Seasonal vertical distribution and diel migration of zooplankton in a temperate stratified lake

The investigation of the vertical distribution of the zooplankton community in the temperate Lake Trichonis during four seasons in 2005, showed the existence of vertical segregation among species, ontogenetic stages and sexes within and between the major groups. In each season, the two or three more abundant rotifer species distributed at separate depth layers, while this feature was maintained during the entire 24 h period, since no diel vertical migrations (DVM) were performed. In contrast, the crustacean community, comprised mainly by the calanoid copepod Eudiaptomus drieschi and the cladoceran Diaphanosoma orghidani, showed various patterns of DVM, being more pronounced in spring and summer. Females of E. drieschi distributed deeper than males, while the copepod nauplii were found mainly in the surface layer in all four seasons. Temperature was the most important abiotic factor affecting directly and indirectly the vertical distribution and migration of various species. During stratification, the metalimnion was the most productive layer in Lake Trichonis, having maximum values of dissolved oxygen and low transparency due to high concentration of organic matter and phytoplankton. The DVM patterns of the crustaceans indicate that the metalimnion acts probably as a daylight refuge against predation by Atherina boyeri, which is the dominant planktivorous fish in the lake.     

Absence of typical diel vertical migration in Daphnia: varying role of water clarity, food, and dissolved oxygen in Lake Eymir, Turkey

Diel vertical migration (DVM) is a complex and dynamic behaviour against predation because the reaction of migrating organisms to light intensity plays a primary role, but is modified by other factors. In the relatively shallow but thermally stratified Lake Eymir, Daphnia pulex de Geers utilized vertical refugia afforded by the hypolimnion during both day and night. Differences in general vulnerability to fish predation determined the differences in their mean residence depths (MRDs) of different population categories such as most conspicuous and vulnerable individuals of adult with eggs inhabited the deepest depth, whereas juveniles stayed close the thermocline. In late spring, profoundly high amplitude of displacement within the hypolimnion, probably due to the hypolimnion being well-lit and relatively well-oxygenated for the fish and rather unsafe for the large-sized daphnids, was recorded. Therefore, the large-sized daphnids daytime refuge was close to the bottom whereas at night they moved upward to benefit from warmer water temperature along with food availability in the presence of fish predation but still remained below the thermocline. In summer, the insignificant amplitude of the hypolimnetic, which later became epilimnetic, displacements were probably due to the near-anoxic condition found below the thermocline. This might have deterred the fish, thus providing a safer refuge for daphnids in the below thermocline, which afterwards became the above thermocline. Low oxygen availability was regarded as the summer proximate factor. The abundant food and warmer water conditions found in the below/above thermocline also accounted for absence of DVM in summer. Consequently, this study suggests that DVM by Daphnia is an adaptation that is plastic to changing environmental conditions.     

Diel vertical migration of the copepod <Emphasis Type="Italic">Thermocyclops inversus</Emphasis> (Kiefer, 1936) in a tropical reservoir: the role of oxygen and the spatial overlap with <Emphasis Type="Italic">Chaoborus</Emphasis>

The effect of water transparency, dissolved oxygen concentration and the invertebrate predator Chaoborus brasiliensis on the day–night vertical distribution of the copepod cyclopoid Thermocyclops inversus was investigated in a shallow tropical reservoir, Nado Reservoir, Belo Horizonte, Brazil. Diel cycles were carried out over a period of 12 consecutive months, between October 1999 and September 2000. The different developmental stages of T. inversus exhibited diel vertical migration (DVM) and displayed a clear ontogenetic trend, with the amplitude of DVM increasing with the age of the organism, and ranging from 0.4 m to 0.8 m for nauplii, 0.4 m to 1.2 m for copepodite, and 1.1 m to 2.1 m for adults. We observed that seasonal changes in dissolved oxygen and C. brasiliensis directly influenced the vertical distribution of the copepod population in this reservoir. Furthermore, it was showed that the diurnal vertical migration is an important predator avoidance behavior since it diminished the spatial overlap between prey and its potential predator. This finding supports the hypothesis that the vertical migration is a defense mechanism against predation. Thus, T. inversus is able to remain in the anoxic layers during day light hours, and at night they move upwards avoiding hypolimnetic waters to escape from predation by Chaoborus.     

