Revisiting Peruvian anchovy (Engraulis ringens) trophodynamics provides a new vision of the Humboldt Current system

The Peruvian anchovy or anchoveta (Engraulis ringens) forages on plankton and is a main prey for marine mammals, seabirds, fish, and fishers, and is therefore a key element of the food web in the Humboldt Current system (HCS). Here, we present results from the analysis of 21,203 anchoveta stomach contents sampled during 23 acoustic surveys over the period 1996–2003. Prey items were identified to the genus level, and the relative dietary importance of different prey was assessed by determination of their carbon content. Variability in stomach fullness was examined relative to the diel cycle, the distance from the coast, sea surface temperature, and latitude, using generalized additive models (GAMs). Whereas phytoplankton largely dominated anchoveta diets in terms of numerical abundance and comprised >99% of ingested prey items, the carbon content of prey items indicated that zooplankton was by far the most important dietary component, with euphausiids contributing 67.5% of dietary carbon followed by copepods (26.3%). Stomach fullness data showed that anchoveta feed mainly during daytime between 07h00 and 18h00, although night-time feeding also made a substantial contribution to total food consumption. Stomach fullness also varied with latitude, distance from the coast, and temperature, but with substantial variability indicating a high degree of plasticity in anchoveta feeding behaviour. The results suggest an ecological role for anchoveta that challenges current understanding of its position in the foodweb, the functioning of the HCS, and trophic models of the HCS.     

东南太平洋秘鲁鳀资源量预报模型的构建

秘鲁鳀（Engraulis ringens）是栖息于东南太平洋沿岸的一种小型中上层鱼类。有效地对秘鲁鳀资源量进行预报将有助于为我国鱼粉进口企业提供决策支撑。为此,本研究结合秘鲁鳀生物（上一个渔汛季度的资源量、渔获物中的幼鱼比例）和环境（渔场水温和nino1+2区的温度距平）因素及捕捞量为预报因子,利用主成分分析和多元线性回归的方法对17个渔汛季度（2006年至2014年第一渔汛季度）秘鲁鳀的资源量建立预报模型并利用主成分分析的结果对影响秘鲁鳀资源变动的因子进行初步评价。研究表明,随着时间的推移样本量的增加,模型拟合资源量与真实资源量的平均相对误差逐渐下降,拟合资源量序列与真实资源量序列的相关系数逐渐增加。最终模型5（建模数据为2006-2013年第二渔汛季度的数据,验证数据为2014年第一渔汛季度的数据）能够很好地拟合出秘鲁鳀资源量的大小及变动趋势拟合资源量序列与真实资源量序列的相关系数为0.861;拟合资源量序列与真实资源量序列的平均相对误差为12%;预报得到的2014年第一渔汛季度的数据与真实值相比,相对误差为1%。结合主成分分析的结果对影响秘鲁鳀资源变动的因素进行评价结果表明,第一主成分的方差贡献率为46%,其中环境因素占据了最大的载荷;第二主成分的方差贡献率为23%,同样环境因素占据了最大载荷,但是,排名第二和第三的因素分别是上一个渔汛季度的资源量和捕捞量,其载荷相当;第三和第四主成分的方差贡献率分别为9%和7%,其中幼鱼比例占的载荷最大。根据各主成分得分序列与资源量序列的相关系数结果,环境因子对秘鲁鳀资源变动有重要影响。     

Geographic structure of European anchovy: A nuclear-DNA study

Atlantic–Mediterranean anchovies were genetically characterized at two polymorphic nuclear loci (intron 6 of two creatine-kinase genes) and compared to reference Engraulis albidus and E. encrasicolus samples from the northern Western Mediterranean to provide new insights into their geographic structure. Northeastern Atlantic anchovy, represented by one sample from the Canary archipelago and one sample from the Alboran Sea, were genetically distinct from Mediterranean E. encrasicolus (Weir and Cockerham's ^θ = 0.027–0.311), indicating geographic isolation from either side of the Almería–Oran oceanographic front. Generally smaller genetic differences were evident among anchovy populations from different sub-basins in the Mediterranean (^θ = − 0.019–0.116), the genetic differences between Black Sea and Ionian Sea/Aegean Sea anchovies being the strongest (^θ = 0.002–0.116). There was no evidence of the presence of E. albidus in our samples outside Camargue (northern shore of the Western Mediterranean). However, a sample from the southern Western Mediterranean appeared to be genetically intermediate between E. albidus and Mediterranean E. encrasicolus, indicating possible hybridization. Anchovy from the Benguela current system off southern Africa possessed allele frequencies characteristic of E. albidus at one locus and Northeastern Atlantic anchovy at the other locus, suggesting past introgression.     

