东海鲐鱼（Scomber japonica）早期生活史过程的生态动力学模拟研究
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摘要
鲐鱼（Scomber japonica）是我国近海重要的中上层鱼类资源，也是我国近海海洋生态系统中重要的种类之一。环境对其补充量的影响很大。以往的研究关注于环境与成鱼资源之间的关系，但每年的补充量主要是由鱼类早期的存活决定，因此海洋环境因素的细微变化将对鱼类生命周期中最为脆弱的鱼卵和仔幼鱼的生长、成活直至种群的补充产生影响。
为此，本文在了解IBM基本理论及应用现状的基础上，综合物理海洋学和渔业资源学等学科，根据鲐鱼生长初期的物理环境，结合其生长特性，利用海洋物理模型（FVCOM）建立了基于个体的鲐鱼早期生活史过程的物理-生物耦合模型；利用该模型证实在正常气候下，台湾东北部产卵场的鱼卵是否向对马海峡海域附近输运，估计从东海南部产卵场到达对马海峡以及太平洋等育肥场的仔幼鱼比例，分析产卵场对各个育肥场补充和联通性，找出影响其输运的动力学因素；探讨极端气候下以及产卵场的变动情况下，对鲐鱼鱼卵仔幼鱼的输运以及丰度分布等影响，从而系统分析物理、生物因素对鲐鱼资源种群变动的内在动力学规律；解释鲐鱼仔幼鱼集群和成鱼渔场形成的动力学因素。本研究将为鲐鱼资源的可持续开发与利用了提供科学依据。主要研究结果如下：
（1）模拟发现，在正常气候流场的驱动下，3月份随着水温的提高，鲐鱼开始产卵，鱼卵仔鱼分布于黑潮与台湾暖流之间的弱流区，并开始向东北方向输送。4月份随着水温的进一步升高，迎来了产卵的高峰期，鱼卵仔鱼数量迅速增加，鱼卵仔鱼在黑潮和台湾暖流的影响下分成两部分，一部分会随台湾暖流向偏北方向输送，另一部分随黑潮向东北方向输运。5月份水温继续升高，绝大部分鲐鱼产卵结束，产卵数量增加减少，由于死亡的原因，数量减少，一些鱼卵仔鱼继续向北的舟山外海输运，其中一些已经被输运到了长江口和杭州湾附近海域，另一部分继续向东北方向输运，最远已经漂移到了五岛列岛西部海域，仔幼鱼绝大部分分布在50-200m等深线之间，但由于黑潮的阻隔作用，鱼卵仔鱼很难冲破锋面进入或穿过黑潮。6月份，随着水温继续升高，向偏北方向输运的仔幼鱼也开始沿着100-200m等深线转向向东北方向输运，6月末绝大部分仔幼鱼被输运到了东海东北部附近海域，只有极少数滞留在长江口和舟山外海。当到达五岛列岛海域时，仔幼鱼被洋流分成了两部分，一部分被对马暖流带进了对马海峡海域，另一部分经过九州岛西部海域被黑潮迅速带进了太平洋里。7月份海水温度继续升高，模拟也即将结束，此时鲐鱼仔幼鱼绝大部分被输运到了济州岛海域、对马海峡海域、九州岛西部海域和太平洋海域，一些由于受黑潮的挟持甚至已经漂出了计算区域，此时东海其他海域内就很少见到鲐鱼的踪迹了。研究认为，黑潮、台湾暖流和对马暖流控制着东海南部产卵场的鱼卵仔鱼的输运。
分析表明，太平洋海域育肥场由于强烈的湍流，仔鱼所处的平均水深比济州岛、对马海峡、九州岛育肥场要深，在7月末，平均水深达400多米，但由于黑潮的高温高盐特性，平均水温并不是最低的，所以进入太平洋海域育肥场的仔幼鱼虽然所处水深较深，基本上也能很好地生长。
研究表明，随着时间的推进，到达各育肥场仔幼鱼的比例情况有所变动。在6月末，大部分的仔幼鱼从东海的南部产卵场被输运到济州岛和九州岛西部海域育肥场。在7月中旬，济州岛和九州岛海域的仔幼鱼因部分输送进了对马海峡和太平洋，比例有所下降。在7月末模拟结束时，仔幼鱼被大部分输送到了济州岛和对马海峡的偏北海域育肥场，而输送到九州岛和太平洋偏南海域育肥场只占了总数的三分之一。
研究发现，东海东南部产卵场是东海西北部邻近日本海域渔场一个十分重要的补充群体，但该产卵场对我国长江口和杭州湾外海的渔场补充量作用却十分有限。鱼卵仔鱼的输运是联通产卵场和育肥场的主要机制，此鱼卵仔鱼的联通关系同该区域的洋流划分系统十分相似，分为两部份，偏北的对马海峡和济州岛育肥场的仔幼鱼主要来自于偏东北A和B部分产卵场，而偏南九州岛和太平洋育肥场则主要来自于偏西南C和D部分产卵场，这证明了物理因素对联通性有很大影响。
（2）在台风风场产生流场驱动下，利用IBM对鲐鱼鱼卵仔鱼输运、分布进行研究。模拟结果显示，在台风过后，鱼卵仔鱼的分布与原来相比并没有明显的差异。
