Seasonal variation in fatty acid composition of seston and the copepod Calanus sinicus(Brodsky,1962)in Jiaozhou Bay and its trophic implications

The fatty acid compositions of seston and Calanus sinicus were investigated to study trophic relationships in Jiaozhou Bay. Principal component analysis was carried out to ordinate the fatty acid patterns of seston in stations and months. The results showed that diatoms were most abundant in the phytoplankton at station A5 (located in the northwest of the bay:36°9′N, 120°20′E) and least abundant at station D7 (located outside of the bay: 35°59′N, 120°26′E). By contrast, dinoflagellates were most abundant at station D7 and least abundant at station A5. According to the annual variations of 16:1ω7 and 18:4ω3/ 16:1ω7, diatoms flourished mainly in spring and summer, while dinoflagellates bloomed exclusively in summer. A distinctive feature of the fatty acid composition of C. sinicus was the prevalence of 20:5ω3 and 22:6ω3. The higher content of 16:1ω7 over 18:4ω3 in females indicated that diatoms contributed more than dinoflagellates to the diet of C. sinicus. The feeding intensity of C. sinicus on diatoms was higher in spring and autumn than in other seasons. The herbivorous indicators 20:1 and 22:1 were comparatively low, suggesting that besides phytoplankton, C. sinicus might feed on a wider range of particles including organic detritus, bacteria and small copepods.

Zooplankton community structure,abundance and biovolume in Jiaozhou Bay and the adjacent coastal Yellow Sea during summers of 2005–2012:relationships with increasing water temperature

Zooplankton abundance, biovolume and taxonomic composition in Jiaozhou Bay and the adjacent coastal Yellow Sea were evaluated using ZooScan measurement of samples collected by net towing every August from 2005 to 2012. Zooplankton abundance and biovolume ranged from 1 938.5 to 24 800 ind./m^3 and 70.8 to 1 480.1 mm^3/m^3 in Jiaozhou Bay and 73.1 to 16 814.3 ind./m^3 and 19.6 to 640.7 mm^3/m^3 in the coastal Yellow Sea. Copepods were the most abundant group in both regions, followed by N octiluca scintillans and appendicularians in Jiaohzou Bay, and chaetognaths and N octiluca scintillans in adjacent coastal Yellow Sea. Over the study period, the most conspicuous hydrographic change was an increase in water temperature. Meanwhile, a general decrease in zooplankton abundance was observed, particularly in copepod populations. Based on redundancy analysis(RDA), the warming trend was the key environmental factor influencing to decrease of copepod abundance. The proportion of small-sized copepods increased while the mean size of all copepods decreased, in significant correlation with water temperature. Our results indicate that zooplankton, particularly copepods, are highly sensitive to change in water temperature, which is consistent with predicted impacts of warming on aquatic ectotherms. Due to their dominance in the zooplankton, the decline in copepod size and abundance could lead to an unfavourable decrease in energy availability for predators, particularly planktivorous fish.

Experimental clearance rates of Aurelia coerulea ephyrae and medusae,and the predation impact on zooplankton in Jiaozhou Bay

The population explosion of jellyfish Aurelia coerulea occurred in Jiaozhou Bay,China in 2009.The potential predation impact of A.coerulea on zooplankton was investigated.Population clearance potential and residence time(t 1/2)for copepods were calculated from laboratory clearance rates and measurements of jellyfish size and abundance from May to August 2009 in Jiaozhou Bay.Clearance rates varied widely with prey organisms,but they were not significantly different among various prey concentrations.Medusae captured rotifers,fish larvae and hydromedusae more efficiently than fish eggs,copepods and chaetognaths.Ephyrae captured rotifers and hydromedusae more efficiently than fish larvae and copepods.Clearance rate linearly increased with the cross sectional area of A.coerulea(size from 0.3 to 7.1 cm).Water temperature also had a marked effect on clearance rate and this was related to the increased beat frequency as water temperature increased.In early May 2009,A.coerulea potentially cleared the volume of water in the Bay less than 0.001 times a day,but this value was estimated to be more than 0.3 times a day in July.The t 1/2 for copepods was less than 6 d in June and July.Abundances of copepods,hydromedusae and chaetognaths were extremely low in 2009 compared to 2008 and 2010(jellyfish non-bloom years).Large predation pressure by the A.coerulea population occurred to control zooplankton communities in Jiaozhou Bay.A.coerulea,when present at a high population level,can be a keystone species in Jiaozhou Bay and control the trophic structure here.

