KOREAN JOURNAL OF ICHTHYOLOGY, Vol. 29, No. 3, 197-24, September 217 ISSN: 1225-8598 (Print), 2288-3371 (Online) Received: August 28, 217 Revised: September 19, 217 Accepted: September 22, 217 한국자생종종개 Orthrias toni (Pisces: Balitoridae) 의생활사 고명훈 * 이화여자대학교에코과학부 Life History of the Siberian Stone Loach Orthrias toni (Pisces: Balitoridae) in the Buk Stream, Goseong, Korea by Myeong-Hun Ko* (Division of EcoScience, Ewha Womans University, Seoul 376, Republic of Korea) ABSTRACT Life history of the Siberian stone loach, Orthrias toni, were investigated using samples collected from Buk Stream, Goseong-gun, Gangwon-do, Korea. Growth was rapid between April and October when water temperature exceeded 1 C, but growth almost stopped from November-March when water temperature was below 1 C. Age groups for O. toni estimated by frequency distribution of total length in spawning season (May) indicated the 6~81 mm is 1-year old, the 82~99 mm group is 2-years old, the 1~119 mm is 3-years old, and the 12~14 mm is more than 4-years old. Total length range by sex was similar, and sex ratio ( / ) was.73. Spawning season according to the gonadosomatic index (GSI) was May with water temperature 14~17 C, and almost all fish were mature after one year old. Fecundity increased rapidly with age with an average of 4,46±3,32 (mean±sd). Size of mature eggs was.87±.3 mm in diameter irrespective of age. Key words: Stone loach, age composition, spawning period, fecundity, mature egg size 서 잉어목 (Cypriniformes), 미꾸리아목 (Cobitoidei) 에속하는 종개과 (Balitoridae) 어류는유럽과아시아에다양하게분화하 여 42 속 618 종이분포하고있으며 (Kottelat, 212; Nelson et al., 216), 우리나라에는 2 속 3 종인종개 Orthrias toni, 대륙 종개 O. nudus, 쌀미꾸리 Lefua costata 가서식하고있다 (Kim, 1997; Kim and Park, 27). Kottelat (212) 은미꾸리아목어류 를정리하면서종개, 대륙종개, 쌀미꾸리 3 종을 Nemachilidae 에포함시키고종개속명을 Barbatula 로사용한바있기때문 에분류학적으로문제가제기되고있다. 종개는대륙종개와 형태적으로유사하나추성의배열및전비공과후비공의형태 에차이가나는것으로보고되었다 (Kim and Park, 27; Cao et al., 212). 분포지에있어서종개는태백산맥을기준으로동 해로흐르는하천에, 대륙종개는서해안으로흐르는한강수계 에서식하는것으로알려져있으며, 국외에서종개는일본의 론 * Corresponding author: Myeong-Hun Ko Tel: 82-2-3277-463, Fax: 82-2-3277-2385, E-mail: hun@jbnu.ac.kr 북해도, 러시아의사할린및시베리아동부등에, 대륙종개는몽고및중국대륙에서식하는것으로알려져있어분포지차이를보인다 (Kim and Park, 27). 우리나라종개속어류는모두자생종으로하천중 상류의돌과자갈등이깔린곳에서식하는대표적저서성어류이며 (Kim, 1997; Kim and Park, 27), 대륙종개의경우수서곤충을주로섭식하는식충성어류로보고되어 (Byeon, 21) 수생태계먹이사슬에서중요한 2차소비자역할을하고있다. 