Vertical distribution of diapausing Calanus pacificus (Copepoda) and implications for transport in the California undercurrent

Migration to deep water during diapause may contribute to the retention of several dominant oceanic calanoid copepod populations in eastern boundary current systems, where the mean flow of poleward undercurrents is in opposition to mean equatorward surface flow. The vertical distributions of Calanus pacificus late copepodid stages were measured at a 1200-m deep, open-ocean station in the Southern California Bight on 13 dates between April 2000 and March 2001 using a MOCNESS (multiple opening and closing net and environmental sensing system). Copepod vertical distribution was compared to the vertical position of the California Undercurrent. Diapausing C. pacificus were primarily found between 300 and 400 m at the beginning of the diapause season, in June and July, and between 250 and 350 at the end of the diapause season, in November and January. Depth distributions were broader from August to October, ranging from about 350 m to the maximum depth sampled, 1100 m, and the median depth of diapausing C. pacificus was deeper, up to 800-900 m, during this period. Maximal depths of diapausing C. pacificus, 1100-1000 m, were greater than have previously been reported. The mean depth of the California Undercurrent was 250 m, and its approximate depth range was 110-430 m. Diapausing C. pacificus CV were abundant in the California Undercurrent at the beginning and end of the diapause season, in June to July and late-November to January, suggesting that poleward transport of diapausing copepods in the California Undercurrent contributes to C. pacificus population retention in the California Current System.     

Life histories of large,grazing copepods in a subarctic ocean gyre: Neocalanus plumchrus,Neocalanus cristatus, and Eucalanus bungii in the Northeast Pacific

Life histories for the dominant, larger copepods of the subartic Pacific have been constructed by sampling from weatherships patrolling Ocean Station P (50°N, 145°W) during 1980 and 1981. Neocalanus plumchrus reproduced at depths below 250 m from July through February. Copepodite stages were present in surface layers from October through August with a large peak in numbers and biomass in spring. Fifth copepodites prepared for diapuse in 38 days during spring and descended to depths below 250 m. They commenced immediately to mature, and the females reproduced without renewed feeding. This schedule contrasts with that of the population in the Strait of Georgia, which remains in diapause from July to January and matures exclusively in January and February. There appears to be a difference between the coastal and oceanic habitats in preparing the diapausing individuals for maturation.Maturation of the diapausing stock of N. plumchrus maintained constant adult populations, averaging 714 males m^?2 from June through October and 1,434 females m^?2 from August through January. This constancy, together with the exponential pattern of decline in the diapause stock from September through February, suggests that density of adults may regulate maturation of fifth copepodites. Offspring of individuals delaying maturation and, thus, reproduction would benefit from the resulting moderation of intraspecific competition, probably that among copepodites.Reproduction of Neocalanus cristatus also occurred below 250 m, and, while spawning was continuous through the year, there was a substantial peak in November. That resulted in a peak of abundance for early copepodite stages in mid-winter, and a peak for the fifth copepodite stage in June. Stocking of the population of fifth copepodites in diapause below 250 m occurred from July through October. Some fifth copepodites were present in surface layers through the entire summer, and some younger copepodites persisted through the summer in progressively declining abundance just below the mixed layer. In autumn 1980 resurgence of early copepodite populations was rapid, occurring during the course of a prolonged October storm. The storm may have improved the habitat either by cooling the mixed layer or by resupplying nutrients to the euphotic zone.Eucalanus bungii reproduced in the mixed layer in early May and in early July. The first event was a spawning by females that had previously spawned in 1979 and then had returned to diapause. The second, heavier spawning (more females, more eggs per female) was by newly matured females from stocks that had overwintered as fifth copepodites. Nauplii peaked sharply in abundance on 19 July, one week after the peak in spawning. First and second copepodites peaked on 1 August, and all had advanced to the third copepodite stage by September. The diapause stock was established by September, principally between 250 and 500 m, and consisted of copepodite stages from third to sixth. Duration of the E. bungii life cycle appears to be typically two years. New nauplii develop as far as the third or fourth copepodite stage during their first summer, then enter diapause. The second summer they advance to the fifth copepodite stage and reenter diapause. Fifth copepodites mature in their third summer at two years of age. The males remain at depth and mate without subsequent feeding. Females migrate at night to the mixed layer where spawning occurs. About 20% of females that had already spawned in 1980 reentered diapause. They would reproduce again in their fourth summer at three years of age. All aspects of the life cycle suggest low mortality rates for copepodite stages, particularly at depth in the habitat occupied during diapuse. There can be no premium on rapid reproduction for E. bungii in the subartic Pacific, and there must even be benefit from spreading reproduction between years. This iteroparity may amount to a “bet-hedging” tactic, the young from a given mother having more than one chance to find sustaining conditions. It also produces gene flow between the year classes of the biennial life cycle.     