Ecological niches and areas of overlap of the squat lobster ‘munida’ (Pleuroncodes monodon) and anchoveta (Engraulis ringens) off Peru

The world’s largest mono-specific fishery, the Peruvian anchovy or anchoveta (Engraulis ringens) fishery, has been the subject of many studies since the 1960s. Details of its relationship with other species have mainly focused on alternations with sardine, Sardinops sagax, and little effort has so far been paid to interactions with other species sharing the same ecosystem. This is the case for Pleuroncodes monodon, the crustacean squat lobster or ’munida’, which has become highly abundant along the Peruvian coast since the mid-1990s. Munida is now an important prey for seabirds, mammals and coastal predatory fish. Knowledge of patterns of distribution and ecological niche of munida is scarce however off Peru. Here we describe and compare spatial patterns of distribution of anchoveta and munida and their ecological niches based on data from 26 acoustic surveys performed along the Peruvian coast between 1998 and 2006. The results indicate that munida and anchoveta share ecological niches but that munida is restricted to the coldest part of the productive cold coastal waters whereas anchoveta do not present any temperature preference over a large range (14–23 °C). The recent increase in munida abundance off Peru is concomitant with colder conditions; with their onset munida extended its range from central Chile northwards. Off Peru the very shallow oxycline keeps munida from its usual bottom habitat and has forced it to adopt pelagic behaviour.     

坛紫菜生活史的研究现状及存在的问题

坛紫菜（Porphyra haitanensis）属暖温性海藻,是中国东南沿海的重要栽培物种,2011年度栽培面积达30余万亩。其生活史过程比较复杂,既有雌雄异体,也有雌雄同体;既存在无性生殖,也存在有性生殖;其能否产生单孢子也常常引起争议。为此,本文总结了坛紫菜的生殖及生活史的研究现状和存在的问题,旨在为坛紫菜的遗传育种提供一定的理论参考。     

文蛤早期发育阶段生长性状遗传参数的估计

采用部分因子设计构建了23个全同胞家系,应用单性状及两性状动物模型对文蛤(Meretrix meretrix)早期不同生长发育时期的生长性状进行遗传力和相关性估计。结果表明,文蛤附着变态期以及稚贝期生长性状的遗传力并无显著差别,在0.11~0.41范围内变化,属于中等遗传力。非遗传的共同环境效应随着年龄的增长而呈现上升趋势,由附着变态期的0.02逐渐增加到稚贝期的0.38,且稚贝期的共同环境效应显著大于其附着变态期(P0.01)。壳长与壳高表型相关和遗传相关的分析结果显示,从附着变态期到稚贝期,壳长与壳高之间的表型相关及遗传相关都存在极显著的正相关性,相关系数的范围分别为0.84~0.95、0.93~0.99。早期发育连续时间段生长性状之间的遗传相关均为正相关,但并不显著,且不同时期的遗传相关也不尽相同(0.34~0.71)。     

Schooling behaviour and environmental forcing in relation to anchoveta distribution: An analysis across multiple spatial scales

