摘要
为了更好地保护和开发利用塔里木河流域叶尔羌高原鳅资源,基于2019-2022年在塔里木河支流车尔臣河采集到的348尾样本,对叶尔羌高原鳅(Triplophysa yarkandensis)的年龄结构与生长特征进行研究。结果显示,采集到的样本体长分布范围为38~290 mm,体质量分布范围为0.8~271.6 g,年龄结构为1~11龄,体长体质量关系式为W=2.47×1
叶尔羌高原鳅(Triplophysa yarkandensis)隶属于鲤形目(Cypriniformes)鳅科(Cobitidae)条鳅亚科(Noemacheilinae)高原鳅属(Triplophysa),是塔里木河流域特有的鱼类。作为高原鳅属鱼类中体型最大的经济鱼类之一,叶尔羌高原鳅被《中国生物多样性红色名录·内陆鱼类》定为易危鱼类,《新疆维吾尔自治区重点保护水生野生动物名录》列为II级重点保护水生动
有关鱼类年龄与生长的研究报道有很多,其研究方法和理论也在不断更新与发
车尔臣河(Qarqan River)是中国第一大内流河塔里木河的重要支流、塔里木河九大水系之一,位于巴音郭楞蒙古自治州境内,全长813 km,是塔里木河受到人类影响相对较小的支流之
1)采样时间。在车尔臣河水域共开展了6次调查,调查时间分别为2019年的5月(春)、8月(夏)、10月(秋)和2020年的6月(夏)、11月(秋)以及2021年的4-5月(春)。
2)采样站点。采用实地考察与历史调查数据相结合的调查布点方式,利用卫星地图进行站位布设,调查站位分布见
站位名称 Station name | 站点 Site | 经度 Longitude | 纬度 Latitude | 海拔/m Altitude |
---|---|---|---|---|
且末水电站 Qiemo Hydropower Station | Q1 | 85°41′43.28"E | 37°43′59"N | 1 750 |
龙口 Longkou | Q2 | 85°35′56.27"E | 38°1′36.74"N | 1 342 |
第二分水枢纽 The second diversion hub | Q3 | 85°30′49.93"E | 38°12′51.44"N | 1 209 |
环境检测站 Environmental testing station | Q4 | 85°46′30.77"E | 38°28′3.65"N | 1 125 |
五苇场 The fifth reed field | Q5 | 86°31′1.16"E | 38°40′34.13"N | 1 015 |
瓦石峡 Washixia | Q6 | 86°40′7.85"E | 38°45′47.37"N | 989 |
台特玛湖 Tetema Lake | Q7 | 88°16′53.54"E | 39°28′46.79"N | 789 |
利用地笼、刺网、抬网等工具进行采样。地笼(长度分别为:5、10、15 m, 网眼直径为4 mm)下在岸边回水湾处;刺网(网眼:1、2、3 cm)斜截河道或横截回水湾;抬网(长8 m、高1.5 m、网眼2 cm)采用2~3人操作,垂直水流方向展开抬网,顺水流方向快速前行,围捕顶水鱼类等方法共捕获叶尔羌高原鳅544尾。现场解剖测量标准体长、全长、体质量、性腺质量、内脏质量和空壳质量等,长度精确到 0.01 cm,体质量精确到0.01 g。对于较大个体,现场摘取微耳石和鳃盖骨;对于当年幼鱼及仔稚鱼,用95%乙醇保存鱼体,带回实验室在解剖镜下摘取微耳石,不取鳃盖骨。利用钙化组织分析法,以微耳石为主,鳃盖骨为辅对叶尔羌高原鳅进行年龄鉴定,并用孵化不到1个月的仔鱼个体的耳石确定首
将采集的耳石用乙醇清洗干净,保存于1.5 mL EP管中加95%乙醇带回实验室并进行固定、打磨和透明等操作:(1)固定:耳石凸面向上,用热熔胶包埋固定于载玻片上,24 h自然风干;(2)打磨:用100

图1 显微镜下叶尔羌高原鳅的耳石(A)和鳃盖骨(B)
Fig.1 The otoliths(A) and operculum(B) of Triplophysa yarkandensis under a microscope
黑色箭头处为年轮标记。The black arrow is the annual ring.
