中国北部海域灾害水母沙蜇(Nemopilema nomurai)及其它钵水母繁殖生物学特征与形态比较

为查清沙蜇大量暴发的原因,采用黄海北部的沙蜇亲体进行自然受精获得了数百万的螅状体,除海月水母和发形霞水母在消化腔和雌体的性腺中完成受精外,上述其余种类均为体外受精,霞水母属的2个种类的浮浪幼虫在形成螅状体前先形成一个具角质的浮浪体囊,沙蜇、海蜇、黄斑海蜇和Stomolophus meleagris螅状体繁殖新螅状体的唯一无性繁殖方式是足囊繁殖,霞水母可通过足囊和由匍匐茎形成囊胞两种方式繁殖新螅状体,海月水母(A.aurita)螅状体繁殖新螅状体的的无性繁殖方式包括足囊繁殖、出芽生殖、匍匐茎生殖、纵向分裂、内繁殖体和外繁殖体生殖,同时有直接发育现象,Rhizostoma pulmo的无性繁殖方式有足囊,出芽生殖,匍匐茎,浮浪体芽。     

红沿河海域常见水母的食物组成分析——基于稳定同位素和脂肪酸标记法

为了探究不同种类水母对浮游食物网的摄食影响,采用碳氮稳定同位素和脂肪酸标记法,研究分析了渤海红沿河海域常见的4种小型水母(伞径直径5 cm:卡玛拉水母Malagazzia carolinae、球形侧腕水母Pleurobrachia globosa、帽铃水母Tiaricodon coeruleus、半球美螅水母Clytia hemisphaerica和3种大型水母(伞径10—200cm:沙海蜇Nemopilema nomurai、海月水母Aurelia aurita)和白色霞水母Cyanea nozakii)的食物组成。结果表明,不同种类水母的食物组成存在差异。其中卡玛拉水母、球形侧腕水母、帽铃水母、半球美螅水母和白色霞水母均偏肉食食性,食物组成中动物性食物所占比例更高,高于海月水母与沙海蜇。在食物粒径谱上,帽铃水母和半球美螅水母较大粒径的食物比例均高于小粒径的悬浮有机物(POM)比例。卡玛拉水母和球形侧腕水母各个粒径食物比例接近。大型水母中,白色霞水母的食物中大粒径的浮游动物的比例高于海月水母,更高于沙海蜇。由此看来,小型水母和大型的白色霞水母的暴发会直接影响大中型浮游动物数量,海月水母的暴发对不同大小的浮游生物均会产生一定的影响。而沙海蜇的暴发会大量摄食1 mm的小型浮游生物和POM,对大中型浮游动物以及更高营养层生物(鱼类等)的影响可能是通过蜇伤以及饵料竞争导致的。因此,不同种类水母暴发对浮游生物的影响存在差异。本研究从摄食角度初步探究了水母对海洋生态系统物质循环和能量流动的影响,为水母暴发的灾害防治提供了数据支持。     

2018年6月渤海大型水母分布特征

2018年6月使用渔业底拖网采样，对渤海大型水母进行了全面调查，调查船舶为"中渔科102"渔业科考船。本研究分析了渤海大型水母的种类组成、渔获密度与伞径大小，并对其源地进行了探讨。结果表明：本次调查共采集到海月水母、沙海蜇、海蜇、多管水母四种大型水母，其中海蜇、多管水母数量较少，各采集到一只。海月水母在渤海三湾均有分布，各海区伞径大小无显著差异且多为幼体（<10cm），密度高值区出现在渤海湾东南侧海域，可达38-221.21ind./（net·h），辽东湾海月水母出现于湾南，密度<5ind./（net·h），湾北未见；作者推测，海月水母在渤海沿岸可能存在多个源头，诸如：莱州湾与渤海湾交界近岸海域、河北近岸、辽东湾大连近岸以及北部近岸。沙海蜇在渤海分布较广，辽东湾为密度高值区，均值为（35.32±21.64）ind./（net·h），但伞径较小，均值为（12.15±6.52）cm；与此相对，渤海湾与莱州湾外侧海域沙海蜇密度虽小[<20ind./（net·h）]，但伞径要显著大于辽东湾，最大伞径均值可达（33.86±7.40）cm；作者推测，沙海蜇在渤海海域发源地主要集中于辽东湾近岸，渤海湾与莱州湾，外海出现的沙海蜇可能源于辽东湾，随海流运输至此。海月水母、沙海蜇在渤海发生时间要晚于黄、东海。本研究结果可为深入分析渤海大型水母的种群动态变化、暴发机理提供基础。     