The Peruvian anchovy or anchoveta (Engraulis ringens) supports the highest worldwide fishery landings and varies in space and time over many scales. Here we present the first comprehensive sub-mesocale study of anchoveta distribution in relation to the environment. During November 2004, we conducted a behavioural ecology survey off central Peru and used a series of observational and sampling tools including SST and CO₂ sensors, Niskin bottles, CTD probes, zooplankton sampling, stomach content analysis, echo-sounder, multibeam sonar, and bird observations. The sub-mesoscale survey areas were chosen from mesoscale acoustic surveys. A routine coast-wide (2000 km) acoustic survey performed just after the sub-mesoscale surveys, provided information at an even larger population scale. The availability of nearly concurrent sub-mesoscale, mesoscale and coast-wide information on anchoveta distribution allowed for a unique multi-scale synthesis. At the sub-mesoscale (100s m to km) physical processes (internal waves and frontogenesis) concentrated plankton into patches and determined anchoveta spatial distribution. At the mesoscale (10s km) location relative to the zone of active upwelling (and age of the upwelled water) and the depth of the oxycline had strong impacts on the anchoveta. Finally, over 100s km the size of the productive area, as defined by the upwelled cold coastal waters, was the determining factor. We propose a conceptual view of the relative importance of social behaviour and environmental (biotic and abiotic) processes on the spatial distribution of anchoveta. Our ecological space has two y-axis; one based on self-organization (social behaviour), and the other based on the environmental processes. At scales from the individual (10s cm), to the nucleus (m), social behaviour (e.g. the need to school) drives spatial organization. At scales larger than the school, environmental forces are the main driver of fish distribution. The conceptual ecosystem models presented in this paper may provide the final links needed to develop accurate forecasts of the spatial distribution of anchoveta over multiple scales.     

海蜇生活史的补充研究

为了更详细地阐述海蜇(Rhopilema esculentum)生活史,作者应用显微观察和对比分析方法,对其各阶段的生长特征进行了研究,继而对已报道的海蜇生活史进行了资料补充。研究结果表明:根据浮浪幼虫纵轴长度及活力将其分为早期、中期和晚期浮浪幼虫;在海蜇浮浪幼虫变态附着和足囊萌发过程中,补充了杯状体发育阶段;根据伞部中胶层变化、中央口发育状况及棒状附着器的出现与否,对碟状体至幼蜇的发育阶段进行了重新划分,将海蜇浮游幼体划分为碟状体、稚蜇和幼蜇3个阶段。实验结果完善了海蜇早期生活史,为深入开展海蜇基础生物学和基础生态学研究提供参考。     

基于个体的东海鲐鱼补充量年际变动模拟研究

东海鲐鱼（Scomber japonicus）是我国近海重要经济鱼种,年际资源量波动较大,物理环境对其资源量的丰歉影响很大。本文运用基于个体的东海鲐鱼生长初期生态模型,模拟1978-2013年东海鲐鱼资源补充量。模拟发现,物理环境的变化确实能导致东海鲐鱼资源补充量的年际差异,并且在90年代后出现了较大的波动,资源补充量最多的2002年（1.8×109尾）与最少的1996年（6.5×108尾）相差1.15×109尾。产卵后的15~30 d,即4月份鱼卵仔鱼所处的海洋物理环境对东海鲐鱼最终的资源补充量起到重要作用,4月底鱼卵仔鱼的存活量基本上决定了东海鲐鱼最终的资源补充量的多寡。4月份的水温是影响鲐鱼资源补充量的物理因素之一,4月份水温高有利于提高鲐鱼的资源补充量,相反,水温偏低不利于鲐鱼的资源补充量。4月份的台湾暖流的流速也是影响鲐鱼资源补充量的另外一个物理因素,4月份台湾暖流的势力强对东海鲐鱼最终资源补充量是有利的,相反,流速小对补充量较不利。4月份台湾暖流的水温、流速决定最终东海鲐鱼资源补充量。     

Modeling fish growth and reproduction in the context of the Dynamic Energy Budget theory to predict environmental impact on anchovy spawning duration

Spawning location and timing are critical for understanding fish larval survival. The impact of a changing environment on spawning patterns is, however, poorly understood. A novel approach is to consider the impact of the environment on individual life histories and subsequent spawnings. In the present work, we extend the Dynamic Energy Budget (DEB) theory to investigate how environment variability impacts the spawning timing and duration of a multiple-batch spawning species. The model is successfully applied to reproduce the growth and reproduction of anchovy (Engraulis encrasicolus) in the Bay of Biscay. The model captures realistically the start and ending of the spawning season, including the timing of the spawning events, and the change in egg number per batch. Using a realistic seasonal forcing of temperature and food availability derived from a bio-physical model, our simulation results show that two thirds of the total spawned mass already accumulates before the start of the spawning season and that the condition factor increases with body length. These simulation results are in accordance with previous estimations and observations on growth and reproduction of anchovy. Furthermore, we show how individuals of equal length can differ in reproductive performance according to the environmental conditions they encounter prior to the spawning season. Hatch date turns out to be key for fecundity at age-1 as it partly controls the ability to build up reserves allocated to reproduction. We suggest the model can be used to realistically predict spawning in spatially and temporally varying environments and provide initial conditions for bio-physical models used to predict larval survival.     