Stage-V copepodites of Calanus sinicus and Calanus jashnovi (Copepoda: Calanoida) in mesopelagic zone of Sagami Bay as identified with genetic markers, with special reference to their vertical distribution

We applied genetic makers to identify Calanus species occurring in Sagami Bay, Japan, in order to investigate their vertical distribution in the upper 1000 m. First, interspecific genetic distances of three gene loci, mitochondrial small ribosomal RNA (srRNA), nuclear internal transcribed spacers 1 (ITS1) and 2 (ITS2), were estimated from morphologically distinguishable adult females of Calanus sinicus, Calanus jashnovi and Calanus pacificus that were collected from Sagami Bay, the Kuroshio Extension and the Oyashio region, respectively. The highest levels of interspecific genetic distance were observed in srRNA, followed by ITS1 and ITS2. The intraspecific genetic distances within C. sinicus were much lower than the interspecific genetic distances, indicating that DNA sequences in these loci are consistent with the morphological differences. This information was used as a criterion for species identification based on DNA sequence variation, and allowed us to identify the fifth copepodites (CVs) or younger stages of these species. Next, the vertical distribution of Calanus species was investigated in Sagami Bay in May 2006, on the basis of a stratified sampling in the upper 1000 m. By applying the genetic markers, 23 individuals comprising all copepodite stages were allocated into either C. sinicus or C. jashnovi, and the small- and large-sized CVs were identified as C. sinicus and C. jashnovi, respectively. The total abundance of C. sinicus was highest at 0-50 m and decreased with depth. On the contrary, CV individuals of C. sinicus were abundant not only in 0-50 m but also below 200 m with minimum occurrences in 150-200 m depth. C. jashnovi was much less abundant than C. sinicus and comprised of only CIV and CV which occurred in the upper 100 m and deeper than 50 m depths, respectively. The abundance of C. sinicus in the 1000-m water column of Sagami Bay was at a level comparable to that in shelf waters, suggesting the importance of off-shelf individuals in the biological production and organic transport in the respective areas.     

Seasonal and depth-dependent variations in the size and lipid contents of stage 5 copepodites of Calanus finmarchicus in the waters of the Newfoundland Shelf and the Labrador Sea

We report on the variation in energy reserves of individual C5 copepodites of Calanus finmarchicus from the Newfoundland continental shelf and the Labrador Sea collected from surface and overwintering (or bottom) depths prior to, during and after the expected timing of the onset of diapause. Overall, the trend was for a decreasing average prosome length as the year progressed for all locations although the decline was smallest in the Labrador Sea and greatest in the deep waters of the continental shelf. The size of the oil sac was closely linked to the weight of the copepodite but the form of this relationship showed substantial variations with depth and season. We show a clear increase in the relative oil sac volume for C. finmarchicus between late spring and late summer, by which time some animals had descended to diapause depths. The progressive decrease in oil sac volume of animals sampled at depth in the Labrador Sea between September and December suggests a significant loss of energy reserves during diapause. From the distribution of volumes and body sizes in December we estimate that 23–53% of individuals would not be able to meet the energetic cost of moulting and early gonad development. Overall, some of our observations appear to invalidate earlier hypotheses concerning the governing role of lipids in the life history of C. finmarchicus. However, assessment of the factors that influence entry into dormancy should be based on the relative probabilities of alternative strategies for successful reproduction (e.g. entering dormancy vs. continuing into a second generation).     

