696
Copyright © 2019 The Korean Society of Fisheries and Aquatic Science pISSN:0374-8111, eISSN:2287-8815
서 론
흉상어목
(Carcharhiniformes)
흉상어과(Carcharhinidae)
어 류는전계적으로12
속60
종(White et al., 2019),
우리나라에5
속12
종이알려져있다(MABIK, 2018).
흉상어과어류의주요 분류형질로주둥이,
이빨,
눈의모양,
지느러미의크기와위치 등이알려져있다(Compagno, 1984).
흉상어과어류에속하는 흉상어속(Carcharhinus)
어류에는 전세계적으로36
종(White et al., 2019),
우리나라에6
종[
무태상어(Carcharhinus brachy- urus),
흰뺨상어(Carcharhinus dussumieri),
남방상어(Carcha- rhinus limbatus),
흑상어(
가칭) (Carcharhinus obscurus),
흉 상어(Carcharhinus plumbeus),
검은꼬리상어(Carcharhinus sorrah)]
이보고되어 있다(MABIK, 2018).
흉상어속 어류는 주둥이가둥글고눈이원형에가까우며,
제2
등지느러미가매 우작고꼬리자루에융기선이없는특징을가진다(White et al.,
2019).
본연구는우리나라에출현기록만있을뿐아직표본에근거하여보고된적없는
Carcharhinus obscurus 5
개체를대상 으로상세한형태및분자특징을제공하고자한다.
재료 및 방법
본연구에서사용된흉상어속어류
5
개체는2010
년9
월경상 남도남해군, 2013
년10
월강원도삼척시, 2014
년10
월전라남 도신안군, 2015
년1
월제주특별자치도제주시, 2019
년9
월부 산광역시에서채집되었다(Fig. 1).
계수및계측은Compagno (1984), Voigt and Weber (2011)
의방법을따랐으며,
줄자및vernia caliper
를이용하여0.1 mm
단위까지측정하였고, 2
개 체(PKU 10234, PKU 11402)
의경우꼬리지느러미앞까지척 추골수를X-ray (Sehwa Medical System SMS CM, Gwangju, Korea)
로 계수하였다.
각부위의 명칭은Compagno (1984), Ebert et al. (2013), Choi (2016)
를따랐다.
표본은실험실로운 반하여사진촬영및측정을마친후15% formalin
에30
일간고한국산 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 Na1
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
한국산 흉상어과 어류 1미기록종, 흑상어 Carcharhinus obscurus
697
정시킨뒤
,
최종적으로70% ethyl alcohol
에담아보존하였다.
측정이끝난표본은부경대학교(Pukyoung National Univer- sity, PKU)
에등록,
보관하였다.
분자분석을 위해
2
개체(PKU 10234, PKU 17878)
의 체측 근육에서Chelex 100 resin (Bio-rad, USA)
을 사용하여ge- nomic DNA
를추출하였다.
미토콘드리아DNA cytochrome c oxidase subunit I
을VF2 (5´-TCAACCAACCACAAAGA- CATTGGCAC-3´)
및FishR2 (5´-ACTTCAGGGTGAC- CGAAGAATCAGAA-3´) universal primer
로 증폭하였다(Ivanova et al., 2007).
이후,
염기서열은BioEdit ver. 7 (Hall, 1999)
의Clustal W (Thompson et al., 1994)
를사용하여정렬하 였으며,
근린결합수(neighbor-joining tree)
는Mega 7 (Kumar et al., 2016)
의Kimura two-parameter model (Kimura, 1980)
을이용하여작성하였다.
근린결합수의신뢰도는bootstrap
방 법으로1,000
번수행하여구하였다.
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,
전장839 mm,
경상남도남해군상주면, 2010
년9
월5
일; PKU 10234, 1
개체,
전장1088 mm,
강원도 삼척시, 2013
년10
월26
일,
정치망; PKU 11402, 1
개체,
전장1135 mm,
전라남도신안군흑산면, 2014
년10
월1
일,
안강망; PKU 53073, 1
개체,
전장1280 mm,
제주특별자치도제주시 추자면, 2015
년1
월5
일,
선망; PKU 17878, 1
개체,
전장1020 mm,
부산광역시해운대구, 2019
년9
월26
일,
자망.
비교표본
무태상어
Carcharhinus brachyurus PKU 10860, 1
개체전장720 mm (Fig. 2B),
경상남도거제시,
선망.
기재
몸은길고방추형이다
.
주둥이는둥글고넙적하다.
콧구멍은 주둥이와입의사이중간에위치하며,
콧구멍의전방콧구멍덮 개는작다.
눈은원형에가깝고아래쪽에흰색순막이있다.
입 은아치형으로길다.
위턱의이빨은넙적하며약간구부러져있 고가장자리가톱니모양을하고있는반면,
아래턱의이빨은가 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 17878Carcharhinus obscurus (MG811817)
SF 512Carcharhinus obscurus (MN723159)
PKU 10234Carcharhinus brachyurus (KX379525)
PKU 10860Squalus suckleyi (KX379519)
PKU 10244 9991
0.020
이우준
ㆍ
김윤지ㆍ
나정현ㆍ
김진구698
늘고길며가장자리가미세한톱니모양을보인다
(Fig. 3A).
분 수공은흔적적이다.
새열은5
쌍이줄지어있고,
모두비슷한크 기이다.
