Reproduction,hatching success,and early naupliar survival in Centropages typicus

The broadcast spawner, Centropages typicus, is a very successful copepod species in many coastal areas of the Mediterranean Sea and North Atlantic Ocean. This review assembles the large amount of information on the reproduction and early life history of C. typicus that has emerged since the 1970s and has made this species one of the best-studied copepods, similar in that regard to species of Acartia and Calanus. Observations on mating behavior and the female gametogenic and oogenic cycles are presented, together with information on seasonal cycles of egg production rates in Mediterranean and Atlantic populations from various regions. These studies indicate a strong latitudinal gradient, with continuous reproduction and the main spawning season occurring earlier (late winter/spring) in warmer waters such as the Mediterranean Sea, compared to northern areas such as the North Sea and in the Kattegat, where C. typicus actively reproduces mainly in late summer and fall with reproduction ceasing altogether in winter in the German Bight. These observations strongly suggest that temperature is the controlling factor for reproductive activity in this species. Egg development times are also temperature dependent but do not vary with latitude, and there is as yet no conclusive evidence that diapause egg production occurs in C. typicus. Laboratory experiments have shown that food quantity and quality both affect fecundity and offspring fitness, but most of these studies have focused on diatom and dinoflagellate diets and non-algal prey have been strongly underrepresented, despite their importance for this omnivorous copepod. Large fluctuations in hatching success and naupliar survival have been reported in field surveys and have subsequently been related to maternal feeding history and food quality or toxicity in laboratory experiments. We identify future lines of research that will help to explain the interannual variability in breeding intensity and recruitment of C. typicus in the field based on better understanding of the species’ reproductive biology.     

Calanoid copepod resting egg abundance and hatching success in the sediment of the Seine estuary (France)

Benthic, viable resting eggs of calanoid copepods were found for the first time in the Seine estuary (France) during July 2008. Vertical distribution of the resting eggs in the sediment was determined up to 10 cm depth. Hatching success of the eggs extracted from different 1-cm thick sediment layers was experimentally tested immediately after extraction and after a long refractory phase (i.e. 11 months) of storage at low temperature (4–5 °C). The hatching success of resting eggs obtained immediately after sediment incubation was lower (0.72%) than the value observed after 11 months (4.50%) with an overall hatching success of 2.37%. The marine, calanoid copepod Temora longicornis was the primary species to hatch from the eggs; however, the estuarine calanoid copepod Eurytemora affinis also hatched from resting eggs. The mean abundance of eggs found in sediment (1.42 × 10⁶ eggs m⁻²) was comparable to that reported for other marine and estuarine calanoid copepods. The Seine estuary sediment had a high variability of egg abundance (between 0.14 and 8.10 × 10⁷ eggs m⁻³) suggesting that the hydrodynamics of this macrotidal estuary are likely responsible for this variability. Significant sediment resuspension occurs in the Seine estuary during flood periods and spring tides leading to resting eggs to contribute along the year to the nauplii recruitment of calanoid copepods. On average, around 400,000 nauplii m⁻³ month⁻¹ of the main calanoid copepods can emerge from the surface layer sediment in the Seine estuary, suggesting that resting eggs could play an important role in the population dynamics of key calanoid copepods in the Seine estuary.     

The contribution of microzooplankton to the diet of mesozooplankton in an upwelling filament off the north west coast of Spain

Incubation experiments were carried out daily during a Lagrangian experiment within an upwelled filament off the Galician coast to determine the importance of microzooplankton in the diet of calanoid copepods. Despite low chlorophyll concentrations the microzooplankton formed the minor component of the diet of the copepod community (7 to 15% of carbon ingested through autotrophic and heterotrophic prey). Ingestion of ciliates was greater than that of heterotrophic dinoflagellates, which reflected a higher abundance of ciliates in the water column. Heterotrophic nanoflagellates appeared also to be consumed by the copepods, although the very small size fraction (2–5μm) was probably not grazed by the larger copepods of Calanus spp. Grazing pressure by the copepods enumerated in the net samples was not sufficient to impact significantly the microzooplankton populations (2 to 51% of daily microzooplankton production was removed). Allometric relationships of grazing on microzooplankton for a range of numerically dominant copepod species are developed from the experimental results. The grazing pressure of the whole copepod community is estimated from these relationships. By considering the total mesozooplankton community we suggest that microzooplankton growth was probably restricted by metazoan grazers.     