Reproductive success of Calanus pacificus during diatom blooms in Dabob Bay, Washington

While numerous laboratory studies in the last decade have shown that diatoms can induce reproductive failure in copepods, field evidence for a negative diatom effect is equivocal. To unambiguously elucidate the effects diatoms have on copepod reproduction in situ, we undertook a study of the abundance, distribution, grazing rates, and reproductive success of Calanus pacificus in Dabob Bay, Washington, USA, during two spring bloom periods. We simultaneously measured the phytoplankton composition, abundance, and distribution. Here we present results for the reproductive success of C. pacificus using four measurements: egg production rate, clutch size, egg hatching success, and naupliar survival (to the first feeding stage). Egg production rate was positively correlated with chlorophyll a concentration, and egg hatching success and naupliar survival were usually greater than 80%. However, in February 2002 and 2003 – during blooms of diatoms of the genus Thalassiosira – either egg hatching success, naupliar survival, or both were significantly depressed compared to other times in spring and summer. These effects, combined with evidence that C. pacificus was grazing aldehyde-producing Thalassiosira at the time of their blooms, indicate that diatoms can negatively affect copepod reproduction in the field, albeit only under specific circumstances and for brief periods.     

太平洋褶柔鱼冬生群资源丰度变动对ENSO事件的响应研究

太平洋褶柔鱼为大洋性经济鱼种,具有一年生命周期,其资源变动受气候和海洋环境条件的显著影响。本研究根据日本提供的2003-2012年太平洋褶柔鱼冬生群体的渔业统计数据,结合产卵场环境数据以及尼诺指数ONI（定义为Niño 3.4区海表温度距平值）,分析不同气候条件下（厄尔尼诺和拉尼娜）太平洋褶柔鱼冬生群体产卵场海表温度（SST）、叶绿素a（Chl-a）浓度以及适宜产卵面积（SSA）的变动情况及对其资源丰度（CPUE）的影响。结果表明,太平洋褶柔鱼冬生群体产卵场SST、Chl-a浓度和SSA具有明显的季节性变化。相关分析表明,各年CPUE与Chl-a浓度以及SSA具有显著的正相关关系（p<0.05）,但与SST相关性不显著（p>0.05）。此外,厄尔尼诺和拉尼娜事件通过驱动太平洋褶柔鱼冬生群体产卵场SSA和关键海域（25°-29°N,122.5°-130.5°E）内的Chl-a空间分布和大小变化,从而改变其资源丰度,但影响作用随各异常事件的强度不同而变化,具体表现为：发生弱强度厄尔尼诺事件时,产卵场SSA较高,Chl-a浓度处于较低水平,导致资源补充量处于较低水平,CPUE降低;发生中等强度厄尔尼诺事件时,产卵场SSA较低,但Chl-a浓度处于较高水平,导致资源补充量增加,CPUE处于上升水平;发生中等强度拉尼娜事件时,产卵场SSA和Chl-a浓度均处于较高水平,资源补充量显著增加,CPUE显著升高。研究表明,厄尔尼诺和拉尼娜事件对太平洋褶柔鱼冬生群体产卵场摄食孵化环境和资源丰度变动具有显著影响。     

Life history and population dynamics of Moerisia sp., a non-native hydrozoan, in the upper San Francisco Estuary (U.S.A.)

Some populations of gelatinous zooplankton appear to be increasing and causing ecological disruptions during large bloom events. The goal of this study was to investigate the life history and population dynamics of the small, invasive Moerisia sp. and its potential ecological impacts in the upper San Francisco Estuary, California. Medusae and polyps were collected from 8 and 2 sites, respectively, during 2007 and 2008. Polyps emerged from a resting stage during June. Asexual reproduction of medusae buds was positively correlated with temperature and negatively related to salinity. The production of polyp buds was positively correlated with dissolved oxygen and water transparency levels. Sexual reproduction, defined by the presence of eggs, was related to the size of medusae. Cessation of reproduction of both polyps and medusae occurred in October, when temperatures dropped below 17 °C. This life history is similar to other hydrozoans and allows Moerisia sp. to reach large numbers seasonally in the San Francisco Estuary, possibly contributing to the recent declines of pelagic fish and zooplankton.     

Tidal and annual variability of the population structure of Eurytemora affinis in the middle part of the Seine Estuary during 2005

