Seasonal variations in copepod body length: a comparison between different species in the Lagoon of Venice

The seasonal variation in length of the most abundant copepod species in the lagoon of Venice were compared and the relative influence of temperature, chlorophyll concentration and salinity examined. Temperature seems to be the primary factor influencing copepod body length in the lagoon of Venice, but the different species vary markedly in their response. Calanoid copepods (Acartia clausi, A. tonsa, Paracalanus parvus and Centropages ponticus) showed a more definite trend of size variation with temperature than the cyclopoids (Oithona nana and O. similis) and the harpacticoid Euterpina acutifrons. The size of the poecilostomatoids Oncaea media and O. subtilis was not affected by temperature, and was almost constant over time. Differences in the metabolic rates, longevity and specialization of calanoid, cyclopoid and harpacticoid copepods could explain their different responses to temperature.     

Space and time distribution of zooplankton in a Mediterranean lagoon (Etang de Berre)

In the Berre lagoon, a large brackish and swallow area near Marseille, the environmental factors (temperature, salinity, oxygen, suspended particulate matter and chlorophyll) generally display strong space and time variations. The rotifer Brachionus plicatilis and the copepod Acartia tonsa constitute the bulk of the zooplankton population during all the year. Their space and time distributions were studied in 23 stations distributed all over the lagoon, during four seasonal cruises (February, June, October, November), at surface and bottom layers. There is no marked difference in the horizontal and vertical distribution of the two species, (except in November when rotifers were prevailing in surface and copepods at depth) and in their time occurence. When the four series of data are pooled, correlation analysis show that A.tonsa is positively correlated with temperature, salinity and seston and negatively to oxygen and chlorophyll. B. plicatilis is positively correlated with temperature and seston, but also with chlorophyll, while salinity has a negative effect. The specific eggs number of both species is chlorophyll dependent. Considering seasonal cruises separately, some differences appear in the sense or the significance of these different correlations. The respective distribution of the two species is only partly dependent on the variation of the environmental factors: most of the variance remains unexplained, as indicated by the result of a stepwise multiple regression analysis using the most significant factors (temperature, salinity and oxygen explain 33 to 42% of the variance in Acartia, while temperature and salinity explain 27 to 28% of the variance in Brachionus). Thus, internal behavioral factors could also play a role in the distribution of organisms, particularly in some cases of aggregations of organisms observed during this study. As the two species occupied the same space habitat most of the year, they are potentially in competition for food. A way to optimize the food utilization could be the time separation of their feeding activity, nocturnal in Acartia and diurnal in Brachionus. Another way could be selective feeding upon food particles depending on their size (Brachionus being able to use finer particles than Acartia) or their quality (Brachionus being more herbivorous than Acartia) as demonstrated in some grazing experiments carried out in parallel.     

Spatial variations in size, weight and condition factor of the females of Acartia clausi (Copepoda: Calanoida) along a salinity gradient in two contrasting estuaries of the Basque coast (Bay of Biscay)

Variations in prosome length and width, dry weight and condition factor of female Acartia clausi copepods were studied at three salinities (35, 34 and 33 psu) in the euhaline region of two estuaries (Bilbao and Urdaibai) of the Basque coast, with different level of anthropogenic impact. Effect of the environmental variables upon the morphology of A. clausi females on a small geographic scale is discussed. In general, biometric variables showed no significant differences between the two estuaries, but dry weight and condition factor were significantly higher in the estuary of Urdaibai at 35 and 34 psu, whilst at 33 psu they were higher in Bilbao. Body dimensions decreased significantly with decreasing salinity in both estuaries, however, no similar trends were observed for dry weight and condition factor. Temperature appeared the main variable to account size variations, but once eliminated seasonal effect of the temperature body size was related with oxygen concentration in the estuary of Bilbao and with salinity in Urdaibai. This study reveals that morphological characteristics of A. clausi not are only dependent on the temperature, but also, within a limited geographical zone, on local differences in environmental variables, mainly salinity and oxygen concentration.     

