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.     

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。     

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.     

Interdecadal variability of anchoveta abundance and overcapacity of the fishery in Peru

Paleontological and historical stock abundance estimates indicate that pelagic fish populations inhabiting upwelling ecosystems undergo large interdecadal variations in abundance with amplitudes equal to, if not larger than, the interannual variability. The interdecadal variability is characterized by periods of high and low abundance, termed “pseudo-cycles”, because of their irregular periodicity. Fisheries targeting small pelagic fish suffer from overall overcapitalization, like many other fisheries, but also from an additional overcapitalization problem: a phase displacement between rapid fish population decreases and a slower disinvestment which follows. This lag produces economic hardship.Here we document the fish:fishery relationship for the Peruvian anchoveta. Anchoveta pseudo-cycles of 20 to >100 years are observed, with the present stock abundance most likely located near upper part of the cycle. Fleet overcapacity expressed as the proportion of unused present capacity is estimated at 72% and processing overcapacity at 89%. A simple bio-economic model demonstrates the risks associated with the pseudo-periodicity in fish stock abundance in conjunction with fishery investment, open access, and overcapacity: a timing bomb for the fishing sector. The lag between disinvestment and decrease in fish abundance is quantified. A reduction of the fishing and processing capacity and measures to decrease the investment lag are recommended to limit the social, economical and political tensions that will result from the expected decrease in stock abundance. Finally, some management options to reduce these risks are discussed.     

不同模态下气候变化对智利竹筴鱼补充量的影响

智利竹筴鱼(Trachurus murphyi)是东南太平洋重要的经济鱼类之一,其资源量受补充量影响明显,了解补充量状况对智利竹筴鱼资源可持续利用和科学管理具有重要意义。本文基于模态分析将1971−2017年间智利竹筴鱼补充量划分不同模态,运用贝叶斯模型平均法,分析海表面温度、海表面盐度、海表面高度、厄尔尼诺和太平洋年代际振荡5个环境因子在不同模态中对补充量的解释能力,并探讨模态变动对补充量预测的影响。结果表明,第1模态(1971−1980年)更多的受捕捞因素的干扰;第2模态(1981−1990年)厄尔尼诺对补充量变动的解释概率最高;第3模态(1991−2001年)解释概率最高的环境因子为太平洋年代际振荡;第4模态(2002−2015年)厄尔尼诺为解释概率最高的环境因子。对比不考虑模态变动的分析结果,两者存在明显差异,基于不同模态的分析结果对智利竹筴鱼补充量变动的解释更为合理。研究认为,智利竹筴鱼补充量变动受到多个环境因子的影响,在不同模态时期起主导作用的环境因子也不同,推测年代际太平洋年代际振荡冷暖期交替与厄尔尼诺现象可能是诱发智利竹筴鱼补充量发生模态转变的重要因素。建议在未来智利竹筴鱼资源评估与管理中,应该考虑不同的模态变化及其影响因子。     

Seasonal and inter-annual variation of mesozooplankton in the coastal upwelling zone off central-southern Chile

Zooplankton sampling at Station 18 off Concepción (36°30′S and 73°07′W), on an average frequency of 30 days (August 2002 to December 2005), allowed the assessment of seasonal and inter-annual variation in zooplankton biomass, its C and N content, and the community structure in relation to upwelling variability. Copepods contributed 79% of the total zooplankton community and were mostly represented by Paracalanus parvus, Oithona similis, Oithona nana, Calanus chilensis, and Rhincalanus nasutus. Other copepod species, euphausiids (mainly Euphausia mucronata), gelatinous zooplankton, and crustacean larvae comprised the rest of the community. Changes in the depth of the upper boundary of the oxygen minimum zone indicated the strongly seasonal upwelling pattern. The bulk of zooplankton biomass and total copepod abundance were both strongly and positively associated with a shallow (<20 m) oxygen minimum zone; these values increased in spring/summer, when upwelling prevailed. Gelatinous zooplankton showed positive abundance anomalies in the spring and winter, whereas euphausiids had no seasonal pattern and a positive anomaly in the fall. The C content and the C/N ratio of zooplankton biomass significantly increased during the spring when chlorophyll-a was high (>5 mg m⁻³). No major changes in zooplankton biomass and species were found from one year to the next. We concluded that upwelling is the key process modulating variability in zooplankton biomass and its community structure in this zone. The spring/summer increase in zooplankton may be largely the result of the aggregation of dominant copepods within the upwelling region; these may reproduce throughout the year, increasing their C content and C/N ratios given high diatom concentrations.     

