Diel periodicity of Subantarctic copepods: relationships between vertical migration, gut fullness and gut evacuation rate

A zooplankton community in the Polar Frontal Zone north of SouthGeorgia was sampled for 5 days in February 1994. Feeding ofvarious copepodite stages of six copepod species was assessedby a series of gut fluorescence/gut evacuation experiments.Feeding periodicity was compared to vertical distribution andmigration patterns, as revealed by Longhurst Hardy PlanktonRecorder (LHPR) and ring net catches. Despite chlorophyll alevels reaching only 0.8 mg m^–3 and daily carbon rationsbased on phytoplankton intake being low, feeding of all thecopepods was restricted mainly to the 8 h night-time perid.During the daytime, the epipelagic community was verticallydispersed within the top 100 m. At night, upward migration bymost species led to a convergence of almost all zooplanktersin the upper half of the surface mixed layer. However, large-amplitudevertical migrations which crossed the thermocline were performedonly by Metridia lucens and Pleuromamma robusta. Although feedingby both migrants and non-migrants was mainly at night, therewas no diel signal in gut evacuation rate. The gut evacuationrates of the nine copepod species and stages differed significantly(5-fold) and were negatively related to the extent of theirdiel vertical migration. The long gut-passage times of the migratoryspecies, M.lucens and P.robusta, would have allowed them timeto defaecate some of the food eaten near the surface at depth,contributing to an active carbon transfer out of the mixed layer.However, their scarcity at this site meant that their grazingcomprised only -<1.4% of total copepod ingestion, so theircombined gut flux is likely to have been negligible (-0.4 mgcarbon day^–1).     

Diel vertical migration of the planktonic copepods at an upwelling station north of Taiwan, western North Pacific

A total of 178 copepod species were identified in an upwellingarea of the Mienhua Canyon off northern Taiwan, western NorthPacific during a spring cruise in 1995. Paracalanus aculeatus,Oncaea venusta and Clausocalanus furcatus were the three dominantspecies, comprising 43% of the total copepod numbers. Most copepodspecies performed normal diel vertical migration, descendingduring daytime and ascending at night to different depth zonesand with different rates. Some dominant copepod species, suchas P. aculeatus, C. furcatus, Temora discaudata and Canthocalanuspauper, apparently congregated in the surface water (between0 and     

Diel feeding and vertical migration of Sagitta serratodentata Krohn tasmanica Thomson (Chaetognatha) in the southern Benguela

The migration and diel feeding of Sagitta serratodentata tasmanicawere investigated at two stations in the southern Benguela.Pronounced diel migration and feeding was observed offshorewhere the food environment was markedly stratified. The inshorepopulation, because of the shallow bottom, did not migrate outof their prey field and did not exhibit a pronounced diel feedingpattern. Copepods comprise almost the entire diet of both populationsand, although Centropages brachiatus was heavily selected for,the data do not differentiate between size or species selection.     

Reproductive response of the copepods Calanoides carinatus and Calanus agulhensis to varying periods of starvation in the southern Benguela upwelling region

Egg production by the calanoid copepods Calanoides carinatusand Calanus agulhensis fed excess Thalassiosira weissflogiiwas monitored in the laboratory following starvation periodsof 1, 3, 5, 7 and 9 days. Following short (1–3 day) periodsof starvation, egg production by C.agulhensis returned to thesatiated rate (51.1 eggs {female} day^–1) more rapidly(after 0.9–2.4 days of excess food) than that of Ca. carinatus(after 2.8–3.1 days). However, following longer (5–9day) periods of starvation, Ca. carinatus regained satiatedlevels of egg production (55.8 eggs {female}^–1 day^–1)more rapidly (after 3.1–4.0 days of excess food) thanC. agulhensis (after 3.8–5.2 days of feeding following5–7 days of starvation). Moreover, many C. agulhensisfemales did not regain normal rates of egg production after9 days of starvation. For both species, the time required foregg production to recover was proportional to the starvationperiod, although only up to 7 days for C. agulhensis, and wasthe same following 4.25 days of starvation. Previously fed Ca.carinatus terminated egg production more rapidly than C. agulhensiswhen starved. The ability of Ca. carinatus to tolerate, andrecover rapidly from, prolonged periods of starvation, combinedwith a comparatively fast development time and high rate ofegg production, provides this species with a strong competitiveadvantage over C. agulhensis in the highly pulsed food environmentof the southern Benguela upwelling region.     