Ecological understanding for fishery management: Condition and growth of anchovy late larvae during different seasons in the Northwestern Mediterranean

The fishery of the European anchovy Engraulis encrasicolus in the Mediterranean needs several ecological approaches to be properly managed. As such, several surveys were carried out to study the ecology of larvae and juveniles of this species, which reproduces during the warmest period of the year (May through September) in the Gulf of Lions. In particular, we studied the late larvae (15 mm total length until metamorphosis), especially as other authors have focused on larvae below that size. Unexpectedly, we also collected anchovy late larvae during the December 2007 survey, whose range in size corresponded to a later spawning period than previously reported. Differences in the nutritional condition of these larvae were assessed by comparing indices of lipid composition and estimating growth rates from otolith measurements to provide information on the probability of survival between the two groups. The analysis of fatty acids, used as tracers of trophic relationships, indicates that these larvae fed mainly on zooplankton. Nutritional conditions of summer and late autumn larvae were very similar. In contrast, growth rates were higher for August larvae, probably due to the different temperatures in the two seasons. Our results are especially relevant in an ecological context where the increasing mean water temperatures in the Western Mediterranean could favor the extension of the anchovy spawning period up to late-Autumn months.     

基于个体的西北太平洋柔鱼冬春生群生活史早期生态模型构建

为了对西北太平洋柔鱼冬春生群生活史早期的生长、死亡、输运和分布等进行研究,将其生长、死亡等早期生物过程进行参数化,利用物理模型(FVCOM-Global)模拟生成北太平洋(10°～60°N,120°E～110°W)三维物理场,采用拉格朗日质点追踪的方法将物理模型和生物模型(基于个体模型)耦合,构建了基于个体的西北太平洋柔鱼冬春生群生活史早期生态模型,并用此模型对1997–2010年冬春生群的输运分布进行数值模拟。模拟结果表明：在一定温度范围内,越靠近最适水温,冬春生群的生长速度越快,当日龄为38 d时,胴长可达到11.76 mm,之后由指数生长逐渐变为线性生长;1997–2010年模拟补充量有年际的波动,在考虑冬春生群亲体量不同的前提下得到的实际补充量最多和最少的年份分别是1999年和2009年;冬春生群在模拟阶段主要位于产卵场内,在25°N以南主要向西部输运,25°N以北逐渐呈向北和向东北方向输运的趋势,尤其在输运后期受强流的影响较大;在垂直方向上,100 m水深以内的仔鱼占比将近一半。本研究能为西北太平洋柔鱼冬春生群的生态动力学深入研究奠定基础。     

2014年夏季南海北部超微型浮游植物分布及环境因子影响

利用流式细胞仪BD Accuri C6对2014年夏季南海北部超微型浮游植物进行了现场的观测研究,发现了3类超微型光合自养浮游植物,聚球藻(Synechococcus,Syn)、原绿球藻(Prochlorococcus,Pro)和超微型真核藻类(pico-eukaryotes,Euk),并对其丰度与分布以及环境因子影响进行了研究。结果表明,Syn、Pro和Euk丰度总平均值分别为5.13×103个/mL,3.27×104个/mL和1.85×103个/mL,碳生物量均值分别为1.19μg/L,1.86μg/L和4.51μg/L。Syn、Pro和Euk的丰度表现出不同的分布特征。Syn、Pro和Euk丰度分布趋势呈现近海低而外海高,Syn和Euk丰度高值区分别出现在沿岸带与陆架和上升流影响海域,Pro丰度高值区出现在沿岸带与陆架,低值区出现在上升流影响海域。Syn、Euk丰度高值区主要分布在次表层,Pro丰度高值区主要分布在真光层底部,Euk丰度垂直变化差异相对Syn和Pro较小。超微型浮游植物与环境因子的相关性分析结果表明,Syn、Pro和Euk的碳生物量均与硝酸盐、硅酸盐浓度和深度呈现负相关关系,Pro的碳生物量与磷酸盐浓度呈现正相关关系。     

