尖裸鲤个体生物学和种群动态学研究
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摘要
2008年8月至2009年8月间,于西藏雅鲁藏布江仁布县至谢通门江段共采集尖裸鲤(Oxygymnocypris stewartii)712尾,通过对采集样本的测定和分析,研究了尖裸鲤的生活史和种群动态学。此外,2008年至2012年在美国密歇根湖的Waukegan. Sturgeon Bay和Sleeping Bear Dunes三个采样点共采集666尾新黑口虾虎鱼(Neogobius melanostomus),通过对采集样本的分析,研究了新黑口虾虎鱼的年龄和生长并初步估算了其死亡率。主要研究结果如下：
1.耳石和脊椎骨均呈现出宽带和窄带交替排列的年轮特征。耳石年轮形成于每年的3月-6月,脊椎骨的年轮形成于每年的3月～5月。耳石上的年轮较为清晰。和脊椎骨相比,耳石作为年龄鉴定材料具有较高的精确性和准确性。因此,耳石是尖裸鲤最佳的年龄鉴定材料,脊椎骨次之。
2.雌性渔获物体长范围为116mm～587mm,年龄组范围为2龄～25龄；雄性渔获物体长范围为167mm～455mm,年龄组范围为3龄～17龄；性别未辨渔获物体长范围为45mm～260mm,年龄组范围为1龄～6龄。体长与体重关系式为：雌性：BW=6.108×10-6SL3.126;雄性.BW=9.872×10-6SL3.052;性别未辨.BW=3.203×10-5SL2.821。 von Bertalanffy生长方程为：雌性：Lt=618.2[1-e-0.106(t-0.315)], Wt=3065.9[1e-0.106(t-0.315)]3.126;雄性：Lt=526.8[1-e-0.141(t-0.491)], Wt=1868.4[1-e-0.141(t-141(t-0.491)]3.052。雌鱼和雄鱼的生长拐点年龄分别为11.1龄和8.4龄。
3.非繁殖季节,尖裸鲤成熟雌雄个体外部形态无明显差别；繁殖季节,成熟雄性个体具珠星,而成熟雌性个体无珠星。根据外部形态特征,性腺发育分为5期；而根据显微结构特征,性腺发育分为6期,卵母细胞则分为7个时相。雌鱼的初次性成熟体长和年龄分别为357mm和7.3龄,雄鱼初次性成熟体长和年龄分别为273mm和5.1龄。尖裸鲤一年产卵一次,繁殖季节为每年的3月～4月,为分批同步产卵鱼类。绝对繁殖力(F)范围为11017粒/尾～56907粒/尾,相对繁殖力(RF)范围为15.8粒/克～40.2粒/克。绝对繁殖力(F)与体长和体重分别呈显著的线性关系,而绝对繁殖力(F)与年龄和卵巢重则无显著性关系。
4.尖裸鲤的摄食强度具有显著性的月变化差异,9月摄食强度最低,2月和3月摄食强度最高。尖裸鲤主要以鱼类和水生昆虫为食。基于相对重要指数、出现率百分比和重量百分比指标,鱼类是尖裸鲤最重要的饵料；而基于个数百分比指标,水生昆虫是尖裸鲤最重要的饵料。尖裸鲤是广食性鱼类,其种群内不同个体间食物组成差异性较大,食物重叠程度较低。尖裸鲤食物组成具有显著的体长和季节性差异,而食物组成的性别差异性不显著。饵料生物的体长范围随着尖裸鲤体长的增大而扩大,饵料生物的体长下限几乎保持恒定而饵料生物的体长的上限则随着尖裸鲤体长的增大而增大。总体上,饵料生物体长与尖裸鲤体长呈线性关系。尖裸鲤摄食消化器官的形态与其食性相适应。
5.估算的尖裸鲤雌性和雄性的周年总瞬时死亡率(Z)分别为0.40/年和0.77/年；雌鱼和雄鱼的自然死亡率(M)分别为0.19/年和0.26/年；雌鱼和雄鱼的捕捞死亡率(F)分别为0.21/年和0.51/年。在当前渔业管理政策下,尖裸鲤雌鱼和雄鱼的繁殖潜力比(SPR)分别为9%和3%,都显著性的低于下限参考点25%,这表明尖裸鲤种群资源正在被过度的开发和利用。通过模拟6种不同的渔业管理政策,我们建议采用渔业管理政策6对尖裸鲤的种群资源进行保护和利用。尖裸鲤的生活史类型偏向k-选择类型,基于其生活史策略,探讨了尖裸鲤种群资源的保护和合理利用措施。
6.采自于Sturgeon Bay新黑口虾虎鱼(Neogobius melanostomus)全长范围为48mm～131mm,估算的年龄范围为2龄～7龄；采自于Waukegan新黑口虾虎鱼全长范围为50mm-125mm,估算的年龄范围为2龄～5龄；采自于Sleeping Bear Dunes新黑口虾虎鱼全长范围为54mm-129mm,估算的年龄范围为2龄～6龄。新黑口虾虎鱼耳石半径对数转换值与其全长的对数转换值呈显著的线性关系。栖息于Sleeping Bear Dunes和Waukegan的新黑口虾虎鱼的生长率显著性高于Sturgeon Bay,而栖息于Sleeping Bear Dunes和Waukegan的新黑口虾虎鱼的生长率则无显著性差异。雄性新黑口虾虎鱼的生长率显著性高于雌性。基于渔获量曲线分析,估算的新黑口虾虎鱼的周年死亡率范围为75%～84%。相对较高的周年死亡率表明,新黑口虾虎鱼种群数量受到密歇根湖捕食者的控制。
To investigate the biology and population dynamics of O. stewartii, a total of712individuals were captured from the Yarlung Zangbo River and its tributary (Xiang Qu) monthly from August2008to August2009. Furthermore, Age and growth of round goby Neogobius melanostomus at three areas of Lake Michigan were studied by otolith analysis from a sample of666specimens collected from2008to2012. The main results are as follows:
1. Otoliths and vertebrae of O. stewartii showed the typical annulus pattern, which was characterized by alternative wide zones and narrow zones. Annuli of otoliths formed from March to June once a year, while annuli of vertebrae formed from March to May once a year. Comparing with vertebrae, annual growth increments in otoliths were more distinct. Otoliths were the more accurate and precise calcified structure for age estimation of O. stewartii, followed by vertebrae.
2. The standard length (SL) of O. stewartii ranged between116mm and587mm for female, between167mm and455mm for male and between45mm and260mm for undetermined, respectively. The estimated age ranged from2years to25year for female, from3year to17year for male and from1year to6year for undetermined, respectively. The standard length-body weight relationships were described as BW=6.108×10-6SL3.126for female, BW=9.872×10-6SL3.052for male and BW=3.203×10-5SL2.821for undetermined. The von Bertalanffy growth equations were Lt=618.2[1-e-0.106(t-0.315)] and Wt=3065.9[1-e-1.106(t-0.315)]3.126for female, and Lt=526.8[1-e-0.141(t-0.491)] and Wt=1868.4[1-e-0.141(t-0.491)]3.052for male. The estimated inflexion points were11.1years for female and8.4years for male, respectively.
3. In non-spawning season, the external morphology of individual were same for matured female and matured male; In spawning season, pearl organs could be observed on matured male, but not on matured female. Gonads were assigned to five macroscopic maturity stages. Based on the histological analysis, gonads were assigned to six microscopic maturity stages and oocytes were assigned to seven stages. The estimated the first maturity size were357mm for female and273mm for male, respectively. The estimated first maturity age were7.3years for female and5.1years for male, respectively. O. stewartii, which spawns once a year from March to April, was a single spawner. The estimated fecundity ranged between11017eggs per fish and56907eggs per fish. The relative fecundity was between15.8eggs/g and40.2eggs/g of fish body weight. The fecundity increased linearly with standard length and body weight, while fecundity was not significantly correlated with age and ovary weight.