최근하천생태계에대한관심이높아지면서생태적적으로교란된하천의경우하천복원사업이진행되고있고 (Park et al., 29; Choi, et al., 211; Lee and Choi, 215), 하천의건강성을평가하여관리하려는사업등 (ME, 27; NIER, 215) 이진행되고있는데, 자생종의생태학적특징은멸종위기종및고유종과함께중요한자료로활용되고있다. 우리나라종개에관한연구는형태와핵형분석, DNA 염기서열등을이용한계통분류학적연구 (Kim et al., 1988; Lee et al., 199; Park and Lee, 1991; Yang et al., 1991) 가있지만생태학적연구는거의연구되지않았으며, 종개과로확대해도대륙종개의피부구조 (Park and Kim, 23) 와생태 (Byeon, 197 http://www.fishkorea.or.kr
198 고명훈 Korea Study station Ganseong-eup East Sea Goseong-gun Number of individuals 15 1 5 5 2+ Female (n=123) 1+ 3+ 4+ 2 4 6 8 1 12 14 4+ 3+ Buk Stream 38 32ʹ 1 15 Male (n=86) 1+ 2+ Total length (mm) Gangwon-do 2 4 km Fig. 2. Frequency distribution of the total length of female and male of Orthrias toni collected from the Buk Stream, Goseong-gun, Gangwon-do, Korea, May, 213. Fig. 1. Study station of Buk Stream, Eocheonri, Ganseong-eup, Goseong-gun, Gangwon-do, Korea. 21) 등이있을뿐이다. 국외의경우도 Barbatula barbatula, Metaschistura cristata, Lefua echigonia 등일부종들에대해서 만생활사와연령및성장, 산란및섭식생태등의연구가진 행되었다 (Smyly, 1955; Sauvonsaari, 1971; Mills et al., 1983; Erös, 21; Hofmann and Fischer, 21; Saat et al., 23; Vinyoles et al., 21; Aoyama and Doi, 211; Patimar et al., 211). 따라서본연구에서는한국자생종종개의생활사특징인성 장및연령, 성비, 산란특성등의생물학적특징을밝히고근 연종들과비교 논의하였다. 1. 조사지점및시기 재료및방법 조사는종개 Orthrias toni 가집단으로서식하는것으로보 고된 (Lee et al., 21) 강원도고성군간성읍간천리의북천 (Fig. 1) 에서 213 년 1 월부터 12 월까지실시하였다. 2. 채집방법 128 37ʹ 128 48ʹ 채집은족대 ( 망목 1 1, 4 4 mm) 를사용하였으며, 채집된 개체는마취제 MS-222 (Sindel, Canada) 로마취하여전장과 체중, 성별등을조사한후생식소분석이필요한일부개체 만 1% 포르말린에고정하고그외모든개체는바로방류하 였다. 또한채집지역의수심, 유속, 하상등을조사하여서식지 특성을파악하였다. 3. 생활사조사방법성장및연령추정은채집된개체를근거로전장빈도분포도 (Ricker, 1971) 를매달작성하여추정하였다. 성비는생식소가성숙하여암수가구별되는 1월부터 5월, 11월부터 12월까지계산하였으며 χ 2 검정을통하여성비 1 : 1 유의성을확인하였다. 산란기는매달 3년생이상인전장 1~14 mm 개체를암컷과수컷으로나누어각각 1~15개체를 1% 포르말린수용액에고정하였으며, 이후실험실로옮겨전장과체중, 생식소무게등을측정한후생식소성숙도 (gonadosomatic index, GSI =gonad weight/body weight 1) 를계산하여추정하였다. 수온변화는매달전자수온계를이용하여측정하였으며, 기온변화는국가수자원관리종합시스템 (WAMIS) 강원도고성군 ( 속초관측소 ) 의기상자료를인용하여일평균기온으로그래프를작성하였다 (WAMIS, 213). 또한포란수는 3월과 4월에채집된개체의난수를계수하였으며, 성숙란의크기는 4월에채집된성숙개체의난을개체별로 3개를측정하여계산하였다. 결과 1. 성장및연령추정종개암 수의전장성적이형을알아보기위해산란기인 5 월에암컷과수컷으로나누어전장빈도분포도를작성하였다. Fig. 2와같이전장분포는암 수가유사한 4개의최빈값을보여전장에대한성적이형은없는것으로판단되었다. 따라서매월전장빈도분포도작성은암 수구분없이하나의도