Contrasting local retention and cross-shore transports of the East Australian Current and the Leeuwin Current and their relative influences on the life histories of small pelagic fishes

Transport between shelf and offshore environments supports a significant proportion of ocean primary productivity and is critical to the life cycle of many marine species. While fundamental differences in the underlying dynamics of eastern and western boundary currents have been recognized and studied for more than half a century, the implications for physical dispersal rates have received much less attention. In this study we explore how Australia’s two major boundary current systems, the East Australian Current and the Leeuwin Current, differ in their local retention and cross-shore transports in the upper water column and how these differences favor contrasting life histories of small pelagic fishes. The results suggest that the East Australian Current forms a partial barrier to onshore transport, but is effective in entraining shelf waters and transporting them offshore, particularly in the region where the current separates from the coast. Blue mackerel (Scomber australasicus) spawn on the outer-shelf in this separation region and may thereby maximize the dispersion of eggs and larvae in the mainly oligotrophic waters of the southern Coral Sea. In contrast, the Leeuwin Current system promotes onshore transport through the combined effects of mean onshore flow and eddy-induced mixing. In the Great Australian Bight, sardine (Sardinops sagax) and anchovy (Engraulis australis) may exploit the high coastal retention of the Leeuwin Current system by spawning on the inner-shelf during summer when the current is weakest and winds assist retention and enhance production through local upwelling.     

Spawning patterns of four species of predominantly temperate pelagic fishes in the sub-tropical waters of southern Queensland

The diverse pelagic fish assemblage of sub-tropical southern Queensland includes fishes with predominantly temperate distributions, such as tailor Pomatomus saltatrix, sardine Sardinops sagax, round herring Etrumeus teres, and Australian anchovy Engraulis australis. The peak spawning seasons of P. saltatrix, S. sagax and E. teres occur during late winter and early spring (June–October). Eggs and larvae of these three species are widely distributed in shelf waters and comprise >50% of the ichthyoplankton assemblage during this period. Mean monthly sea surface temperatures (SSTs) during late winter and early spring range from 21 to 23 °C, and are thus similar to those recorded in southern Australia during summer and autumn, which is the spawning season of these three species in those temperate waters. E. australis eggs occur mainly in inshore waters, and comprise >50% of fish eggs collected during summer and autumn when mean monthly SSTs in southern Queensland exceed 27 °C. E. australis also spawns mainly during summer and autumn in temperate Australia. Hence, water temperature may be less important as a determinant of the spawning season of E. australis than it is for the other three species. The suitability of southern Queensland for spawning by predominantly temperate species during late winter and early spring may contribute to the high diversity of the region's pelagic fish assemblage. Adult P. saltatrix, S. sagax and E. teres appear to migrate northwards into southern Queensland during early winter to spawn, and larvae may be transported southwards into temperate waters by the East Australian Current. This dispersal-migration pattern is similar to those observed for several species, including P. saltatrix, in the western boundary current systems off the east coasts of North America and Africa. Hence, pelagic fishes in ecosystems off the east coast of three continents migrate into sub-tropical waters to spawn, and larvae are transported back into temperate nursery areas by the prevailing current.     

基于灰色系统西南大西洋阿根廷滑柔鱼资源丰度预测模型的构建

阿根廷滑柔鱼（Illex argentinus）是西南大西洋鱿钓渔业的主要作业鱼种,对资源丰度进行准确的预测可指导企业合理安排渔业生产。因此,本研究根据2000-2016年我国西南大西洋阿根廷滑柔鱼的生产数据,以单位捕捞努力量的渔获量（Catch per unit effort, CPUE）为阿根廷滑柔鱼资源丰度的指标,利用灰色绝对关联分析和灰色预测建模的方法（GM（0, N））,计算2001-2015年CPUE的时间序列值与产卵期（6-8月）产卵场海表面温度（Sea surface temperature, SST）时间序列值的灰色绝对关联度,选取产卵场海域中灰色绝对关联度大于0.90的海区SST建立资源丰度预测模型,并用2016年实际CPUE进行验证。灰色绝对关联分析表明,6-8月,30°~40°S,45°~60°W海域内存在若干海区的SST与次年对数CPUE时间序列呈现较强的关联度,可作为预报因子。GM（0, N）模型结果表明,以6-8月产卵场SST作为环境因子建立的模型4能较好地拟合出阿根廷滑柔鱼资源丰度变动趋势,与2016年真实值相比,相对误差为7%,该模型可较好地作为阿根廷滑柔鱼资源丰度的预测模型。相反,包含6月和7月SST的模型1效果优于不包含6月SST的模型2或不包含7月SST的模型3,拟合得到的2016年的数据与真实值相比,相对误差分别为128%和289%,这说明6月和7月是西南大西洋阿根廷滑柔鱼的主要产卵月份。     