为了验证水平分布结果的真实性，消除生物个体的随机性和所处水深的影响，做了理想化的实验和理论验证。研究分析结果显示，确实台风对仔幼鱼分布的影响不大，原因是鲐鱼鱼卵仔幼鱼输运路径正处在台风影响最小的区域内，台风作用的时间较短以及台风气旋与潮流的反气旋旋转相互抵消减少了台风对仔幼鱼输运区域低频率潮流的影响。
研究发现，台风会使丰度分布和输运速度在输运路径的后端有所差异，使高密度的鱼卵仔鱼范围向东北方向上扩散和增大输运路径后端鱼卵仔鱼输运速度的趋势。又作了加长台风作用时间的数值实验，发现台风对仔幼鱼分布有一定影响。更改了台风作用时间的实验，得到了同样的结果。台风使鱼卵仔鱼向深水移动的趋势，并使仔幼鱼的死亡率增大。
（3）产卵场位置的变动，对鱼卵仔幼鱼的输运分布产生很大的影响，偏西的产卵场由于受台湾暖流的影响较大，输运整体偏西北，在输运的过程中有大量仔幼鱼在中国沿海滞留和过境，最后绝大部分输运到了偏北的济州岛育肥场。偏东的产卵场由于受黑潮影响较大，整体输运偏东南，几乎就没有鱼卵仔鱼漂移到中国沿海附近，并且漂移速度很快，最后输运到偏南太平洋育肥场数量增多。台湾暖流和黑潮影响偏西和偏东产卵场鱼卵仔鱼的输运速度。正常产卵位置在3个产卵场中的存活机率和生长方面都是最佳的。
产卵深度的变动对鱼卵仔鱼的输运和丰度分布没有太大的影响，但5m和15m深产卵的死亡率有所增加，正常产卵深度（10m）是最佳产卵深度。
（4）通过确定了鲐鱼仔幼鱼的运动规则，首次建立起了具有游泳能力的鲐鱼IBM模型。结果显示，具有游泳能力仔幼鱼前期对输运的影响不大，后期随着增强的游泳能力，逐渐有集群现象出现，使死亡率降低，向东北输运速度降低，输运到太平洋和日本海的幼鱼数量都有所下降，平均所处水深降低，更适应生长。
研究发现，产卵位置的变动使偏西产卵场的集群受台湾暖流影响较大，导致集群偏西。偏东产卵场受黑潮影响较大，集群偏东。说明物理环境和生物因素同样会对具有游泳能力仔幼鱼的输运产生影响。
（4）对成体产卵鲐鱼进行洄游分析发现，鲐鱼处在20m适宜水层，台湾东北部鲐鱼集群主要受台湾暖流、大陆沿岸水、黑潮影响。在台湾暖流和沿岸水交汇的锋面附近有大量的鲐鱼分布，在黑潮与中国大陆沿岸水形成的潮境区域也有大量的鲐鱼分布。
研究表明，产卵场位置的变动使偏西产卵场由于受台湾暖流影响较大，在台湾暖流和沿岸水的锋面附近有大量聚集，而偏东的产卵受黑潮影响较大，会在黑潮形成的锋面附近进行大量的集群。所以鲐鱼所处位置的不同，会影响其集群的位置，说明物理环境和生物因素也会对成鱼的洄游和集群产生影响。
Chub mackerel (Scomber japonica) is important pelagic fish resources inthe China Sea. Changes of fish resources are not entirely influenced byfishing, but the environment has a big effect on recruitment. Past researchhas focused on relatinship between environment and resource, but annualrecruitment is mainly determined by the survival in fish early life stage.Minor change of marine environment will have an impact on growth,surivival and recruitment of eggs and larvae, which is the most vulnerable inthe fish life stage. Application of based on individual-based model to studythe growth of fish early history has been relatively developed at abroad, butin China, relative research is not well developed.