Genome-wide SNP markers provided insights into the reproductive strategy and genetic diversity of the green tide causative species Ulva prolifera in China

Ulva prolifera is the causative species of the annually occurring large-scale green tides in China since 2007.Its specific biological features on reproductivity strategies,as well as intra-species genetic diversity,are still largely unknown,especially at the genome level,despite their importance in understanding the formation and outbreak of massive green tides.In the present study,the restriction site-associated DNA genotyping approach(2b-RAD)was adopted to identify the genome-wide single-nucleotide polymorphisms(SNPs)of 54 individual thalli including samples collected from Subei Shoal in 2019 and Qingdao coast from 2019 to 2021.SNPs genotype results revealed that most of the thalli in 2019 and 2020 were haploid gametophytes,while only half of the thalli were gametophytes in 2021,indicating flexibility in the reproductive strategies for the formation of the green tides among different years and the dominance of asexual and vegetative reproductive mode for the floating period.Besides,population analysis was conducted,and it revealed a very low genetic diversity among samples from Subei Shoal and the Qingdao coast in the same year and a higher divergence among samples in different years.The results showed the efficiency of 2b-RAD in the exploration of SNPs in U.prolifera and provided the first genome-wide scale evidence for the origin of the large-scale green tides on the Qingdao coast.This study improved our understanding of the reproductive strategy and genetic diversity of the green tide causative species and will help further reveal the biological causes of the green tide in China.

Experimental clearance rate and intraguild predation of jellyfish Cyanea nozakii

Cyanea nozakii,a common jellyfish distributed in offshore China,has a complex trophic relationship with other zooplankton groups.However,few studies have reported the predation rates and prey selection patterns of C.nozakii medusae on different prey items.Research is also lacking on the intraguild predation of Aurelia coerulea(another common bloom jellyfish in offshore China)by C.nozakii.To address the knowledge gaps,the clearance rates of C.nozakii for different prey items,including copepods(small<1000μm and large>1000μm),fish larvae,and gelatinous prey(hydromedusae,A.coerulea ephyrae,and chaetognaths),were measured.The influence of predator size on the clearance rate was also determined.Additionally,we examined the intraguild predation of C.nozakii on A.coerulea medusae.The clearance rates of C.nozakii varied widely with prey organisms,being independent of prey concentrations.Gelatinous organisms,except for chaetognaths,were captured with considerably high efficiency,followed by fish larvae and copepods,indicating the preferential prey selection of gelatinous organisms by C.nozakii.The clearance rate increased linearly with the cross-sectional area of C.nozakii.Body size in medusae may,to some extents,underpin their capacity to capture more prey by increasing the encounter rate and capture success through ontogeny.C.nozakii preyed voraciously on A.coerulea in high feeding efficiency,but the clearance rate decreased with increasing A.coerulea(as prey)size.This phenomenon of intraguild predation suggests a speculative hypothesis of potential population regulation of A.coerulea by C.nozakii.The information regarding the feeding ecology of C.nozakii reported in this study is important for understanding plankton dynamics in marine ecosystems with extensive occurrences of this jellyfish.

Photobehaviors ofthe calanoid copepod Calanus sinicus from the Yellow Sea to visible and UV-B radiation as a function of wavelength and intensity

The effects of wavelength-specific visible light, white light, and ultraviolet-B (UV-B, 280- 315 nm) on selected behaviors, grazing rate, spawning rate, and hatching rate of the marine copepod Calanus sinicus collected from the Yellow Sea were studied. Calanus sinicus placed in a partitioned experimental system responded positively phototaxis to blue-cyan and yellow light but negatively to orange light and UVB. No obvious dodge activity was found among C. sinicus irradiated with <0.005 mW/cm^2 UV-B. Under 0.20, 0.30 and 0.50 mW/cm^2 UV-B radiation, the lethal half times of individuals were 30.47, 2.86, and 1.96 h, respectively. Grazing of C. sinicus was restrained at >0.10 mW/cm^2 UV-B, whereas yellow-red light stimulated grazing. Egg production rate was highest at a white-light intensity of 1.58 mW/cm^2, with an average rate of 10.04 eggs/(female·d). These results are consistent with the observed phenomenon that C. sinicus in the Yellow Sea mostly spawn near dawn. Our results indicate that light intensity and spectrum are important factors affecting the diel vertical migration of C. sinicus under natural conditions in the Yellow Sea.