4. The feeding intensity of O. stewartii varied significantly over month (P<0.05), which indicated that the feeding intensity was low in Septermber and high in February and March. According to the IRJ,%W and%F, fish was the most important prey, while based on the%N, aquatic insect was the most important prey. O. stewartii is a generalized predator, which relies upon a wide trophic spectrum at the population level, and is characterized by a strong individual feeding specialization. The noticeable variations were observed in O. stewartii diet composition with fish size and season, but not with gender. The range of absolute prey sizes eaten expanded with increasing body size of O. stewartii. The lower limit of prey-size remained at almost the same level, but the upper limit increased with fish size. O. stewartii showed a significant increase in maximum prey size consumed. Overall, prey size increased linearly with increasing predator size. The structure of digestive organs was fit to the feeding habits of O. stewartii.
5. The estimated annual total mortality were0.40year-1for female and0.77year-1for male, respectively; The natural mortality were0.19year-1for female and0.26year-1for male, respectively; The fishing mortality were0.21year-1for female and0.51year-1for male, respectively. Under the current management policy, the spawning potential ratio of O. stewartii were9%for female and3%for male, indicating that O. stewartii population was overfished, with a high chance of recruitment failure in the future. Through evaluating six management scenarios, the sixth management scenario might be the better management policy for O. stewartii. The life history type of O. stewartii tends to the k-selection model. Based on the life history type of O. stewartii, we suggested some management policies.
6. Total length (TL) of round goby ranged from48mm to131mm for Sturgeon Bay, from50mm to125mm for Waukegan, and from54mm to129mm for Sleeping Bear Dunes. Estimated ages of round goby ranged from2years to7years for Sturgeon Bay, from2years to5years for Waukegan, and from2years to6years for Sleeping Bear Dunes. Body-otolith relationships were calculated for each combination of area and sex. Logarithm of TL was accurately described as a linear function of logarithm of otolith radius. Back-calculated TL at age from the otolith analysis was used to fit a von Bertalanffy growth model for each combination of area and sex. For both sexes, round gobies at Sleeping Bear Dunes and Waukegan grew significantly faster than those at Sturgeon Bay. However, round goby growth did not significantly differ between Sleeping Bear Dunes and Waukegan for either sex. At all three areas of Lake Michigan, males grew significantly faster than females. Based on catch curve analysis, estimates of annual mortality rates ranged from75%to84%. These relatively high mortality rates suggested that round gobies may be under predatory control in Lake Michigan.