Vertical, lateral and longitudinal movement of zooplankton in a large river

1. The spatial distribution and movement patterns of zooplankton in large rivers are little known compared with those in lake environments. We conducted a series of studies in the Ohio River (U.S.A.) during the low flow period to assess diel vertical (DVM), longitudinal and lateral movement of crustacean zooplankton. 2. The dominant large zooplankter, the copepod Eurytemora affinis, showed a consistent vertical migration pattern of daytime ascent and night‐time descent during all sampling periods – the reverse of the most common migratory pattern of zooplankton in lakes. The cladoceran Bosmina migrated in a similar way in two of the three sampling periods. Surveys taken longitudinally in the river showed similar trends for both taxa. 3. During the lateral surveys, E. affinis was significantly more abundant in the shallow littoral zone during the night than in the daytime. The combination of vertical and lateral movement patterns along with the diel distribution of zooplanktivorous fish suggest that these movements are a predator‐avoidance mechanism. 4. Sampling programmes in large rivers should consider that larger zooplankton such as E. affinis may not be randomly distributed in the river channel and behaviours such as diel vertical migration may be just as evident in river habitats as in lakes.     

The hypolimnetic protozoan plankton of a eutrophic lake

The seasonal distribution of benthic species in the water column above and below the thermocline in a small eutrophic lake is described. During summer stratification populations of Spirostomum spp, Loxodes spp., Plagiopyla and Deltopylum become established in the plankton on or below the oxycline/thermocline. At shallow sites no migration occurred and populations of the migratory species in the benthos were sparse, with the exception of Plagiopyla which occurred in high densities in the sediment. Two distinct planktonic populations are established during stratification: an epilimnetic community of obligate planktonic ciliates and a hypolimnetic community of benthic migrants.     

Coexistence of two similar copepod species, <Emphasis Type="Italic">Eudiaptomus gracilis</Emphasis> and <Emphasis Type="Italic">E. graciloides</Emphasis>: the role of differential predator avoidance

Habitat choice in relation to environmental factors of two coexisting calanoid copepod species, Eudiaptomus gracilis and E. graciloides, was studied in a mesotrophic lake and in large indoor mesocosms. Both species and sexes showed pronounced diel vertical migration (DVM) in the field. In 12 m deep mesocosms with free ranging fish DVM was observed and species increased day depth over time. No changes were observed in copepod day depth over time in experiments with fish kairomone. It is hypothesized that fish kairomone acts as an early warning system to copepods which respond by moving deeper, but only as far as the thermocline. For full DVM, a nearby mechanical stimulus is necessary. Thus, as fish go deeper to feed, copepods retreat. The response of copepods to fish predation, in the presence of low and high numbers of Daphnia,shows that copepods effectively use Daphniaas living shields to avoid predation. The two species adopt different vertical migration strategies depending on whether there are high or low numbers of Daphniapresent. A dominant feature of mesocosm experiments was the night time aggregating (lekking) of E. gracilis males at the surface. When the spring and autumn percentages of risk takers in the epilimnion were compared, E. gracilis, particularly males, suffered the greatest cost.     

Diurnal vertical migration of Daphnia hyalina Leydig, 1860 (Crustacea: Cladocera) in Lake Bled (Slovenia) in relation to temperature and predation

Diurnal vertical migration (DVM) of Daphnia hyalina in Lake Bled was most intense during summer stratification. The extent of DVM varied with the size of the animal and its reproductive state. Migration distances were shortest in immature specimens and longest in ovigerous females. During daytime, ovigerous females stayed deeper in the water column than females without ova or immatures. The daytime temperature of water at the median depth of the ovigerous females did not exceed 10 °C, even in the warmest season. At night they migrated upward to an environment which was warmer by as much as 9 °C.Laboratory observations indicate that specimen's size and water temperature determine the velocity of passive sinking, such that morning descent of the different groups of Daphnia can be explained by passive sinking alone.Our hypothesis is that the distribution of different groups of D. hyalina in Lake Bled is influenced by two types of predators: fish (Perca fluviatilis L. and Rutilus rutilus (L.)) and larvae of Chaoborus flavicans (Meig.), the latter appearing in the epilimnion during the night. Fish predation has a key-role at the beginning of thermal stratification. Supposing that in spring the gene pool of Daphnia consists of a mix of different genotypes, distributed at different depths during the day, fish predation combined with a presence of fish chemicals favored genotypes with a lower day-depth during the spring/summer period.     