Annual variability in abundance and population structure of the copepod Eurytemora affinis was studied in the maximum turbidity zone of the Seine Estuary in 2005. An Eulerian sampling strategy was applied monthly from March to July and from September to December. Chlorophyll a and suspended particulate matter (SPM) concentration, copepod abundance and stage distribution, and phytoplankton abundance were measured in sub-surface and near-bottom water during the ebb phase. Total E. affinis abundance was at a maximum in March and April (>200 × 10³ ind. m⁻³), and decreased from May to September (<25 × 10³ ind. m⁻³). This decrease corresponds to annual increases in temperature, salinity, chlorophyll a concentration and phytoplankton abundance, which was dominated by large diatoms, and decreases in SPM and river discharge. The phenology observed in 2005 was almost two months earlier compared to previous studies in the 1990s, when E. affinis reached maximum abundance in May and June. The low proportion of nauplii (<50%) in the population and high abundance of ovigerous females suggests that low recruitment is probably related to anomalously low temperatures in late winter (<5 °C). Whatever the horizontal position of the population in the estuary, adult and late copepodid stages are distributed in higher salinity than naupliar stages. Overall E. affinis population abundance was driven by parameters that characterize water masses at the tidal scale and by river discharge and chlorophyll a at the annual scale. By integrating the tidal effect, the high-frequency sampling protocol used appears to be optimal for investigating annual variability of planktonic communities in megatidal estuaries.     

西北冰洋陆坡区极北哲水蚤种群快速补充研究

在北冰洋的高纬度海区,陆坡—海盆之间的交换对极北哲水蚤（Calanus hyperboreus）的种群补充具有非常重要的意义。为了研究极北哲水蚤在西北冰洋种群补充的地理差异,我们利用2003年夏季所采集的样品,分析了该物种的丰度、种群结构和体长分布。从总丰度的地理分布来看,极北哲水蚤主要分布在楚科奇海与楚科奇深海平原之间的陆坡区（CS-slope）,而在水深较浅的楚科奇海并没有记录。在CS-slope区域,极北哲水蚤的总丰度在1 110.0—5 815.0个/m³之间,而其他海区的总丰度在40.0—950.0个/m³之间。从不同的发育期分布上来看,早期幼体（CI-CIV）在CS-slope区域占优势,而CV期幼体和成体在深水海盆区占优势。从体长的地理分布上来看,差异最为明显的是CⅢ期幼体,其在CS-slope区域的前体长在2.48—2.61 mm之间,而在其他海区的前体长在2.16—2.37 mm之间。与环境因子相关性的分析结果显示,早期幼体（CI-CIV）的丰度与叶绿素a的浓度呈正相关关系,而CV期幼体和成体却与叶绿素a的浓度呈负相关关系。我们的结果表明,极北哲水蚤可以通过加快第一个生长季节的发育速度而受益于初级生产力的增加,并且高生产力的CS-slope区域是陆坡-海盆之间种群补充的潜在来源。     

Comparison between amphipod assemblages associated with Caulerpa racemosa var. cylindracea and those of other Mediterranean habitats on soft substrate

The spread of the invasive alga Caulerpa racemosa var. cylindracea in shallow-water habitats can present different faunal assemblage composition. We compared the amphipod assemblages associated with C. racemosa and natural habitats found on shallow-water Mediterranean soft substrata. Four vegetated habitats were compared: C. racemosa, Caulerpa prolifera, Cymodocea nodosa and Posidonia oceanica with unvegetated substrata. Samples were collected during two sampling periods (September 2004 and March 2005). A total of 63 amphipod species were recorded. The results showed that the vegetated habitats sampled, including C. racemosa stands, supported a higher abundance and species richness of amphipods. Furthermore, the assemblage structure differed between the different habitats, while the abundance of some species was significantly different, depending on habitat. For example, Microdeutopus obtusatus was favoured by C. racemosa habitat; Ampelisca diadema was associated with C. prolifera beds; and Hyale schmidti was more abundant in P. oceanica meadows. Habitat invasion by C. racemosa can exert an important influence on biotic assemblages, modifying habitat structure and associated fauna.     

Temporal variation in abundance and fecundity of the invading copepod Eurytemora americana in Bahía Blanca Estuary during an unusual year

