拉萨裂腹鱼个体生物学和种群动态研究
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摘要
拉萨裂腹鱼(Schizothorax waltoni)隶属于鲤形目(Cypriniformes),鲤科(Cyprinidae),裂腹鱼亚科(Schizothoracinae),裂腹鱼属(Schizothorax),仅分布于西藏地区雅鲁藏布江中上游干、支流及附属水体,为我国的特有种。为研究拉萨裂腹鱼的生物学特性及其种群动态,于2008年8月至2009年8月在雅鲁藏布江谢通门至仁布江段及其支流香曲和年楚河,采集拉萨裂腹鱼样本1145尾。本文对所采样本进行了测量分析,主要研究结果如下：
1.拉萨裂腹鱼的微耳石、脊椎骨和鳃盖骨上均有年轮存在,年轮清晰度为耳石>脊椎骨>鳃盖骨。拉萨裂腹鱼的年轮每年形成一轮,形成时间为每年的3月～5月。脊椎骨和鳃盖骨与耳石之间的IAPE分别为25.33%和23.35%。1龄～20龄样本中,脊椎骨鉴定的平均年龄同耳石鉴定的平均年龄较为接近,分别为10.57龄和10.64龄,显著高于鳃盖骨所鉴定的平均年龄9.65龄；大于20龄时,耳石鉴定的平均年龄(27.75龄)显著高于脊椎骨(18.50龄)和鳃盖骨(19.50龄)所鉴定的年龄。相比较而言,拉萨裂腹鱼年龄鉴定的最佳材料是微耳石,脊椎骨次之,鳃盖骨最差。
2.1145尾渔获物的体长范围为41mm～642mm,体重范围为1.1g～3788.1g,年龄组成为1龄～40龄,雌鱼的最大年龄为40龄,雄鱼的最大年龄为37龄。拉萨裂腹鱼的丰满度和含脂量随月份变化显著(p<0.05)。微耳石各规格参数(长度、宽度、高度和重量)与其体长的相关性均高于与年龄的相关性。
拉萨裂腹鱼的体长-体重关系式为：
性别未辨群体：W=3.664×10-5SL2.800
雌性群体：W=1.365×10-5SL2.984
雄性群体：W=1.238×10-5SL2.999
von Bertalanffy生长方程为：
雌鱼：Lt=644.3{1-exp[-0.084(t-0.247)]} Wt=3291.9{1-exp[-O.084(t-O.247)]}2.984
雄鱼：Lt=586.2{1-exp[-0.084(t+2.250)]}Wt=2477.9{1-exp[-0.084(t+2.250)]}2.999
雌鱼的生长拐点年龄为13.3龄,雄鱼的生长拐点年龄为10.8龄。
3.在繁殖季节,拉萨裂腹鱼成熟雄鱼体表具珠星；非繁殖季节,雌雄间无外形差异；拉萨裂腹鱼种群的雌雄比为1.19：1,与1：1存在显著性差异(p<0.05)。从性腺组织切片来看,拉萨裂腹鱼精巢和卵巢各分为6个时期。拉萨裂腹鱼属同步产卵类型,一年产卵一次,繁殖期较短,集中于3月～4月。拉萨裂腹鱼的绝对繁殖力为8338～50021(21693±9870)粒,相对繁殖力为5.1-22.4(13.4±4.0)粒/g；随着体长、体重和性腺重量的增加,其绝对繁殖力呈增加趋势,但与年龄的相关性不大。拉萨裂腹鱼雌鱼的最小性成熟个体体长为243mm,年龄为6龄,初次性成熟体长为479.0mm,对应年龄为13.5龄；雄鱼的最小性成熟个体体长为244mm,年龄为5龄,初次性成熟体长为369mm,对应年龄为10.2龄。
4.拉萨裂腹鱼的摄食强度随月份和体长变化差异显著(p<0.05)。拉萨裂腹鱼主要以大型无脊椎动物为食,尤其是摇蚊幼虫和纹石蛾幼虫；大型无脊椎动物在拉萨裂腹鱼肠含物中的出现率、个数百分比、重量百分比和相对重要性百分比都较其他种类食物高。拉萨裂腹鱼是广食性鱼类,其食物组成的多样性(H’)和均匀性(J)随性别的不同、体长的不同和季节的不同而有所差别；但不同个体间的食物组成差异较小,食物组成重叠程度较高。总体来看,拉萨裂腹鱼对大型无脊椎动物和小型无脊椎动物的选择指数均较高,选食程度较强,其摄食消化器官的形态与其食性是相适应的。
5.拉萨裂腹鱼雌鱼的总死亡系数、自然死亡系数和捕捞死亡系数分别为0.13/年、0.09/年和0.05/年,雄鱼分别为0.14/年、0.09/年和0.05/年；开发率雌雄均为0.35/年。雌鱼的补充年龄和起捕年龄分别为3.43龄和6龄,雄鱼分别为2.00龄和8龄。拉萨裂腹鱼属比较典型的K-选择类型鱼类。目前雅鲁藏布江谢通门至仁布江段的拉萨裂腹鱼种群资源的开发程度不高,有利于种群的可持续发展。
Schizothorax waltoni (Regan) belongs to the genus of Schizothorax, the subfamily of Schizothoracinae, the family of Cyprinidae and the order of Cypriniformes. S. waltoni, the unique species in China, only distributed in the upper and middle reaches of the Yarlung Tsangpo River and its tributaries. To study the biological characteristics of S. waltoni and its population ecology,1145individuals were collected from the Yarlung Tsangpo River and its tributaries (Xiang Qu and Nyang Qu) monthly from August2008to August2009. The main results are as follows:
1. The typical pattern with translucent zones that alternated with opaque zones was found in otoliths, vertebrae and opercular bones of S. waltoni. The definition for annulus: otoliths> vertebrae> opercular bones. Annuli formed between March and May once a year in all of the three calcified structures. Between the vertebrae and otoliths, the I APE value was25.33%. Between the opercular bones and otoliths, the IAPE value was23.35%. For age classes1-20yeras, the average age is relatively close between vertebrae (10.57years) and otolith (10.64years), significantly higher than the opercular bones (9.65years)(p<0.05). For age classes>20years, the average age of the otolith (27.75years) significantly higher than that of vertebrae (18.50years) and opercular bone (19.50years)(p<0.05). In comparison, otolith is the best material for the age estimation of S. waltoni.
