한국고유종북방종개의연령및산란기특징 73 Fig. 1. Frequency distribution of the total length of Iksookimia pacifica in the Bukcheon (stream), Ganseong-eup, Gangwon-do, Ko
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1 72 Korean 고명훈 Journal 원용진of Ichthyology, Vol. 28, No. 2, 72-78, June 2016 ISSN: (Print), (Online) Received: June 11, 2016 Revised: June 22, 2016 Accepted: June 27, 2016 한국고유종북방종개 Iksookimia pacifica (Pisces: Cobitidae) 의연령및산란기특징 고명훈 원용진 * 이화여자대학교에코과학부 The Age and Spawning Characteristics of the Korean Endemic Spine Loach, Iksookimia pacifica (Pisces: Cobitidae) in the Bukcheon (stream), Korea by Myeong-Hun Ko and Yong-Jin Won* (Division of EcoScience, Ewha Womans University, Seoul 03760, Korea) ABSTRACT The age and spawning characteristics of Iksookimia pacifica were investigated in this study to provide baseline data for its ecological characteristics in the Bukcheon (stream), Goseonggun, Gangwon-do, Korea from January to December, Age groups of I. pacifica (female) estimated by the frequency distribution of total length in the spawning season indicated that the 30~49 mm group is 1 year old, the 50~74 mm group is 2 years old, the 75~99 mm group is 3 years old, the 100~124 mm group is 4 years old, and the 130~160 mm group is 5 years old. The sex ratio ( / ) was 0.63, and females were 30~60 mm larger than males. The lamina circularis at the base of pectoral fins in males as a secondary sexual character was created at the 13th month after hatching. The spawning season according to the gonadosomatic index (GSI) was July to August (main spawning season in July) with water temperatures of 20~25. The average number of eggs was 2,503±1,337 with a diameter of 1.11±0.04 mm in mature eggs. Key words: Spine loach, Iksookimia pacifica, age groups, spawning characteristics 서 미꾸리과 (Cobitidae) 어류는잉어목 (Cypriniformes) 에속하 며유라시아와아프리카북부에널리서식하는담수저서성어류로 26속 177종이보고되었고 (Nelson, 2006), 우리나라에는 5속 16종이서식한다 (Kim, 2009). 이들은우리나라전역의하천에서식하고있는데, 이소적종분화를통하여분화하였기때문에지역적고유성및높은고유성 (16종중 13종이고유종 ) 을가지며독특한생태적특징을갖는것으로보고되고있다 (Kim, 1997; Kim and Park, 2007; Kim, 2009). 북방종개 Iksookimia pacifica는 Kim et al. (1999) 에의해 Cobitis pacifica로신종보고되었고이후 Kim (2009) 에의해반문및형태적특징등을근거로기름종개속 Cobitis에서참 론 * Corresponding author: Yong-Jin Won Tel: , Fax: , won@ewha.ac.kr 종개속 Iksookimia로전속되었다. 