将采集的鳃盖骨经过开水煮沸,用毛刷清理干净,烘干后装入牛皮纸袋中带回实验室并进行以下处理:(1)漂白:将鳃盖骨置于1%双氧水中漂白24~48 h;(2)烘干:将漂白后的鳃盖骨清洗干净,60 ℃烘干;(3):观察:将处理好的鳃盖骨在解剖镜入射光下观察,并拍照记录(
采用Keys公式对叶尔羌高原鳅的体长体质量关系进行回归分析,公式如下:
W=a | (1) |
将体长和体质量数据进行对数化处理,以lnW为协变量进行协方差分析(ANCOVA),检验不同性别体长体质量关系是否存在显著性差异,并采用Pauly-t检验比较异速生长指数(b)值和3(等速生长状态下的b值)是否存在显著性差异。Pauly-t检验公式为:
(2) |
采用von Bertalanffy生长方程拟合叶尔羌高原鳅的生长特征,公式如下:
Lt =L∞[1- | (3) |
Wt=W∞[1- | (4) |
式(
采用表观生长指数()对同属鱼类生长性能进行比较,公式如下:
lgk+2lgL∞ | (5) |
生长拐点年龄(ti)计算公式如下:
(6) |
式(
采用Excel 2019、SPSS 24.0、Origin 2017进行数据统计分析,采用ArcMap 10.6、GoogleEarth Pro、Lightools进行图片处理。统计显著性标准设为0.05。
2019-2021年在塔里木河支流车尔臣河共采集到544尾叶尔羌高原鳅,其中348尾用于生长年龄研究,雌性157尾,雄性148尾,性别未辨43尾。叶尔羌高原鳅雌性个体体长范围为39~275 mm,平均体长为(165.9±47.42) mm,优势体长范围为120~210 mm,占总数的29.88%;体质量范围为0.87~266.96 g,平均体质量为(64.59±52.24) g,优势体质量范围为0~90 g,占总数的34.20%。雄性个体体长范围为62~290 mm,平均体长为(176.46±39.61) mm,优势体长范围为120~210 mm,占总数的32.18%;体质量范围为2.17~271.60 g,平均体质量为(74.89±48.57) g,优势体质量范围为30~120 g,占总数的31.61%。性别未辨鳅体长范围为38~145 mm,平均体长为(84.00±27.69) mm,优势体长范围为60~120 mm,占总数的7.76%;体质量范围为0.8~32.99 g,平均体质量为(8.91±7.74) g,优势体质量范围为0~30 g,占总数的12.07%(

图2 叶尔羌高原鳅体长(A)和体质量(B)频数分布
Fig.2 Standard length(A) and body weight(B) frequency distribution of Triplophysa yarkandensis
叶尔羌高原鳅总体年龄范围为1~11龄,雌性个体年龄范围为1~10龄,2~5龄为优势年龄组,占总个体数的36.21%;雄性个体年龄范围为1~11龄,3~6龄为优势年龄组,占总个体数的36.49%;性别未辨个体年龄范围为1~4龄,1~2龄为优势年龄组,占总个体数的11.21%(

图3 叶尔羌高原鳅群体年龄结构分布
Fig.3 Age structure distribution of Triplophysa yarkandensis
用实测法,对车尔臣河采集到的叶尔羌高原鳅雌性个体、雄性个体、性别未辨和种群总体的实测体长和体质量数据分别进行拟合(

图4 叶尔羌高原鳅群体体长体质量关系
Fig.4 Length-weight relationship ofTriplophysa yarkandensis
雌性群体:W=2.18×1
雄性群体:W=2.82×1
性别未辨:W=1.95×1
协方差分析表明,叶尔羌高原鳅雌性和雄性体长与体质量方程差异不显著(F=1.459,P>0.05),故将其合并为种群总体的幂函数方程。
种群总体:W=2.47×1
经Pauly-t检验,叶尔羌高原鳅种群总体b值显著小于3(t=3.917>t0.05,P<0.05),表明叶尔羌高原鳅群体为负异速生长。
年龄Age | 雌性Female | 雄性Male | 性别未辨Undetermined | ||||||
---|---|---|---|---|---|---|---|---|---|
数量 n | 平均值±标准差/mm Mean±SD | 范围/mm Range | 数量 n | 平均值±标准差/mm Mean±S.D. | 范围/mm Range | 数量 n | 平均值±标准差/mm Mean±S.D. | 范围/mm Range | |
1 | 7 | 73.07±27.21 | 39~126 | 2 | 64.50±2.50 | 62~67 | 31 | 71.83±21.36 | 38~101 |
2 | 25 | 122.58±18.43 | 99~189 | 8 | 124.75±16.03 | 100~155 | 8 | 108.75±9.62 | 96~124 |
3 | 34 | 137.05±15.11 | 105~165 | 29 | 144.60±20.78 | 116~225 | 3 | 123.33±8.65 | 112~133 |
4 | 43 | 173.37±26.37 | 141~270 | 51 | 168.14±21.02 | 130~220 | 1 | 145.00 | 145 |
5 | 24 | 193.50±20.65 | 147.5~242 | 35 | 196.97±19.79 | 153~245 | |||
6 | 16 | 228.31±18.27 | 177~255 | 12 | 226.00±19.69 | 195~265 | |||
7 | 4 | 247.00±10.65 | 230~257 | 5 | 211.20±25.19 | 167~245 | |||
8 | 1 | 269.00 | 269 | 2 | 241.50±16.50 | 225~258 | |||