2015年5月黄海及东海北部大型水母分布及生物量估算

2015年5月搭载“北斗”渔业调查船、使用渔拖网的采样方式,在整个黄海及东海北部进行系统的走航式大面调查,记录了30°N—39°N海区内的大型水母种类组成、伞径大小及生物量分布,估算和比较了大型水母与其他渔业生物的生物量。结果表明,5月整个调查区,大型水母的总生物量估算值5.9万t,绝大部分由黄海中部的多管水母和洋须水母生物量贡献所致。出现的大型水母种类伞径分布呈单峰型。不同种类的水母分布具有明显地理区域和水文偏好性。其中,洋须水母主要分布于黄海中、北部50m水深以深水域,多管水母主要分布于黄海中部50m水深以浅的西侧以及整个东海北部;沙海蜇多为幼体,分布于黄、东海交汇区31°N—33°N间;霞水母较为集中出现于31°N以南、123°N以西近海。各水母种类的高密区的底层水温按洋须水母、沙海蜇、四叶小舌水母、霞水母呈升高趋势。东海多管水母分布区的底层水温与沙海蜇相近;黄海多管水母分布区的底层水温较洋须水母略高。沙海蜇和四叶小舌水母较其他水母的适温范围宽。霞水母和洋须水母处于相对高盐区域。     

沙海蜇(Nemopilema nomurai)代谢及分解过程对水环境理化因子及浮游植物影响的研究

从上行控制角度,通过野外采样和围隔培养实验,研究了水母的代谢及分解过程对水体环境中pH、溶解氧、营养盐组成的影响,以及该过程中浮游植物的变化。实验结果表明,沙海蜇在代谢过程中短时间内会大量消耗水体中的溶解氧（dissolved oxygen,DO）,使水体出现低氧和轻度酸化。代谢过程释放出大量营养盐,使水体中的溶解无机氮（dissolved inorganic nitrogen,DIN）浓度在24h内增加为原来的12倍,溶解无机磷（dissolved inorganic phosphorus,DIP）浓度增加了40多倍,进而引起水体中叶绿素a（chlorophyll a,chl a）浓度的增加。沙海蜇的分解过程使水体表现出明显的低氧（缺氧）和酸化现象。沙海蜇生物量越大,分解时间越长,对水体的改变程度越明显,此外,还释放出大量的营养盐并改变原有的营养盐结构,可以刺激甲藻和绿藻的生长,甚至可能引发藻华。     

黄海水温与沙海蜇丰度年际变化的相关分析

近十年来,我国黄、东海沙海蜇的数量呈上下波动趋势,除2008、2010、2011、2013年为不暴发年外,其余年份均为暴发年或弱暴发年(本文界定沙海蜇平均丰度范围为2—10ind./100m2为暴发年,1—2ind./100m2为弱暴发年,0—1ind./100m2为不暴发年)。为研究沙海蜇数量年际变化的原因,本文借助同化的海洋模式结果,分析了2006—2013年南黄海沙海蜇平均丰度与表底层海水温度的关系、与不同温度持续时间的关系。研究结果发现,在海州湾附近,对于暴发年2007年和2009年,春季底层海水10—18°C持续时间为130天,比不暴发年2010年和2011年多近15天。在长江口区域,不暴发年2008年和2011年夏秋季底层海水18—25°C持续时间较长,约80天,比暴发年2007年多20天。在长江口、苏北近岸以及海州湾区域,春季底层海水10—18°C持续时间越长,南黄海水母丰度呈现越大的趋势;夏秋季底层海水18—25°C持续时间越长,第二年水母生物量则越大。结果支持和验证了春季底层10—18°C持续时间长有利于当年水母暴发及夏秋季底层18—25°C持续时间长有利于来年水母暴发的推论。本文通过分析沙海蜇丰度和温度变化的关系,可以为将来预测该水母数量提供基础。     