북방종개는참종개속어류중유일하게동해안북부강릉남대천이북의하천에만서식하여담수어류분포구계상동북한아지역 (Eastnorth Korea subdistrict) 에속하며 (Kim, 1997), 서식지론주로모래바닥에서식하여자갈과돌바닥에주로서식하는다른참종개속어류와차이를보인다 (Kim, 1997; Kim and Park, 2007; Ko, 2015). 미꾸리과어류의생태학적연구는참종개 Iksookimia koreensis (Kim, 1978) 를시작으로부안종개 I. pumila (Kim and Lee, 1984), 수수미꾸리 Kichulchoia multifasciata (Chong, 1986) 남방종개 I. hugowolfeldi (Choi, 2003), 왕종개 I. longicorpa (Kim and Ko, 2005), 줄종개 Cobitis tetralineata (Kim et al., 2006), 새코미꾸리 Korocobitis rotundicaudata (Byeon, 2007), 점줄종개 C. lutheri (Ko and Park, 2011), 얼룩새코미꾸리 K. naktongensis (Hong et al., 2011) 등비교적많이이루어졌다. 북방종개에대한연구는생태적특징 (Choi and Byeon, 2009) 과난발생및초기생활사 (Lee et al., 2011), 서식지특성 72
2 한국고유종북방종개의연령및산란기특징 73 Fig. 1. Frequency distribution of the total length of Iksookimia pacifica in the Bukcheon (stream), Ganseong-eup, Gangwon-do, Korea from January to December, Gray: juvenile, Black: Female, Dark gray: male. 및섭식생태 (Ko, 2015), 난막구조 (Kim and Park, 1995) 등이있으나, 아직까지연령및산란기특성에대해서는간략한보고만이있었다. 따라서본연구에서는참종개속에서독특한분포및생태적특성을가지는우리나라고유종북방종개의연령및산란기특징을조사하여생태학적특성을밝히고기존에연구된결과및유연종과비교 논의하고자한다. 재료및방법 조사는북방종개 Iksookimia pacifica 개체군크기가큰것으로알려진 (Lee et al., 2010) 강원도고성군간성읍어천리일대의북천에서 2013년 1월부터 12월까지조사를실시하였다. 채 집은족대 ( 망목 1 1, 4 4 mm) 를사용하여매달 14~16일사이에실시하였으며, 기온과수온은 12시를기준으로디지털온도계 (T-250A, ASAHI, Japan) 를사용하여측정하였다. 채집된개체는마취제 MS-222 (Sindel, Canada) 로마취하여전장과체중, 성비등을조사한후대부분방류하였으며, 일부생식소분석이필요한개체만 10% 포르말린용액에고정하여실험실로옮겨분석하였다. 전장은 0.01 mm까지, 체중은 0.01 g까지측정하였으며, 성비는수컷의골질반 (lamina circularis) 의유무로판단하였다. 성장도와연령은매달채집된개체의전장빈도분포도를작성하여추정하였으며 (Ricker, 1971), 전장과체중과의관계식은산란기이후인 9월에채집된개체를근거로작성하였다. 산란기는매달생식소가발달한암컷 ( 전장 120~160 mm) 과수컷 ( 전장 80~110 mm) 을각각 5~10개씩채집하여생식소성숙도 (gonadosomatic index, GSI=gonad weight/body
3 74 고명훈 원용진 weight 100) 를계산하여추정하였다. 포란수는 6월과 7월에채집된개체의난수를계수하였으며, 성숙란의크기는 7월에채집된성숙개체의난을개체별 30개를측정하여계산하였다. 결과 1. 성장및연령추정매월채집된전장빈도분포는 Fig. 1과같이나타났다 년에태어난당년생치어는 8월에처음채집되었으며전장 15~39 (28.4±4.03) mm였고, 이후성장을하여 10월에는 20~44 (33.6±3.75) mm였으며, 이후월동기로접어들면서 12 월까지유사한전장범위를보였다. 2012년에태어난당년생치어는 1월부터 4월까지 20~40 mm로거의성장을하지않았으나 5월이후부터급격한성장을하여 6월 25~49 mm, 8 월 40~69 mm, 10월 45~74 mm로성장하였으며, 8월이후부터는수컷가슴지느러미에골질반이형성되면서암 수가구별되었다. 2년생이상의연령군도공통적으로 4월부터 9 월까지는비교적빠른성장을보였으나그외의계절은느린성장을보이거나성장하지않았다. 산란기인 7월을기준으로연령을추정하면암 수가구별되지않은만 1년생은전장 30~49 mm이었고, 만 2년생이상은암 수가구별되었는데, 암컷은만 2년생 50~74 mm, 만 3년생 75~99 mm, 만 4 년생 100~124 mm, 만 5년생이상 130~160 mm로, 수컷은만 2년생 50~74 mm, 만 3년생이상 75~110 mm로추정되었다. 