Ecological niches of Arctic deep-sea copepods: Vertical partitioning,dietary preferences and different trophic levels minimize inter-specific competition

The biodiversity of pelagic deep-sea ecosystems has received growing scientific interest in the last decade, especially in the framework of international marine biodiversity initiatives, such as Census of Marine Life (CoML). While a growing number of deep-sea zooplankton species has been identified and genetically characterized, little information is available on the mechanisms minimizing inter-specific competition and thus allowing closely related species to co-occur in the deep-sea pelagic realm. Focussing on the two dominant calanoid copepod families Euchaetidae and Aetideidae in Fram Strait, Arctic Ocean, the present study strives to characterize ecological niches of co-occurring species, with regard to vertical distribution, dietary composition as derived from lipid biomarkers, and trophic level on the basis of stable isotope signatures. Closely related species were usually restricted to different depth layers, resulting in a multi-layered vertical distribution pattern. Thus, vertical partitioning was an important mechanism to avoid inter-specific competition. Species occurring in the same depth strata usually belonged to different genera. They differed in fatty acid composition and trophic level, indicating different food preferences. Herbivorous Calanus represent major prey items for many omnivorous and carnivorous species throughout the water column. The seasonal and ontogenetic vertical migration of Calanus acts as a short-cut in food supply for pelagic deep-sea ecosystems in the Arctic.     

Reproduction of the giant jellyfish, <Emphasis Type="Italic">Nemopilema nomurai</Emphasis> (Scyphozoa: Rhizostomeae), in 2006–2008 as peripherally-transported populations

This study investigated the sexual maturation process, release of spermatozoa or eggs and oocyte diameter of the rhizostomid medusae Nemopilema nomurai using samples collected from August 2006 to June 2008 from the waters around Korea and Japan, including peripheral areas outside the species’ usual habitat. Immature medusae were observed from June to October only in the western sector of the study area. The onset of spermatozoa and egg release occurred in September and October, respectively, and peaked in December and January. Medusae migrated eastward from source areas with the Tsushima Warm Current, where they formed gametes and spawned. Peak position and maximum oocyte diameter increased as the gonads developed according to the size-frequency distribution of oocytes. No fertilized eggs or embryos were found in the gonads. The correlation was analyzed with bell diameter, maximum oocyte diameter, sampling date, surface water temperature and gonad color to estimate which environmental factors and maturation indices were related to the maturation stage of females. Maturation stage correlated well with maximum oocyte diameter, which correlated negatively with surface water temperature. There was no significant correlation between bell diameter and maturation stage. Therefore, bell diameter was inappropriate for determining maturation index. Sex could not be distinguished clearly by gonad color. However, light pink gonads were more prevalent in males and various deep colors such as orange and brown were more frequent in female medusae.     

Experimental studies on the feeding and reproduction of Calanoides patagoniensis (Copepoda,Calanoid) in a southern upwelling ecosystem of the Humboldt Current

Seasonal cohorts of the large-sized copepod Calanoides patagoniensis allowed testing the relative ability of this species to exploit food resources that prevail during winter time in southern upwelling ecosystems of the Humboldt Current. This was achieved by considering a local winter flagellate assemblage (WFA), Thalassiosira rotula isolated from the local spring phytoplankton and a laboratory culture of Prorocentrum minimum, as food offers in consecutive 96 h experiments. Ingestion rates (IR) varied between 7 and 14 μg C f⁻¹ d⁻¹, egg production reached a peak of 70 eggs while egg production rates (EPR) varied between 27 ± 6 and 31 ± 4 eggs f⁻¹ d⁻¹. Feeding and reproductive traits were dependent on food offer, and after 72 h both IR and EPR decreased by 28% and 40% respectively, when copepods were steadily fed with the diatom. The relatively high reproductive performance supported by WFA was notable; showing the feeding behavior of this species can couple with food availability in the field with successful reproductive outcomes. Migration strategies allowing remain in upper food-rich coastal waters along with this flexible trophic behavior may better explain why this species is among the most recurrent ones in these variable yet productive upwelling areas.     