2019年5月黄海及东海北部大型水母分布特征

2019年5月,利用渔业底拖网,对我国黄海以及东海北部海域进行了全面系统的大型水母调查,分析了大型水母的种类组成、伞径大小和生物量以及与温度、盐度的关系。结果表明,本次调查主要捕获到沙海蛰、霞水母、洋须水母、多管水母四种大型水母,沙海蛰生物量最高,多管水母分布范围最广、数量最大。沙海蛰集中分布在调查海域南部,各海域伞径差异显著,在黄东海交界海域采集到幼水母体(10cm),生物量高值区出现在东海北部离岸海域,可达6422.16kg/km2;白色霞水母集中分布在东海北部,在近岸海域采集到幼水母体(6—7cm),生物量高值区位于离岸海域,可达7417.49kg/km2;洋须水母集中分布在黄海水深较深海域,北部海域个体较大,在黄海中部、南部交界处采集到幼水母体(10cm),生物量较低,高值区出现在黄海中部与南部,可达449.94kg/km2;多管水母分布范围较广,东海北部海域个体伞径较大,在山东半岛东部发现幼水母体(5cm),生物量高值区出现在黄海中部近岸海域,可达4901.42kg/km2。对比文献资料,发现整个调查海域,大型水母总体生物量比2015年同期有所增加。本文为研究该海域大型水母的年际变化规律提供数据基础。     

辽东湾北部近海大型水母数量分布和温度盐度特征

通过2009-2011年5月下旬-7月下旬辽东湾北部近海10m等深线内的大型水母调查数据,分析了辽东湾北部近海近三年中大型水母资源状况,并探讨了辽东湾大型水母的生态类型。结果显示:辽东湾北部近海大型水母种类主要有海蜇(Rhopilema esculentum)、沙蜇(Nemopilema nomurai)、白色霞水母(Cyanea nozakii)、海月水母(Aurelia sp.1),海蜇和沙蜇是优势种。海蜇幼水母阶段主要集中分布在5m等深线以内的近岸河口水域,随着个体增大有略向深水或密度较小的水域扩散的趋势,仍主要分布在5m等深线两侧水域,属于高温低盐种类。6月份调查海区中发现大量的沙蜇幼水母,随着沙蜇个体增大,7月份调查海区中沙蜇数量大幅度减少。辽东湾海月水母在南部海域出现较多,2010、2011年在北部近海部分海域出现。白色霞水母近几年来辽东湾出现较少,栖息在盐度较高的水域。辽东湾各种大型水母中,沙蜇的生长速度最快。辽东湾海蜇幼水母、沙蜇幼水母的海区出现时间要晚于黄、东海。     

Feeding ecology of the banded wrasse Notolabrus fucicola (Labridae) in southern New Zealand: Prey items,seasonal differences,and ontogenetic variation

Notolabrus fucicola Richardson, a large common labrid inhabiting shallow waters around New Zealand and southern Australia, were collected monthly (December 1996‐February 1998) around Kaikoura on the east coast of the South Island, New Zealand. Their diet, temporal variation in prey and gut fullness, and ontogenetic differences were examined. N. fucicola is a generalist predator, the major prey items being bivalves, amphipods, and crabs, which varied temporally in their diet. There were size specific changes in their diet. Small fish (100–180 mm) ate mostly amphipods and isopods, whereas larger fish (> 180 mm) ate mainly bivalves, crabs, and gastropods. There was a temporal variation in gut fullness but no significant difference between sexes.     

Abundance and distribution of Nemopilema nomurai (Scyphozoa,Rhizostomeae), in Korean waters in 2005–2013