Seasonal changes in partial,reverse diel vertical migrations of cisco Coregonus artedi

The objectives of this study were to (1) document changes in partial, reverse diel vertical migrations (DVM) patterns of cisco Coregonus artedi in Ten Mile Lake, MN, U.S.A., throughout the year and (2) evaluate the mechanisms that may cause shifts in migration behaviour. Results indicated that C. artedi vertical distributions remained deep in the water column during the day and night of the spring and autumn, which was related to a low risk, low reward strategy. During summer, a partial migration occurred where a portion of the population remained deeper according to the low risk, low reward strategy, while the other portion performed a more extensive high risk, high reward reverse DVM. In winter, C. artedi did not migrate because there were only low risk, low reward conditions present at all depths. The extensive partial, reverse DVM during summer probably increased the growth potential of C. artedi, helping individuals survive in a lake with low zooplankton prey resources.     

Diel vertical migration of larval and early‐juvenile burbot optimises survival and growth in a deep,pre‐alpine lake

1. Burbot larvae (Lota lota) perform a substantial diel vertical migration (DVM) of increasing amplitude in the pelagic zone during a 3‐month period before migrating to the littoral zone as early‐juveniles. We hypothesised that feeding in the warm surface layers at night and then spending the day in cold water below the thermocline reduces metabolic costs and earns burbot larvae an energetic advantage. 2. To test our hypothesis, we mimicked the temperature conditions experienced by vertically migrating burbot in the pelagic zone. We also simulated three further scenarios, in which temperature remained constant. 3. Burbot showed the best performance (defined as specific growth rate multiplied by the probability of survival) in the treatments simulating DVM. The high temperature treatment, simulating permanent residence in the warm epilimnion, resulted in high growth combined with high mortality. At a permanently low temperature, simulating life in the hypolimnion, growth was poor and activity reduced. 4. In a deep, temperature‐stratified lake, where the apparently beneficial overall medium temperature is found in a restricted layer within the thermocline, DVM optimises performance in young burbot. Various ultimate factors might act synergistically in selecting for DVM in larval and early‐juvenile burbot.     

Diel vertical migration by males,females, copepodids and nauplii in a limnetic population of diaptomus (Copepoda)

Diel vertical migration (DVM) by the sexes and life history stages of the copepod Diaptomus novamexicanus (Herrick) was investigated during two summers in a subalpine lake (Castle Lake, California, USA). Migratory behavior was quantified repeatedly by 1300 and 21–2200 hr sampling in the epilimnion. Population density and vertical distribution were estimated routinely at 1300 hr and at 2200 hr on selected dates. Males had greater night to day biomass differentials and ratios than females. The sexes showed comparable DVM participation (% of total biomass migrating) and amplitudes of migration (distance of ascent). Copepodids displayed higher participation, night/day ratios and amplitudes than younger copepods, but adult participation differed only from that of nauplii. Active participation in high amplitude ascents thus increased with age and was greatest in adult males. Analysis suggests that such relative behaviors probably occur in most limnetic Diaptomus, and that the magnitude of ascent into the epilimnion is associated with daytime vertical distribution.     

Salinity tolerance of the gemmules of Eunapius fragilis (Leidy) and the inhibition of germination by various salts

We report results of a field test of the predator avoidance hypothesis as an explanation of the adaptive significance of diel vertical migration in zooplankton. We determined the vertical distribution and diel migration of the planktonic copepod Acartia hudsonica, concurrently with the abundance of pelagic fish, transparency and thermal stratification of the water column, on six cruises over a one year period in a temperate marine lagoon (Jakles Lagoon, San Juan Island, Washington, USA). Striking seasonal variability was observed in all biological and environmental variables. Linear regressions of the strength of diel vertical migration in A. hudsonica on these environmental variables resulted in only one statistically significant relationship, that between copepod diel vertical migration and predator abundance. These results, together with those of previous studies, point to diel vertical migration as a widespread behavioral response of planktonic prey to the presence of their predators.     