가슴지느러미기점은4
번째새열에위치하며모서리가 둥근삼각형으로길고크다(
전장의12.5-13.9%).
제1
등지느러 미는가슴지느러미기저의말단보다뒤쪽에위치하며모서리가 둥근삼각형이다.
배지느러미는몸의중앙보다뒤쪽에위치하 며크기가작다(
전장의5.2-5.6%).
제2
등지느러미는꼬리자루 에가깝게위치하며매우작다(
전장의2.0-2.7%).
뒷지느러미 는배지느러미와꼬리지느러미사이중간에위치한다.
등지느 러미앞에는가시가없다.
양등지느러미사이에융기선이있다(Fig. 4A).
꼬리자루의측면에융기선이없다.
꼬리지느러미상 엽은하엽에비해길며꼬리지느러미말단각이있고말단은뾰 족하다.
꼬리지느러미앞까지척추골수는93-94
개이다(Fig. 5).
몸은전체적으로방패비늘로덮여있다
. 체색
신선할때
,
머리와몸의등쪽은짙은회색또는회갈색을,
측면 은옅은회색또는옅은회갈색을,
배쪽은흰색을띤다.
가슴지 느러미,
제1
등지느러미,
제2
등지느러미,
배지느러미,
꼬리지느 러미는회색및회갈색을띠며뒷지느러미는흰색을띤다.
포르말린고정후에도신선할때의체색이유지되었다
. 분자분석
본 연구에서흉상어속
2
개체(PKU 10234, PKU 17878)
의 미토콘드리아DNA cytochrome c oxidase subunit I (COI)
염 기서열479 bp
는NCBI
에등록하였다(
등록번호, MN723159- MN723160).
흉상어속2
개체의염기서열은호주서부에서채 집된C. obscurus (MG 811817)
와99.7-100%
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 17878Carcharhinus obscurus (MG811817)
SF 512Carcharhinus obscurus (MN723159)
PKU 10234Carcharhinus brachyurus (KX379525)
PKU 10860Squalus suckleyi (KX379519)
PKU 10244 9991
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 17878Carcharhinus obscurus (MG811817)
SF 512Carcharhinus obscurus (MN723159)
PKU 10234Carcharhinus brachyurus (KX379525)
PKU 10860Squalus suckleyi (KX379519)
PKU 10244 9991
0.020
Fig. 5. X-ray photo showing vertebrae of Carcharhinus obscurus, PKU 10234.
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)
와는24.9-25.2%
차이로명확히구분되었다(Fig. 6). 분포
한국전해역
(
본연구),
전세계의온대및열대해역의400 m
수이우준
ㆍ
김윤지ㆍ
나정현ㆍ
김진구700
심까지분포한다
(Compagno, 1984; Ebert et al., 2013).
고 찰
2010
년부터2019
년까지우리나라서해,
남해,
동해에서채 집된흉상어과흉상어속에속하는5
개체는양등지느러미사이 에융기선이뚜렷한점,
꼬리지느러미앞까지척추골이93-94
개인점등에근거하여Carcharhinus obscurus
로동정하였다(Lesueur, 1818; Garrick, 1982; Compagno, 1984; Voigt and Weber, 2011; Aonuma et al., 2013; Ebert et al., 2013).
본종은Garrick (1982)
의흉상어속의분류학적연구와비교하면주둥 이에서눈까지의길이,
주둥이에서새열까지의길이,
입의길이,
입술주름길이,
아가미길이,
제2
등지느러미높이,
꼬리지느러 미길이에서근소한차이를보이나대부분일치하였다(Table
1).
본종은한국에보고되어있는흉상어속어류가운데무태상어
Carcharhinus brachyurus
와형태적으로유사하나위턱의이 빨모양(C. obscurus
는넙적하다vs. C. brachyurus
는가늘다)
와양등지느러미사이의융기선의유무(C. obscurus
는뚜렷한 융기선을가진다vs. C. brachyurus
는융기선이없거나흔적적 이다)
로잘구분된다.
또한,
국내에는없으나형태적으로유사 한동속의C. galapagensis
와는꼬리지느러미앞까지척추골수(C. obscurus
는86-94
개vs. C. galapagensis
는103-109
개)
에 서잘구분된다(Garrick, 1982; Ebert et al., 2013).
나아가,
한 국산C. obscurus 2
개체(PKU 10234, PKU 17878)
의미토콘드 리아DNA COI
염기서열479 bp
을비교한결과, NCBI
에등록 된C. obscurus (MG 811817)
와99.7-100%
일치하였으며, C.
brachyurus
와는3%
차이를보여형태분류결과를잘지지해 주었다. C. obscurus
의국명으로Kim and Ryu (2017)
가제안한
“
흑상어”
를따른다.
사 사
이논문은
2019
년도한국연구재단바이오의료기술개발사업(NRF-2019M3A9G4067132)
및2019
년도해양수산과학기술 진흥원해양수산생명공학기술개발사업(No. 20170431)
의예 산지원을받아수행된연구입니다.
이논문의완성도를높여주 신세분심사위원께감사드립니다.
References
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(B)
(A)
(B)
(A)
(B)
Carcharhinus obscurus (MN723160)
PKU 17878Carcharhinus obscurus (MG811817)
SF 512Carcharhinus obscurus (MN723159)
PKU 10234Carcharhinus brachyurus (KX379525)
PKU 10860Squalus suckleyi (KX379519)
PKU 10244 9991
0.020
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.
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