Interactive effects of temperature and predation on an estuarine zooplankton community

The winter planktonic copepod community of the estuaries near Beaufort N.C. underwent a consistent transition from dominance by Acartia tonsa Dana to dominance by Centropages spp., which was associated with unpredictable decreases in water temperature and increases in abundance of predatory mysids. Centropages subsequently remained dominant until spring. Experimental manipulations of water temperature and mysid abundance in enclosures showed that copepod species composition was determined by an interaction of direct temperature effects with predatory interactions among copepods and mysids. Low temperatures stimulated recruitment of Centropages spp. and inhibited recruitment of Acartia tonsa. Temperature decreases also apparently stimulated migration of mysids into the study area. While A. tonsa was dominant, selective predation by mysids reduced the relative abundance of A. tonsa, but after Centropages became dominant mysid predation had no effect on copepod species composition. Predation by Centropages on the nauplii of other species probably contributed to its persistent dominance in the estuary. Rising temperatures in the spring favored recruitment of Acartia tonsa and inhibited Centropages recruitment. Selective predation by fish entering the estuary in spring may have contributed to the decline in abundance of Centropages spp. and mysids.     

Impacts of very warm temperature on egg production rates of three Acartiidae (Crustacea,Copepoda) in a Northern African lagoon

The daily Egg Production Rate (EPR) of the three Acartiidae copepods namely Acartia clausi, Paracartia latisetosa and Paracartia grani in the North Lagoon of Tunis was compared to temperature and food availability every season from March, 2003 to February, 2004 corresponding to the major seasonal peaks of their abundance and over a 20-days period. The daily EPR was evaluated by assuming a direct correlation with chlorophyll a, particulate organic carbon (POC), and some easily extractable macromolecular compounds from the seston, such as proteins, carbohydrates and lipids. The results showed significant differences in EPR between seasons and species. Temperature and salinity negatively affected the fecundity of A. clausi and positively that of P. latisetosa and P. grani. The A. clausi EPR was supported by chlorophyll a and sestonic proteins while that of P. latisetosa and P. grani correlated with POC suggesting that these two taxa feed on microzooplankton and detritus more than on phytoplankton. Overall, EPR data showed potential population recruitment evidently the highest in P. latisetosa in summer, P. grani in autumn and A. clausi in winter. Our results indicate that temperature and food quality are the determining factors of both abundance and reproduction of the copepods under study.     

Predation, production and the organization of an estuarine copepod community

The copepod community of the estuaries near Beaufort, NC underwenta consistent seasonal succession from a spring assemblage dominatedby the medium-sized copepod Acartia tonsa (1 mm) to a summer—fallassemblage dominated by the small-bodied copepods Parvocalanuscrizsrirostris and Oithona colcarva (both 0.5 mm). However,in enclosure experiments during this period, A. tonsa dominatedthe community, due to higher growth rates and its predationon the nauplii of other species. Nutrient additions enhancedthe dominance by A. tonsa. The decline in abundance of A. tonsain the estuary was associated with increased abundance of planktivorousanchovies and silversides. In other enclosure experiments, planktivorousfish eliminated A. tonsa and other large copepods, althoughthey persisted in enclosures lacking fish. I conclude that predationby size-selective planktivorous fish prevents dominance by A.tonsa during summer—fall. ¹Present address: Institute of Marine Sciences, University ofNorth Carolina, 3407 Arendell Street, Morehead City, NC 28557,USA     

Reproductive response of Euterpina acutifrons in two estuaries of the Basque Country (Bay of Biscay) with contrasting nutritional environment

The reproductive response involving egg number and size was studied for the egg carrying copepod Euterpina acutifrons in two estuaries with different nutritional environment (Bilbao and Mundaka). To characterise the available food for copepods, the concentration of particulate organic matter (POM), the combined concentration (PGL) and proportion of particulate protein, carbohydrate and lipids and the concentration of chlorophyll a were measured in the <20 μm seston fraction. Data on female size (prosome length) and abiotic factors, such as temperature, salinity and oxygen saturation in water, were also obtained to relate them to reproductive and nutritional variables. Correlation and stepwise regression analyses indicated that temperature via female size was the main factor controlling brood volume in a temporal context, whilst the nutritional environment had a secondary role. Nevertheless, the nutritional environment caused noticeable between-estuary differences in the reproductive response of this species. ANOVA tests showed that E. acutifrons clutch size and brood volume were significantly bigger in the estuary of Bilbao, which was richer in both total (POM) and labile (PGL) organic particulate matter than the estuary of Mundaka. In contrast, egg size, female size and phytoplanktonic food quantity did not differ significantly between estuaries. These results suggest that the organic enrichment with particles of detritical-heterotrophic origin enhanced E. acutifrons clutch size and brood volume in Bilbao. Egg size was closely related to phytoplankton in both estuaries, but a clear trade-off between egg size and number per clutch was observed only in Mundaka. Thus, we suggest that the reduction of egg number in order to increase egg size might happen only under food-limiting conditions, but not in organically rich and quite constant nutritional environments such as found those in Bilbao.     