Short-term variation in copepod community and physical environment in the waters adjacent to the Kuroshio Current

A continuous survey examined short-term variations in the zooplankton community and physical ocean environment from the northeastern Izu Islands to Boso Peninsula in Japan. High copepod abundance and small upwellings in the surface layer and salinity minimum layer in the subsurface were observed on the north side of coastal fronts in the westernmost transect, moving southward as the Kuroshio Current left the Boso Peninsula. Thus, the salinity minimum layer might be a key factor forming upwelling and the fronts, leading to large abundance of coastal copepods off the northeastern Izu Islands. A community structure analysis of calanoid copepods revealed an intermediate belt assemblage between coastal and offshore (Kuroshio) assemblages. Copepod abundance was remarkably low and Ctenocalanus vanus dominated (nearly 37%) in the intermediate belt zone, indicating that C. vanus has a relatively high tolerance to adverse environments for calanoid copepods. As the Kuroshio Current left the Boso Peninsula, the coastal assemblage expanded in the same direction, and the intermediate belt assemblage off the northeastern Izu Islands disappeared. The largest population of Calanus sinicus was found along the two western transects off the northeastern Izu Islands (>1000 m depth), which was assumed to be transported from Sagami Bay and advanced southwestward while growing from copepodite stages CIII to CV. Larvae of C. sinicus would be an important food for fish larvae in addition to Paracalanus parvus s.l., the numerically dominant species in the coastal assemblage, and C. vanus under the adverse conditions for coastal copepods.     

Reproductive resilience of ice‐dependent Antarctic silverfish in a rapidly changing system along the Western Antarctic Peninsula

The Western Antarctic Peninsula (wAP) is globally one of the systems most heavily impacted by climate change, notably steep declines in sea ice extent. In forage species, reproductive resilience to change is particularly important because population fluctuations are rapidly communicated through the system via trophic interactions. The reproductive traits of the ice‐dependent forage species Antarctic silverfish (Pleuragramma antarctica) from different areas along the wAP and at the tip of the Antarctic Peninsula were investigated through macroscopic and histological analyses of gonads, with the aim to assess its reproductive potential and to test for spatial differences in fecundity and spawning season. Fish samples were collected in late summer off Charcot Island, in Marguerite Bay and off Joinville Island; no fish were caught in the central wAP. Samples from Charcot Island and Marguerite Bay consisted of adults in developing gonad stage, whereas those from Joinville consisted almost exclusively of juveniles. Mean GSI was relatively low (2–3%) and similar in both sexes, as specimens were still far from being actively reproducing. Developing females exhibited two discrete, though partially overlapping modes of oocytes of different size, with vitellogenic oocytes measuring 0.5–1.0 mm. Absolute and relative fecundity ranged between 3000 and 12,000 eggs per female and between 80 and 190 eggs·g^?1, with a strong relationship between absolute fecundity and body size. These results were consistent with a single population at Charcot Island and Marguerite Bay and indicated substantial reproductive potential, which may mitigate population isolation and reductions in habitat availability but cannot ultimately offset catastrophic loss of spawning habitat linked to sea‐ice retreat.     