9 | 1 | 252.00 | 252 | 1 | 250.00 | 250 | |||
10 | 2 | 272.50±2.50 | 270~275 | 1 | 265.00 | 265 | |||
11 | 2 | 280.00±10.00 | 270~290 |
雌性:Lt=407.2572[1-
雄性:Lt=327.9114[1-
将体长体质量关系式代入生长方程,得到体质量生长方程:
雌性:Wt=697.2606[1-
雄性:Wt=373.9090[1-
由

图5 叶尔羌高原鳅的年龄与体长关系
Fig.5 The relationship between age and standard length of Triplophysa yarkandensis
体长生长不具有拐点,而体质量生长速度呈现先升后降的趋势,当体质量生长加速度为0时,体质量生长速度达到最大值,此点为体质量的生长拐点(ti):
雌性的拐点年龄为8.5龄,此时的体长为265.63 mm,体质量为204.04 g;雄性的拐点年龄为5.9龄,此时的体长为212.26 mm,体质量为108.94 g。
鱼类种群特征可以从侧面反映鱼类生存现状,同种鱼类会因不同的环境而表现出不同生长速度、个体大小等,一般在相对较恶劣、捕捞强度大的条件下,鱼类个体会趋于小型化,性腺提前发育成熟,以保证可以利用更少的资源而达到延续种群的目的,而在较好的条件下,鱼类个体就会相对成长得更大,寿命更
鱼类在自然界中生存,要不断适应环境,最终经过自然选择幸存下来的个体才能继续繁衍以壮大种群,然而受到自身遗传、环境指标、空间大小、食物数量等多方面的因素制约,鱼类也会通过调节自身繁殖、摄食、洄游等行为来应对自然环境的变化,其中年龄结构与生长特性最能表现出鱼类的生存状
车尔臣河叶尔羌高原鳅群体年龄结构为1~11龄,这与克孜
鱼类的生长特性能直观地表现出鱼类种群的发展情况,其生长是由自身基因和自然环境共同作用的结果,同一物种对于不同的环境条件会做出不同的反应,从而会表现出不同的生长状态而出现种间差
由于水利工程建设、修桥筑路工程、挖沙以及过度的农业用水,破坏鱼类生境,引起断流造成干支流连通性以及生境的片段化,导致叶尔羌高原鳅的种群隔离、交流受阻,导致补充群体数量不足,种群数量减少的速度加快。同时,在与外来物种与竞争中,土著种占据劣势,使生长缓慢、成熟年龄较迟的鱼类资源量逐年下降,已经临近濒危。针对上述问题,并结合叶尔羌高原鳅生物学特性提出以下保护措施及建议:(1)加大对野生鱼类保护的宣传力度,严厉打击非法捕捞行为,增强人们对这一珍贵野生动物资源的自觉保护意识。(2)就地建立叶尔羌高原鳅种质资源保护区,对这幸存的高质量栖息地加以保护,加强其种质资源的保护。(3)积极开展对叶尔羌高原鳅的科学研究和人工繁殖工作,以人工授精、人工流水孵化的形式进行濒临物种的人工繁殖,对一些濒危的鱼类进行抢救性保护,建立活体种质资源库,开展人工增殖放流活动,在研究分析的基础上维持车尔臣河的生态流量以满足叶尔羌高原鳅繁殖所需。
参考文献References
陈生熬. 叶尔羌高原鳅早期发育及盐碱适应生理机制[D].武汉:华中农业大学,2019.CHEN S A.Early development and physiological mechanism of salinity-alkalinity adaption of Triplophysa yarkandensis(Day)[D].Wuhan: Huazhong Agricultural University,2019(in Chinese with English abstract). [百度学术]
阿达可白克·可尔江,霍堂斌,姜作发,等.塔里木河干流叶尔羌高原鳅种群生物学研究[J].水生态学杂志,2011,32(1):34-39.Adakbek·Kar jan,HUO T B,JIANG Z F,et al.Study on biological characters of Triyplophysa(Hedinichthys)yarkandensis(Day)from the main stream of Tarim River[J].Journal of hydroecology,2011,32(1):34-39 (in Chinese with English abstract). [百度学术]
陈生熬.塔里木河叶尔羌高原鳅种群生态学研究[D].武汉:华中农业大学,2012.CHEN S A.Population ecology of loach in Yarkant Plateau of Tarim River[D].Wuhan:Huazhong Agricultural University,2012(in Chinese with English abstract). [百度学术]
曾霖,唐文乔.叶尔羌高原鳅的年龄、生长与繁殖特征[J].动物学杂志,2010,45(5):29-38.ZENG L,TANG W Q.Age,body growth and reproductive characteristics of Triplophysa yarkandensis[J].Chinese journal of zoology,2010,45(5):29-38 (in Chinese with English abstract). [百度学术]
朱国平,魏联.南极鱼类年龄与生长研究进展[J].水产学报,2017,41(10):1638-1647.ZHU G P,WEI L.Age and growth of Antarctic fish species:a review[J].Journal of fisheries of China,2017,41(10):1638-1647 (in Chinese with English abstract). [百度学术]