암컷과수컷은만 1년생과만 2년생에서유사한전장범위를보였으나 3년생이후부터는암컷이수컷보다더많이성장하여 10~30 mm가더컸다. 조사기간중가장큰개체는암컷 165 mm, 수컷 121 mm였으며, 전장과체중과의관계식은수컷 y =3E-06x (R 2 =0.994), 암컷 y =4E-06x (R 2 =0.996) 이였다. Table 1. The sex ratio of Iksookimia pacifica in the Bukcheon (stream), Ganseong-eup, Gangwon-do, Korea from January to December, 2013 Month Female Male Total Sex ratio ( / ) χ 2 Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total 1, , The critical value for χ 2 goodness-of-fit test of equal numbers of females and males (1 df) at 95% significance is 성비 성비는 1월부터 12월까지채집된개체중암 수가구별되는 1년생이상의개체는암컷 1,457개체, 수컷 917개체로성비 ( / ) 는 0.63으로나타났으며 (Table 1), 암 수간의성비는유의한차이를보였다 (χ 2 =122.83, P<0.05). 월별로는 5월과 9월이유의한차이를보이지않았으나 (χ 2 <3.84, P>0.05), 그외의모든달은유의한차이를보였다 (χ 2 >3.84, P<0.05). Fig. 2. Relationship between the total length and body weight of Iksookimia pacifica in the Bukcheon (stream), Ganseong-eup, Gangwon-do, Korea, September, 산란기특징 1) 생식소성숙도 (GSI) 산란기를추정하기위하여매월생식소성숙도 (GSI) 를분석하였다. 암컷은 1월부터 3월까지는 3.1~3.6% 로낮았으나이 후 4월 5.5±0.59%, 5월 13.3±3.05%, 6월 20.8±3.23% 로급격히높아졌으며 7월은 22.2±3.27% 로정점을보였다. 이후 8 월에는 2.8±1.48% 로급격히낮아졌으며이후 10월까지낮게유지되다 11월부터 12월까지는 3.7~4.0% 로약간높아지는
4 한국고유종북방종개의연령및산란기특징 75 A. Temperature B. Female Table 2. Mature egg size of Iksookimia pacifica in the Bukcheon (stream), Ganseong-eup, Gangwon-do, Korea, July, 2013 Total length (mm) Body weight (g) GSI (%) Fecundity Mature egg size (mm) ± , ± , ± , ± , ± , ± , ± , ± , ± , ± , ± , ± , ± , ±0.08 Mean±SD 1.11±0.04 C. Male Fig. 3. Temperature and gonadosomatic index (GSI) of Iksookimia pacifica in the Bukcheon (stream), Ganseong-eup, Gangwon-do, Korea from January to December, 경향을보였다 (Fig. 3B). 수컷은 1 월부터 3 월까지는 1.2~1.8% 로낮았으나이후 4 월 2.7±1.23%, 5 월 3.4±1.02% 로급격 히높아졌으며 7 월까지 3.3~3.4% 로높게유지되었다. 하지 만 8 월에는 1.3±0.63% 로급격히낮아졌으며이후 12 월까지 1.1~1.4% 로낮게유지되었다 (Fig. 3C). 산란기인 7 월에채집 된암컷과수컷은생식소성숙도가모두높게나타났지만 8 월 에채집된암컷과수컷모두는생식소성숙도가낮게나타났다. 8월에전장 15~39 mm의당년생치어가채집된것으로볼때산란시기는 7월부터 8월로추정되고산란성기는 7월로추정되었으며이때의수온은 20~25 였다 (Fig. 3A). 생식소가성숙하여산란에참여하는개체는 7월을기준으로암컷은전장 100 mm 이상 (4년생이상 ), 수컷은 75 mm 이상 (3년생이상 ) 이었다 (Table 2, Fig. 4) Fig. 4. Fecundity of Iksookimia pacifica in the Bukcheon (stream), Ganseong-eup, Gangwon-do, Korea from June to July, ) 포란수와성숙난의크기포란수는만 4년생 (n=14) 이 1,505±481개, 5년생이상 (n=13) 이 3,577±1100개로나타났으며전체 (n=27) 는 2,503±1337개였다 (Fig. 4). 산란기성숙란은전장과생식소성숙도, 포란수와상관없이 1.11±0.04 (1.09~1.13) mm로대부분균일하였다 (Table 2). 고 어류의연령을추정하는방법은여러가지방법이있으나산란시기가짧고성장이빠른어류에서는전장빈도분포법 (Petersen method) 을많이이용하여왔다 (Ricker, 1971; Bagenal and Tesch, 1978; Moyle and Cech, 2000). 특히미꾸 찰