Intraguild predation by polyps of three scyphozoan jellyfish:Nemopilema nomurai,Aurelia coerulea,and Rhopilema esculentum

Jellyfish blooms occur worldwide and have resulted in serious problems in tourism,fisheries,coastal industries,and the marine ecosystem.The life cycle of scyphozoan jellyfish consists of a pelagic medusa stage and a benthic polyp stage.Success of asexual reproduction of the polyps determines directly the number of medusae;thus,the polyp stage is the key to understanding the population dynamics of medusae.Nemopilema nomurai,Aurelia coerulea,and Rhopilema esculentum are three scyphozoan jellyfish commonly inhabit in Chinese coastal waters.Polyps of A.coerulea are easily visible,while those of N.nomurai and R.esculentum remain yet to be found in the wild.However,distribution of the medusa indicates that the polyps of all three species may occur together.To evaluate the distribution pattern of polyps of the three species and explore intraguild predation by the polyps,we conducted a laboratory experiment that considered the attachment sequence and size relationship of calyx diameter of the polyps.We found that the polyps of A.coerulea preyed on polyps of the other two species in all treatments,except when polyps of R.esculentum were bigger than those of A.coerulea.The polyps of R.esculentum preyed on the polyps of N.nomurai only when polyps of R.esculentum attached first and were bigger than those of N.nomurai.Colonies of N.nomurai polyps were rarely found in the places inhabited by A.coerulea polyps.In addition,A.coerulea polyps are known to inhabit at depths of less than 20 m in coastal sea,thus,we speculate that N.nomurai polyp colonies might occur at depths of more than 20 m.Therefore,our finding that polyps of A.coerulea aggressively preyed on polyps of other species may help understand other such systems of jellyfish bloom in the world.

Carbon biomass,production rates and export flux of copepods fecal pellets in the Changjiang(Yangtze) River estuary

Copepod fecal pellets are ubiquitous throughout the oceans. Their production and export can represent a highly efficient pathway of carbon export. However, the role these fecal pellets play in carbon export in the Changjiang(Yangtze) River estuary is not well known. Two cruises were carried out in the Changjiang estuary in the spring and summer of 2013, during which time carbon biomass, production, and export of copepod fecal pellets were studied. Spring and summer fecal pellet carbon biomass ranged 0.30–1.01 mg C/m^3(mean=0.56±0.20 mg C/m^3) and 0.31–1.18 mg C/m^3(mean=0.64±0.24 mg C/m^3), respectively, significantly lower than phytoplankton. At most stations, fecal pellet carbon biomass was higher in surface or subsurface layers than deeper layers. Production rates ranged 0.65–1.49 pellets/(ind.?h)(mean=1.02±0.27 pellets/(ind.?h)) in spring and 0.62–1.34 pellets/(ind.?h)(mean=0.98±0.22 pellets/(ind.?h)) in summer, within the range reported in previous studies. Higher production rates of fecal pellets occurred at stations with higher chlorophyll a concentrations, and production rates of copepods of size 500–1 000 μm greater than copepods >1 000 μm during both cruises. The potential export flux of fecal pellets was slightly higher in summer(mean=68.95±14.37 mg C/(m^2 ?d)) than spring(mean=52.08±11.33 mg C/(m^2 ?d)) owing to higher summer copepod abundances. To our knowledge, this study is the first of its kind in the Changjiang estuary, and it confirms the significant role of copepod fecal pellets in local carbon export.