Nemopilema nomurai have appeared massively since 2003 in Korean waters, and regular offshore monitoring has been performed since 2005 in the eastern Yellow Sea (YS), northwestern East China Sea (ECS), Korean southern waters (Ksw), and western East Sea (ES). Korea Jellyfish Monitoring Network (KoJEM) was established in 2006 for coastal jellyfish monitoring. From these, sight survey monitoring data were gathered and analyzed to extract the yearly and seasonal fluctuation of N. nomurai’s abundance and distribution. The first massive annual appearance of N. nomurai occurred in the northwestern ECS, off Shanghai, China, in mid May of 2005–2008 and of 2012–2013. In other years or concurrently with the appearance in the northwestern ECS in the same year, several individuals were found in the mid-eastern YS and Ksw, and southwestern ES. The population of N. nomurai at the northwestern ECS moved to the southeastern YS and Jeju Strait in June, and from the July to August it extended its distribution all around the Korean peninsula and persisted till October. Since October, the distribution was retracted into the southern YS and Ksw, and disappeared slowly. N. nomurai’s yearly abundance varied a lot, from nearly 0 inds 10^?4m^?2 in 2010 to 62.4 inds 10^?4m^?2 in 2005. The highest abundance was recorded in 2005, followed by 2007 and 2009, and the least abundance was in 2008 and 2010–2013. The results were compared with those of the Liaoning Bay, Bohai Sea, which led to speculation about a new seeding place of N. nomurai, and mortality during the early planktonic phase of N. nomurai was proposed as the determinant of the yearly variation in abundance.     

Altered feeding habits and strategies of a benthic forage fish (Fundulus heteroclitus) in chronically polluted tidal salt marshes

Responses in feeding ecology of a benthic forage fish, mummichogs (Fundulus heteroclitus), to altered prey resources were investigated in chronically polluted salt marshes (the Arthur Kill-AK, New York, USA). The diet niche breadth of the AK populations of mummichogs was significantly lower than that of the reference population, reflecting reduced benthic macroinfaunal species diversity. Most of the AK populations also had 2-3 times less food in their gut than the reference population. This disparity in gut fullness among the populations appeared to be partly due to ingested prey size shifts; some of the AK populations ingested fewer large prey than the reference population. Furthermore, benthic assemblages were strongly associated with sediment-associated mercury; gut fullness of the AK populations also significantly decreased with increasing mercury body burdens. These results indicate that chronic pollution may have directly (chemical bioaccumulation) and indirectly (reduced prey availability) altered the feeding ecology of mummichogs.     

海蜇(Rhopilema esculentum)浮游幼体对褐牙鲆(Paralichthys olivaceus)卵和初孵仔鱼捕食的比较研究

利用实验生态学方法研究不同个体大小的海蜇浮游幼体[平均伞径(2.5±0.1),(4.1±0.2),(11.7±0.5),(21.1±0.6)mm]对不同密度(10,30,50,80ind/L)的褐牙鲆卵[卵径(0.92±0.01)mm]和初孵仔鱼[全长(3.01±0.08)mm]的捕食率,解析海蜇浮游幼体对初孵仔鱼的捕食率随捕食时间(0.5,1,2,3,4,5h)的变化特征。结果表明,各个体组海蜇浮游幼体对卵的捕食率均显著低于对仔鱼的捕食率;它们对卵的捕食率与卵密度和海蜇个体大小的关系不显著,但对仔鱼的捕食率随海蜇个体大小及仔鱼密度的增大而显著升高;伞径21.1mm的个体对仔鱼的捕食率在开始捕食后1h时达到最大值[17.3ind/(predator.h)],此后随捕食时间的延长而逐渐下降。在自然水域中,如果二者发生时空上的匹配关系,海蜇浮游幼体对仔鱼的捕食可能影响褐牙鲆的早期存活及其资源补充量的变动。     