A model of copepod population dynamics in the southern Benguela upwelling region

A simple population dynamics model was constructed to simulate temporalvariability in the biomass of a dominant copepod Calanoidescarinatus (Copepoda: Calanoida) along the West Coast region ofSouth Africa. Calanoides carinatus is extensivelypreyed upon by the commercially important anchovy Engrauliscapensis, thus variability in zooplankton production may serveas a useful predictor of variability in anchovy recruitment levels. Themodel developed here circumvents the need to include a large number ofparameters because it uses satellite-derived estimates of chlorophylla concentration and sea surface temperature as primaryinputs. Abundance estimates necessary to initialize the model are readilyobtainable from biannual research cruises. The model successfully simulatesobserved features of a copepod population's response to pulses of upwellingand is robust with respect to most of its parameters because minor changesin their values result in predictable changes in model output. The modelshowed greatest sensitivity to parameters that are difficult to determineempirically, such as predator-induced mortality rates. Gaps in our presentunderstanding of the nature and scale of processes affecting copepod eggabundance, survival and viability in the southern Benguela system wereidentified as the dominant impediment to simulating copepod populationdynamics in the region.     

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.     

In situ grazing pressure and diel vertical migration of female Calanus euxinus in the Black Sea

Gut pigment and abundance of the female Calanus euxinus (Hulsemann) weremeasured from several water layers (defined by density values), with3–5 h intervals during 30 h and 21 h at a station in the southwesternBlack Sea in April and in September 1995, respectively. The female C.euxinus was observed to begin migration to the upper phytoplankton-richlayer approximately 3 or 4 hours before the sunset. Only a fraction of thefemale Calanus population (0.2% in April and 3.6% inSeptember) did not migrate but remained at the depth of the oxygen minimumzone during the nighttime. The migrating population was determined to havespent 7.5 h in the euphotic zone in April and 10.5 h in September. Thegrazing rate of female Calanus euxinus was measured from the gut contentdata collected from the layers which contain the euphotic zone. Thepercentage of primary production grazed by the female C. euxinus wascalculated as 14.5% in April and 9.5% in September. This revised version was published online in July 2006 with corrections to the Cover Date.     

Visual predators and the diel vertical migration of copepods under Arctic sea ice during the midnight sun

Despite the midnight sun, herbivore copepods Calanus hyperboreus,C. glacialis and Pseudocalanus acuspes displayed a normal dielvertical migration (NDVM) under the ice cover of Barrow Straitin spring, ascending into the chlorophyll-rich under-ice surfacelayer around maximum relative rate of change in irradiance (     

Vertical migration and habitat partitioning of six euphausiid species in the northern Benguela upwelling system

The vertical distribution of the adults of six species of euphausiids,Thysanöessa gregaria, Nematoscelis megalops, Euphausiaamericana, E.gibboides, E.hanseni, Stylocheiron longicorne,as well as the larval stages of E.hanseni, were studied duringa 48 h cycle at a fixed station in the northern Benguela upwellingsystem. All the species, except T.gregaria and S.longicorne,proved to perform diel vertical migration, but both migratoryand non-migratory species appeared to be segregated in spaceduring night-time hours, regardless of potential prey (phytoplanktonand copepods). It is suggested that water column structure andhydrographic discontinuities caused by a warm, depth-localizedintrusion, as well as the reproductive strategies of particularspecies, are responsible for this pattern. Eggs and larval stagesof E.hanseni were concentrated near the surface, and the effectof the short-term pulses of the intrusion on their abundanceis discussed.     