Factors affecting feeding selectivity of visual predators on the copepod Acartia tonsa: locomotion,visibility and escape responses

Visual predation by fish on copepods involves prey encounter, attack and capture; during any of these processes prey selection can occur. Developmental changes in copepods, including increases in swimming speed, size and image contrast increase the encounter rate and distance at which they can be detected by predators. Copepods compensate for this increase vulnerability with age through diel vertical migration and improved escape capabilities. This study quantifies the changes in swimming speed and movement pattern with developmental stage of the copepod Acartia tonsa, using a video-computer system for motion analysis. Changes in visible size and image contrast with developmental stage were quantified under simulated natural illumination conditions using a video based image analysis system. The escape responses of the naupliar stages of the copepod Acartia tonsa were quantified in response to a stationary pipette sucking in water at a constant speed. Accurate quantification of the parameters that affect feeding selectivity of planktivorous fish will provide the basis for evaluation of their relative importance in future studies.     

Morphological variability of geographically distinct populations of the estuarine copepod Acartia Tonsa

Variations in the number of spines on the left and right posterior dorsal and posterior margins of the prosome as well as the length of the prosome of Acartia tonsa from three estuaries, the upper western side of the Chesapeake Bay, Montauk Bay near the eastern end of Long Island Sound and the coast of Peru were determined. The length of the prosome and number of spines in each of the four locations were used as an indication of morphological similarity between the populations.     

Feeding rates and productivity of the copepod Acartia bifilosa in a highly turbid estuary; the Gironde (SW France)

Acartia spp. are the dominant copepod species in the Gironde estuary, seaward of the turbidity maximum area. Acartia bifilosa develop a large population in spring and early summer whereas Acartia tonsa appear in late summer. High values and high variability of chlorophyll a/suspended particulate matter ratio are found seaward of the turbidity maximum area. Feeding rates of A. bifilosa were measured by fluorometry. Phytoplankton ingestion was found to be highly variable, between 8 to 80% of copepod carbon body weight. Except for adult females, copepods were heavier in summer than in winter. PB ratios, estimated by the instantaneous growth rate method, varied from 0.03 d^–1 to 0.14 d^–1. The gut contents and P/B ratios of Acartia bifilosa were related to chl a/SPM ratio. From those data, and a few obtained for A. tonsa, it is concluded that only in summer months phytoplankton ingestion is enough to maintain secondary production.     

Copepod production estimated by combining in situ data and specific temperature-dependent somatic growth models

Although growth in adult copepods is frequently assumed to be similar to juvenile growth, some evidence have pointed out that under in situ conditions, it can be lower, with, as a consequence, underestimation of secondary production. In addition, under field conditions, juvenile growth in copepods is close to maximum rates estimated at food-saturated conditions. Based on previous assumptions, this study aimed to test the applicability of a new approach for copepod production estimate, derived from temperature-dependent growth models and in situ data, such as seawater temperature and copepod biomass. For this purpose, site-specific copepod juvenile growth models, defined for Acartia tonsa and A. clausi populations from a Southern European estuary (Canal de Mira, Ria de Aveiro, Portugal), were used and copepod biomass was taken from zooplankton samples collected during 2 years at six sampling stations. By comparing the obtained results with published data, the feasibility of the approach for copepod secondary production estimates and its applicability in worldwide marine ecosystems was confirmed. Future studies should combine the estimates of adult and juvenile production in order to evaluate its relative contribution and to obtain a more precise estimate of secondary production.     

Bacteria Associated with the Surface and Gut of Marine Copepods

Little is known about the nature of bacteria associated with the surface and gut of marine copepods, either in laboratory-reared animals or in the natural environment. Nor is it known whether such animals possess a gut flora. The present report deals with studies of microorganisms isolated from healthy, laboratory-reared copepods of the species Acartia tonsa Dana, from several species of wild copepods collected from a marine or estuarine environment, and from laboratory dishes containing moribund copepods. Evidence for a unique gut flora in laboratory-reared animals is presented; the predominant bacteria were represented by the genus Vibrio. Other organisms such as Pseudomonas and Cytophaga were found less abundantly associated with the copepods and not specifically associated with the gut.     