水母(沙海蜇)降解与营养盐(氮、磷)释放的模拟与实证

The aim of this study was to investigate nitrogen and phosphorus released in the process of the decomposition of giant jellyfish in the laboratory and found the evidence to verify the influence of nutrients released by the decomposition of jellyfish on the ecosystem in the field. The release of nitrogen and phosphorus from the decomposition of Nemopilema nomurai was examined in a series of experiments under different incubation conditions such as different p H values, salinity values, temperatures and nitrogen and phosphorus concentrations. The results showed that the complete decomposition of Nemopilema nomurai generally took about 4–8 d. The release of nitrogen and phosphorus from the decomposition of Nemopilema nomurai could be divided into two stages: the early stage and the later stage, although the efflux rate of nitrogen was one order more than phosphorus. In the early stage of the decomposition of Nemopilema nomurai, the concentrations of dissolved nitrogen, dissolved phosphorus, total nitrogen and total phosphorus in seawater increased rapidly, and the concentration of nitrogen could reach the highest level in the whole degradation process. In the later stage of the decomposition, the concentrations of dissolved nitrogen and total nitrogen declined slowly, while the concentration of phosphorus in water could reach a maximum in the degradation process. High p H, low salinity,high temperature and N/P will promote the release of nitrogen; low p H is unfavorable to the release of nitrogen but favorable to the release of phosphorus. In addition, we found the concentrations of ammonium and phosphate in the bottom water were higher than those in the surface water during the period of jellyfish bloom in the Jiaozhou Bay, proving that nutrients released by the decomposition of jellyfish have significant influence on nitrogen and phosphorus in the field. For the whole Yellow Sea, nutrients released by jellyfish carcasses may reach up to(2.63±2.98)×107 mol/d of dissolved nitrogen(DN) and(0.74±0.84)×106 mol/d of dissolved phosphorus(DP) during the period of jellyfish bloom. The values are comparable to riverine inputs in a day, but much higher than sediment–water exchange flux in the Yellow Sea. The great amounts of nutrients must have significant influence on the nutrients balance of the Yellow Sea during the period of jellyfish dead and decomposition. Both the experimental data and field observations proved that the decomposition of jellyfish may release a great amount of nutrient to the surrounding environment during the period of jellyfish decomposition.     

Prey characteristics and dietary composition in intertidal Astrostole scabra (Echinodermata: Asteroidea)

Dietary composition in juvenile Astrostole scabra at Kaikoura, New Zealand (42°25′S, 173°42′E), changed qualitatively with increased seastar size so that food was partitioned amongst starfish size classes according to a combination of prey species and size. Small A. scabra fed more frequently than large specimens. Size‐discriminant feeding was evident as large Turbo smaragdus and Risellopsis varia, small Melagraphia aethiops, and intermediate sized Micrelenchus dilatatus, Ischnochiton maorianus, Onithochiton neglectus, and Amaurochiton glaucus were consumed in proportions greater than those found in the habitat. Major prey species have both spatial and size refuges from predation.     

Variability in the nutritional value of the major copepods in Cape Cod Bay (Massachusetts, USA) with implications for right whales

The North Atlantic right whale, a seriously endangered species, is found in Cape Cod Bay (Massachusetts, USA) during the winter and early spring. During their residency in these waters, these whales are frequently observed feeding. This study evaluated spatial and temporal changes in the chemical composition (carbon weight and C/N ratio) of the food resource targeted by the right whales in Cape Cod Bay. The three taxa measured (Centropages typicus, Pseudocalanus spp., and Calanus finmarchicus) had highly variable chemical compositions resulting from the different life strategies and from fluctuations in their surrounding environment. The impact of seasonal variability in the energy densities of the food resource of right whales was calculated and compared to the energetic requirements of these whales. Calculations indicated that differences in the nutritional content of the zooplankton prey in Cape Cod Bay could have a considerable effect on the nutrition available to the right whales. Therefore, it is likely that using more precise estimates of the energetic densities of the prey of right whales would lead to a re‐evaluation of the adequacy of the food resource available to these whales in the North Atlantic.     

Two new species of the marine genus Limnodriloides and a record of Tubificoides fraseri Brinkhurst (Oligochaeta: Tubificidae) from New Zealand

We characterised seasonal and ontogenetic changes in diet and prey energy density of rainbow trout (Oncorhynchus mykiss) in Lake Rotoiti, New Zealand, to better understand the prey requirements of trout in central North Island lakes. Common smelt (Retropinna retropinna) was the dominant prey item of rainbow trout larger than 200 mm (77.8% of diet by weight), followed by kōura (freshwater crayfish Paranephrops planifrons; 6.3%), common bully (Gobiomorphus cotidianus; 5.5%), and kōaro (Galaxias brevipinnis; 3.4%). Juvenile rainbow trout (<200 mm) consumed amphipods, aquatic and terrestrial insects, oligochaetes, tanaid shrimps, and smelt. Trout consumed kōaro only in autumn and winter; consumption of other species did not vary seasonally. The maximum size of smelt consumed increased with increasing trout size, but trout continued to consume small smelt even as large adults. Consumption of larger prey items (kōaro and kōura) also increased with increasing trout size. This study indicates the importance of smelt for sustaining rainbow trout populations, as predation on other species was relatively low. These findings provide a basis for bioenergetic modelling of rainbow trout populations in lakes of the central North Island of New Zealand.     