This project to make the interdisciplinary research, according to thephysical environment of the early growth mackerel, combined with itsgrowth characteristics, using ocean physical model, and developsphysical-ecological coupling model of early life history of chub mackerelbased on individual, and the research on the growth, transport, migration,connectivity and recruitment of chub mackerel eggs and larvae, explore theinner dynamics factor of population resources fluctuation, in order to accurately predict mackerel resource and realize biological resource recoveryin China Sea provide the scientific basis.
Based on introduced individual-based model, this paper develops abio-physical dynamic model of early life history of chub mackerel in the EastChina Sea (ECS). The physical model is developed based on the FVCOM(Finite Volume Coast and Ocean Model) simulates the3-D physical fields,using3d,10km resolution march average temperature and salinity initialfields, eight main tidal constituent on the open boundaries. The biologicalmodel uses on individual-based models (IBM). Early life of chub mackerel isdefined by five stages according to age and length, parameterized early lifebiological process (spawn, growth, survival, etc) of chub mackerel of ECS.The eggs hatch time and larval metamorphosis time is the function of watertemperature in the model. Growth is the function of temperature and food.The mortality rate was a negative correlation with the length and a positivecorrelation with the growth rate. Food availability is calculated from thecoastal upwelling index. Estimated spawning ground is the southwest of theECS (Taiwan northeast). Spawning period is from March to June. Batchspawning eggs is released in the10m water layer. The physical field duringMarch-July yielded from the physical model is coupled with the biologicalmodel through the Lagrange particle tracking, considering the passive driftby current and random walk by turbulence. Using the super-individualtechnology, the individuals are grown when the particle drifts passively. The application of the model for normal climate, extreme climate and the changeof spawning ground study transport of eggs and larvae, to study dynamicsfactors of the changes effect. The main research results are as follows:
The results of flow field under normal climatological forcing conditionshow that with the increase of water temperature in March, chub mackerelbegan to spawn. Eggs are distribution in between Taiwan Warm Current(TWC) and the Kuroshio, and transported to the northeast. In April with thewater temperature further raise, the spawning peak is coming. Eggs increasedrapidly. Eggs and larvae were divided into two parts by the influence ofTWC and kuroshio. One can follow TWC to northward transportation. Theother part can follow the Kuroshio to northeast direction to transport. Watertemperature continues to rise in May, the most mackerel spawn over. Thenumber of spawning increase is reduced, because of death. The number ofeggs and larvae is decrease. Some continue to transport north for ZhoushanIsland. There is very little has been transported to the Yangtze Estuary andthe Hangzhou Bay. The other part also continues to transport northeast. Thefarthest drift is in the western Goto-retto islands. Most of larvae are thedistribution in between50m and200m isobaths. But because of the kuroshiobarrier, it is difficult to break into, or through the kuroshio to larvea. Watertemperature continues to rise in June. Part of the larvae to transport northalso began to turn to the northeast transport with other along the100-200misoaths. Most of larvae was transported to northeast of ECS in end of June only a little of larvae stranded in the Yangtze Estuary and Zhoushan Sea.When larvae get to the Goto-retto islands, larvae are divided into two partsby tidal currents, and a part was broughted into Tsushima Strait by TsushimaStrait Warm Current (TSWC), the other part to go through western sea ofKyushu was quickly broughted into the Pacific by the Kroshio. In July, watertemperatures continue to rise. The simulation is coming to the end. The mostof larvae was transported to Cheju, Tsushima Strait, Kyushu Island and thePacific Ocean. Some has drift out calculation domain by the kuroshio.Larvae were rarely found in this time in other ECS. The transportation oflarvae is significantly influenced by complex hydrodynamic TWC, Kuroshioand TSWC.
Statistical analysis showed that the mean water depth of larvae is deeperthan other three nursery grounds because of strong turbulence of nurseryground in the Pacific Ocean,. In end of July, average depth exceeds400m.Due to characteristics of the high temperature and salinity of the Kuroshio,the average water temperature is not the lowest. Juvenile basically can also isvery good to grow, that was transported into nursery ground in the PacificOcean.