Sampling and analysis of gut contents in relation to environmental variability and diel vertical migration by herbivorous zooplankton

The accuracy of standard sampling and analysis procedures forestimating ingestion by herbivorous zooplankton was assessedusing models. Artificial environments were created in a computermodel, allowing for depth-dependent variability in temperature,chlorophyll and primary production. Model zooplankton were simulatedwithin these artificial environments using individual-basedmodels. The model zooplankton feed and defaecate at rates determinedby temperature and food concentrations, and also exhibit dielvertical migration (DVM) according to a variety of migrationmodels. The computer model was run for different combinationsof these nine environmental and five DVM models. Data were ‘sampled’from the model output, similar to field sampling of mesozooplanktongrazing. Daily ingestion was calculated from the gut ‘samples’using standard procedures for analysing gut fluorescence. Thesample results were compared with the actual ingestion valuesin the model, and some causes of discrepancies were noted. (i)If incorrect temperatures were assumed when calculating thegut evacuation rate (K), then estimates of ingestion were wrongby up to 40%. (ii) Non-uniform food environments gave errorsof up to 30% because of the large variability of measured gutcontents among individuals. (iii) Sampling from only part ofthe total depth range (e.g. at the chlorophyll maximum) resultedin estimates of ingestion being only 5% of the real value. Thissampling practice should be discouraged, because the sampleis not random. (iv) If sampling is not frequent enough, errorscan be as large as 45%, but more usually were 10% for realisticsampling frequencies. We describe an analysis procedure thatuses Monte Carlo-type simulations in a computer spreadsheetto estimate population consumption. These calculations takeinto account natural variability due to populations, samplesand assumptions. We urge that results should be presented asranges of possible values, rather than as single ‘mean’values, to allow for easier recognition of meaningful differencesamong samples and systems.     

Distribution and diel vertical migration of the euphausiid Thysanoessa macrura in Gerlache Strait,Antarctica

Thysanoessa macrura was found throughout Gerlache Strait, Antarctica, during four surveys carried out from 30 October to 23 November 1989, with the highest abundance being 332 individuals m^–2 (0–290 m). Reproduction had begun just before the surveys took place, as indicated by the presence of females with attached spermatophores and of larvae. Thirteen-month old females were reproductive. Larvae in 9 depth strata between 0–290 m were dominated by calyptopis stages, and developed from calyptopis 1 to furcilia 1 during November. Larval abundance was not correlated to chlorophyll a concentration, which showed a consistent east-west gradient in Gerlache Strait with highest concentrations (>30 mg chlorophyll a m^–3) in bays of the Antarctic Peninsula. Survival of larvae appeared to not be affected by phytoplankton abundance. Older T. macrura showed strong diel vertical migration between the surface at night and depths to 120 m during mid-day. Larvae were consistently found in the chlorophyll a-rich upper 50 m during night (90%) and day (81%), while adults and juveniles were found in the upper 50m at night (83%), but only 16% remained there during the day.     

Diel vertical migration and feeding of chaetognaths in coastal waters of the eastern Mediterranean

This study investigates the diel vertical distribution and the diet of the most important chaetognath species found in the 0–50 m surface layer of a coastal area in the eastern Mediterranean during a 24-hour period in July 2004. Among the recorded chaetognaths, Sagitta enflata was the most abundant species (41.6%), followed by S. minima (32.5%) and S. serratodentata (20.8%). Those three species exhibited a “twilight migration” pattern, with only small differences among them. Vertical separation was found between S. enflata and S. minima. Both species preyed mainly on cladocerans, although copepods were the most abundant group in the zooplankton assemblage. The chaetognath species followed partially the diel vertical migration of their prey. S. enflata showed high feeding intensity at different times in both day and night, while S. minima fed more intensively at midday (12:00) and at dusk (20:00), and S. serratodentata in the morning (08:00). It seems that in order to coexist in an area of low productivity the chaetognath species follow the basic ecological rules of space, time and food-type separation, in order to reduce the inter- and intra-specific competition. The high preference of S. minima and especially of S. enflata for the cladocerans made them probably the most important predators of cladocerans during summer.     