廖传松,陈思宝,叶少文,等.三峡水库鱼类群落结构及主要鱼类种群年龄与生长[J].中国水产科学,2021,28(6):695-702.LIAO C S,CHEN S B,YE S W,et al.Fish assemblage structure and age and growth of the primary fish populations in the Three Gorges Reservoir[J].Journal of fishery sciences of China,2021,28(6):695-702 (in Chinese with English abstract). [百度学术]
刘乐乐,刘海平,王纤纤,等.西藏双须叶须鱼年龄与生长特点研究[J].生物资源,2020,42(6):617-628.LIU L L,LIU H P,WANG Q Q,et al.Age and growth characteristics of Ptychobarbus dipogon in Tibet[J].Biotic resources,2020,42(6):617-628 (in Chinese with English abstract). [百度学术]
李筱芹, 郑江, 万卓坤, 等.雅砻江上游厚唇裸重唇鱼种群结构与生长研究 [J].生态学报, 2023(16): 1-18.LI X Q,ZHENG J,WAN Z K,et al.Population structure and growth characteristics of Gymnodiptychus pachycheilus in the upper reach of Yalong River[J].Acta ecologica sinica,2023(16): 1-18 (in Chinese with English abstract). [百度学术]
达伟,王书峰,沈永平,等.1957—2019年昆仑山北麓车尔臣河流域水文情势及其对气候变化的响应[J].冰川冻土,2022,44(1):46-55.DA W,WANG S F,SHEN Y P,et al.Hydrological response to the climatic changes in the Qarqan River Basin at the northern slope of Kunlun Mountains during 1957—2019[J].Journal of glaciology and geocryology,2022,44(1):46-55 (in Chinese with English abstract). [百度学术]
HOU Y F,CHEN Y N,DING J L,et al.Ecological impacts of land use change in the arid Tarim River Basin of China[J/OL].Remote sensing,2022,14(8):1894[2023-06-16].https://doi.org/10.3390/rs14081894. [百度学术]
马燕武,郭焱,张人铭,等.新疆塔里木河水系土著鱼类区系组成与分布[J].水产学报,2009,33(6):949-956.MA Y W,GUO Y,ZHANG R M,et al.Fauna composition and distribution of aboriginal fish in the Tarim River of Xinjiang Uygur Autonomous Region[J].Journal of fisheries of China,2009,33(6):949-956 (in Chinese with English abstract). [百度学术]
王智超,姜作发,张长征,等.塔里木河流域土著鱼类名录文献考究及检索表的编制[J].渔业科学进展,2010,31(1):15-18.WANG Z C,JIANG Z F,ZHANG C Z,et al.Checklist literature study and compilation of bracketed keys for indigenous fishes Tarim River Basin[J].Progress in fishery sciences,2010,31(1):15-18 (in Chinese with English abstract). [百度学术]
刘其根,沈建忠,陈马康,等.天然经济鱼类小型化问题的研究进展[J].上海水产大学学报,2005,14(1):79-83.LIU Q G,SHEN J Z,CHEN M K,et al.Advances of the study on the miniaturization of natural economical fish resources[J].Journal of Shanghai Fisheries University,2005,14(1):79-83 (in Chinese with English abstract). [百度学术]
VIEIRA A R,SEQUEIRA V,NEVES A,et al.Reproductive strategy of forkbeard,Phycis phycis,from the Portuguese coast[J].Helgoland marine research,2016,70(1):1-15. [百度学术]
陈生熬,马春晖,丁慧萍,等.塔里木河叶尔羌高原鳅繁殖生物学研究[J].水生生物学报,2013,37(5):810-816.CHEN S A,MA C H,DING H P,et al.The reproductive biology of Triplophysa(hedinichthys) yarkandensis(day) in Tarim River[J].Acta hydrobiologica sinica,2013,37(5):810-816 (in Chinese with English abstract). [百度学术]