With the advance of time, the proportion of larvae reached each parts ofnursery ground will has variation. In the end of June, most of larvae weretransported from spawning ground in south ECS to nursery ground of ChejuIsland and Kyushu Island. In mid-july, the proportion of nursery of Cheju Island and Kyushu Island is decreased becase of transport into nurseryground of the Pacific Ocean and Tsushima Strait. In the end of the simulationin July, most of larvae were transported to north nursery ground of Kyushuand Tsushima Strait. But only a third of the total larvae were transported tothe south nursery ground of Pacific Ocean snd Kyushu.
The spawning ground in the southern East China Sea made morecontributions to the recruitment to the fishing ground in northeast East ChinaSea, but less to the Yangtze estuary and Zhou-Shan Island. The transport ofeggs and larvae is the main mechanism of connection between spawningground and nursery ground. Connectivity is very similar to division of theocean currents system in ECS, divided into two parts. Northwest part(Section A) and Southwest part (Section B) of spawning ground had strongconnectivity with the nursery grounds of Cheju and Tsushima Straits.Northeast part (Section C) and Southeast part (Section D) of spawningground had strong connectivity with the nursery grounds of Kyushu andPacific Oceans. This is proof that physical factors impacts on connectivity.
Flow field under the typhoon forcing drives IBM to study transportdistribution of eggs and larvae of chub mackerel. The result showed that nosignificant difference distribution of eggs and larvae is found forclimatological forcing condition after a typhoon pass. But the typhooninfluence on water layer and survival of larvae. In order to verify theauthenticity of the horizontal distribution, eliminate the stochastic and individual water depth. We do the idealistic experiment. The results showedthat the little impact of larval distribution is really. The reason is transportpath of lavae is in the area of the typhoon minimum affect. Typhoon sweptthe ECS is a short period. The cancellation of tide-induced anti-cyclonic andtyphoon-driven cyclonic currents reduced its influence on the low-frequencycurrents and thus on larval transport in this region. Some difference is foundin larval abundance and velocity of transport in back end of transport path,where high density larvae northeast drift and velocity of transport increased.The typhoon caused higth mortality, to move to deeper water layer. Addingan experiment of the longer the typhoon duration, we observe that typhooncertain effect to larval distribution.
The change of position has great influence on transport distribution ofeggs and larvae. The west spawning ground was great influenced by Taiwanwarm current. Larval transport was northwest. A large amount of eggs andlarvae was retention and transit in coastal China in the process of transport.The most of larvae were finally transported to north nursery ground of ChejuIsland. The east spawning ground was great influenced by the Kuroshio.Larval transport was southeast. Almost no eggs and larvae was drift incoastal China. Speed of drift is very fast. A large amount of larvae werefinally transported to south nursery ground of Pacific Ocean. Transport speedof wast and east spawning ground was influenced by Taiwan warm currentand the Kuroshio. Survival and growth of normal spawning ground are the best in three spawning ground. The the change of spawning depth has noeffect on larval distribution and transport. But mortality rate of5m and15mdeep increases. Normal spawning depth (10m) is the best spawning depth.
We set the movement rule and develop IBM including a swimmingabilities submodel. The result shows that larval swimming ability had littleinfluence in the early transport stage. With the increase swimming ability inthe later stage, larvae have appeared to shcool. Larval mortality is reduced.Speed of northeast transport is also reduced. The amount of transport to thePacific Ocean and the Janpan Sea is decline. Average water depth is lower,which is more suitable to grow. The change of spawning ground make thewest spawning ground was influenced by Taiwan warm current. Larvaeschool in west. East spawning ground was influenced by the Kuroshio.Larvae school in eest. It is to explain the physical environment and biologicalfactors will also have effect to transport of swimming ability of larvae.
We study migration of adult spawning mackerel. The result show suitabledepth of mackerel is20m. School of mackerel in northeastern Taiwan wasmainly influenced by TWC, mainland coast water, the Kuroshio. There are alarge number of mackerel distributionn in the intersection front of TWC andcoast water. There are aslo a large number of mackerel distributionn in tidearea of coast water and the Kuroshio. The change position of spawningground is west spawning ground have large accumulation in front of TWCand coast water becase of large influenced by TWC, and east spawning ground have large accumulation in front of the Kuroshio becase of largeinfluenced by the kuroshio. Different locations of mackerel will affect theposition of school, it that explain the physical environment and biologicalfactors can influence on the migration and school of mackerel.