The relative importance of food and temperature to copepod egg production and somatic growth in the southern Benguela upwelling system

The fecundity and somatic growth rates of Calanus agulhensisand Calanoides carinatus, the dominant large calanoid copepodsin the southern Benguela upwelling system, as well as the fecundityof several other common copepods, were measured between Septemberand March of 1993/94 and 1994/95. Mean egg production of mostcopepods was low at >30 eggs female^-1 day^-1 {Calanoides carinatus23.7, Calanus agulhensis 19.0, Neocalanus tonsus 16.1 and Rhincalanusnasutus 26.1), whereas the mean fecundity of Centropages brachiatuswas significantly greater (83.6 eggs female^–1 day^-1).This study also presents the first comprehensive field estimatesof the fecundity of Nanno-calanus minor (mean: 26.1 eggs female^–1day^–1, range: 0.0–96.2 eggs female^–1 day^–1)and of somatic growth of N6 and all copepodite stages of Calanoidescarinatus (decreasing from 0.58 day^–1 for N6 to 0.04 day^–1for C5). Somatic growth rates of Calanus agulhensis also declinedwith age: from 0.57 day1 for N6 to 0.09 day1 for C5. Data ongrowth rates were used to assess the relative importance offood [as measured by total chlorophyll (Chi) a concentration],phytoplankton cell size (proportion of cells >10 µm)and temperature to the growth of copepods. Multiple regressionresults suggested that fecundity and somatic growth rates werepositively related to both Chi a concentration and phytoplanktoncell size, but not to temperature. Although it was not possibleto separate the effects of Chi a concentration and phytoplanktoncell size, data from previous laboratory experiments suggestthat copepod growth is not limited by small cells per se, butby the low Chi a concentrations that are associated with theseparticles in the field. Despite growth not being directly relatedto temperature, a dome-shaped relationship was evident in somespecies, with slower growth rates at cool (<13°C) andwarm (>18°C) temperatures. The shape of this relationshipmirrors that of Chi a versus temperature, where poor Chi a concentrationsare associated with cool and warm temperatures. It is concludedthat the effect of food limitation on growth of copepods outweighsthat of temperature in the southern Benguela region. Sourcesof variability in relationships between growth and Chi a concentrationare discussed.     

Diel cycles of reproduction and vertical migration in the marine cladocerans Pseudevadne tergestina and Penilia avirostris

The marine cladocerans Pseudevadne tergestina and Penilia avirostrisuse different strategies to avoid visual predators. Pseudevadnetergestina exhibited a pronounced diel cycle in reproduction,but did not perform diel vertical migration. Parthenogeneticfemales with fully developed embryos were absent during theday in both fishless enclosures and in the natural environment.This observation suggests that the diel cycle was due to nocturnalrelease of neonates and not the result of selective predationby fish. The need to use the compound eye to locate food wouldforce P. tergestina to remain in the illuminated surface waterduring the day and release their offspring in darkness thusdecreasing their visibility to fish. The diel cycle in reproductionin P. avirostris was not as pronounced as that in P. tergestina,but females with mature embryos were still more common at nightthan during the day. As a grazer of phytoplankton, P. avirostrisperformed diel vertical migration and retreated to dark, deeperwaters during the day, avoiding visual predators.     

Small-scale distribution and diel vertical migration of zooplankton in a shallow lake (Lake Naardermeer,the Netherlands)

Groundwater is a major influence on the hydrological, chemical and thermal regime of chalk streams in the southern U.K. However, little is currently known about the nature of the sediment delivery system within these chalk stream systems, even though sediment-related problems have been increasingly cited as a cause of habitat degradation and of declining salmonid stocks. To address this knowledge gap, suspended sediment fluxes were monitored at 4 sites within the Hampshire Avon catchment between February 1999 and August 2000. Maximum suspended sediment concentrations ranged from nearly 45 mg l^–1 to 260 mg l^–1. Over the study period, annual suspended sediment loads ranged from 644 to 6215 t yr^–1 and annual specific sediment yields ranged from 1.4 to 12.5 t km^–2 yr^–1. The results show that, relative to other U.K. rivers, the study chalk streams are characterised by low suspended sediment concentrations and loads and less episodic behaviour.     