5 76 고명훈 원용진 리과 (Cobitidae) 어류는다른잉어목어류에비해체고가낮고전장이긴특징을가지며, 암 수에있어성장도차이를보이는것으로알려져왔다 (Uchida, 1939; Kim, 1997, 2009). 본조사결과북방종개 Iksookimia pacifica도수컷에골질반 (lamuna circularis) 이나타나암 수가구별되었고, 구분된암컷과수컷은연령수및전장범위에서큰차이를보여이전에보고된미꾸리과참종개속 Iksookimia (Kim, 1978; Kim and Lee, 1984; Kim and Ko, 2005) 및기름종개속 Cobitis (Kim et al., 2006; Ko, 2009; Ko and Park, 2011) 어류와유사하였다. 한편 Choi and Byeon (2009) 은 7월의연곡천집단을암컷과수컷구별없이체장빈도분도를통해만 1년생체장 40~59 mm, 만 2년생 60~89 mm, 만 3년생 90~109 mm, 만 4년생 110~126 mm으로추정하였다. 본조사결과와비교하면, 측정방법및암 수를구별하지않은점을감안하더라도체장 60 mm 이하에서 Choi and Byeon (2009) 은 1개의연령군이있다고하였으나본연구에서는 2개의연령군이나타나차이를보였다. 북방종개의연령수및전장범위를미꾸리과어류와비교하면, 산란기를기준으로연령은북방종개가최소암컷만 5 세이상, 수컷만 3세이상으로추정되어왕종개 I. longicorpa (Kim and Ko, 2005) 와유사하였지만, 암컷만 3세이상, 수컷만 2세이상인참종개 I. koreensis (Kim, 1978) 와부안종개 I. pumila (Kim and Lee, 1984), 기름종개 Cobitis hankugensis (Ko, 2009), 점줄종개 C. lutheri (Ko and Park, 2011), 줄종개 C. tetralineata (Kim et al., 2006) 보다는많았다. 전장에있어서북방종개는만 2년생까지암 수가거의유사하였으나만 3 년생이상에서는암컷이수컷보다전장 30~60 mm가더크게나타나다른미꾸리과어류와유사하였다. 연령별크기에있어서는만 1년생의길이가전장 30~49 mm로비교적작은편이었지만다성장한 5년생이상의크기는암컷 130~160 mm 로, 연령수가비슷한왕종개 (Kim and Ko, 2005) 와비교하여보면북방종개는낮은연령대에서는왕종개보다전장범위가작으나만 5년생이상에서는거의유사하게나타나초기성장은느리나지속적인성장을통하여큰개체까지성장하는것으로추정된다. 전장빈도분포법을통한연령추정은연령이높아지면서성장도가느려지고개체변이폭이커지기때문에 5 년생이상의연령에서는뚜렷이구분되지않는다. 일부미꾸리과어류에서는연륜을가진형질을분석하여연령을추정하였는데, 이석을이용한경우유럽의 C. paludica는암컷 5세, 수컷은 3세 (Przybylski and Valladolid, 2000; Oliva-Paterna et al., 2002), C. taenia는암컷 4세, 수컷 3세 (Robotham, 1981) 까지, 비늘을이용한참종개는암 수모두 3세까지 (Kim et al., 2008) 추정되어본결과의연령수와비교적유사하였으나, 안하극을이용한일본의 Niwaella delicata는암컷 12.5세, 수컷은 11.5세까지추정된경우도있어 (Kano, 2000), 추후연륜형질을이용한연구결과와비교검토가필요하다고생각된다. 미꾸리과어류중기름종개속 Cobitis과참종개속 Iksookimia, 미꾸리속 Misgurnus, 새코미꾸리속 Koreocobitis 등은이차성징으로수컷가슴지느러미제2 기조에골질반을가지며 (Kim, 1997, 2009) 산란시암컷을감싸는지지대역할을한다 (Kawanabe et al., 1989; Bohlen, 1999; Lee, 2009). 북방종개의골질반은부화후 13개월이후에형성되어왕종개 (Kim and Ko, 2005) 와줄종개 (Kim et al., 2006), 점줄종개 (Ko and Park, 2011) 등과비교적유사하였다. 그러나줄종개 (Kim et al., 2006) 와점줄종개 (Kim and Jeong, 1988; Ko and Park, 2011), 기름종개 (Ko, 2009) 는산란기에수컷의반문이변형되어암컷과구별되는것으로보고되었는데, 북방종개에서는이러한반문변화는관찰되지않았다. 