为此，本文在了解IBM基本理论及应用现状的基础上，综合物理海洋学和渔业资源学等学科，根据鲐鱼生长初期的物理环境，结合其生长特性，利用海洋物理模型（FVCOM）建立了基于个体的鲐鱼早期生活史过程的物理-生物耦合模型；利用该模型证实在正常气候下，台湾东北部产卵场的鱼卵是否向对马海峡海域附近输运，估计从东海南部产卵场到达对马海峡以及太平洋等育肥场的仔幼鱼比例，分析产卵场对各个育肥场补充和联通性，找出影响其输运的动力学因素；探讨极端气候下以及产卵场的变动情况下，对鲐鱼鱼卵仔幼鱼的输运以及丰度分布等影响，从而系统分析物理、生物因素对鲐鱼资源种群变动的内在动力学规律；解释鲐鱼仔幼鱼集群和成鱼渔场形成的动力学因素。本研究将为鲐鱼资源的可持续开发与利用了提供科学依据。主要研究结果如下：
（1）模拟发现，在正常气候流场的驱动下，3月份随着水温的提高，鲐鱼开始产卵，鱼卵仔鱼分布于黑潮与台湾暖流之间的弱流区，并开始向东北方向输送。4月份随着水温的进一步升高，迎来了产卵的高峰期，鱼卵仔鱼数量迅速增加，鱼卵仔鱼在黑潮和台湾暖流的影响下分成两部分，一部分会随台湾暖流向偏北方向输送，另一部分随黑潮向东北方向输运。5月份水温继续升高，绝大部分鲐鱼产卵结束，产卵数量增加减少，由于死亡的原因，数量减少，一些鱼卵仔鱼继续向北的舟山外海输运，其中一些已经被输运到了长江口和杭州湾附近海域，另一部分继续向东北方向输运，最远已经漂移到了五岛列岛西部海域，仔幼鱼绝大部分分布在50-200m等深线之间，但由于黑潮的阻隔作用，鱼卵仔鱼很难冲破锋面进入或穿过黑潮。6月份，随着水温继续升高，向偏北方向输运的仔幼鱼也开始沿着100-200m等深线转向向东北方向输运，6月末绝大部分仔幼鱼被输运到了东海东北部附近海域，只有极少数滞留在长江口和舟山外海。当到达五岛列岛海域时，仔幼鱼被洋流分成了两部分，一部分被对马暖流带进了对马海峡海域，另一部分经过九州岛西部海域被黑潮迅速带进了太平洋里。7月份海水温度继续升高，模拟也即将结束，此时鲐鱼仔幼鱼绝大部分被输运到了济州岛海域、对马海峡海域、九州岛西部海域和太平洋海域，一些由于受黑潮的挟持甚至已经漂出了计算区域，此时东海其他海域内就很少见到鲐鱼的踪迹了。研究认为，黑潮、台湾暖流和对马暖流控制着东海南部产卵场的鱼卵仔鱼的输运。
分析表明，太平洋海域育肥场由于强烈的湍流，仔鱼所处的平均水深比济州岛、对马海峡、九州岛育肥场要深，在7月末，平均水深达400多米，但由于黑潮的高温高盐特性，平均水温并不是最低的，所以进入太平洋海域育肥场的仔幼鱼虽然所处水深较深，基本上也能很好地生长。
研究表明，随着时间的推进，到达各育肥场仔幼鱼的比例情况有所变动。在6月末，大部分的仔幼鱼从东海的南部产卵场被输运到济州岛和九州岛西部海域育肥场。在7月中旬，济州岛和九州岛海域的仔幼鱼因部分输送进了对马海峡和太平洋，比例有所下降。在7月末模拟结束时，仔幼鱼被大部分输送到了济州岛和对马海峡的偏北海域育肥场，而输送到九州岛和太平洋偏南海域育肥场只占了总数的三分之一。
研究发现，东海东南部产卵场是东海西北部邻近日本海域渔场一个十分重要的补充群体，但该产卵场对我国长江口和杭州湾外海的渔场补充量作用却十分有限。鱼卵仔鱼的输运是联通产卵场和育肥场的主要机制，此鱼卵仔鱼的联通关系同该区域的洋流划分系统十分相似，分为两部份，偏北的对马海峡和济州岛育肥场的仔幼鱼主要来自于偏东北A和B部分产卵场，而偏南九州岛和太平洋育肥场则主要来自于偏西南C和D部分产卵场，这证明了物理因素对联通性有很大影响。
（2）在台风风场产生流场驱动下，利用IBM对鲐鱼鱼卵仔鱼输运、分布进行研究。模拟结果显示，在台风过后，鱼卵仔鱼的分布与原来相比并没有明显的差异。
为了验证水平分布结果的真实性，消除生物个体的随机性和所处水深的影响，做了理想化的实验和理论验证。研究分析结果显示，确实台风对仔幼鱼分布的影响不大，原因是鲐鱼鱼卵仔幼鱼输运路径正处在台风影响最小的区域内，台风作用的时间较短以及台风气旋与潮流的反气旋旋转相互抵消减少了台风对仔幼鱼输运区域低频率潮流的影响。