Eurytemora americana has been only reported as invader in Bahía Blanca Estuary, Argentina within the South Hemisphere. There are a few experimental researches under laboratory conditions done with this species and its reproductive behaviour around the world is very scarce. Consequently, it is still not possible to completely understand its population dynamics. In the present study, E. americana reproductive temporal behaviour and relationships among abundance, female size, egg production and hatching success were examined in the Bahía Blanca Estuary, during 2007 pulse. In order to determine the potential relationships between these variables and the environmental variables, experimental incubations were conducted in the laboratory simulating natural conditions. Spearman’s rank correlation was used to analyze the relationships among all variables. Temporal change of biotic and environmental variables was corroborated by a Mann–Whitney/Kruskal–Wallis non-parametric tests, with significant differences (p ? 0.01) in all variables throughout the study. Abundance population results showed very high values in relation to those recorded in recent years in Bahía Blanca Estuary. This response could be due to the unusual combination of environmental factors (polar wave with temperatures ≤6 °C and a drought period with high salinities, 32.7–36.6) recorded during the studied winter period. Significant positive correlations between abundance and salinity (p < 0.01, n = 226), and hatching success (p < 0.01, n = 25) as well as a significant negative correlation between abundance and chlorophyll a (p < 0.01, n = 226) were found. Although E. americana shows a k-strategy within its annual pulse, it presented two markedly distinct behaviours depending on temporal environmental variability. From July to early september, when the estuary evidenced high salinity, low temperature and high food availability, E. Americana showed large females, large clutch size and high hatching success. When environmental conditions became unfavorable from September to October, small females, small clutch size and very low hatching success were observed. The latter is associated with diapause egg laying which ensures population recruitment. According to our findings the particular combination of low temperatures, high salinities and high available food (i.e. variables which each year modulate its pulse) during 2007 winter–spring, favored the great development of E. americana. This invading species in its opportunistic role has managed to exploit a vacant niche in the estuary, developing two different behaviours within the k-strategy depending on change in environmental conditions.     

Spatial and temporal variability in coccolithophore abundance and production of PIC and POC in the NE subarctic Pacific during El Niño (1998), La Niña (1999) and 2000

Seasonal variations in coccolithophore abundance, chlorophyll, nutrients and production of particulate organic and inorganic carbon (POC and PIC) were determined along a coastal to oceanic east-west transect (Line P) culminating at Ocean Station Papa in the northeastern subarctic Pacific between 1998 and 2000. Offshore stations generally exhibited low seasonality in chlorophyll concentrations, with moderate seasonality in POC production. Near shelf stations showed a similar pattern to offshore stations, but were also characterized by sporadic events of higher POC productivity. During the 1998 El Niño, June was characterized by low chlorophyll and POC productivity along the transect, presumably as a result of depleted surface nitrate. In contrast, during the 1999 La Niña, and in 2000, higher POC productivity and surface nitrate occurred along the transect in June. Chlorophyll and POC productivity were similar in late summer in all 3 years. The coccolithophore population was usually numerically dominated by Emiliania huxleyi, particularly in June. Along the transect, abundance of coccolithophores was much higher in June during the 1998 El Niño (mean of 221 cells ml⁻¹) than in the 1999 La Niña (mean of 40 cells ml⁻¹), with their abundance in late summers of both years being very low. Abundances were even higher along the transect in June and the late summer of 2000 with sporadic ‘blooms’ of >1000 cells ml⁻¹ at some stations (cruise averages 395 and 552 cell ml⁻¹, respectively). Production rates of PIC did not consistently correlate with areas of high coccolithophore abundance. PIC production was high (100-250 mg C m⁻² d⁻¹) along the transect during June 1998, and low (1-40 mg C m⁻² d⁻¹) during both winters, June 1999 and during late summers of 1998 and 1999. The year 2000 was more complicated, with high rates of PIC production accompanying high abundance of coccolithophores in late summer, but lower rates of PIC production accompanying high coccolithophore numbers in June. Our data suggest that the abundance of coccolithophores and the production rates of PIC in the subarctic are higher than previously thought. Occasional PIC:POC production ratios of 1 or greater in 1998 and 2000 suggest that coccolithophores in this region could have a significant impact on the efficiency of the biological carbon pump.     

Oceanographic influences on the distribution and relative abundance of market squid paralarvae (Doryteuthis opalescens) off the Southern and Central California coast

Market squid (Doryteuthis opalescens) are ecologically and economically important to the California Current Ecosystem, but populations undergo dramatic fluctuations that greatly affect food web dynamics and fishing communities. These population fluctuations are broadly attributed to 5–7‐years trends that can affect the oceanography across 1,000 km areas; however, monthly patterns over kilometer scales remain elusive. To investigate the population dynamics of market squid, we analysed the density and distribution of paralarvae in coastal waters from San Diego to Half Moon Bay, California, from 2011 to 2016. Warming local ocean conditions and a strong El Niño event drove a dramatic decline in relative paralarval abundance during the study period. Paralarval abundance was high during cool and productive La Niña conditions from 2011 to 2013, and extraordinarily low during warm and eutrophic El Niño conditions from 2015 to 2016 over the traditional spawning grounds in Southern and Central California. Market squid spawned earlier in the season and shifted northward during the transition from cool to warm ocean conditions. We used a general additive model to assess the variability in paralarval density and found that sea surface temperature (SST), zooplankton displacement volume, the log of surface chlorophyll‐a, and spatial and temporal predictor variables explained >40% of the deviance (adjusted r² of .29). Greatest paralarval densities were associated with cool SST, moderate zooplankton concentrations and low chlorophyll‐a concentrations. In this paper we explore yearly and monthly trends in nearshore spawning for an economically important squid species and identify the major environmental influences that control their population variability.     