Patterns in the spatial distribution of Peruvian anchovy (Engraulis ringens) revealed by spatially explicit fishing data

Peruvian anchovy (Engraulis ringens) stock abundance is tightly driven by the high and unpredictable variability of the Humboldt Current Ecosystem. Management of the fishery therefore cannot rely on mid- or long-term management policy alone but needs to be adaptive at relatively short time scales. Regular acoustic surveys are performed on the stock at intervals of 2 to 4 times a year, but there is a need for more time continuous monitoring indicators to ensure that management can respond at suitable time scales. Existing literature suggests that spatially explicit data on the location of fishing activities could be used as a proxy for target stock distribution. Spatially explicit commercial fishing data could therefore guide adaptive management decisions at shorter time scales than is possible through scientific stock surveys. In this study we therefore aim to (1) estimate the position of fishing operations for the entire fleet of Peruvian anchovy purse–seiners using the Peruvian satellite vessel monitoring system (VMS), and (2) quantify the extent to which the distribution of purse–seine sets describes anchovy distribution. To estimate fishing set positions from vessel tracks derived from VMS data we developed a methodology based on artificial neural networks (ANN) trained on a sample of fishing trips with known fishing set positions (exact fishing positions are known for approximately 1.5% of the fleet from an at-sea observer program). The ANN correctly identified 83% of the real fishing sets and largely outperformed comparative linear models. This network is then used to forecast fishing operations for those trips where no observers were onboard. To quantify the extent to which fishing set distribution was correlated to stock distribution we compared three metrics describing features of the distributions (the mean distance to the coast, the total area of distribution, and a clustering index) for concomitant acoustic survey observations and fishing set positions identified from VMS. For two of these metrics (mean distance to the coast and clustering index), fishing and survey data were significantly correlated. We conclude that the location of purse–seine fishing sets yields significant and valuable information on the distribution of the Peruvian anchovy stock and ultimately on its vulnerability to the fishery. For example, a high concentration of sets in the near coastal zone could potentially be used as a warning signal of high levels of stock vulnerability and trigger appropriate management measures aimed at reducing fishing effort.     

A time series of prokaryote secondary production in the oxygen minimum zone of the Humboldt current system, off central Chile

Because the marine picoplanktonic communities are made up of phylogenetically different microbial groups, the re-evaluation of key processes such as bacterial secondary production (BSP) has become an important contemporary issue. The difficulty of differentiating the metabolic processes of Bacteria from the rest of the microorganisms in the water column (i.e., Archaea and Eukarya) has made it difficult to estimate in situ BSP. This work presents the seasonal variability of the prokaryote secondary production (PSP) measured by the incorporation of ¹⁴C-leucine in the oxygen minimum zone (OMZ) off central-southern Chile. The BSP and potential archaeal secondary production (PASP) were determined through the combined use of ¹⁴C-leucine and N¹-guanyl-1, 7-diaminoheptane (GC₇), an efficient inhibitor of archaeal and eukaryote cell growth. BSP accounted for the majority of the PSP (total average, 59 ± 7.5%); maximum values were ∼600 μg C m⁻³ h⁻¹ and, on several dates, BSP represented 100% of the PSP. Similarly, PASP was also an important fraction of the PSP (total average, 42.4 ± 8.5%), although with levels that ranged from not detectable (on given dates) to levels that represented up to ∼97% of PSP (winter 2003). Our results showed that both Bacteria and Archaea accounted for almost equal portions of the prokaryote heterotrophic metabolism in the OMZ, and that PASP is notoriously enhanced through temporal pulses of heterotrophy. This indicates that, at least in marine systems with high abundance of Archaea (e.g., mesopelagic realm), the secondary production obtained through methods measuring the uptake of radiolabeled substrates should be considered as PSP and not as BSP. If the latter is the target measurement, then the use of an inhibitor of both archaeal and eukaryote cell growth such as GC₇ is recommended.     

High resolution vertical distribution of the copepod Calanus chilensis in relation to the shallow oxygen minimum zone off northern Peru using LOKI,a new plankton imaging system

The vertical distribution of copepodite stage V and adult Calanus chilensis was studied on two transects across the Humboldt Current System off northern Peru using the LOKI system. LOKI is an optical plankton recorder, which simultaneously collects images of zooplankton and environmental data such as temperature, salinity, oxygen, and fluorescence. Image quality allowed determination of CV, females and males and identification of C. chilensis from 3 co-occurring Calanid copepods. C. chilensis was inhabiting the upper 250 m. Highest abundances with a maximum of ca. 44.000 Ind. m⁻² were observed in a narrow band within Cold Coastal Water at stations closest to the coast, coinciding with the Poleward Undercurrent. This raises questions for the life cycle closure within the Humboldt Current system. In contrast to observations in the southern part of the Humboldt Current System, the three stages studied were most abundant in hypoxic waters at oxygen concentrations between 5 and 50 µM. Thus C. chilensis seems to be the only species of the family Calanidae where not only a resting stage can tolerate hypoxia, but also both adult stages. This impacts availability to predators, as despite a locally high biomass only part of the population is available to anchovy and other important fish species which are restricted to waters with higher oxygen concentrations.     