Vertical zonation and distributions of calanoid copepods through the lower oxycline of the Arabian Sea oxygen minimum zone

This paper provides the first comprehensive analysis of calanoid copepod vertical zonation and community structure at midwater depths (300–1000 m) through the lower oxygen gradient (oxycline) (0.02 to 0.3 ml/L) of an oxygen minimum zone (OMZ). Feeding ecology was also analyzed. Zooplankton were collected with a double 1 m² MOCNESS plankton net in day and night vertically-stratified oblique tows from 1000 m to the surface at six stations during four seasons as part of the 1995 US Joint Global Ocean Flux Study (JGOFS) Arabian Sea project. The geographic comparison between a eutrophic more oxygenated onshore station and an offshore station with a strong OMZ served as a natural experiment to elucidate the influence of depth, oxygen concentration, season, food resources, and predators on the copepod distributions.Copepod species and species assemblages of the Arabian Sea OMZ differed in their spatial and vertical distributions relative to environmental and ecological characteristics of the water column and region. The extent and intensity of the oxycline at the lower boundary of the OMZ, and its spatial and temporal variability over the year of sampling, was an important factor affecting distributional patterns. Calanoid copepod species showed vertical zonation through the lower OMZ oxycline. Clustering analyses defined sample groups with similar copepod assemblages and species groups with similar distributions. No apparent diel vertical migration for either calanoid or non-calanoid copepods at these midwater depths was observed, but some species had age-related differences in vertical distributions. Subzones of the OMZ, termed the OMZ Core, the Lower Oxycline, and the Sub-Oxycline, had different copepod communities and ecological interactions. Major distributional and ecological changes were associated with surprisingly small oxygen gradients at low oxygen concentrations. The calanoid copepod community was most diverse in the most oxygenated environments (oxygen >0.14 ml/L), but the rank order of abundance of species was similar in the Lower Oxycline and Sub-Oxycline. Some species were absent or much scarcer in the OMZ Core. Two copepod species common in the Lower Oxycline were primarily detritivorous but showed dietary differences suggesting feeding specialization. The copepod Spinocalanus antarcticus fed primarily on components of the vertical particulate flux and suspended material, a less versatile diet than the co-occurring copepod Lucicutia grandis. Vertical zonation of copepod species through the lower OMZ oxycline is probably a complex interplay between physiological limitation by low oxygen, potential predator control, and potential food resources. Pelagic OMZ and oxycline communities, and their ecological interactions in the water column and with the benthos, may become even more widespread and significant in the future ocean, if global warming increases the extent and intensity of OMZs as predicted.     

Batch fecundity and spawning frequency of southern blue whiting (Micromesistius australis) in the southwest Atlantic Ocean

The reproductive biology of southern blue whiting (Micromesistius australis) inhabiting waters of the southwest Atlantic Ocean (52°‐53°S and 57°‐62°W) was studied through histological analysis of the ovaries. During 2003, the spawning peak occurred at the beginning of September south of the Malvinas Islands. Batch fecundity of this species ranged from 35 000 (41 cm total length, TL) to 245 000 (62 cm TL) oocytes and relative batch fecundity was 122 ± 43 (Mean ± SD, n = 24) oocytes per gram somatic weight. The average proportion of mature females with stage‐0 post‐ovulatory follicles was 0.25 for September 2003, which indicated a spawning frequency of 4 days during the peak of the season. Large females produced more egg batches than small spawners during the same period, but no significant relationship was found between oocyte dry weight and female size of southern blue whiting.     

Factors affecting the elimination of PCBs in the marine copepod Acartia tonsa

The effects of feeding, egg laying, and fecal pellet production on the elimination of polychlorinated biphenyls (PCBs) from the marine copepod Acartia tonsa were studied in a series of experiments. Copepods were exposed to ¹⁴C-labelled Aroclor 1254 and allowed to depurate in clean seawater. Copepods fed during depuration eliminated PCBs more rapidly than unfed copepods whether or not the original PCB exposure medium had contained food. Both eggs and fecal pellets contained PCBs during depuration, with the weight specific concentration of PCB in the eggs (up to 407 ppm, dry weight) exceeding four times that in the females that produced them. Female copepods eliminated PCBs twice as rapidly as males, indicating that egg production is an important route for PCB elimination.     