2. The standard length (SL) of S. waltoni ranged from41mm to642mm, and weight (W) ranged from1.1g to3788.1g. The estimated age for females was4years to40years, male was4years to37years and undetermined was1years to9years. The fullness and coefficient of fat differed monthly in significantly (p<0.05). The correlations of otolith length/breadth/thickness/weight and fish length were higher than those of age.
The SL-W relationship of S. waltoni were described as:W=1.365×10-5SL2984for females, W=1.238×10-5SL2.999for males, and W=3.664×10-5SL2.800for undetermined.
The von Bertalanffy growth function for S. waltoni were described as:Lt=644.3{1-exp [-0.084(t-0.247)]} and Wt=3291.9{1-exp [-0.084(t-0.247)]}2.984for females, and Lt=586.2{1-exp [-0.084(t+2.250)]} and Wt=2477.9{1-exp [-0.084(t+2.250)]}2.999for males.
The age inflexion point for the growth of female was13.3years, while it for male was10.8years.
3. There is no shape difference between male and female except the spawning seasoa In spawing season, pearl organs cound be found in males but not in females. The ratio for female and male was1.19:1, which differed from1:1significantly (p<0.05). The gonad development of S. waltoni could be divided into6stages. S. waltoni were single spawners with a short-term spawning, they spawned once a year during March and April. The absolute fecundity of S. waltoni ranged from8338eggs to50021eggs, the relative fecundity ranged from5.1/g to22.4/g body weight. With the increase of standard length, body weight, and ovary weight, the absolute fecundity increased, but not correlation with ages. The standard length and age for the smallest sexual maturation female were243mm and6years, the first maturity of female were479.0mm and13.5years. The standard length and age for the smallest sexual maturation male were244mm and5years, the first maturity male were369.0mm and10.2years.
4. The ingest intension of S. waltoni varied significantly over month and length classes (p<0.05). Macroinvertebrates was the main food for S. waltoni, especially the chironomid larvae and hydropsychidae larvae. The occurrence rate, the number percentage, weight percentage and relative importance percentage for macroinvertebrates were higher than other kinds of food. S. waltoni was a generalized food fish with high diet overlap within individuals. The feeding diversity (H') and evenness (J) were different with season, fish size and sex. For S. waltoni, the selection index of macroinvertebrates and small invertebrates were higher, the shape of digestive organs could meet the needs of its feeding habits.