Distribution of Calanus finmarchicus in the northern North Atlantic and Arctic Ocean—Expatriation and potential colonization

The distribution of Calanus finmarchicus was studied on a transect across the central Greenland Sea, and on five transects from the Eurasian shelves across the Atlantic Inflow in the Arctic Ocean. Stage composition was used as an indicator for successful growth; gonad maturity and egg production were taken as indicators for reproductive activity. On the Arctic Ocean transects, these parameters were measured simultaneously from the sibling species Calanus glacialis. Response of egg production rate to different temperatures at optimal food conditions was very similar between both species in the laboratory. C. finmarchicus was present at all stations studied, but young developmental stages were only present close to the regions of submergence of Atlantic water under the Polar water. This together with a decreasing abundance and biomass from west to east along the Atlantic Inflow in the Arctic Ocean and reproductive failure indicates that C. finmarchicus is expatriated in the Arctic Ocean. We hypothesize that the late availability of food in the Arctic Ocean, rather than low temperature per se, limits reproductive success. Better reproductive success in the very low temperature regions of the Return Atlantic Current and the marginal ice zone in the Greenland Sea supports this hypothesis. The possibility for a replacement of C. glacialis by C. finmarchicus and consequences for the ecosystem after increasing warming of the Arctic are discussed.     

Cross-shelf distribution of copepods and the role of event-scale winds in a northern California upwelling zone

Cross-shelf distribution and abundance of copepod nauplii and copepodids were measured during three summer upwelling seasons (2000–2002) in a coastal upwelling zone off northern California. These 3 years varied considerably in the intensity of winds, abundance of chlorophyll, and water temperature. The cruises in 2000 were characterized by relaxation conditions, with generally high levels of chlorophyll and high water temperature. The cruises in 2001 and 2002 were dominated by strong and persistent upwelling events, leading to lower chlorophyll and water temperatures. The copepod assemblage was dominated by Oithona spp., Acartia spp. and Pseudocalanus spp., with Metridia pacifica (lucens), Microsetella rosea, Oncaea spp. and Tortanus discaudatus also common during all 3 years. The cross-shelf distribution of copepods was generally shifted offshore during upwelling and onshore during relaxation events, although some variability between species occurred. Abundance of all life stages generally exhibited a negative correlation with cross-shelf transport averaged over at least 1–4 days and lagged by 0–3 days, indicating lower abundances during and immediately after active upwelling. However, copepod nauplii seemed to respond positively to wind events lasting 1–5 days followed by a period of relaxation lasting 6 or 7 days. These rapid rates of change in abundance are probably too great to be due to in situ growth and reproduction alone; physical processes must also play a role. These results suggest a highly dynamic relationship between copepods and upwelling events off northern California, with species-specific responses to upwelling to be expected.     

黄海浒苔绿潮防灾减灾现状与早期防控展望

截至2019年,浒苔绿潮连续12年大规模暴发,对近海生态系统、沿岸环境与社会经济造成严重影响,已经成为黄海最严重的生态环境问题。本文总结了黄海浒苔绿潮防灾减灾现状与成效,分析了存在的问题,然后基于对该绿潮起源与成因的认识,将其早期分为3个关键过程,即浒苔微观繁殖体在养殖设施上的着生与生长过程,定生浒苔脱离附着基形成漂浮浒苔过程,浅滩漂浮浒苔进入深水区形成大面积绿潮过程。最后分别从加强新材料与技术研发防控绿藻着生、强化养殖设施回收管理严控定生绿藻落滩、浅滩汇聚通道拦截打捞等3种途径提出了早期防控措施建议,以期为黄海浒苔绿潮的源头防控提供科学依据。