Vertical distribution of pelagial zooplankton in a middle-sized dimictic valley reservoir

Our observations indicate the vertical distribution of zooplankton and its seasonal changes in Dubník II reservoir (Slovakia) are determined mainly by the thermal regime of the reservoir, by transparency, and by fish and invertebrate predation. During periods of circulation, zooplankton vertical distribution in the whole water column was more homogeneous, whilst during summer temperature stratification zooplankton concentrated in the epilimnion — rotifers in higher layers than crustaceans. During summer stagnation a steep thermal gradient occurred at the boundary of the epi-and hypolimnion and low temperature and low dissolved oxygen in hypolimnion offered a refuge for Chaoborus flavicans larvae against fish, enabling coexistence of vertebrate and invertebrate predation. This evidence supports our previous findings concerning dominance of rotifers in zooplankton and representation of crustaceans by small-bodied species in the study reservoir. Steep thermal gradient and the presence of Chaoborus larvae caused very low zooplankton abundance in the lower part of the water column and a reduction of cladocerans refuges against fish to layers of thermocline or closely under thermocline where Daphnia cucullata and Daphnia parvula were found. Our previous assumptions about the high density of zooplanktivorous fish in Dubník II reservoir are supported by the fact that these small cladocerans are represented by smaller individuals in the upper layers and bigger individuals in deeper layers.     

Habitat Characteristics of Eurytemora lacustris (Poppe, 1887) (Copepoda,Calanoida): The Role of Lake Depth,Temperature,Oxygen Concentration and Light Intensity

Field observations, laboratory experiments and a literature survey were conducted to evaluate the habitat characteristics of Eurytemora lacustris (Poppe 1887), a freshwater calanoid copepod species. Combined effects of temperature and oxygen concentration in the deep water of thermally stratifying lakes seem to be the ultimate factors governing the occurrence of the species throughout its home‐territory. E. lacustris is largely restricted to relatively deep lakes (>30 m) providing a hypolimnetic refuge characterised by low temperatures (<∼10 °C) and oxygenated water during summer. Therefore, although the species is spread over much of Europe it was only found in a small number of lakes. Long‐term records in different lakes revealed E. lacustris to be perennial with relatively high biomasses occurring from May to September. During thermal stratification on average 87% of the nauplii and 72% of the copepodite biomass was found in hypolimnetic waters colder than 10 °C. Diurnal vertical migration was observed for the copepodid stages, but the migration amplitude clearly decreased from May to September. The migration amplitude was significantly related to light intensity. According to its special habitat requirements, E. lacustris might be considered a glacial relict sensitive to temperature increase and oxygen depletion. (© 2005 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Ovigerity,selective predation,and variable diel vertical migration in Euchaeta elongata (Copepoda: Calanoida)

Summary We present a statistical analysis of a previously published (Yen, 1983) but heretofore unanalyzed data set on the vertical distributions and diel vertical migration (DVM) of adult females of the marine planktonic copepod Euchaeta elongata in Dabob Bay, Washington, USA. Non-ovigerous females were strongly migratory on all four dates sampled, residing between 75–175 m during the day and at shallower depths during the night, commonly entering the upper 50 m of the water column. In contrast, ovigerous females were non-migratory or weakly migratory, largely remaining between 100–175 m both day and night, and entering the upper 50 m of the water column only rarely. Thus non-ovigerous females always migrated much more strongly, as measured by both amplitude of migration and the proportion of animals migrating, than did ovigerous females. These results led us to hypothesize that differential susceptibility to visually orienting predators was the cause of these differences in DVM behavior in female E. elongata, and we subsequently undertook an experimental study of the feeding selectivity of the copepod's natural predator, Pacific herring (Clupea harengus pallasi). Pacific herring exhibited a highly significant preference for ovigerous over nonovigerous adult female E. elongata. The demographic consequences of variable DVM in adult female E. elongata were investigated by way of life table analyses. Results indicated that under conditions of thermal stratification of the water column there is a distinct demographic disadvantage (reduced rate of realized population growth) incurred by non-migratory or weakly migratory ovigerous females due to delayed egg development at cooler subsurface temperatures. We conclude that ovigerous female E. elongata remain at depth both day and night to avoid visually orienting predators, and that such behavior must afford the copepod a demographic advantage of no less than a 26% reduction in adult mortality to offset the demographic cost of delayed egg development.     