5. For females, the total mortality was0.13/year, natural mortality was0.09/year, fishing mortality was0.05/year and exploitation level was0.35/year. For males, the total mortality was0.14/year, natural mortality was0.09/year, fishing mortality was0.05/year and exploitation level was0.35/year. The tr and tcfor females were3.43years and6years, those for males were2years and8years. The life history pattern of S. waltoni belonged to k-selection. At present, the resource of S. waltoni had been fished properly in the Yarlung Tsangpo River.
1.拉萨裂腹鱼的微耳石、脊椎骨和鳃盖骨上均有年轮存在,年轮清晰度为耳石>脊椎骨>鳃盖骨。拉萨裂腹鱼的年轮每年形成一轮,形成时间为每年的3月～5月。脊椎骨和鳃盖骨与耳石之间的IAPE分别为25.33%和23.35%。1龄～20龄样本中,脊椎骨鉴定的平均年龄同耳石鉴定的平均年龄较为接近,分别为10.57龄和10.64龄,显著高于鳃盖骨所鉴定的平均年龄9.65龄；大于20龄时,耳石鉴定的平均年龄(27.75龄)显著高于脊椎骨(18.50龄)和鳃盖骨(19.50龄)所鉴定的年龄。相比较而言,拉萨裂腹鱼年龄鉴定的最佳材料是微耳石,脊椎骨次之,鳃盖骨最差。
2.1145尾渔获物的体长范围为41mm～642mm,体重范围为1.1g～3788.1g,年龄组成为1龄～40龄,雌鱼的最大年龄为40龄,雄鱼的最大年龄为37龄。拉萨裂腹鱼的丰满度和含脂量随月份变化显著(p<0.05)。微耳石各规格参数(长度、宽度、高度和重量)与其体长的相关性均高于与年龄的相关性。
拉萨裂腹鱼的体长-体重关系式为：
性别未辨群体：W=3.664×10-5SL2.800
雌性群体：W=1.365×10-5SL2.984
雄性群体：W=1.238×10-5SL2.999
von Bertalanffy生长方程为：
雌鱼：Lt=644.3{1-exp[-0.084(t-0.247)]} Wt=3291.9{1-exp[-O.084(t-O.247)]}2.984
雄鱼：Lt=586.2{1-exp[-0.084(t+2.250)]}Wt=2477.9{1-exp[-0.084(t+2.250)]}2.999
雌鱼的生长拐点年龄为13.3龄,雄鱼的生长拐点年龄为10.8龄。
3.在繁殖季节,拉萨裂腹鱼成熟雄鱼体表具珠星；非繁殖季节,雌雄间无外形差异；拉萨裂腹鱼种群的雌雄比为1.19：1,与1：1存在显著性差异(p<0.05)。从性腺组织切片来看,拉萨裂腹鱼精巢和卵巢各分为6个时期。拉萨裂腹鱼属同步产卵类型,一年产卵一次,繁殖期较短,集中于3月～4月。拉萨裂腹鱼的绝对繁殖力为8338～50021(21693±9870)粒,相对繁殖力为5.1-22.4(13.4±4.0)粒/g；随着体长、体重和性腺重量的增加,其绝对繁殖力呈增加趋势,但与年龄的相关性不大。拉萨裂腹鱼雌鱼的最小性成熟个体体长为243mm,年龄为6龄,初次性成熟体长为479.0mm,对应年龄为13.5龄；雄鱼的最小性成熟个体体长为244mm,年龄为5龄,初次性成熟体长为369mm,对应年龄为10.2龄。
4.拉萨裂腹鱼的摄食强度随月份和体长变化差异显著(p<0.05)。拉萨裂腹鱼主要以大型无脊椎动物为食,尤其是摇蚊幼虫和纹石蛾幼虫；大型无脊椎动物在拉萨裂腹鱼肠含物中的出现率、个数百分比、重量百分比和相对重要性百分比都较其他种类食物高。拉萨裂腹鱼是广食性鱼类,其食物组成的多样性(H’)和均匀性(J)随性别的不同、体长的不同和季节的不同而有所差别；但不同个体间的食物组成差异较小,食物组成重叠程度较高。总体来看,拉萨裂腹鱼对大型无脊椎动物和小型无脊椎动物的选择指数均较高,选食程度较强,其摄食消化器官的形态与其食性是相适应的。
5.拉萨裂腹鱼雌鱼的总死亡系数、自然死亡系数和捕捞死亡系数分别为0.13/年、0.09/年和0.05/年,雄鱼分别为0.14/年、0.09/年和0.05/年；开发率雌雄均为0.35/年。雌鱼的补充年龄和起捕年龄分别为3.43龄和6龄,雄鱼分别为2.00龄和8龄。拉萨裂腹鱼属比较典型的K-选择类型鱼类。目前雅鲁藏布江谢通门至仁布江段的拉萨裂腹鱼种群资源的开发程度不高,有利于种群的可持续发展。
Schizothorax waltoni (Regan) belongs to the genus of Schizothorax, the subfamily of Schizothoracinae, the family of Cyprinidae and the order of Cypriniformes. S. waltoni, the unique species in China, only distributed in the upper and middle reaches of the Yarlung Tsangpo River and its tributaries. To study the biological characteristics of S. waltoni and its population ecology,1145individuals were collected from the Yarlung Tsangpo River and its tributaries (Xiang Qu and Nyang Qu) monthly from August2008to August2009. The main results are as follows:
1. The typical pattern with translucent zones that alternated with opaque zones was found in otoliths, vertebrae and opercular bones of S. waltoni. The definition for annulus: otoliths> vertebrae> opercular bones. Annuli formed between March and May once a year in all of the three calcified structures. Between the vertebrae and otoliths, the I APE value was25.33%. Between the opercular bones and otoliths, the IAPE value was23.35%. For age classes1-20yeras, the average age is relatively close between vertebrae (10.57years) and otolith (10.64years), significantly higher than the opercular bones (9.65years)(p<0.05). For age classes>20years, the average age of the otolith (27.75years) significantly higher than that of vertebrae (18.50years) and opercular bone (19.50years)(p<0.05). In comparison, otolith is the best material for the age estimation of S. waltoni.