1.耳石和脊椎骨均呈现出宽带和窄带交替排列的年轮特征。耳石年轮形成于每年的3月-6月,脊椎骨的年轮形成于每年的3月～5月。耳石上的年轮较为清晰。和脊椎骨相比,耳石作为年龄鉴定材料具有较高的精确性和准确性。因此,耳石是尖裸鲤最佳的年龄鉴定材料,脊椎骨次之。
2.雌性渔获物体长范围为116mm～587mm,年龄组范围为2龄～25龄；雄性渔获物体长范围为167mm～455mm,年龄组范围为3龄～17龄；性别未辨渔获物体长范围为45mm～260mm,年龄组范围为1龄～6龄。体长与体重关系式为：雌性：BW=6.108×10-6SL3.126;雄性.BW=9.872×10-6SL3.052;性别未辨.BW=3.203×10-5SL2.821。 von Bertalanffy生长方程为：雌性：Lt=618.2[1-e-0.106(t-0.315)], Wt=3065.9[1e-0.106(t-0.315)]3.126;雄性：Lt=526.8[1-e-0.141(t-0.491)], Wt=1868.4[1-e-0.141(t-141(t-0.491)]3.052。雌鱼和雄鱼的生长拐点年龄分别为11.1龄和8.4龄。
3.非繁殖季节,尖裸鲤成熟雌雄个体外部形态无明显差别；繁殖季节,成熟雄性个体具珠星,而成熟雌性个体无珠星。根据外部形态特征,性腺发育分为5期；而根据显微结构特征,性腺发育分为6期,卵母细胞则分为7个时相。雌鱼的初次性成熟体长和年龄分别为357mm和7.3龄,雄鱼初次性成熟体长和年龄分别为273mm和5.1龄。尖裸鲤一年产卵一次,繁殖季节为每年的3月～4月,为分批同步产卵鱼类。绝对繁殖力(F)范围为11017粒/尾～56907粒/尾,相对繁殖力(RF)范围为15.8粒/克～40.2粒/克。绝对繁殖力(F)与体长和体重分别呈显著的线性关系,而绝对繁殖力(F)与年龄和卵巢重则无显著性关系。
4.尖裸鲤的摄食强度具有显著性的月变化差异,9月摄食强度最低,2月和3月摄食强度最高。尖裸鲤主要以鱼类和水生昆虫为食。基于相对重要指数、出现率百分比和重量百分比指标,鱼类是尖裸鲤最重要的饵料；而基于个数百分比指标,水生昆虫是尖裸鲤最重要的饵料。尖裸鲤是广食性鱼类,其种群内不同个体间食物组成差异性较大,食物重叠程度较低。尖裸鲤食物组成具有显著的体长和季节性差异,而食物组成的性别差异性不显著。饵料生物的体长范围随着尖裸鲤体长的增大而扩大,饵料生物的体长下限几乎保持恒定而饵料生物的体长的上限则随着尖裸鲤体长的增大而增大。总体上,饵料生物体长与尖裸鲤体长呈线性关系。尖裸鲤摄食消化器官的形态与其食性相适应。
5.估算的尖裸鲤雌性和雄性的周年总瞬时死亡率(Z)分别为0.40/年和0.77/年；雌鱼和雄鱼的自然死亡率(M)分别为0.19/年和0.26/年；雌鱼和雄鱼的捕捞死亡率(F)分别为0.21/年和0.51/年。在当前渔业管理政策下,尖裸鲤雌鱼和雄鱼的繁殖潜力比(SPR)分别为9%和3%,都显著性的低于下限参考点25%,这表明尖裸鲤种群资源正在被过度的开发和利用。通过模拟6种不同的渔业管理政策,我们建议采用渔业管理政策6对尖裸鲤的种群资源进行保护和利用。尖裸鲤的生活史类型偏向k-选择类型,基于其生活史策略,探讨了尖裸鲤种群资源的保护和合理利用措施。
6.采自于Sturgeon Bay新黑口虾虎鱼(Neogobius melanostomus)全长范围为48mm～131mm,估算的年龄范围为2龄～7龄；采自于Waukegan新黑口虾虎鱼全长范围为50mm-125mm,估算的年龄范围为2龄～5龄；采自于Sleeping Bear Dunes新黑口虾虎鱼全长范围为54mm-129mm,估算的年龄范围为2龄～6龄。新黑口虾虎鱼耳石半径对数转换值与其全长的对数转换值呈显著的线性关系。栖息于Sleeping Bear Dunes和Waukegan的新黑口虾虎鱼的生长率显著性高于Sturgeon Bay,而栖息于Sleeping Bear Dunes和Waukegan的新黑口虾虎鱼的生长率则无显著性差异。雄性新黑口虾虎鱼的生长率显著性高于雌性。基于渔获量曲线分析,估算的新黑口虾虎鱼的周年死亡率范围为75%～84%。相对较高的周年死亡率表明,新黑口虾虎鱼种群数量受到密歇根湖捕食者的控制。
To investigate the biology and population dynamics of O. stewartii, a total of712individuals were captured from the Yarlung Zangbo River and its tributary (Xiang Qu) monthly from August2008to August2009. Furthermore, Age and growth of round goby Neogobius melanostomus at three areas of Lake Michigan were studied by otolith analysis from a sample of666specimens collected from2008to2012. The main results are as follows:
1. Otoliths and vertebrae of O. stewartii showed the typical annulus pattern, which was characterized by alternative wide zones and narrow zones. Annuli of otoliths formed from March to June once a year, while annuli of vertebrae formed from March to May once a year. Comparing with vertebrae, annual growth increments in otoliths were more distinct. Otoliths were the more accurate and precise calcified structure for age estimation of O. stewartii, followed by vertebrae.