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。     

日本鳗鲡补充群体的日龄、全长、出生时间及耳石生长的变化

在中国大陆沿岸6个取样床搜集了日本鳗鲡白仔鳗,191尾用于长度测定,112尾鉴定了日龄及测定了耳石半径,近中心核的标记轮直径、耳石生长轮的宽度.补充群体长度平均值从(53.4±2.2)mm到(58.1±1.7)mm,长度与补充群体的地理分布之间无内在联系.补充群体日龄从(138.2±6.7)d至(155.2±9.0)d,长江口和温州地区补充群的日龄最低,以此为中心,向高纬度和低纬度走向分布,补充群的日龄都逐渐增大,补充群出生时间从1993年8月15日至1994年1月10日.低纬度的群体补充的时间早,其出生时间也早.纬度越高,补充的时间越迟,出生时间也晚.补充群耳石半径平均值从(142.5±8.4)μm至(155.8±14.5)μm,具有显着性差异(p<0.01).标记轮直径从(10.36±0.85)μm至(10.81±1.26)μm,其差异不具显著性(p>1.01).耳石生长轮平均宽度为(0.996-1.097) μm,其差异具有显著性(p<0.01),生长轮平均宽度以温州和长江口群体最大,以此为中心,向高纬度和低纬度分布的补充群的平均值分别逐渐减小.补充群耳石从中心至边缘生长轮宽度变化的共同点是在第90轮处平均宽度值最高.     

秘鲁外海茎柔鱼栖息地适宜性年代际变动

茎柔鱼广泛分布于东南太平洋海域,是我国重要的远洋捕捞对象之一,其种群易受气候和栖息地环境的影响。利用海表面温度(SST)和海表面高度距平(SSHA)两个关键环境因子,计算1950−2015年1−12月秘鲁外海茎柔鱼栖息地适宜性指数(HSI),对比分析太平洋年代际涛动(PDO)位于冷暖位相下茎柔鱼渔场环境以及栖息地质量的变动。结果显示,1950−2015年PDO呈现冷、暖、冷3个位相变化,其中PDO冷位相内的SST距平(SSTA)和SSHA明显低于PDO暖位相。交相关分析结果表明,PDO指数与SSTA和SSHA均呈显著正相关,而HSI与PDO指数、SSTA和SSHA均呈显著负相关。PDO位于冷位相时,茎柔鱼渔场内水温变冷,海面高度下降,适宜的SST和SSHA范围增加,因此茎柔鱼有利的栖息地面积增大;而PDO位于暖位相时,水温增暖,海面高度上升,适宜的SST和SSHA范围缩减,导致茎柔鱼适宜的栖息地面积缩小。研究认为,太平洋年代际涛动调控了茎柔鱼渔场内的环境变化,进而对茎柔鱼栖息地质量及适宜栖息地范围产生显著影响。     

Impact of climate and hydrology on juvenile fish recruitment towards estuarine nursery grounds in the context of climate change

Larvae of various coastal fish undergo immigration from spawning grounds towards estuarine nurseries. Several environmental factors can have an important impact on survival at this stage making it crucial for recruitment. Generalized linear models were applied in order to investigate the relation between river drainage, sea surface temperature (SST), NAO index and the North-South wind component intensity, over the two months prior to the estuarine colonization peak, and the densities of Platichthys flesus, Dicentrarchus labrax, Diplodus vulgaris and Diplodus bellottii in the nursery grounds based on a discontinuous historical dataset (from 1978 to 2006), for the Tagus estuary. The relation between SST over the 12 months prior to the estuarine colonization peak and fish densities in the nurseries was also investigated, as it integrates the periods of spawning stock maturation, spawning and larval immigration. While SST over the prior 12 months was negatively correlated with the abundance of P. flesus, it was positively correlated with the abundance of D. bellottii. Abundance of D. vulgaris was positively correlated with SST in the two months prior to the estuarine colonization peak, while the abundance of D. labrax was positively correlated with river drainage. The relations between SST and the abundance of P. flesus, a cold-water species with declining densities, and the subtropical species D. bellottii and D. vulgaris, which are increasing in abundance, are indicative of species abundance alterations related to climate warming. Dicentrarchus labrax will probably also be affected by climate change because of lowered precipitation and consequently river drainage.     