研究发现，台风会使丰度分布和输运速度在输运路径的后端有所差异，使高密度的鱼卵仔鱼范围向东北方向上扩散和增大输运路径后端鱼卵仔鱼输运速度的趋势。又作了加长台风作用时间的数值实验，发现台风对仔幼鱼分布有一定影响。更改了台风作用时间的实验，得到了同样的结果。台风使鱼卵仔鱼向深水移动的趋势，并使仔幼鱼的死亡率增大。
（3）产卵场位置的变动，对鱼卵仔幼鱼的输运分布产生很大的影响，偏西的产卵场由于受台湾暖流的影响较大，输运整体偏西北，在输运的过程中有大量仔幼鱼在中国沿海滞留和过境，最后绝大部分输运到了偏北的济州岛育肥场。偏东的产卵场由于受黑潮影响较大，整体输运偏东南，几乎就没有鱼卵仔鱼漂移到中国沿海附近，并且漂移速度很快，最后输运到偏南太平洋育肥场数量增多。台湾暖流和黑潮影响偏西和偏东产卵场鱼卵仔鱼的输运速度。正常产卵位置在3个产卵场中的存活机率和生长方面都是最佳的。
产卵深度的变动对鱼卵仔鱼的输运和丰度分布没有太大的影响，但5m和15m深产卵的死亡率有所增加，正常产卵深度（10m）是最佳产卵深度。
（4）通过确定了鲐鱼仔幼鱼的运动规则，首次建立起了具有游泳能力的鲐鱼IBM模型。结果显示，具有游泳能力仔幼鱼前期对输运的影响不大，后期随着增强的游泳能力，逐渐有集群现象出现，使死亡率降低，向东北输运速度降低，输运到太平洋和日本海的幼鱼数量都有所下降，平均所处水深降低，更适应生长。
研究发现，产卵位置的变动使偏西产卵场的集群受台湾暖流影响较大，导致集群偏西。偏东产卵场受黑潮影响较大，集群偏东。说明物理环境和生物因素同样会对具有游泳能力仔幼鱼的输运产生影响。
（4）对成体产卵鲐鱼进行洄游分析发现，鲐鱼处在20m适宜水层，台湾东北部鲐鱼集群主要受台湾暖流、大陆沿岸水、黑潮影响。在台湾暖流和沿岸水交汇的锋面附近有大量的鲐鱼分布，在黑潮与中国大陆沿岸水形成的潮境区域也有大量的鲐鱼分布。
研究表明，产卵场位置的变动使偏西产卵场由于受台湾暖流影响较大，在台湾暖流和沿岸水的锋面附近有大量聚集，而偏东的产卵受黑潮影响较大，会在黑潮形成的锋面附近进行大量的集群。所以鲐鱼所处位置的不同，会影响其集群的位置，说明物理环境和生物因素也会对成鱼的洄游和集群产生影响。
Chub mackerel (Scomber japonica) is important pelagic fish resources inthe China Sea. Changes of fish resources are not entirely influenced byfishing, but the environment has a big effect on recruitment. Past researchhas focused on relatinship between environment and resource, but annualrecruitment is mainly determined by the survival in fish early life stage.Minor change of marine environment will have an impact on growth,surivival and recruitment of eggs and larvae, which is the most vulnerable inthe fish life stage. Application of based on individual-based model to studythe growth of fish early history has been relatively developed at abroad, butin China, relative research is not well developed.