Planktonic ciliates as food for the scyphozoan <Emphasis Type="Italic">Aurelia coerulea</Emphasis>: feeding and growth responses of ephyra and metephyra stages

Blooms of the moon jellyfish, Aurelia coerulea, often occur in eutrophic embayments where microzooplankton are abundant. Hence, microzooplankton may play a role in A. coerulea population dynamics. In this study, feeding and growth responses of the ephyra and metephyra stages of A. coerulea were examined on a diet of planktonic ciliates, the main components of microzooplankton. Laboratory experiments indicated that ephyrae actively fed on ciliates and that the central disc diameter increased over the 5-day incubation period. Carbon-specific growth rates estimated from daily increases of central disc diameter increased with carbon-specific prey consumption rates until about 4 µg C ind day^?1 and then became saturated. By fitting these results to rectangular hyperbolic equations, the maximum specific growth rates were estimated to be 0.32 day^?1 for ciliate prey. However, the central disc diameter of the metephyrae did not increase on a diet of ciliates during the 5-day incubation, even though consumption of prey ciliates was observed. These results suggest that planktonic ciliates serve as a food source for growth at the ephyra stage of A. coerulea and then the contribution to growth decreases at and after the metephyra stage. This prey switch-dependency on the development stage of A. coerulea may be associated with the frequent occurrence of jellyfish blooms in eutrophic embayments.     

Feeding ecology of the mud shrimp Solenocera melantho in the East China Sea

The feeding habits of the mud shrimp Solenocera melantho were studied in the East China Sea by analysing 720 stomach contents and comparing diet composition and prey diversity among seasons, sexes and size classes. Feeding activity was linked to season and ovarian condition. The stomach fullness was high during spring and summer and low in fall. Females with advanced ovaries also displayed low stomach fullness, which suggests that feeding activity is affected by the reproductive cycle. The diet of S. melantho consisted of 42 different prey categories, primarily smaller crustaceans, Foraminifera, polychaetes, mollusks and fishes (larvae and eggs). Crustacea,Foraminifera and polychaetes together constituted the dominant prey groups, accounting for 75% of total prey abundance and >50% of total prey occurrence. Diet composition differed among seasons between sexes and size classes. Females and males consumed the same prey items but with different orders of importance. In terms of body size, the small shrimps preferred relatively small food items, such as Foraminifera and algae, whereas larger shrimps preferred polychaetes and crustaceans. Trophic diversity varied among the seasons within both sexes and size classes, with the highest values being observed in fall. Niche overlap was high within both sexes and size classes among seasons (0.6051–0.8740).     

Predatory Promesostoma species (Plathelminthes, Rhabdocoela) in the Wadden Sea

Laboratory experiments were performed on the food ecology of four congeneric species of free-living plathelminths, Promesostoma caligulatum, P. marmoratum, P. rostratum, and P. meixneri, all inhabiting an intertidal sandflat near the island of Sylt (North Sea). Their prey spectrum is within the microcrustaceans: P. caligulatum preferred ostracods, while the other three species favoured copepods, with species-specific differences for copepod species and size classes. Daily consumption of prey species varied with the size of both the predator and the prey. On average, P. marmoratum consumed 0.76 Harpacticus flexus per day while this rate decreased to 0.06 in P. meixneri, the smallest predator. When these Promesostoma species were fed with Tachidius discipes, a smaller prey species, their predation rates were about 25% higher. While the larger predators preferred the larger harpacticoids as prey, the small P. meixneri preferred small cyclopoids over larger harpacticoids. In terms of biomass, P. marmoratum's mean consumption of T. discipes per day was about half the predator's own weight. This average varied with prey density and temperature. A comparison of these consumption rates with the field densities of the predators and their prey shows that the plathelminth predators may consume as much as 10% per day of their copepod prey populations, thus strongly influencing these prey populations on these sandflats. The predation pressure of P. caligulatum on ostracods was about 1% per day of the prey population. Since ostracods usually have fewer generations per year, the total effect on the population dynamics may be similar to that on copepods. Therefore, nocturnal swimming of copepods in the water column may be interpreted as an attempt to escape plathelminth predators.