Interannual variations in abundance and body size in Neocalanus copepods in the central North Pacific

As the integral components of zooplankton in the subarctic NorthPacific, the three Neocalanus species (N. cristatus, N. plumchrusand N. flemingeri) are characterized by an annual life cycleand rapid development in the surface layer during spring–summer.Patterns of interannual variation of abundance and body sizeof these Neocalanus species were analyzed using the time-seriesdata collected during the summers of 1979–1998 (20 years)at stations along the longitudinal transect line in the centralNorth Pacific, crossing five sub-areas (Alaska Current System,Subarctic Current System, Northern Transition Domain, SouthernTransition Domain and Subtropical Current System). In the southernsub-areas, quasi-decadal oscillation was observed for the 3-yearrunning mean of abundance and prosome length for copepoditestage 5 (C5) of the three Neocalanus species. Although the oscillationsignal diminished towards northern waters, it showed a positivephase during the early 1980s and 1990s and a negative phaseduring the late 1980s. In the northern waters, a biennial patternwas pronounced for anomalies of C5 prosome length for N. plumchrusand N. flemingeri, which was large in odd years and small ineven years. Significantly positive covariations among the threespecies were found for both abundance and prosome length aroundmid-latitude, where they were abundant. In the correlation analysis,these observed yearly patterns showed a statistically insignificantcorrelation with most environmental (integrated mean temperaturein surface waters, water column stability and chlorophyll aconcentration) or climatological (North Pacific Index and SouthernOscillation Index) variables. The regional difference of theoscillation signal and the synchronized covariation among thesespecies suggest that interannual variations of their abundanceand body size are mediated by common environmental force(s)with some spatial and temporal scales in the subarctic NorthPacific.     

Population dynamics of calanoid copepods in the subtropical mesohaline Danshuei Estuary (Taiwan) and typhoon effects

We studied the populations of the dominant calanoid copepods Pseudodiaptomus annandalei and Acartia spp. in the mesohaline Danshuei Estuary for two successive springs and summers. We analyzed environmental factors (i.e., temperature, salinity, chlorophyll a, light attenuation coefficient, and suspended particulate matter) and the population densities and structures at the surface and near the estuarine bed. The population of P. annandalei dominated the zooplankton community from surface to bottom, except during a medusa bloom in 2009 and during the post-typhoon period, when Acartia spinicauda dominated. The relationships between environmental factors and densities differed between and within populations of P. annandalei and Acartia spp. We concluded that the primary determinants of the P. annandalei population are predation and the typhoon-related freshwater runoff, which can strongly influence the copepod succession in this part of the estuary.     

Reproductive trade-off of the copepod Acartia tonsa in a hypersaline estuary of the Southwestern Atlantic. Temporal variations in the morphology of eggs

The purpose of the present work was to study the seasonal variations in egg production, morphology and hatching success in the cryptic species Acartia tonsa, taking into account variations in female size, population abundance and environmental factors in a turbid and hypersaline estuary. Sampling was performed during the austral warm (18–23°C and 32–36 salinity) and cold seasons (5–7°C; 32–38) in Bahía Blanca Estuary (BBE), Argentina, during 2007 and 2009. Field-collected females were incubated in the laboratory simulating in situ environmental conditions, and specimens from fixed samples were measured using optical and scanning electronic microscopy. Acartia tonsa’s marked seasonality in its reproductive traits was found to ensure its permanence in the water column all over the year. During the warm season, small-sized females were observed to invest their energy in the production of subitaneous eggs with high hatching success and smooth appearance (12.95?±?2.38 eggs f^?1 day^?1 and specific egg production rate (SEP) of 16.57%C f^?1 day^?1). During the cold season, females invested C in body mass as well as in the production of resting eggs of three different morphotypes (6.56?±?3.2 eggs f^?1 day^?1 and SEP of 7.37%C f^?1 day^?1). Although these morphotypes were found to show differences in surface ornamentation, they exhibited the same delayed hatching behaviour. The eggs with shorter spines were found to integrate the resting egg bank in BBE. Our findings confirming a delayed egg hatching behaviour and a great tolerance to low temperatures and high salinities in the A. tonsa population in BBE suggest that this possible strain is a valuable phenotype for aquaculture.     