殷名称.鱼类生态学[M].北京:中国农业出版社,1995.YIN M C.Fish ecology[M].Beijing:China Agriculture Press,1995(in Chinese). [百度学术]
陈生熬,姚娜,任道全,等.塔里木河上游叶尔羌高原鳅种群结构研究[J].东北农业大学学报,2014,45(8):90-97.CHEN S A,YAO N,REN D Q,et al.Study on the population structure of Triplophysa(Hedinichthys) yarkandensis(Day) in the upper reaches of Tarim River[J].Journal of Northeast Agricultural University,2014,45(8):90-97 (in Chinese with English abstract). [百度学术]
王新月.叶尔羌高原鳅年龄、生长、繁殖和种群判别的研究[D].阿拉尔:塔里木大学,2022.WANG X Y.Age,growth,reproduction and population discrimination of Triplophysa yarkandensis[D].Alar:Tarim University,2022 (in Chinese with English abstract). [百度学术]
王德忠.塔里木河鱼类区系变化及土著鱼类保护问题[J].干旱区研究,1995,12(3):54-59.WANG D Z.The changes of fishes fauna and protections of aboriginal fishes in the Tarim River[J].Arid zone research,1995,12(3):54-59 (in Chinese with English abstract). [百度学术]
HUO T B,JIANG Z F,KARJAN A,et al.Length-weight relationships of 16 fish species from the Tarim River,China[J].Journal of applied ichthyology,2012,28(1):152-153. [百度学术]
姚娜,马良,金珊珊,等.青海北川河流域拟鲶高原鳅生长特性研究[J].内蒙古农业大学学报(自然科学版),2019,40(6):5-10.YAO N,MA L,JIN S S,et al.Growth characteristics of Triplophysa siluroides (herzenstein) in Beichuanhe Basin of Qinghai[J].Journal of Inner Mongolia Agricultural University (natural science edition),2019,40(6):5-10 (in Chinese with English abstract). [百度学术]
李亮涛.雅鲁藏布江中游东方高原鳅年龄生长和种群动态研究[D].武汉:华中农业大学,2016.LI L T.Study on age,growth and population dynamics of Triplophysa orientalis in the middle reaches of Yarlung ZangboTsangpo River,Tibet[D].Wuhan:Huazhong Agricultural University,2016(in Chinese with English abstract). [百度学术]
张雪飞,何春林,宋昭彬.大渡河上游麻尔柯河高原鳅的年龄与生长[J].动物学杂志,2010,45(4):11-20.ZHANG X F,HE C L,SONG Z B.Age and growth of Triplophysa markehenensis from the markehe river in upper reaches of the Dadu River[J].Chinese journal of zoology,2010,45(4):11-20 (in Chinese with English abstract). [百度学术]
刘成杰,张志明,马徐发,等.新疆额尔齐斯河银鲫年龄鉴定比较与生长特征研究[J].水生态学杂志,2015,36(6):51-58.LIU C J,ZHANG Z M,MA X F,et al.Comparitive study of age determination and growth characteristics for Carassius auratus gibelio in the Irtysh River,Xinjiang[J].Journal of hydroecology,2015,36(6):51-58 (in Chinese with English abstract). [百度学术]
BOISCLAIR D,LEGGETT W C.Among-population variability of fish growth:II.Influence of prey type[J].Canadian journal of fisheries and aquatic sciences,1989,46(3):468-482. [百度学术]