Diel vertical migration of Cryptomonas and Chromatium in the deep chlorophyll maximum of a eutrophic lake

Large populations of the phytoflagellates Cryptomonas rostratiformis,Cryptomonas phaseolus, Cryplomonas undulata and the phototrophicbacterium Chromatium cf. okenii have been observed in the oxic/anoxicboundary layer of the slightly eutrophic, dimictic lake Schlachtenseeduring the summer stratification. Vertical distribution of thesepopulations was studied with the help of a new close-intervalsampler and by in Situ fluorescence measurements with fine spatial(cm to dm) and temporal (h) resolution on 4 days in 1991, 1994and 1995. All populations lived close to the chemo dine butshowed a regular diel vertical migration with daytime ascentand night-time descent and a low migration amplitude. At leastfor one species— C. rostratiformis—the pattern ofmigration suggests that this behaviour has an endogenous origin.As a result of diel vertical migration, the population of C.rostratiformiswas spatially separated from the other dominating populationsduring daytime. Ultimate cause of diel vertical migration wasa better light supply of all populations. Another probable advantagewas the reduction in grazing pressure, as large percentagesof all populations moved into the anoxic, hydrogen sulphidecontaining water layers during night-time. ¹Present address Department of Lowland Rivers and Shallow Lakes,Institute of Freshwater Ecology and Inland Fisheries, Müggelseedamm260, D-12587 Berlin, FRG     

Diel vertical migration and feeding rhythm of Daphnia longispina and Bosmina corexoni in Lake Toya, Hokkaido, Japan

Diel vertical migration (DVM) and diel feeding rhythm of two cladocerans, Daphnia longispina and Bosmina coregoni were investigated at the pelagic area of Lake Toya (Hokkaido, Japan) in May, August and October 1992. Both species performed nocturnal DVM. The amplitude of DVM, however, became smaller from May to October. Such seasonal variations in DVM could not be explained by light penetration and/or water temperature. The two species had a clear feeding rhythm; they fed at night in May and October but also after sunrise in August. These feeding rhythms appeared to be related to the light-dark cycle, but were not necessarily associated with their DVM. We suggest that the diel feeding rhythm and DVM are regulated independently by light cues.     

Diel migration and vertical distribution of Cladocera in Lake D. Helvecio (Minas Gerais,Brazil)

Six genera of Clad ocera (Diaphanosoma, Daphnia, Ceriodaphnia, Moina, Bosmina, Bosminopsis), each of them usually with only one species were found in Lake D. Helvecio, a natural valley lake located in the eastern part of Brazil. Diurnal migratory movements of the organisms observed in this lake showed a different pattern in different species. Closely related species, which explore the same food source, live in different layers, thus avoiding interspecific competition. The migratory behaviour of the species was studied mainly in relation to temperature and oxygen distribution in the lake. Thus, analyses were made in the summer (January, 1978) when a strong stratification occurs with the establishment of a thermocline and an oxycline. Comparisons were made also with the data obtained in winter (July, 1978), when a complete mixing of water occurs.     

Differential behaviour and shifts in genotype composition during the beginning of a seasonal period of diel vertical migration

During the first few weeks of a recurring seasonalperiod of diel vertical migration in Lake Maarsseveen(The Netherlands), part of the hybrid Daphniagaleata × hyalina population migrated, whileanother part remained in the epilimnion. In theepilimnion, 0+ perch prey upon daphnids duringdaytime. Gradually, the number of adult Daphniain the epilimnion decrease until the epilimnion isnearly devoid of daphnids. The population as a wholemay decrease, as in 1991, or may increase asin 1992. Genotype composition, as determined byallozyme analysis, changed substantially within afortnight in 1992, and one genotype became dominant.Our data are in agreement with the hypothesis thatpredation on different genotypes (clones)occurs during the beginning of a seasonal period ofdiel vertical migration, though our data do not allowto exclude alternativeexplanations.     

Population structure, vertical distribution and diel migration of Sagitta setosa (Chaetognatha) in the south-western part of the Black Sea

The population structure of Sagitta setosa from the south-westernpart of the Black Sea was examined between May 1994 and December1996. In addition, the vertical distribution and diel verticalmigration of S.setosa were studied for juveniles (