북방종개의성비 ( / ) 는 0.63으로나타나동종연곡천집단 0.93보다낮았으며 (Choi and Byeon, 2009), 미꾸리과어류왕종개 0.65 (Kim and Ko, 2005), 남방종개 I. hugowolfeldi 0.60 (Choi, 2003), 참종개 0.74 (Kim, 1978), 점줄종개 0.65 (Kim and Jeong, 1988), 0.78 (Ko and Park, 2011), 줄종개 0.58 (Kim et al., 2006), 새코미꾸리 K. rotundicaudata 0.78 (Byeon, 2007) 등과대체로유사하였지만부안종개 1.01보다는크게낮았다 (Kim and Lee, 1984). 한편점줄종개의경우성장하면서수컷에서암컷으로성전환을통해성비가변한다는보고가있고 (Kim and Park, 1992), 단성생식을하는기름종개-왕종개 complex는성비가 0.06으로수컷의비율이극단적으로낮으며동소적으로서식하는이들의부모종인기름종개 1.55, 왕종개 0.51로보고되어독특한양상을보이는것으로알려진바있다 (Ko, 2009). 온대지역에서식하는담수어류의산란은수온과광주기등의계절적변화영향을받으며 (Bye, 1984), 크게춘하산란형과추동산란형으로구분되며춘하산란형은주로수온의영향을받는것으로알려져있다 (Shimzu and Hanyu, 1982; Kaneko et al., 1986). 북방종개산란기는수온 20~25 인 7~8월 ( 산란성기 7월 ) 로추정되어 Choi and Byeon (2009) 와동일하게춘하산란형에속하였으며, 줄종개 6~7월 (Kim et al., 2006), 점줄종개 7월 (Ko and Park, 2011) 과비교적유사하였으나, 수수미꾸리 Kichulchoia multifasciata 5월 (Chong, 1986), 부안종개 5월 (Kim and Lee, 1984), 참종개 5~6월 (Kim, 1978), 새코미꾸리 5~6월 (Byeon, 2007), 얼룩새코미꾸리 K. naktongensis 5~6월 (Hong et al., 2011), 왕종개 6월보다는 (Kim and Ko, 2005) 느린편이었다. 포란수와성숙난의크기는종특이성을밝히는형질로사용되어왔는데, 포란수는 2,503±1,337개로나타나 Lee et al. (2011) 의 2,968±502개와유사하였으나 Choi and Byeon (2009) 의 809±295개보다는많은편이었으며, 왕종개 (2,402±944개; Kim and Ko, 2005) 와남방종개 (2,084개; Choi, 2003), 기름종개 (2,783±1,543개; Ko, 2009) 와유사하였으나
6 한국고유종북방종개의연령및산란기특징 77 좀수수치 (69±35개; Ko and Bang, 2014), 수수미꾸리 (820± 207; Chong, 1986), 부안종개 (329개; Kim and Lee, 1984), 참종개 (1,138±431개; Kim, 1978), 점줄종개 (1,127±453개; Ko and Park, 2011), 줄종개 (1,288±583개; Kim et al., 2006), 새코미꾸리보다는 (1,365±720개; Byeon, 2007) 많고미꾸라지 (8,500~1,3500개; Kim et al., 1992), 미꾸리 (16,430~40,000 개 ; Uchida, 1939), 얼룩새코미꾸리보다는 (22,643±4,629 개 ; Hong et al., 2011) 적은편이었다. 난경은 1.11±0.04 mm 로 Lee et al. (2011) 의 1.09±0.04 mm와같았으나 Choi and Byeon (2009) 의 0.76±0.01 mm 보다는컸고, 미꾸리과내에서는 1.0~1.2 mm인참종개 (Kim, 1978), 남방종개 (Choi, 2003), 점줄종개 (Ko and Park, 2011), 줄종개 (Kim et al., 2006), 기름종개 (Ko, 2009), 새코미꾸리 (Byeon, 2007), 미꾸리 (Uchida, 1939) 와비교적유사하였으나 0.8~0.9 mm인얼룩새코미꾸리 (Hong et al., 2011) 와미꾸라지보다는 (Kim et al., 1987) 크고, 1.3~1.8 mm인부안종개 (Kim and Lee, 1984) 와왕종개 (Kim and Ko, 2005), 좀수수치 (Ko and Bang, 2014), 수수미꾸리보다는 (Chong, 1986) 작은편이었다. 따라서북방종개의연령구조는 5세이상으로구분되어같은참종개속의왕종개와유사하며비교적연령수가많은편이었다. 성비는대체적으로참종개속과기름종개속어류와유사하였으나, 산란기와난경은참종개속어류보다기름종개속어류와유사하였으며, 포란수는참종개속의왕종개와남방종개, 기름종개속의기름종개와비교적유사한특징을보였다. 