한국자생종종개의생활사 199 표로작성하였다. 당년생 (213년부화 ) 치어는전장 26~49 (mean±sd, 38.6± 4.93) mm로 6월에처음관찰되었으며, 7월 28~59 (47.2±5.44) mm, 8월 4~75 (57.9±6.95) mm, 9월 38~69 (62.±6.76) mm 로급격한성장을보였으며, 이후수온이내려가면서성장은둔화되어 1월 5~71 (65.2±5.83) mm였고수온이 1 이하로내려가는 11월부터 12월까지는 52~79 mm로성장을멈추었다. 당년생 11~12월의전장범위는 1월의첫번째최빈값범위 62~84 mm와유사하여같은당년생으로판단되었다. 1월부터 4월까지나타난당년생 ( 첫번째최빈값, 212년에부화 ) 은 1월부터 4월까지는수온이낮아거의성장을하지않았으나 5월부터성장을시작하여 6월 68~9 mm, 7월 73~92 mm, 8월 77~94 mm, 9월 8~96 mm로빠른성장을보였으나 11월이후에는 8~12 mm로온도가급격히내려가면서성장을멈추었다. 이들의 11~12월전장범위는 1월의두번째최빈값의범위와유사하여동일한 1년생으로판단되었다. 2년생이상의연령군도공통적으로수온 1 이상되는 4월에서 1월까지는성장이이루어졌으나수온이 1 이하인 1월부 3 Jan (n=132) 3 Jul (n=27) 2 1 212 211 21 29 2 1 3 Feb (n=137) 3 Aug (n=161) 2 2 1 1 3 Mar (n=182) 3 Sep (n=159) 2 2 Number of individuals 1 3 2 Apr (n=248) 1 3 2 Oct (n=212) 1 1 3 2 May (n=29) 1+ 2+ 3 2 Mov (n=186) 1 3+ 4+ 1 3 Jun (n=292) 3 Dec (n=121) 2 2 1 213 + 1 2 4 6 8 1 12 14 2 4 6 8 1 12 14 Total length (mm) Fig. 3. Monthly variation of age and year class in frequency distribution of total length of Orthrias toni collected from the Buk Stream, Goseonggun, Gangwon-do, Korea from January to December, 213.
2 고명훈 Table 1. The sex ratio of Orthrias toni in the Buk Stream, Goseonggun, Gangwon-do, Korea, from January to December, 213. Month Female Male Total Jan. Feb. Mar. Apr. May Nov. Dec. 터 3 월, 11 월부터 12 월은거의성장하지않았고 1 월의최빈값 단계가 12 월에한단계씩이동한것으로나타났다. 따라서이 러한최빈값의이동으로볼때각각의최빈값이각각의연령 으로판단되었다. 산란기 (5 월 ) 를기준으로연령은암 수구 분없이 1 년생은전장 6~81 mm, 만 2 년생은 82~99 mm, 만 3 년생은 1~119 mm, 만 4 년생이상은 12~14 mm 로추정 되었다 (Fig. 3). 서식지는당년생치어가빠른여울부 ( 유속 2~6 cm/sec) 의자갈 ( 입자크기 5~15 cm) 바닥에서식하였으나 1 년생이상 은공통적으로느린여울부 ( 유속 ~2 cm/sce) 의돌또는큰 돌바닥 ( 입자크기 1~4 cm) 에주로서식하여차이를보였다. 2. 성비 1 월부터 12 월중생식소가성숙하여암 수가구별되는달 은 1 월부터 5 월, 11 월부터 12 월까지 7 개월이었다 (Table 1). 이 기간중에채집된암컷은 71 개체, 수컷은 514 개체로성비 ( / ) 는.73 으로나타나암 수간의성비는유의한차이를보여 1 : 1 이아니었다 (χ 2 =28.78, P<.5). 월별로는 2 월과 11 월 12 월이유의한차이를보이지않아 1 : 1 이었으나 (χ 2 <3.84, P>.5), 1 월과 3 월, 4 월, 5 월은유의한차이를보여 1 : 1 이 아니었다 (χ 2 >3.84, P<.5). 3. 산란기특징 1) 산란기추정 77 74 112 151 123 13 61 55 63 7 97 86 83 6 132 137 182 248 29 186 121 Sex ratio ( / ).71.85.63.64.7.81.98 산란기를추정하기위하여매월생식소성숙도 (GSI) 와온도 변화를조사하였다 (Fig. 4). 암컷은 1 월부터 4 월까지생식소 성숙도가 2% 이상이었으며약간씩증가하는경향을보이다 가 4 월에 23.2±5.2% 로정점을보였다. 하지만 5 월에 8.4± 3.41% 급격히감소하고이후 8 월까지조금씩감소하여 8 월에 2.7±.34% 로최소치를보였다. 이후다시생식소성숙도는급 격히증가하여 1 월에 13.4±.9%, 12 월에 18.4±1.91% 로 χ 2 3.67.88 9.69 11.76 6.55 2.15.1 Total 71 514 1215.73 28.78 The critical value for χ 2 goodness-of-fit test of equal numbers of females and males (df = 1) at 95% significance is 3.84. Temperature ( ) Gonadosomatic index (%) Gonadosomatic index (%) 3 2 1-1 3 25 2 15 1 5 2. 