2. The standard length (SL) of O. stewartii ranged between116mm and587mm for female, between167mm and455mm for male and between45mm and260mm for undetermined, respectively. The estimated age ranged from2years to25year for female, from3year to17year for male and from1year to6year for undetermined, respectively. The standard length-body weight relationships were described as BW=6.108×10-6SL3.126for female, BW=9.872×10-6SL3.052for male and BW=3.203×10-5SL2.821for undetermined. The von Bertalanffy growth equations were Lt=618.2[1-e-0.106(t-0.315)] and Wt=3065.9[1-e-1.106(t-0.315)]3.126for female, and Lt=526.8[1-e-0.141(t-0.491)] and Wt=1868.4[1-e-0.141(t-0.491)]3.052for male. The estimated inflexion points were11.1years for female and8.4years for male, respectively.
3. In non-spawning season, the external morphology of individual were same for matured female and matured male; In spawning season, pearl organs could be observed on matured male, but not on matured female. Gonads were assigned to five macroscopic maturity stages. Based on the histological analysis, gonads were assigned to six microscopic maturity stages and oocytes were assigned to seven stages. The estimated the first maturity size were357mm for female and273mm for male, respectively. The estimated first maturity age were7.3years for female and5.1years for male, respectively. O. stewartii, which spawns once a year from March to April, was a single spawner. The estimated fecundity ranged between11017eggs per fish and56907eggs per fish. The relative fecundity was between15.8eggs/g and40.2eggs/g of fish body weight. The fecundity increased linearly with standard length and body weight, while fecundity was not significantly correlated with age and ovary weight.
4. The feeding intensity of O. stewartii varied significantly over month (P<0.05), which indicated that the feeding intensity was low in Septermber and high in February and March. According to the IRJ,%W and%F, fish was the most important prey, while based on the%N, aquatic insect was the most important prey. O. stewartii is a generalized predator, which relies upon a wide trophic spectrum at the population level, and is characterized by a strong individual feeding specialization. The noticeable variations were observed in O. stewartii diet composition with fish size and season, but not with gender. The range of absolute prey sizes eaten expanded with increasing body size of O. stewartii. The lower limit of prey-size remained at almost the same level, but the upper limit increased with fish size. O. stewartii showed a significant increase in maximum prey size consumed. Overall, prey size increased linearly with increasing predator size. The structure of digestive organs was fit to the feeding habits of O. stewartii.
5. The estimated annual total mortality were0.40year-1for female and0.77year-1for male, respectively; The natural mortality were0.19year-1for female and0.26year-1for male, respectively; The fishing mortality were0.21year-1for female and0.51year-1for male, respectively. Under the current management policy, the spawning potential ratio of O. stewartii were9%for female and3%for male, indicating that O. stewartii population was overfished, with a high chance of recruitment failure in the future. Through evaluating six management scenarios, the sixth management scenario might be the better management policy for O. stewartii. The life history type of O. stewartii tends to the k-selection model. Based on the life history type of O. stewartii, we suggested some management policies.
6. Total length (TL) of round goby ranged from48mm to131mm for Sturgeon Bay, from50mm to125mm for Waukegan, and from54mm to129mm for Sleeping Bear Dunes. Estimated ages of round goby ranged from2years to7years for Sturgeon Bay, from2years to5years for Waukegan, and from2years to6years for Sleeping Bear Dunes. Body-otolith relationships were calculated for each combination of area and sex. Logarithm of TL was accurately described as a linear function of logarithm of otolith radius. Back-calculated TL at age from the otolith analysis was used to fit a von Bertalanffy growth model for each combination of area and sex. For both sexes, round gobies at Sleeping Bear Dunes and Waukegan grew significantly faster than those at Sturgeon Bay. However, round goby growth did not significantly differ between Sleeping Bear Dunes and Waukegan for either sex. At all three areas of Lake Michigan, males grew significantly faster than females. Based on catch curve analysis, estimates of annual mortality rates ranged from75%to84%. These relatively high mortality rates suggested that round gobies may be under predatory control in Lake Michigan.
引文
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