요약 북방종개 Iksookimia pacifica의연령및산란기특징을밝히기위해 2013년 1월부터 12월까지강원도고성군북천에서조사를실시하였다. 전장빈도분포도를이용하여연령을추정한결과암컷은산란기 (7월) 를기준으로전장 30~49 mm 는만 1년생, 50~74 mm는만 2년생, 75~99 mm는만 3년생, 100~124 mm는만 4년생, 130~160 mm는만 5년생이상으로추정되었다. 성비 ( / ) 는 0.63이었으며암컷이수컷보다 30~60 mm가더컸다. 이차성징인수컷가슴지느러미의골질반은부화후 13개월후에생성되어암 수가구별되었다. 산란기는생식소성숙도로추정한결과 7~8월 ( 산란성기 7월 ) 로추정되었으며, 포란수는 2,503±1,337개, 성숙난의크기는 1.11±0.04 mm였다. 사사 이논문은 2013 년도정부 ( 교육부 ) 의재원으로한국연구 재단의지원을받아수행된기초연구사업임 ( 과제번호 : 2013 R1A1A ). REFERENCES Bagenal, T.B. and F.W. Tesch Age and growth. In: Bangenal, T. (ed.), Methods for Assesment of Fish Production in Fresh Waters, 3th edition. Blackwell Scientific Publications, Oxford, London, Edinburgh, Melbourne, pp Bohlen, J Reproduction of spined loach, Cobitis taenia, (Cypriniformes; Cobitidae) under laboratory conditions. J. Appl. Ichthyol., 15: Bye, V.J The role of environmental factors in the timing of reproductive cycle. Fish reproduction: Strategies and tacties, Academic Press, London: Byeon, H.K Ecology of Koreocobitis rotundicaudata (Cobitidae) in the Naerin Stream, Korea. Korean J. Ichthyol., 19: (in Korean) Choi, E.K Biology of the Southern spined loach, Iksookimia hugowolfeldi (Pisces, Cobitidae). Master Thesis, Chonbuk Nat. Univ., 42pp. (in Korean) Choi, J.K. and H.K. Byeon Ecological characteristics of Cobitis pacifica (Cobitidae) in the Yeongok Stream. Korean J. Limnol., 42: (in Korean) Chong, D.S Morphological and bionomical studies of Niwaella multifasciata (Wakiya et Mori). Master Thesis, Chon buk Nat. Univ., 37pp. (in Korean) Hong, Y.K., H. Yang and I.C. Bang Habitat, reproduction and feeding habit of endangered fish Koreocobitis naktongensis (Cobitidae) in the Jaho Stream, Korea. Korean J. Ichthyol., 23: (in Korean) Kaneko, T., K. Aida and I. Hanyu Changes in ovarian activity and fine structure of pituitary gonadotoropin during spawning cycle of thachichibugoby, Tridentiger obscurus. Nippon Suisan Gakkaishi, 52: Kano, Y Age and growth of the Ajime-loach, Niwaella delicata, in the Yura river, Kyoto, Japan. Ichthyol. Res., 47: Kawanabe, H., N. Mizuno and K. Hosoya Freshwater Fishes of Japan. YAMA-KEI Publishers Co., Ltd., Tokyo, 720pp. Kim, D.H., H.Y. Jo and H.J. Lee Study on the reproduction and growth of Iksookimia koreensis Kim (Pisces: Cobitida) in the Namdae