This project to make the interdisciplinary research, according to thephysical environment of the early growth mackerel, combined with itsgrowth characteristics, using ocean physical model, and developsphysical-ecological coupling model of early life history of chub mackerelbased on individual, and the research on the growth, transport, migration,connectivity and recruitment of chub mackerel eggs and larvae, explore theinner dynamics factor of population resources fluctuation, in order to accurately predict mackerel resource and realize biological resource recoveryin China Sea provide the scientific basis.
Based on introduced individual-based model, this paper develops abio-physical dynamic model of early life history of chub mackerel in the EastChina Sea (ECS). The physical model is developed based on the FVCOM(Finite Volume Coast and Ocean Model) simulates the3-D physical fields,using3d,10km resolution march average temperature and salinity initialfields, eight main tidal constituent on the open boundaries. The biologicalmodel uses on individual-based models (IBM). Early life of chub mackerel isdefined by five stages according to age and length, parameterized early lifebiological process (spawn, growth, survival, etc) of chub mackerel of ECS.The eggs hatch time and larval metamorphosis time is the function of watertemperature in the model. Growth is the function of temperature and food.The mortality rate was a negative correlation with the length and a positivecorrelation with the growth rate. Food availability is calculated from thecoastal upwelling index. Estimated spawning ground is the southwest of theECS (Taiwan northeast). Spawning period is from March to June. Batchspawning eggs is released in the10m water layer. The physical field duringMarch-July yielded from the physical model is coupled with the biologicalmodel through the Lagrange particle tracking, considering the passive driftby current and random walk by turbulence. Using the super-individualtechnology, the individuals are grown when the particle drifts passively. The application of the model for normal climate, extreme climate and the changeof spawning ground study transport of eggs and larvae, to study dynamicsfactors of the changes effect. The main research results are as follows:
The results of flow field under normal climatological forcing conditionshow that with the increase of water temperature in March, chub mackerelbegan to spawn. Eggs are distribution in between Taiwan Warm Current(TWC) and the Kuroshio, and transported to the northeast. In April with thewater temperature further raise, the spawning peak is coming. Eggs increasedrapidly. Eggs and larvae were divided into two parts by the influence ofTWC and kuroshio. One can follow TWC to northward transportation. Theother part can follow the Kuroshio to northeast direction to transport. Watertemperature continues to rise in May, the most mackerel spawn over. Thenumber of spawning increase is reduced, because of death. The number ofeggs and larvae is decrease. Some continue to transport north for ZhoushanIsland. There is very little has been transported to the Yangtze Estuary andthe Hangzhou Bay. The other part also continues to transport northeast. Thefarthest drift is in the western Goto-retto islands. Most of larvae are thedistribution in between50m and200m isobaths. But because of the kuroshiobarrier, it is difficult to break into, or through the kuroshio to larvea. Watertemperature continues to rise in June. Part of the larvae to transport northalso began to turn to the northeast transport with other along the100-200misoaths. Most of larvae was transported to northeast of ECS in end of June only a little of larvae stranded in the Yangtze Estuary and Zhoushan Sea.When larvae get to the Goto-retto islands, larvae are divided into two partsby tidal currents, and a part was broughted into Tsushima Strait by TsushimaStrait Warm Current (TSWC), the other part to go through western sea ofKyushu was quickly broughted into the Pacific by the Kroshio. In July, watertemperatures continue to rise. The simulation is coming to the end. The mostof larvae was transported to Cheju, Tsushima Strait, Kyushu Island and thePacific Ocean. Some has drift out calculation domain by the kuroshio.Larvae were rarely found in this time in other ECS. The transportation oflarvae is significantly influenced by complex hydrodynamic TWC, Kuroshioand TSWC.
Statistical analysis showed that the mean water depth of larvae is deeperthan other three nursery grounds because of strong turbulence of nurseryground in the Pacific Ocean,. In end of July, average depth exceeds400m.Due to characteristics of the high temperature and salinity of the Kuroshio,the average water temperature is not the lowest. Juvenile basically can also isvery good to grow, that was transported into nursery ground in the PacificOcean.