Geographical implications of seasonal reproduction in the bat star Asterina stellifera

The reproductive cycle and environmental cues that regulate gonad production in Asterina stellifera were studied from April 2009 to April 2011 in a rocky subtidal habitat at the southernmost limit of its distribution (Mar del Plata, Argentina). The geographic variation in reproductive traits between latitudinal range limits was analyzed. The gonadal and pyloric caeca weight varied with sea star size and time in both sexes. Despite a previous study which suggested the absence of recruitment in a 4 year period, our data of the same period demonstrated that spawning happens from early spring to early summer. The gonad and pyloric caeca weight did not show an inverse relationship, this suggested that there is no dependence on energy transfer between the organs and that the bat star presented a good nutritional state. Seawater temperature appears to be the variable explaining gonad proliferation at the range limits of A. stellifera distribution. Furthermore, differences in sex ratio, oocyte production, oogenesis duration and capability of energy transformation into ova were found between range limits.     

Zooplankton distribution across Fram Strait in autumn: Are small copepods and protozooplankton important?

We investigated zooplankton distribution in September 2006/2007 at eight stations across Fram Strait in contrasting water masses ranging from cold Polar water to warm Atlantic water. Our main objectives were: (1) to describe the plankton community in the upper 200 m during autumn, and (2) to investigate the importance of small-sized copepods and protozooplankton in an arctic ecosystem when the majority of the large Calanus species had entered diapause. We sampled both with a WP-2 net and Go-Flo bottle and show that small copepods <1 mm are significantly undersampled using a WP-2 net with 90 μm mesh.Small copepods and protozooplankton made a significant contribution both in terms of abundance and total zooplankton biomass at all stations in September, when the large calanoid copepods had left the upper 200 m. The dominating group in the upper 60 m at all stations was Oithona spp. nauplii and their daily estimated grazing potential on the <10 μm phytoplankton ranged from 0.1% to 82% of the standing stock. Both Oithona copepodites and nauplii biomass showed a significantly positive relation with temperature, but not with potential food. Heterotrophic protozooplankton, on the other hand, were most likely bottom-up regulated by the availability of phytoplankton <10 μm. We hypothesise that Oithona nauplii and protozooplankton compete for food and conclude that there was a strong link between the zooplankton community and the microbial food web in Fram Strait.     

Aggregation of the Arctic copepod Calanus hyperboreus over the ocean floor of the Greenland Sea

According to combined observations from vertical plankton tows, dredging with epibenthic nets 1 m above the ocean floor, video recordings and acoustic data from a scanning sonar obtained during descent and during deployment on the ocean floor, the calanoid copepod Calanus hyperboreus was aggregated in high concentrations near the ocean floor of the Greenland Sea between 2300 and 2500 m during late July and August. Concentrations were highest very close to the ocean floor and decreased rapidly further upward. These nearly mono-specific aggregations were apparently drifting in cloud-like formations with a horizontal extension of ca. 270 m with the near-bottom currents. Maximum abundances observed were up to 2 orders of magnitude higher than in the water column. The biomass in the bottom 20 m layer was around 18% of the biomass in the rest of the water column. Stage composition, reduced metabolic rates and insensibility to mechanical stimuli indicate that these C. hyperboreus were representing the resting population. The fact that high concentrations were observed during deployments lasting >1 d and in 3 years suggests that aggregation near the ocean floor is a regular, rather than an extraordinary, pattern in the life history of C. hyperboreus in the Greenland Sea, but there is need for comparison with other seas and eventually other Calanus species.     