1.5 1..5. Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Fig. 4. Change in temperature and gonadosomatic index (GSI) of Orthrias toni in the Buk Stream, Goseong-gun, Gangwon-do, Korea, from January to December, 213. Air temperature data were obtained from WAMIS (213). 증가하였다. 수컷은 1 월부터 4 월까지 1.2% 이상으로높았으 나 5 월.9±.27%, 6 월.6±.24% 로급격히감소하여최소 치를보였다. 이후 7 월부터다시증가하기시작하여 8 월.9±.47% 를보였고 9 월부터 12 월까지는 1.3% 이상으로높았다. 따라서생식소성숙도가급격히감소하는점과 6 월에당년생 치어 ( 전장 27~48 mm) 가채집된점으로볼때산란기는 5 월 로추정되었다. 또한이때채집된수컷의두부에추성이관찰 되었으며, 수온은 14~17, 기온은 1~2 범위였다. 2) 포란수와성숙난의크기 Month Air T. Water T. Female (n=113) Male (n=93) 암컷의연령별생식소성숙도와포란수, 성숙란의크기는 Table 2 와같았다. 생식소는만 1 년생소수의개체를제외하 고만 1 년생부터만 4 년생이상까지모두성숙하였으며, 생식 소성숙도는연령이증가함에따라각각약 2 배가까이증가하 는경향을보였다. 난의크기는균일하였으며, 포란수는 1 년생
한국자생종종개의생활사 21 Table 2. Gonadosomatic index, fecundity and matured egg diameter of Orthrias toni by age in the Buk Stream, Goseong-gun, Gangwon-do, Korea, from fish collected in March and April, 213. Total length (mm) Number of individuals* Body weight (g) Gonadosomatic index of female (%) Fecundity Matured egg diameter (mm)** Age 1+ 2+ 3 + 4+ Total 6~81 9 3.2±.63 9.7±2.68 955±451.85±.3 (n = 6) 82~99 13 5.6±1.28 18.6±5.88 2,863±1,294.88±.1 (n=5) 1~119 21 9.1±1.3 2.3±6.45 5,342±1,983.86±.2 (n = 6) 12~14 5 18.8±4.43 27.4±4.74 11,211±2,546.88±.3 (n = 3) *Data from sample collected in March and April; **Data from sample collected only in April. n is the number of individuals examined. - 48 8.±4.71 18.6±7.31 4,46±3,32.87±.3 (n = 2) 955±451 개, 2 년생 2,863±1,294 개, 3 년생 5,342±1,983 개, 4 16 년생이상 11,211±2,546개로연령이높아지면서포란수는급격히증가하였고전체평균은 4,46±3,32개였다 (Fig. 5). 4 월의성숙난은거의완숙되어있었으며크기는연령과관계없이균일한크기로평균.87±.3 mm였다. 5월에배란된성숙란은회색으로불투명하였으며기질에달라붙는점착란이었다. Fecundity 14 12 1 8 6 2+ 3+ 4+ 고찰 미꾸리아목 (Cobitoidei) 어류는종또는속에따라독특한성적이형을보이는것으로알려져있는데, 미꾸리과 (Cobitidae) 어류대부분이수컷가슴지느러미에골질반 (lamina circularis) 이나타나고공통적으로전장크기에있어암컷이수컷보다큰경향을보이며 (Kim and Park, 27; Kim, 29; Kottelat, 212), 기름종개속 Cobitis 어류대부분은산란기전후로수컷의반문에성적이형이나타난다 (Kim and Jeong, 1988; Kim et al., 26; Ko, 29). 또한종개과 (Balitoridae) 의쌀미꾸리 Lefua costata 수컷도두개의짙은검은줄이나타나암컷과구별된다 (Kim and Park, 27). 그러나종개 Orthrias toni의성적이형을조사한결과산란기에수컷두부에추성이관찰된것이외는특별한성적이형이관찰되지않았다. 또한암 수의연령별전장에대한성적이형은없었다 (Fig. 2). 근연종인 Barbatula barbatula도암컷과수컷의전장에큰차이가없는것으로보고된바있다 (Vinyoles et al., 21). 