An enhanced underwater camera apparatus for seabed observation of megabenthic epifauna in the northern Yellow Sea

Seabed photographing has been applied with various underwater camera apparatuses(UCAs)for observations of megabenthic epifauna,which reveals more details than traditional sampling tools do.In this study,we improved a UCA named a towed underwater video-camera system(TUV system)with image processing software for seabed photographing in coastal areas up to 100 m.In May 2017,the TUV system was tested at 4 stations in the Zhangzi Island marine area in the northern Yellow Sea to investigate local megabenthic epifauna,especially brittle stars.At each station,more than 500 good seabed photographs each in area of 0.1550 m2 were obtained in just 10 min.Almost all of the epifauna larger than 1 mm could be identified from the photographs,including echinoderms,bivalves,cnidarians,and crustaceans.Three dominant brittle stars(Ophiopholis mirabilis,Ophiura sarsii vadicola,and Stegophiura sladeni)were spotted,and their abundance,disc diameter,cluster size,and coverage area were calculated and analyzed from the seabed photographs.The results show that the TUV system could be applied in coastal waters of hard sandy bottoms and could be used for quantitative investigations of megabenthic epifauna.

Reproductive cycle of Ophiopholis mirabilis (Echinodermata: Ophiuroidea) in Zhangzi Island area, northern Yellow Sea

Ophiopholis mirabilis is a common species with a high population density on the western coasts of the northern Pacifi c Ocean.The number of O.mirabilis has been increasing recently in the scallop aquaculture zone(the Zhangzi Island area,northern Yellow Sea)in China.To explore the mechanism of its population variation,the reproductive cycle of O.mirabilis was investigated in this area(39°04′N;122°51′E)from February 2017 through January 2018 and determined by the monthly gonad index(GI),histological examinations of the gonads and the oocyte size-frequency distribution.O.mirabilis had a clear annual reproductive cycle that was synchronous between males and females.Sea temperature and food availability played important roles in O.mirabilis reproduction.The GI value was less reliable for determining reproductive activity in O.mirabilis because the nutritive tissues within the gonads may be utilized to synthesize gametes,leading to a decrease in GI during maturation.The histological results also show that abundant nutritive phagocytes were present in the gonads of O.mirabilis,which,together with the germ cells,aff ected the weight of the gonads.In addition,the mature oocytes of O.mirabilis were relatively small(75–150μm),indicating that the larval development was planktotrophic.This study provided insights into the reproductive patterns and biology of O.mirabilis and is an essential basis for the quantity control of this species in aquaculture areas.

Natural predators of polyps of three scyphozoans:Nemopilema nomurai,Aurelia coerulea,and Rhopilema esculentum

Jellyfish blooms have become a hot research topic in recent decades because they pose a serious threat to fisheries,coastal industries,tourism,and the marine ecosystem.The life cycle of scyphozoan jellyfish consists of a pelagic medusa stage and a benthic polyp stage,where asexual reproduction and strobilation of the polyps directly affect the abundance of ephyra and subsequently medusa abundance.The dynamics of polyps are affected by both environmental and biological factors,and predation by natural predators is one of the most important biological factors.Nemopilema nomurai,Aurelia coerulea,and Rhopilema esculentum are three scyphozoan species that are commonly found in Chinese coastal waters,and previous studies reported that the survivorship of polyps differs among the three species when they are exposed to the same benthic community.To identify potential natural predators of polyps of these three species in Chinese coastal waters and to determine whether the predation rates on polyps of the three species differ,we collected 39 species of macrozoobenthos from the Bohai Sea,Yellow Sea,and East China Sea from May 2014 to June 2016 and conducted predation tests and predation rate measurements.We found that the nudibranchs Pleurobranchaea novaezealandiae,Okenia plana,and Chromodoris tinctoria and the sea anemones Paracalliactis sinica,Calliactis japonica,Anthopleura incerta,and Anthopleura midori could prey on the polyps of all three scyphozoan species.The predation rates increased with the body length of the predators.The predations rates were also related to the polyp species,although the different predators showed no consistent preference for a particular species of polyp.Our results indicate that introducing predators to locations inhabited by polyps might be a way to control the benthic polyp populations and prevent subsequent jellyfish blooms.