Reproductive biology of horse mackerel Trachurus capensis and Trachurus trecae in Namibia and Angola

Harvest control measures for Namibia and Angola are defined using reference points for spawning biomass, which requires knowledge of spawning potential and dynamics of fishery stocks. The reproductive biology of two species of horse mackerel, Trachurus capensis and Trachurus trecae, in these waters was investigated for spawning distribution in time and space. Samples were collected during two scientific cruises conducted in February and July/August 2007 off the coast of Namibia and Angola. A total of 396 fish ovaries was examined to determine the spawning dynamics of the two species. The ovaries were histologically analysed and their oocyte distributions mapped. The length (TL)–weight relationships of the two species differed significantly. The observed development of oocyte recruitment and absence of a hiatus in the oocyte distribution indicated that these species are indeterminate spawners. Analyses of the data from the NatMIRC database (2001–2004) indicated annual changes in gonadosomatic index (GSI) for T. capensis at different latitudes, but these changes were small. GSI also fluctuated between years and areas, but there were no clear spatial or temporal trends, with averages ≤4.0% at all stations. The GSI of both species was not indicative of the histologically determined development of gonad maturity. Oocyte maturation for T. capensis indicated that smaller and younger fish spawn first. Batch fecundity (F) for T. trecae was best described by F = 4 986 TL (cm) – 99 645 (r ² = 0.83). This study provided the first analyses of oocyte development in time and space for the two species in the region.     

The zooplankton community in the Greenland Sea: Composition and role in carbon turnover

The micro- and mesozooplankton communities in surface waters of the Greenland Sea are described based on data from five cruises covering an annual cycle. Special emphasis is given to the summer period (June and August), prior to and after the descent of Calanus spp. Calanus spp. dominated the copepod community during the spring bloom and in the beginning of the summer. However, during the summer, there was a pronounced shift in the zooplankton composition in the euphotic zone. In contrast to what has been observed in other Arctic systems, smaller genera such as Pseudocalanus spp., Oncaea spp. and Oithona spp. became abundant and the total copepod biomass remained high after the Calanus spp. descended for hibernation. The peak protozooplankton biomass in the Greenland Sea (June) co-occurred with the peak in Calanus spp. Protozooplankton biomass then decreased during the summer. Growth of protozooplankton and grazing rates of the two dominating non-Calanus genera, Oithona and Pseudocalanus, were measured. For both copepod genera, protozooplankton constituted 40% or more of the diet, and maximum clearance was on prey items with an equivalent spherical diameter between 15 and 30 μm. The non-Calanus components of the zooplankton community were responsible for 70–99% of the total zooplankton grazing on phytoplankton during summer and were crucial for the recycling and respiration of primary production.     

Living on the edge: feeding of subtropical open ocean copepods

The objective of this study was to provide quantitative information on environmental feeding rates of warm water oceanic epipelagic copepods. We determined clearance rates at 23 °C for various particle size ranges in shipboard studies in the western oligotrophic subtropical Atlantic Ocean for females of the calanoid species Clausocalanus furcatus and Mecynocera clausii. These in situ clearance rates were then applied to the various particle size ranges of environmental particle spectra of auto‐ and heterotrophs at different depths from three stations in the western Atlantic. After calculating the metabolic demands of each of these two copepod species and applying an assimilation efficiency of 90%, we determined that C. furcatus meets its metabolic demands in all six cases, and M. clausii in two of six cases. Clausocalanus furcatus would also meet its energy demands at 25 °C, where it is often found, while M. clausii at 20 °C, where it is regularly found, would cover its metabolic needs in four of six cases. It is hypothesized that these species, and most likely most of the other co‐occurring copepod species, are limited in their abundance by food availability, or, better said, are ‘living on the edge’ in relation to food abundance.     