BOISCLAIR D,LEGGETT W C.Among-population variability of fish growth:III.Influence of fish community[J].Canadian journal of fisheries and aquatic sciences,1989,46(9):1539-1550. [百度学术]
SMOLIŃSKI S,BERG F.Varying relationships between fish length and scale size under changing environmental conditions-multidecadal perspective in Atlantic herring [J/OL].Ecological indicators,2022,134[2023-06-16].https://doi.org/10.1016/j.ecolind.2021.108494. [百度学术]
田波,吴金明,梁孟,等.长江中游武汉江段铜鱼的年龄与生长[J].水产学报,2021,45(1):68-78.TIAN B,WU J M,LIANG M,et al.Age and growth of Coreius heterodon from Wuhan stretch of the middle Yangtze River[J].Journal of fisheries of China,2021,45(1):68-78 (in Chinese with English abstract). [百度学术]
任波,马燕武,吐尔逊,等.阿克苏河叶尔羌高原鳅Triplophysa(hedinichthgs)yarkandensis(Day)生物学研究[J].水产学杂志,2004,17(1):46-52.REN B,MA Y W,Turson,et al.The study on the biology of Triplophysa(hedinichthgs) yarkandensis(Day)in Akesu River[J].Chinese journal of fisheries,2004,17(1):46-52 (in Chinese with English abstract). [百度学术]
陈生熬,谢从新,宋勇,等.塔里木河沙雅段叶尔羌高原鳅形态生物学研究[J].新疆农业科学,2011,48(8):1499-1504.CHEN S A,XIE C X,SONG Y,et al.Morphological biology of Triplophysa(hedinichthys) yarkandensis(day) in Shaya section of Tarim River[J].Xinjiang agricultural sciences,2011,48(8):1499-1504 (in Chinese with English abstract). [百度学术]
王锦秀,任道全,王新月,等.塔里木河下游河段叶尔羌高原鳅形态生物学研究[J].家畜生态学报,2021,42(8):58-63.WANG J X,REN D Q,WANG X Y,et al.Morphological biology of Triplophysa yarkandensis in the lower reachers of Tarim River[J].Journal of domestic animal ecology,2021,42(8):58-63 (in Chinese with English abstract). [百度学术]
谢从新.鱼类学[M].北京:中国农业出版社,2010.XIE C X.Ichthyology[M].Beijing:China Agriculture Press,2010 (in Chinese with English abstract). [百度学术]
BRANSTETTER S G.Biological parameters of the sharks of the northwestern gulf of mexico in relation to their potential as a commercial fishery resource (age,growth)[D].Texas: Texas A&M University,1986. [百度学术]
王健,张富斌,胡华明,等.雅鲁藏布江下游弧唇裂腹鱼的年龄结构与生长特性[J].水生生物学报,2022,46(12):1770-1779.WANG J,ZHANG F B,HU H M,et al.Characteristics of age and growth of Schizothorax curvilabiatus in the lower reaches of Yarlung Zangbo River[J].Acta hydrobiologica sinica,2022,46(12):1770-1779 (in Chinese with English abstract). [百度学术]
CAMPANA S E.Accuracy,precision and quality control in age determination,including a review of the use and abuse of age validation methods[J].Journal of fish biology,2001,59(2):197-242. [百度学术]
TAO H,GEMMER M,BAI Y G,et al.Trends of streamflow in the Tarim River Basin during the past 50 years:human impact or climate change?[J].Journal of hydrology,2011,400(1/2):1-9. [百度学术]