

),

전세계의온대열대해역의

400 m

(5)

이우준

김윤지

나정현

김진구

700

심까지분포한다

(Compagno, 1984; Ebert et al., 2013).

고 찰

2010

년부터

2019

까지우리나라서해

,

남해

,

동해에서 집된흉상어과흉상어속에속하는

5

개체는등지느러미사이 융기선이뚜렷한

,

꼬리지느러미앞까지척추골이

93-94

개인등에근거하여

Carcharhinus obscurus

동정하였다

(Lesueur, 1818; Garrick, 1982; Compagno, 1984; Voigt and Weber, 2011; Aonuma et al., 2013; Ebert et al., 2013).

종은

Garrick (1982)

흉상어속의분류학적연구와비교하면주둥 이에서눈까지의길이

,

주둥이에서새열까지의길이

,

입의길이

,

입술주름길이

,

아가미길이

,

2

등지느러미높이

,

꼬리지느러 길이에서근소한차이를보이나대부분일치하였다

(Table

1).

종은한국에보고되어있는흉상어속어류가운데무태상

Carcharhinus brachyurus

형태적으로유사하나위턱의 모양

(C. obscurus

넙적하다

vs. C. brachyurus

가늘다

)

등지느러미사이의융기선의유무

(C. obscurus

뚜렷한 융기선을가진다

vs. C. brachyurus

융기선이없거나흔적적 이다

)

구분된다

.

또한

,

국내에는없으나형태적으로유사 동속의

C. galapagensis

와는꼬리지느러미앞까지척추골수

(C. obscurus

86-94

vs. C. galapagensis

103-109

)

구분된다

(Garrick, 1982; Ebert et al., 2013).

나아가

,

국산

C. obscurus 2

개체

(PKU 10234, PKU 17878)

미토콘드 리아

DNA COI

염기서열

479 bp

비교한결과

, NCBI

등록

C. obscurus (MG 811817)

99.7-100%

일치하였으며

, C.

brachyurus

와는

3%

차이를보여형태분류결과를지지해 주었다

. C. obscurus

국명으로

Kim and Ryu (2017)

제안

흑상어

따른다

.

사 사

논문은

2019

년도한국연구재단바이오의료기술개발사업

(NRF-2019M3A9G4067132)

2019

년도해양수산과학기술 진흥원해양수산생명공학기술개발사업

(No. 20170431)

지원을받아수행된연구입니다

.

논문의완성도를높여 세분심사위원께감사드립니다

.

References

Aonuma Y, Yamaguchi A, Yagishita N and Yoshino T. 2013.

Carcharhinidae. In: Fishes of Japan with Pictorial Keys to the Species. Nakabo T, ed. Tokai University Press, Tokyo, Japan, 171-176.

Choi Y. 2016. Sharks of Korea. National Marine Biodiversity Institute of Korea, Seocheon, Korea, 1-223.

Compagno LJV. 1984. FAO species catalogue. Vol.4, Sharks of the World. An annotated and illustrated catalogue of shark species known to date. FAO Fisheries Synopsis 125, 1-655.

Ebert DA, Fowler S and Compagno L. 2013. Sharks of the World. Wild Nature Press, Plymouth, U.K., 456-476.

Garrick JAF. 1982. Sharks of the genus Carcharhinus. NOAA Technical Report 445, 1-194.

Hall TA. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/

NT. Nucleic Acids Symp Ser 41, 95-98.

Ivanova NV, Zemlak TS, Hanner RH and Hebert PDN. 2007.

Universal primer cocktails for fish DNA barcoding. Mol Ecol Notes 7, 544-548. https://doi.org/10.1111/j.1471- 8286.2007.01748.x.

Kim JK and Ryu JH. 2017. Distribution map of sea fishes in Korea. Maple design, Busan, Korea, 43-47.

Kimura M. 1980. A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. J Mol Evol 16, 111-120. https://doi.

org/10.1007 /BF01731581.

Kumar S, Stecher G and Tamura K. 2016. MEGA7: molecular evolutionary genetics analysis version 7.0 for bigger datas- ets. Mol Biol Evol 33, 1870-1874. https://doi.org/10.1093/

molbev/ msw054.

Lesueur CA. 1818. Description of several new species of North American fishes. J Acad Nat Sci Philadelphia 1, 222-235.

MABIK (Marine Biodiversity Institute of Korea). 2018. Nation- al list of marine species. Namu Press, Seocheon, Korea, 21- Masuda H, Amaoka K, Araga C, Uyeno T and Yoshino T. 1984. 22.

The fishes of the Japanese archipelago. Tokai Univeristy Press, Tokyo, Japan, 5-7.

Thompson JD, Higgins DG and Gibson TJ. 1994. CLUSTALW:

improving the sensitivity of progressive multiple sequence Fig. 6. Neighbor-joining tree constructed by the mitochondrial

DNA COI sequences for two Carcharhinus species, with one out- group Squalus suckleyi. Parenthesis and superscript indicate NCBI registration number and voucher number, respectively. Numbers at branches indicate bootstrap probabilities in 1,000 bootstrap repli- cations. Bar indicates genetic distance of 0.02. COI, cytochrome c oxidase subunit I.

(B)

(A)

(B)

(A)

(B)

Carcharhinus obscurus (MN723160)

PKU 17878

Carcharhinus obscurus (MG811817)

SF 512

Carcharhinus obscurus (MN723159)

PKU 10234

Carcharhinus brachyurus (KX379525)

PKU 10860

Squalus suckleyi (KX379519)

PKU 10244 99

91

0.020

(6)

한국산 흉상어과 어류 1미기록종, 흑상어 Carcharhinus obscurus

701

alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucl Acids Res 22, 4673-4680. http://dx.doi.org/10.1093/nar/ 22.22.4673.

Voigt M and Weber D. 2011. Field guide for sharks of the genus

Carcharhinus. Verlag Dr. Friedrich Pfeil, Munchen, Ger-

many, 1-151.

White WT, Kyne PM and Harris M. 2019. Lost before found:

A new species of whaler shark Carcharhinus obsolerus from the Western Central Pacific known only from historic records. PloS ONE 14, e0209387. https://doi.org/10.1371/

journal.pone.0209387.

수치

Fig. 2. Carcharhinus obscurus, PKU 17878, female, 1020 mm TL,  Busan, Korea (A), and Carcharhinus brachyurus, PKU 10860,  fe-male, 720 mm TL, Geoje, Korea (B)
Fig. 5. X-ray photo showing vertebrae of Carcharhinus obscurus,  PKU 10234.
Table 1. Comparison of the morphological characters of Carcharhinus obscurus and Carcharhinus brachyurus

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