With the advance of time, the proportion of larvae reached each parts ofnursery ground will has variation. In the end of June, most of larvae weretransported from spawning ground in south ECS to nursery ground of ChejuIsland and Kyushu Island. In mid-july, the proportion of nursery of Cheju Island and Kyushu Island is decreased becase of transport into nurseryground of the Pacific Ocean and Tsushima Strait. In the end of the simulationin July, most of larvae were transported to north nursery ground of Kyushuand Tsushima Strait. But only a third of the total larvae were transported tothe south nursery ground of Pacific Ocean snd Kyushu.
The spawning ground in the southern East China Sea made morecontributions to the recruitment to the fishing ground in northeast East ChinaSea, but less to the Yangtze estuary and Zhou-Shan Island. The transport ofeggs and larvae is the main mechanism of connection between spawningground and nursery ground. Connectivity is very similar to division of theocean currents system in ECS, divided into two parts. Northwest part(Section A) and Southwest part (Section B) of spawning ground had strongconnectivity with the nursery grounds of Cheju and Tsushima Straits.Northeast part (Section C) and Southeast part (Section D) of spawningground had strong connectivity with the nursery grounds of Kyushu andPacific Oceans. This is proof that physical factors impacts on connectivity.
Flow field under the typhoon forcing drives IBM to study transportdistribution of eggs and larvae of chub mackerel. The result showed that nosignificant difference distribution of eggs and larvae is found forclimatological forcing condition after a typhoon pass. But the typhooninfluence on water layer and survival of larvae. In order to verify theauthenticity of the horizontal distribution, eliminate the stochastic and individual water depth. We do the idealistic experiment. The results showedthat the little impact of larval distribution is really. The reason is transportpath of lavae is in the area of the typhoon minimum affect. Typhoon sweptthe ECS is a short period. The cancellation of tide-induced anti-cyclonic andtyphoon-driven cyclonic currents reduced its influence on the low-frequencycurrents and thus on larval transport in this region. Some difference is foundin larval abundance and velocity of transport in back end of transport path,where high density larvae northeast drift and velocity of transport increased.The typhoon caused higth mortality, to move to deeper water layer. Addingan experiment of the longer the typhoon duration, we observe that typhooncertain effect to larval distribution.
The change of position has great influence on transport distribution ofeggs and larvae. The west spawning ground was great influenced by Taiwanwarm current. Larval transport was northwest. A large amount of eggs andlarvae was retention and transit in coastal China in the process of transport.The most of larvae were finally transported to north nursery ground of ChejuIsland. The east spawning ground was great influenced by the Kuroshio.Larval transport was southeast. Almost no eggs and larvae was drift incoastal China. Speed of drift is very fast. A large amount of larvae werefinally transported to south nursery ground of Pacific Ocean. Transport speedof wast and east spawning ground was influenced by Taiwan warm currentand the Kuroshio. Survival and growth of normal spawning ground are the best in three spawning ground. The the change of spawning depth has noeffect on larval distribution and transport. But mortality rate of5m and15mdeep increases. Normal spawning depth (10m) is the best spawning depth.
We set the movement rule and develop IBM including a swimmingabilities submodel. The result shows that larval swimming ability had littleinfluence in the early transport stage. With the increase swimming ability inthe later stage, larvae have appeared to shcool. Larval mortality is reduced.Speed of northeast transport is also reduced. The amount of transport to thePacific Ocean and the Janpan Sea is decline. Average water depth is lower,which is more suitable to grow. The change of spawning ground make thewest spawning ground was influenced by Taiwan warm current. Larvaeschool in west. East spawning ground was influenced by the Kuroshio.Larvae school in eest. It is to explain the physical environment and biologicalfactors will also have effect to transport of swimming ability of larvae.
We study migration of adult spawning mackerel. The result show suitabledepth of mackerel is20m. School of mackerel in northeastern Taiwan wasmainly influenced by TWC, mainland coast water, the Kuroshio. There are alarge number of mackerel distributionn in the intersection front of TWC andcoast water. There are aslo a large number of mackerel distributionn in tidearea of coast water and the Kuroshio. The change position of spawningground is west spawning ground have large accumulation in front of TWCand coast water becase of large influenced by TWC, and east spawning ground have large accumulation in front of the Kuroshio becase of largeinfluenced by the kuroshio. Different locations of mackerel will affect theposition of school, it that explain the physical environment and biologicalfactors can influence on the migration and school of mackerel.
引文
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