2. The standard length (SL) of S. waltoni ranged from41mm to642mm, and weight (W) ranged from1.1g to3788.1g. The estimated age for females was4years to40years, male was4years to37years and undetermined was1years to9years. The fullness and coefficient of fat differed monthly in significantly (p<0.05). The correlations of otolith length/breadth/thickness/weight and fish length were higher than those of age.
The SL-W relationship of S. waltoni were described as:W=1.365×10-5SL2984for females, W=1.238×10-5SL2.999for males, and W=3.664×10-5SL2.800for undetermined.
The von Bertalanffy growth function for S. waltoni were described as:Lt=644.3{1-exp [-0.084(t-0.247)]} and Wt=3291.9{1-exp [-0.084(t-0.247)]}2.984for females, and Lt=586.2{1-exp [-0.084(t+2.250)]} and Wt=2477.9{1-exp [-0.084(t+2.250)]}2.999for males.
The age inflexion point for the growth of female was13.3years, while it for male was10.8years.
3. There is no shape difference between male and female except the spawning seasoa In spawing season, pearl organs cound be found in males but not in females. The ratio for female and male was1.19:1, which differed from1:1significantly (p<0.05). The gonad development of S. waltoni could be divided into6stages. S. waltoni were single spawners with a short-term spawning, they spawned once a year during March and April. The absolute fecundity of S. waltoni ranged from8338eggs to50021eggs, the relative fecundity ranged from5.1/g to22.4/g body weight. With the increase of standard length, body weight, and ovary weight, the absolute fecundity increased, but not correlation with ages. The standard length and age for the smallest sexual maturation female were243mm and6years, the first maturity of female were479.0mm and13.5years. The standard length and age for the smallest sexual maturation male were244mm and5years, the first maturity male were369.0mm and10.2years.
4. The ingest intension of S. waltoni varied significantly over month and length classes (p<0.05). Macroinvertebrates was the main food for S. waltoni, especially the chironomid larvae and hydropsychidae larvae. The occurrence rate, the number percentage, weight percentage and relative importance percentage for macroinvertebrates were higher than other kinds of food. S. waltoni was a generalized food fish with high diet overlap within individuals. The feeding diversity (H') and evenness (J) were different with season, fish size and sex. For S. waltoni, the selection index of macroinvertebrates and small invertebrates were higher, the shape of digestive organs could meet the needs of its feeding habits.
5. For females, the total mortality was0.13/year, natural mortality was0.09/year, fishing mortality was0.05/year and exploitation level was0.35/year. For males, the total mortality was0.14/year, natural mortality was0.09/year, fishing mortality was0.05/year and exploitation level was0.35/year. The tr and tcfor females were3.43years and6years, those for males were2years and8years. The life history pattern of S. waltoni belonged to k-selection. At present, the resource of S. waltoni had been fished properly in the Yarlung Tsangpo River.
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