Stream, Cheorwon-gun, Gangwon-do, Korea. Korean J. Ichthyol., 20: (in Korean) Kim, D.S., J.H. Kim and I.S. Park Induced and multiple spawnings by Human Chorionic Gonadotropin injection of the loach, Misgurnus mizolepis (Teleostomi; Cobitidae). J. Aquacult., 5: (in Korean) Kim, I.S Ecological studies of cobitid fish, Cobitis koreensis in Jeonju-cheon Creek, Jeonrabug-do province, Korea. Korean J. Ecol., 2: (in Korean)
7 78 고명훈 원용진 Kim, I.S Illustrated Encyclopedia of Fauna & Flora of Korea, Vol. 37, Freshwater Fishes. Ministry of Education, Yeongi, 518pp. (in Korean) Kim, I.S A review of the spined loaches, family Cobitidae (Cypriniformes) in Korea. Korean J. Ichthyol., 21 (supplement): Kim, I.S. and M.H. Ko Ecology of Iksookimia longicorpa (Cobitidae) in the Seomjin River, Korea. Korean J. Icthyol., 17: (in Korean) Kim, I.S., M.H. Ko and J.Y. Park Population ecology of Korean sand loach Cobitis tetralineata (Pisces; Cobitidae) in the Seomjin River, Korea. J. Ecol. Field Biol., 29: (in Korean) Kim, I.S. and M.T. Jeong Seasonal variation of the color pattern in the Cobitid fish Cobitis taenia lutheri form Korea. Korean J. Ecol., 11: (in Korean) Kim, I.S. and J.Y. Park Adhesive membrane of oocyte in korean cobitid species (Pisces, Cobitidae). Korean J. Zool., 38: Kim, I.S. and J.Y. Park Sex ratios and hermaphroditism of Cobitis lutheri (Pisces, Cobitidae) from Korea. Korean J. Ichthyol., 4: Kim, I.S. and J.Y. Park Freshwater Fishes of Korea. Kyohak Publishing Co., Ltd., Seoul, 467pp. (in Korean) Kim, I.S., J.Y. Park and T.T. Nalbant The far-east speceis of the genus Cobitis with description of three new taxa (Pisces; Ostairophysi; Cobitidae). Trav. Mus. Nat. Hist. Nat. ʻGriore Antipaʼ, 44: Kim, I.S. and W.O. Lee Morphological and ecological aspects on the population of Cobitis koreensis Kim (Pisces: Cobitidae) in the Begchon Stream, Puan-gun, Cholla-bugdo, Korea. Korean J. Ecology, 7: (in Korean) Kim, Y.U., D.S. Kim and Y.S. Park Development of eggs, Larvae and Juveniles of loach, Misgurnus mizolepis Günther. Bull. Korean Fish. Soc., 20: (in Korean) Ko, M.H Reproductive mechanisms of the unisexual diploid-triploid hybrid complex between the spined loach Cobitis hankugensis and Iksookimia longicorpa (Teleostei, Cobitidae) in Korea. Doctoral Thesis, Chonbuk Nat. Univ., 160pp. (in