Zooplankton Migration in Three Lakes of Western New York

We examined the diel vertical migration (DVM) of zooplankton in three lakes of western New York; Lakes Conesus, Lime, and McCargo. In all three lakes, the dipteran predator Chaoborus was a more dramatic migrator than any of the cladocerans or copepods. In contrast, another invertebrate predator, Leptodora, seemed to have the least vertical dispersal. In Conesus Lake, following the evening ascent and upper-water night-positioning of Chaoborus flavicans, Daphnia pulicaria expanded its own vertical range of dispersal thereby decreasing its degree of overlap with Chaoborus. The vertical distribution of Diaptomus sicilis was mostly below that of Daphnia pulicaria, thus reducing possible competition for food resources. Poor oxygen conditions in the lower waters of Lakes Lime and McCargo restricted all zooplankton (except Chaoborus) to an epilimnetic zone where the regions of niche overlap and predation were narrowed. Such annually-induced restrictions on vertical dispersal are probably common features of numerous stratified lakes with hypolimnetic reducing conditions. No fixed pattern of vertical dispersal or migration is likely in lakes where stratification and non-stratification follow each other seasonally. Vertical migration does provide some flexibility in niche separation, but the potential for chaotic behavior in interacting predators and prey may confound simple explanations of DVM.     

From polar night to midnight sun: photoperiod, seal predation, and the diel vertical migrations of polar cod (Boreogadus saida) under landfast ice in the Arctic Ocean

The winter/spring vertical distributions of polar cod, copepods, and ringed seal were monitored at a 230-m station in ice-covered Franklin Bay. In daytime, polar cod of all sizes (7–95 g) formed a dense aggregation in the deep inverse thermocline (160–230 m, −1.0 to 0°C). From December (polar night) to April (18-h daylight), small polar cod <25 g migrated into the isothermal cold intermediate layer (90–150 m, −1.4°C) at night to avoid visual predation by shallow-diving immature seals. By contrast, large polar cod (25–95 g), with large livers, remained below 180 m at all times, presumably to minimize predation by deep-diving mature seals. The diel vertical migration (DVM) of small polar cod was precisely synchronized with the light/dark cycle and its duration tracked the seasonal lengthening of the photoperiod. The DVM stopped in May coincident with the midnight sun and increased schooling and feeding. We propose that foraging interference and a limited prey supply in the deep aggregation drove the upward re-distribution of small polar cod at night. The bioluminescent copepod Metridia longa could have provided the light needed by polar cod to feed on copepods in the deep aphotic layers.     

Habitat selection and diel distribution of the crustacean zooplankton from a shallow Mediterranean lake during the turbid and clear water phases

1. The fish fauna of many shallow Mediterranean Lakes is dominated by small‐bodied exotic omnivores, with potential implications for fish–zooplankton interactions still largely unknown. Here we studied diel variation in the vertical and horizontal distribution of the crustacean plankton in Lake Vela, a shallow polymictic and eutrophic lake. Diel sampling was carried out on three consecutive days along a horizontal transect, including an open‐water station and a macrophyte (Nymphaea alba) bed. Since transparency is a key determinant of the predation risk posed by fish, the zooplankton sampling campaigns were conducted in both the turbid (autumn) and clear water (spring) phases. 2. In the turbid phase, most taxa were homogeneously distributed along the vertical and horizontal axes in the three consecutive days. The only exception was for copepod nauplii, which showed vertical heterogeneity, possibly as a response to invertebrate predators. 3. In the clear water phase, most zooplankton taxa displayed habitat selection. Vertically, the general response consisted of a daily vertical migration (DVM), despite the limited depth (1.6 m). Horizontally, zooplankters showed an overall preference for the pelagic zone, independent of the time of the day. Such evidence is contrary to the postulated role of macrophytes as an anti‐predator refuge for the zooplankton. 4. These vertical (DVM) and horizontal (macrophyte‐avoidance) patterns were particularly conspicuous for large Daphnia, suggesting that predation risk from size‐selective predators (fish) was the main factor behind the spatial heterogeneity of zooplankton in the spring. Thus, the difference in the zooplankton spatial distribution pattern and habitat selection among seasons (turbid and clear water phases) seems to be mediated the predation risk from fish, which is directly related to water transparency. 5. The zooplankton in Lake Vela have anti‐predator behaviour that minimises predation from fish. We hypothesise that, due to the distinct fish community of shallow Mediterranean lakes, aquatic macrophytes may not provide adequate refuge to zooplankters, as seen in northern temperate lakes.     