종개의성비 ( / ) 는.73으로암컷이수컷에비해유의하게많았는데 (Table 1), 같은속의대륙종개 O. nudus는.98 (Byeon, 21), 유럽에서식하는 B. barbatula도 1.9 (Erös, 21) 로암 수차이가없었다. 같은미꾸리상과에속하는우리나라미꾸리과어류의성비는참종개 Iksookimia koreensis.74 (Kim, 1978), 왕종개 I. longicorpa.65 (Kim and Ko, 25), 북방종개 I. pacifica.63 (Ko and Won, 216), 동방종개 Fig. 5. Fecundity of Orthrias toni by total length and age in the Buk Stream, Goseong-gun, Gangwon-do, Korea from fish collected in March and April, 213. I. yongdokensis.69 (Ko et al., 216), 점줄종개 C. nalbanti.65 (Ko and Park, 211), 새코미꾸리 Koreocobitis rotundicaudata.78 (Byeon, 27), 수수미꾸리 Kichulchoia multifasciata.73 (Chong, 1986) 등다수가.6~.8 사이로나타나본종과 유사하였다. 종개의성장은수온이 1 이상되는 4 월부터 1 월까지 성장을하였으나수온이 1 이하인 11 월부터이듬해 3 월까 지는거의성장을하지않았다. 이러한월별성장주기는이미 보고된우리나라미꾸리과어류들인왕종개, 북방종개, 동방종 개, 줄종개 C. tetralineata, 점줄종개, 기름종개 C. hankugensis 등과비교적유사하였다 (Kim and Ko, 25; Kim et al., 26; Ko, 29; Ko and Park, 211; Ko and Won, 216; Ko et al., 216). 4 2 1+ 5 6 7 8 9 1 11 12 13 14 15 Total length (mm) 연령은월별로전장빈도분포도를조사하여추정한결과, 산 란기를기준으로 1 년생은전장 6~81 mm, 만 2 년생은 82~99 mm, 만 3 년생은 1~119 mm, 만 4 년생이상은 12~14 mm 로추정되었으며, 연령군은월에따라 4~5 개의연령군이관
22 고명훈 Table 3. Comparison of total length by age in spawning season of the stone loach Species Total length (mm) by age 1+ 2+ 3 + 4+ 5+ Reference Orthrias toni 6~81 82~99 1~119 12~14 Present study O. nudus 58~99 1~12 Byeon, 21 Barbatula barbatula 51~53 75~77 91 14 Erös, 21 Metaschistura cristata 49 83 12 19 Smyly, 1955 46 72 89 11 112 Sauvonsaari, 1971 65 93 13 Mills et al., 1983 Table 4. Comparison of egg size and fecundity of Balitoridae and Cobitidae in Korea Balitoridae Cobitidae Orthrias toni O. nudus Species Egg size Fecundity Reference Iksookimia yongdokensis I. koreensis I. pumila I. longicorpa I. hugowolfeldi I. pacifica Cobitis hankugensis C. lutheri C. tetralineata Koreocobitis rotundicaudata K. naktongensis Kichulchoia brevifasciata K. multifasciata Misgurnus anguillicaudatus M. mizolepis.87±.3.75±.4 1.45±.3 1.1±.8 1.35±.8 1.32±.12 1.3±.7 1.11±.4.98±.5 1.9±.2.98±.1 1.18±.9.87±.5 1.46±.7 1.78±.6 1.1 1.12 4,46±33 4,119±2,416 2,292±618 1,138±431 328±23 2,42±944 1,933±53 2,53±1,337 2,783±1,543 1,127±453 1,288±583 1,365±72 22,643±4,629 69±35 82±27 16,43~4, 8,5~13,5 Present study Byeon, 21 Ko et al., 216 Kim, 1978 Kim and Lee, 1984 Kim and Ko, 25 Choi, 23; Park, 216 Ko and Won, 216 Ko, 29 Ko and Park, 211 Kim et al., 26 Byeon, 27 Hong et al., 211 Ko and Bang, 214 Chong, 1986 Uchida, 1939 Kim et al., 1987; 1992 찰되었다. 