Korean) Ko, M.H Habitat characteristics and feeding ecology of the korean endemic species, Iksookimia pacifica (Pisces: Cobitidae) in the Bukcheon (stream), Korea. Korean J. Ichthyol., 27: (in Korean) Ko, M.H. and J.Y. Park Growth and spawning ecology of Cobitis lutheri (Teleostei: Cobitidae) in the Mangyeong river, Korea. Korean J. Ichthyol., 23: (in Korean) Lee, I.R Studies on the conservation biology of an endangered freshwater fish, Iksookimia choii. Doctoral dissertation, Soonchunhyang Univ., 167pp. (in Korean) Lee, W.O., M.H. Ko, J.M. Bak, D.H. Kim, H.J. Jeon and K.H. Kim Characteristics of fish fauna and community structure in Buk Stream of Goseong, Korea. Korean J. Ichthyol., 22: (in Korean) Lee, W.O, K.H. Kim, J.M. Baek, Y.J. Kang, H.Z. Jeon and C.H. Kim Embryonic development and early life history of the northern loach, Cobitis pacifica (Pisces: Cobitidae). Korean J. Limnol., 44: 1-8. (in Korean) Moyle, P.B. and J.J.Jr., Cech Fishes: An Introduction to Ichthyology, 4th edition. Prentice Hall Inc., New Jersey, 744pp. Nelson, J.S Fishes of the World. John Wiley & Sons, Inc., Hoboken, pp Oliva-Paterna, F.j., M.M. Torralva and C. Fernández-Delgado Age, growth and reproduction of Cobitis paludica in a seasonal stream. J. Fish Biol., 60: Przybylski, M. and M. Valladolid Age and growth of Cobitis paludica in the Lozoya River (Central Spain). Folia Zoo., 49 (Suppl.): Ricker, W.E Methods for assessment of fish production in freshwater. IBP hand book, 3: Robotham, P.W.J Age, growth and reproduction of a population of spined loach, Cobitis taenia (L.). Hydrobiologia, Shimizu, A. and I. Hanyu Environmental regulation of annual reproductive cycle in a spring-spawning bitterling Acheilognathus tabira. Nippon Suisan Gakkaishi, 48: Uchida, K The Fishes of Tyosen. Part I. Numatognathi, Eventognathi. Bull. Fish Exp. Sta. Gov. Gener. Tyôsen, pp (in Japanese)
94 김형수 양현 Park (213) 은생식소성숙도와조직학적분석을근거로본종의산란시기를 4~월로제안한바있다. 본연구는일반적인어류의산란기추정에이용되는생식소성숙도지수를 Kim and Park (213) 의연구보다더많은개체를대상으로매월분석하여정확한산란기를추정하고수컷의생식소성숙
Korean Journal of Ichthyology, Vol. 28, No. 2, 93-99, June 216 ISSN: 122-898 (Print), 2288-3371 (Online) Received: March, 216 Revised: May 23, 216 Accepted: June 23, 216 낙동강유곡천에서식하는수수미꾸리 Kichulchoia multifasciata
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