Vertical distribution of planktonic rotifers in a karstic meromictic lake

Vertical distribution of planktonic rotifers is described in relation to temperature and oxygen in Lake La Cruz, a single-doline, closed karstic lake (121 m diameter and 25 m maximum depth) which shows iron meromixis. Samples were taken by peristaltic pumping at 10 cm depth intervals in the oxycline zone from June 1987 to September 1988. A model of rotifer vertical structure in stratified lakes is proposed. Rotifers concentrate their populations at the depths with intense gradients. As stratification develops some rotifer populations show a downward migration following the thermocline and some others show an upward migration following the oxycline. The production-respiration balance in the lake, and so the position of the oxycline with respect to the thermocline and the layer of maximum production, depends on meteorological conditions. A shift in the dominance of congeneric or related species can occur in consecutive years. In Lake La Cruz, mixing conditions and subterranean inflow in spring were much more intense in 1988 than 1987, and the distance between production and decomposition depths was smaller in 1988. Anuraeopsis miraclei, an oxycline-bound species with high abundance in 1987, was displaced by A. fissa in 1988. A. fissa, which was a metalimnetic species during early summer, reached peak densities (3 × 10⁴ ind l^–1) at the oxycline, equaling the abundance of A. miraclei the preceeding year.     

Metabolism and foraging strategies of mid‐latitude mesozooplankton during cyanobacterial blooms as revealed by fatty acids,amino acids,and their stable carbon isotopes

Increasing sea surface temperatures (SST) and blooms of lipid‐poor, filamentous cyanobacteria can change mesozooplankton metabolism and foraging strategies in marine systems. Lipid shortage and imbalanced diet may challenge the build‐up of energy pools of lipids and proteins, and access to essential fatty acids (FAs) and amino acids (AAs) by copepods. The impact of cyanobacterial blooms on individual energy pools was assessed for key species temperate Temora longicornis and boreal Pseudo‐/Paracalanus spp. that dominated field mesozooplankton communities isolated by seasonal stratification in the central Baltic Sea during the hot and the cold summer. We looked at (a) total lipid and protein levels, (b) FA trophic markers and AA composition, and (c) compound‐specific stable carbon isotopes (δ¹³C) in bulk mesozooplankton and in a subset of parameters in particulate organic matter. Despite lipid‐poor cyanobacterial blooms, the key species were largely able to cover both energy pools, yet a tendency of lipid reduction was observed in surface animals. Omni‐ and carnivory feeding modes, FA trophic makers, and δ¹³C patterns in essential compounds emphasized that cyanobacterial FAs and AAs have been incorporated into mesozooplankton mainly via feeding on mixo‐ and heterotrophic (dino‐) flagellates and detrital complexes during summer. Foraging for essential highly unsaturated FAs from (dino‐) flagellates may have caused night migration of Pseudo‐/Paracalanus spp. from the deep subhalocline waters into the upper waters. Only in the hot summer (SST>19.0°C) was T. longicornis submerged in the colder subthermocline water (~4°C). Thus, the continuous warming trend and simultaneous feeding can eventually lead to competition on the preferred diet by key copepod species below the thermocline in stratified systems. A comparison of δ¹³C patterns of essential AAs in surface mesozooplankton across sub‐basins of low and high cyanobacterial biomasses revealed the potential of δ¹³C‐AA isoscapes for studies of commercial fish feeding trails across the Baltic Sea food webs.     

Observations on the vertical distribution of zooplankton, including post-larval teleosts, off Plymouth in the presence of a thermocline and a chlorophyll-dense layer

During 1973 – 1977, when Sagitta elegans was returningto dominance over S. setosa in Plymouth waters, there was often,in summer in stratified conditions, a community characterizedby S. setosa in the warm water above the thermocline, and atypical S. elegans community in the colder layer below the thermocline.This vertical separation persisted at night even though somespecies from the S. elegans community migrated up into the warmlayer. Such vertical migration was not stopped by a temperaturediscontinuity of from 3° to 6°C, but fewer individuals,or a lesser proportion of species, took part at the higher temperaturedifference. Only a small part of the zooplankton, comprisingsmall individuals of Limacina retroversa, the appendicularians,and, possibly, post-larvae of Arnoglossus laterna, showed signsof aggregation at the thermocline or the chlorophyll maximumclose to it.