이러한결과는이석과아가미뚜껑 (operculae) 의연륜으로분석된근연종 Metaschistura cristata의 3~5개연령군 (Smyly, 1955; Sauvonsaari, 1971; Mills et al., 1983), B. barbatula의 4개연령군 (Erös, 21) 과비교적유사하였다. 하지만우리나라에서식하는대륙종개는산란기에 2개의연령군만나타나는것으로보고되어차이를보였다 (Byeon, 21) (Table 3). 종개의생식소성숙도는산란이후 9월부터급격히발달하여월동기에 2% 안팎까지성숙한후 3~4월에정점을보인후 5월에감소하여 (Fig. 4), 근연종대륙종개및 B. barbatula 와유사한생식주기를보였으나산란기는 6월로차이를보였다 (Byeon, 21; Vinyoles et al., 21). 우리나라미꾸리과어류중에서수수미꾸리는월동기까지생식소성숙도가 2% 이상성숙하고 5~6월에산란한다고보고되어생식주기및산란기가본종과매우유사하였지만 (Chong, 1986; Kim and Yang, 216), 참종개속및기름종개속어류는월동기까지생식소성숙도가 5% 미만으로낮다가봄에급격히성숙한후 6월부터 7월에산란하여 (Kim, 1978; Kim and Lee, 1984; Kim and Ko, 25; Kim et al., 26; Ko, 29; Ko and Park, 211; Ko and Won, 216; Ko et al., 216) 본종과생식주기가거의반대경 향이었고산란기도큰차이를보였다. 포란수는평균 4,46±3,32개, 성숙란크기는.87±.3 mm로대륙종개의포란수 (4,119±2,416개) 와는유사하였으나성숙란의크기 (.75±.4 mm) 는대륙종개보다큰편이었다. 이러한포란수와난경을우리나라미꾸리과어류와비교하면 (Table 4), 포란수는얼룩새코미꾸리 (Hong et al., 211) 와미꾸리 Misgurnus anguillicaudatus (Uchida, 1939), 미꾸라지 M. mizolepis (Kim et al., 1992) 보다는적은편이었으나그외의종들보다는많은편이었고, 난경은미꾸리과모두보다작았다. 또한성숙란은회색으로불투명하고강한점착성을가지고있어비교적투명하고약한점착성을가지는미꾸리과어류와차이를보였다. 요약 강원도고성군북천에서종개 Orthrias toni의생활사를조사하여생물학적특징을밝혔다. 성장은수온이 1 이상되는 4월부터 1월까지빨랐으나수온이 1 이하인 11월부터이듬해 3월까지는거의멈추었다. 연령은전장빈도분포도로
한국자생종종개의생활사 23 볼때산란기인 5 월을기준으로전장 6~81 mm 는만 1 년생, 82~99 mm 는만 2 년생, 1~119 mm 는만 3 년생, 12~14 mm 는만 4 년생이상으로추정되었다. 암컷과수컷의연령별 전장범위는거의동일하였고, 성비 ( / ) 는.73 으로암컷이 많았다. 산란기는생식소성숙도 (GSI) 로추정한결과 5 월이었 으며, 이때의수온은 14~17 였다. 생식소는모든연령이성 숙하였고, 연령이높아지면서포란수는급격히증가하였으며, 평균 4,46±3,32 개였다. 성숙란의크기는연령과상관없이 직경.87±.3 mm 로비슷하였다. REFERENCES Aoyama, S. and T. Doi. 211. Morphological comparison of early stages of two Japanese species of eight-barbel loaches: Lefua echigonia and Lefua sp. (Nemacheilidae). Folia Zool., 6: 355-361. Byeon, H.K. 27. Ecology of Koreocobitis rotundicaudata (Cobitidae) in the Naerin Stream, Korea. Korean J. Ichthyol., 19: 299-35. (in Korean) Byeon, H.K. 21. Ecological study of Orthrias nudus (Balitoridae) in the Eoron Stream of Korea. Korean J. Ichthyol., 22: 162-167. (in Korean) Cao, L., R. Causse and E. Zhang. 212. Revision of the loach species Barbatula nuda (Bleeker 1865) (Pisces: Balitoridae) from North China, with a description of a new species from Inner Mongolia. 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