Calanoid copepods assemblages in Pearl River Estuary of China in summer: Relationships between species distribution and environmental variables

To study the relationships between species distribution and environmental variables, calanoid copepods were classified and enumerated, and environmental variables were analyzed along 91 sites of the Pearl River Estuary (China) and adjacent waters. The 91 sites were divided into 3 groups based on surface salinity: the Inner area had salinity level below 10‰ and was mostly located in the Pearl River Estuary, the Coastal area had salinity between 10‰ and 32‰ and was almost located along the coastline, while the Offshore area had salinity level higher than 32‰ and was mainly located in the open area. Indicator Species Analysis was conducted to identify the indicator species within each group from among all calanoid copepods taxa. We found that the Inner area was characterized by Acartiella sinensis, Pseudodiaptomus poplesia and Pseudodiaptomus inopinus, the coastal area was characterized by Temora turbinata and Subeucalanus subcrassus, and the Offshore area was characterized by Paracandacia truncata, Subeucalanus subtenuis, Euchaeta rimana, Pareucalanus attenuatus, Rhincalanus cornutus, Cosmocalanus darwini, Centropages gracilis, Undinula vulgaris, Nannocalanus minor and Paraeuchaeta russelli. Canonical Correspondence Analysis (CCA) was used to identify relevant biotic and abiotic parameters that can best explain the distribution of calanoid copepods. CCA showed that during the summer of 2006, salinity, nutrient variables, especially SiO₃–Si, NO₃–N and DTN, Depth and Chl a were the environmental variables that strongly impacted the distribution and community structure of calanoid copepods.     

Feeding selectivity of calanoid copepods on phytoplankton in Jangmok Bay,south coast of Korea

Grazing impacts of calanoid copepods on size-fractionated phytoplankton biomass [chlorophyll (Chl)-a] were measured in Jangmok Bay, Geoje Island, Korea, monthly from November 2004 to October 2005. The ingestion rate of calanoid copepods on total phytoplankton biomass ranged between 1 and 215 ng Chl-a copepod^?1 day^?1 during bottle incubations. Results indicated that microphytoplankton (> 20 μm) was the primary food source for calanoid copepods in grazing experiments on 3 phytoplankton size categories (< 3 μm, 3–20 μm, and > 20 μm). The ingestion rate on microphytoplankton showed a significant increase (r = 0.93, p < 0.01) with Chl-a concentration. Nanophytoplankton (3–20 μm) showed a negative ingestion rate from June 2005 to October 2005, but the reason is not completely understood. Calanoid copepods were unable to feed efficiently on picophytoplankton (< 3 μm) due to unfavorable size. Calanoid copepods removed between 0.1% and 27.7% (average, 3.6 ± 15.8%) of the phytoplankton biomass daily during grazing experiments. Grazing pressure was high in winter and early spring (January–March: 15.6–27.7%), while low in summer (June–August: ?33.1–0.0%) and autumn (September–November: ?1.4–5.1%). Results suggest that calanoid copepods play an important role in controlling the biomass and size structure of phytoplankton in winter and early spring.     

Dietary separation between co-occurring copepods in a food-limited tropical coral reef of the Sanya Bay

Food differentiation among coexistent species in the field is important strategy for copepods to acquire materials and maintain population stabilization.In situ diet analysis of co-occurring six copepod species in coral waters of the Sanya Bay was conducted using a PCR protocol based on 18 S ribosomal gene.Various prey organisms were uncovered,including dinoflagellate,diatom,green algae and plant,protozoa and metazoan.All these spatially coexisting six species showed different dietary diversity,with the food niche breadth(B)ranging from 1.00(Temora turbinate in morning)to 10.68(Calanopia elliptica in night).While food overlap between all these copepods were low,with the average value of the diet niche overlap index being approximately 0.09.Even temporally co-existing species sampled from the same time point fed on different groups of prey items with the food overlap index of 0.04 to 0.07 in midday and night but 0 in morning.As the most important dominant copepod in the Sanya Bay,Subeucalanus subcrassus seems to be capable to regulate its feeding,by exhibiting a rhythm of herbivorous feeding in midday and carnivorous feeding in morning and night,to better coordinate with other competitors for utilization of food resources.For most copepods,none of the prey items belonged to the dominant phytoplankton in the ambient water,indicating that copepod can better their survival by widening the choice of potential food resources in food limited environment.The dietary separation observed here might be important strategy for copepod to maintain population stabilization and thriving in the Sanya coastal waters.