Seasonal variations in upwelling and in the grazing impact of copepods on phytoplankton off A Coruña (Galicia, NW Spain)

The impact of grazing by copepods on phytoplankton was studied during a seasonal cycle on the Galician shelf off A Coruña (NW Spain). Grazing was estimated by measuring the chlorophyll gut content and the evacuation rates of copepods from three mesh-size classes: 200-500 (small), 500-1000 (medium), and 1000-2000 μm (large). Between February 1996 and June 1997, monthly measurements of water temperature, chlorophyll concentration, primary production rates, and copepod abundance, chlorophyll gut content, and evacuation rates were taken at an 80-m-deep, fixed shelf station. Additionally, the same measurements were collected daily during two bloom events in March and in July 1996. Small copepods were the most abundant through the seasonal cycle. The highest grazing impact, however, was due to the medium and large size classes. Grazing by small copepods exceeded grazing by medium and large copepods only during phytoplankton spring blooms. The impact of copepod grazing (considering all size fractions) was generally low. On average, 2% of the phytoplankton biomass and 6% of the primary production were removed daily by the copepod community. Maximum grazing impact values (9% of the phytoplankton biomass and 39% of the primary production) were found in mid-summer. These results suggest that most of the phytoplankton biomass would escape direct copepod grazing in this upwelling area.     

Effects of food on bacterial community composition associated with the copepod Acartia tonsa Dana

The estuarine copepod Acartia tonsa naturally carried diverse strains of bacteria on its body. The bacterial community composition (BCC) remained very conservative even when the copepod was fed different axenic algal species, indicating that the food per se did not much affect BCC associated with the copepod. In xenic algal treatments, however, copepod-associated BCC differed with each alga fed, even though the same bacterial source was used to inoculate the algae. In addition, starved copepods taken at the same location but at different times significantly differed in their BCC. Algal species composition and copepod life history therefore serve to regulate BCC associated with copepods, and spatial and temporal variations in algal species composition and copepod origin would alter bacteria–copepod interactions.     

Zooplankton-associated and free-living bacteria in the York River, Chesapeake Bay: comparison of seasonal variations and controlling factors

Zooplankton provide microhabitats for bacteria, but factors which influence zooplankton-associated bacterial abundance are not well known. Through a year-long field study, we measured the concentration of free-living bacteria and bacteria associated with the dominant mesozooplankters Acartia tonsa and Balanus sp. Free-living bacteria peaked in the summer, while zooplankton-associated bacteria peaked in summer and winter. No relationships were found between bacterial abundance per individual and zooplankter width, length, surface area or body volume. Multiple regression analyses indicated that free-living and Acartia-associated bacterial concentrations were explained by temperature, salinity, ammonium, chl a, and all term interactions. Balanus-associated bacterial concentration was explained by ammonium and phosphate. Ammonium significantly influenced all sampled bacterial communities. In laboratory experiments, copepods raised under high ammonium concentration had higher bacterial concentrations than those raised under low ammonium condition. Transplant experiments showed that high ammonium favored loosely attached bacteria, whereas low ammonium selected for firmly attached bacteria, suggesting greater exchange between free-living and zooplankton-associated bacterial communities in nutrient-rich systems. Additional sampling of other zooplankton taxa all showed high bacterial concentrations, supporting the notion that zooplankton function as microbial hotspots and may play an important, yet overlooked, role in marine biogeochemical cycles.     

Seasonal feeding and fecundity of the calanoid copepod Acartia tonsa in Long Island Sound: is omnivory important to egg production?

Many suspension-feeding copepods show omnivorous feeding behavior. However, the relative contribution to egg production of herbivorous and heterotrophic feeding in copepods remains an open question. In this study, we quantified pigment ingestion rates and egg production rates of the planktonic calanoid copepod Acartia tonsa from July to November of 1986 in Long Island Sound, a large temperate estuary. Pigment ingestion and egg production rates were better correlated to the > 10-µm chlorophyll size fraction than total chlorophyll. Maximum pigment ingestion and egg production rates were observed during the fall bloom in September. Pigment ingestion and egg production rates were linearly related. However, pigment ingestion rates accounted for only 48% of the variance in egg production rates. If female A. tonsa had fed entirely as herbivores over the course of the season, the observed gross efficiency of egg production, K₁ = egg production ratelpigment ingestion rate, in terms of nitrogen, would have been 0.68. This growth efficiency is considerably higher than the expected value, K ₁ = 0.38, for this species based on laboratory studies of herbivorous feeding. We suggest that the ratio H = K ₁/K₁ is a measure of the fraction of egg production that is due to herbivorous feeding. Thus, we infer that herbivory accounted for 56% (0.38/0.68) of egg production in A. tonsa in this study. The ratio H is an useful tool in examining the relative contribution to egg production of herbivorous and heterotrophic feeding in copepods.