几种保存方法对中华哲水蚤和强壮箭虫体型的影响

选取5%的甲醛,–20℃冰冻和液氮为保存方法,以中国近海的2种优势浮游动物中华哲水蚤(Calanus sinicus)和强壮箭虫(Sagitta crassa)为研究对象,以体长、体宽和面积为体型参数,研究了上述几种保存方法在2个月的时间内对浮游动物体型的影响。研究结果显示,对于中华哲水蚤,甲醛保存2个月后,体长下降为原始体长的99.4%,体宽和面积都增加为107%;冰冻和液氮保存2个月后,体长分别增加为原始体长的101%和106.8%,体宽下降为90%和81.6%,面积下降为92.2%和88.8%。对于强壮箭虫,在甲醛、冰冻和液氮3种保存条件下,保存2个月后体长有所下降,分别为原始体长的92.8%,50.7%和48.4%。体宽和面积有所增加,体宽分别变为原始体宽的139.9%,159.4%和263.8%,面积分别变为原始面积的119%,97.8%和107%。综合分析发现,胶质类浮游动物在冰冻和液氮保存条件下,身体变形严重,因此用ZooScan测得的结果误差较大,相较而言,甲醛对胶质类浮游动物体型的影响较小;中华哲水蚤比强壮箭虫在各种保存条件下体型的变化要小;保存初期,浮游动物体型的变化幅度较大,后期趋于稳定;在3种保存条件下,中华哲水蚤和强壮箭虫体长的变化最稳定,其次是面积。对浮游动物的体型进行研究时,推荐使用甲醛保存条件下的体长或面积。     

不同饵料对真蛸亲体产罗同量、受精卵孵化率及初孵幼体大小的影响

分别采用文蛤（Meretrix meretrix）、菲律宾蛤仔（Ruditapes philippinarum）、缢蛏（Sinonovacula constrzcta）、许氏平鲌（Sebastodes fuscescens）投喂处于产卵、护卵期的真蛸（Octopus vulgaris）亲体,通过测量亲蛸摄食量、产卵量、...     

Spawning strategies and transport of early stages of the two Cape hake species,Merluccius paradoxus and M. capensis,in the southern Benguela upwelling system

Seasonal and short-term variability of environmental parameters influence the spawning strategies of fish species. In this study, the spawning strategies and the transport of early stages of the two Cape hake species off South Africa were investigated. Distribution of eggs and larvae of Merluccius paradoxus and M. capensis was analysed in order to derive more detailed and species-specific information on spawning season, spawning location, and transport of early stages. Samples were collected during three pilot surveys between January and October 2007 and during an extensive survey in September/October 2008 in the southern Benguela upwelling system off South Africa. Eggs and larvae of M. paradoxus were found in greater numbers than those of M. capensis during all surveys. Highest abundances were found from September to October, indicating one spawning peak for M. paradoxus during late austral winter to spring. The western Agulhas Bank was identified as the primary spawning ground, and smaller spawning events occurred on the West Coast. Larvae of both species were mainly distributed in subsurface waters between 25 and 100 m. More than 50% of all larvae caught had a total length between 3 and 4 mm and size increased significantly with decreasing latitude. Merluccius capensis were found closer inshore than M. paradoxus, indicating that early stages of the two species followed separate drift routes. We assume that this distribution pattern most likely evolved from differences in spawning location and phenology. The spawning strategies of M. paradoxus and M. capensis are well adapted to a time-frame of optimal transport conditions favourable for larval survival in the highly variable environment of the southern Benguela upwelling system, but the peak spawning of the two species is separated in time and space.     

Carbon, nitrogen and phosphorus stoichiometry in Japanese anchovy (Engraulis japonicus) from the Huanghai Sea, China

Generally, nutrient cycle is closely related to the element distribution in biomass and the population dynamics in ecosystems. Carbon, nitrogen and phosphorus in Japanese anchovy (Engraulis japonicus) of different body lengths from the Huanghai Sea (Yellow Sea) were determined to better understand their variability and reasons during its life history. The mean content was 45.12%, 10.12% and 2.02% for C, N and P, respectively. Significant differences in C, N and P elemental composition were found among different sizes, which could be explained by varying proportions of storage compounds in whole body fish, and varying degrees of ossification. Considering abundant resources in Japanese anchovy, it was an important P-pool in the cycle of P. Moreover, the excreted N/P ratio was significantly different in fishes of different sizes, especially at high gross growth efficiency. In the past two decades, overexploitation tended to cause smaller body length in the community structure of anchovy, which presumably changed the nutrient cycle in food webs of the Huanghai Sea. Exptrapolation of the results indicates that Japanese anchovy may be important for conveying nutrient in the Huanghai Sea.