Diel vertical migration of the copepod Calanus sinicus before and during formation of the Yellow Sea Cold Bottom Water in the Yellow Sea

To understand the effects of the Yellow Sea Cold Bottom Water(YSCBW)on the diel vertical migration(DVM)of the copepod Calanus sinicus,we surveyed vertical distribution of C.sinicus at a fixed station in the Yellow Sea before(spring)and during(summer)formation of the YSCBW.Cold water(<10 C)was observed in the bottom layer when the water column was thermally stratified in summer,but the water column was thermally well-mixed in spring 2010.Samples were collected from five different layers at 3-h intervals using an opening-closing net.Adult females(1–155 ind./m3)showed a clear normal DVM pattern throughout the entire water column in spring,whereas adult males did not migrate.DVM of copepodite V(CV)individuals was not clear,but the maximum abundance of CI–CIV occurred consistently in the upper 10–20 m layer,where there was a high concentration of chlorophyll-a(Chl-a)(0.49–1.19μg/L).In summer,weak DVM was limited to cold waters beneath the thermocline for adult females(<30 ind./m3),but not for adult males.The maximum abundance of CI–CIV also occurred consistently in the subsurface layer(20–40 m)together with high concentrations of Chl-a(0.81–2.36μg/L).CV individuals(1–272 ind./m3)moved slightly upward nocturnally to the near-surface layer(10–20 m),where the average temperature was 25.74 C,but they were not found in the surface layer(0–10 m;28.31 C).These results indicate that the existence of the YSBCW affected food availability at depth and the vertical temperature distribution,leading to variation in the amplitude and shape of stage-specific vertical distributions(CI to adults)in C.sinicus before and during the formation of cold waters in the Yellow Sea during the study period.     

Diel variations of marine snow concentration in surface waters and implications for particle flux in the sea

Successive measurements of the size distribution and abundance of marine snow in the upper 100 m of the Santa Barbara Channel, California, with an in situ still camera system following 11 tagged water masses revealed a consistent pattern of nighttime decreases in the abundance of large particles. A net nocturnal reduction in particulate flux from the upper 100 m as calculated from camera profiles occurred in 75% of the day–night comparisons, and nighttime aggregate carbon losses resulted in a 38% average reduction in camera-derived aggregate flux. Intensive investigation of three stations for 24–48 h each indicated that nighttime decreases in aggregate concentrations and derived aggregate flux could be registered throughout the observed water column. Nocturnal decreases in marine snow concentration are unlikely to result from diel variations in the production of marine snow either as feeding webs of zooplankton or through variations in aggregation rates of smaller particles. Moreover, measured diel variations in the intensity of surface mixing and convective overturn during one of the 24 h deployments were not intense enough to produce aggregate fragmentation and reduced aggregate flux. Nighttime increases in large crustacean zooplankton (i.e., euphausiids and the large copepod Calanus pacificus) could explain some or all of the reduction in aggregate abundance at most stations. Fragmentation and consumption of marine snow by migrating macrozooplankton could produce our observed synchronous diel cycles in marine snow concentration. This is the first empirical evidence of a diel pattern in the concentration and calculated particulate flux of large sinking particles in near-surface waters. The data presented here are consistent with the only other existing diel study, which also reported decreases in marine snow abundance at night at 270 m depths in the oceanic north Atlantic. Diel variations in the sizes and concentrations of marine snow may impact water column processes dependent upon particle availability and size, such as grazing and remineralization, and may generate a diel cycle of food availability to the benthos.     

Seasonal changes in zooplankton abundance,biomass, size structure and dominant copepods in the Oyashio region analysed by an optical plankton counter

To identify seasonal patterns of change in zooplankton communities, an optical plankton counter (OPC) and microscopic analysis were utilised to characterise zooplankton samples collected from 0 to 150 m and 0 to 500 m in the Oyashio region every one to three months from 2002 to 2007. Based on the OPC measurements, the abundance and biomass of zooplankton peaked in June (0–150 m) or August (150–500 m), depending on the depth stratum. The peak periods of the copepod species that were dominant in terms of abundance and biomass indicated species-specific patterns. Three Neocalanus species (Neocalanus cristatus, Neocalanus flemingeri and Neocalanus plumchrus) exhibited abundance peaks that occurred before their biomass peaks, whereas Eucalanus bungii and Metridia pacifica experienced biomass peaks before their abundance peaks. The abundance peaks corresponded to the recruitment periods of early copepodid stages, whereas the biomass peaks corresponded to the periods when the dominant populations reached the late copepodid stages (C5 or C6). Because the reproduction of Neocalanus spp. occurred in the deep layer (>500 m), their biomass peaks were observed when the major populations reached stage C5 after the abundance peaks of the early copepodid stages. The reproduction of E. bungii and M. pacifica occurred near the surface layer. These species first formed biomass peaks of C6 and later developed abundance peaks of newly recruited early copepodid stages. From the comparison between OPC measurements and microscopic analyses, seasonal changes in zooplankton biomass at depths of 0–150 m were governed primarily by E. bungii and M. pacifica, whereas those at depths of 150–500 m were primarily caused by the three Neocalanus species.     

黄海春季和秋季中华哲水蚤脂类含量及组成分析

脂类物质在哲水蚤的休眠、繁殖以及个体发育过程中发挥着重要作用,通过分析2010年11月(秋季)和2011年4月(春季)黄海中华哲水蚤Calanus sinicus的脂类组成,对脂类各组分的生理功能进行了初步探讨。结果表明,春季、秋季中华哲水蚤雌体脂类平均含量占干质量的10.6%和12%,C5期个体脂类含量偏高,占到干质量的17%。中华哲水蚤所含脂类主要由蜡脂和磷脂组成,两种脂类约占总脂的90%,三酰甘油含量约占总脂的6%。结合种群结构对脂类的生理作用进行了初步探讨,发现:(1)蜡脂作为主要的能量储存方式,其含量变化反映出不同发育期、不同季节中华哲水蚤的能量需求变化;(2)磷脂作为结构性脂类在中华哲水蚤个体生长与生殖过程中具有重要作用;(3)三脂酰甘油在中华哲水蚤中的含量不高,但是可能在脂类转化利用过程中发挥作用。     

Distributions of Calanus spp. and other mesozooplankton in the Labrador Sea in relation to hydrography in spring and summer (1995-2000)

We collected mesozooplankton samples in the upper 100 m in spring or early summer each year between 1995 and 2000 along a section from Hamilton Bank (Labrador) to Cape Desolation (Greenland), and along additional sections in spring 1997 and early summer 1995. The North Atlantic waters of the central basin were characterised by the presence of the copepods Calanus finmarchicus, Euchaeta norvegica and Scolecithrocella minor and euphausiids. Calanus glacialis, Calanus hyperboreus and Pseudocalanus spp. were associated with the Arctic waters over the shelves. Amongst the other enumerated groups larvaceans were concentrated over the shelves and around the margins. Amphipods, pteropods and the copepods Oithona spp. and Oncaea spp. showed no definable relationships with water masses or bathymetry, while the diel migrant ostracods and chaetognaths were confined to deep water. Metrida longa, also a strong diel migrant, and Microcalanus spp., a mainly deep water species and possible diel migrant, were both sometimes quite abundant on the shelves as well as in the central basin, consistent with their likely Arctic origins.Analysis of community structure along the section across the Labrador Sea indicated that stations could be grouped into five different zones corresponding to: the Labrador Shelf; the Labrador Slope; the western and central Labrador Sea; the eastern Labrador Sea and Greenland Slope; and, the Greenland Shelf. The boundaries between zones varied spatially between years, but community composition was relatively consistent within a given zone and a given season (spring versus early summer). The relationship between community composition and water masses was not entirely straightforward. For example, Labrador Shelf water was generally confined to the shelf, but in spring 2000 when it also dominated the adjacent slope zone, the community in the Labrador Slope zone was similar to those found in other years. Conversely, in spring 1997, when Arctic organisms were unusually abundant in the Labrador Slope zone, there was no increased contribution of shelf water. In addition, North Atlantic organisms were often found on the shelves when no slope or central basin water was present.Although other organisms were sometimes very abundant, the mesozooplankton preserved dry weight biomass was dominated everywhere by the three species of Calanus, which together always accounted for ≥70%. One species, C. finmarchicus, comprised >60% of the total mesozooplankton biomass and >80% of the abundance of large copepods in spring and summer throughout the central Labrador Sea. In western and central regions of the central basin average C. finmarchicus biomass was ca 4 g dry weight m⁻² and average abundance, ca 17?000 m⁻² over both seasons. Highest levels (ca 7 g dry weight m⁻², >100?000 m⁻²) occurred in the northern Labrador Sea in spring and in eastern and southwest regions in early summer. C. hyperboreus contributed ca 20% of the total mesozooplankton biomass in the central basin in spring and <5% in early summer, while C. glacialis accounted for <1%. Over the shelves, C. hyperboreus contributed a maximum of 54% and 3.6 g dry weight m⁻², and C. glacialis, a maximum of 29% and 1 g dry weight m⁻², to the total mesozooplankton biomass.     

An overview of Calanus helgolandicus ecology in European waters

We review current knowledge and understanding of the biology and ecology of the calanoid copepod Calanus helgolandicus in European waters, as well as provide a collaborative synthesis of data from 18 laboratories and 26 sampling stations in areas distributed from the northern North Sea to the Aegean and Levantine Seas. This network of zooplankton time-series stations has enabled us to collect and synthesise seasonal and multi-annual data on abundance, body size, fecundity, hatching success and vertical distribution of C. helgolandicus. An aim was to enable comparison with its congener Calanus finmarchicus, which has been studied intensively as a key component of European and north east Atlantic marine ecosystems. C. finmarchicus is known to over-winter at depth, whereas the life-cycle of C. helgolandicus is less well understood. Overwintering populations of C. helgolandicus have been observed off the Atlantic coast between 400 and 800 m, while in the Mediterranean there is evidence of significant deep-water populations at depths as great as 4200 m. The biogeographical distribution of C. helgolandicus in European coastal waters covers a wide range of habitats, from open ocean to coastal environments, and its contribution to mesozooplankton biomass ranges from 6% to 93%. Highest abundances were recorded in the Adriatic and off the west coast of Spain. C. helgolandicus is generally found in 9-20 °C water, with maximum abundances from 13-17 °C. In contrast, C. finmarchicus is found in cooler water between 0 and 15 °C, with peak abundances from 0 to 9 °C. As water has warmed in the North Atlantic over recent decades, the range of C. helgolandicus and its abundance on the fringes of its expanding range have increased. This review will facilitate development of population models of C. helgolandicus. This will not only help answer remaining questions but will improve our ability to forecast future changes, in response to a warming climate, in the abundance and distribution of this important species.     

Juvenile bottlenecks and salinity shape grey mullet assemblages in Mediterranean estuaries

Previous research has suggested that competitive bottlenecks may exist for the Mediterranean grey mullets (Osteichthyes, Mugilidae) at the fry stage with the exotic Cyprinus carpio (Osteichthyes, Cyprinidae) playing a central role. As a consequence, the structure of grey mullet assemblages at later stages is thought to reflect previous competition as well as differences in osmoregulatory skills. This paper tests that hypothesis by examining four predictions about the relative abundance of five grey mullet species in 42 Western Mediterranean estuary sites from three areas (Aiguamolls de l'Empordà, Ebro Delta and Minorca) differing in the salinity level and occurrence of C. carpio. Field data confirmed the predictions as: (1) Liza aurata and Mugil cephalus were scarce everywhere and never dominated the assemblage; (2) Liza saliens dominated the assemblage where the salinity level was higher than 13; (3) Liza ramado always dominated the assemblage where the salinity level was lower than 13 and C. carpio was present; and (4) Chelon labrosus dominated the assemblage only where the salinity level was lower than 13 and C. carpio was absent. The catch per unit effort of C. labrosus of any size was smaller in the presence of C. carpio than where it had not been introduced, which is in agreement with the juvenile competitive bottleneck hypothesis. Discriminant analysis confirmed that the assemblage structure was linked to the salinity level and the occurrence of C. carpio for both early juveniles and late juveniles as well as adults. The data reported here reveal that the structure of grey mullet assemblages inhabiting Mediterranean estuaries is determined by salinity and competitive interactions at the fry stage.