83 Comparative analysis of vertically migrating euglena viridis populations in tidal and non‐tidal benthic environments

Benthic populations of Euglena viridis exhibit vertical migration behavior on high energy intertidal beaches and along the sand banks of freshwater streams. This study examines similarities and differences in the migratory behavior and cell morphology of populations of E. viridis inhabiting Scripps Beach, La Jolla, California and Coble Brook, Burlington, North Carolina. The timing of migration was measured by counting the number of cells in samples collected from the sediment surface throughout the day. Sediment cores were extracted and sectioned to determine the vertical distribution of the population. Neutral density filters and opaque canisters were used to shade the substratum to 56%, 22%, 2%, and 0% of incident irradiance (Io) to examine the effect of light on cell morphology and migratory behavior. On intertidal beaches, E. viridis exhibited a tidal rhythm in vertical migration with cells migrating below the sediment surface at night (>15 cm) and during daytime high tides. In this habitat, the upward migration response was enhanced at irradiances lower than 100% Io but cell morphology was not altered by shading. On the banks of freshwater streams, E. viridis exhibited a diurnal migratory rhythm with both tear‐drop and spherical morphologies observed throughout the day. The population was most concentrated at the surface around solar noon and at night it was located between 1 and 2 cm below the surface. Shading did not enhance upward migration but it did affect cell morphology. These results will be interpreted in the context of the dominant selection pressures in each environment.     

VERTICAL MIGRATION OF A MIXED‐SPECIES EUGLENA (EUGLENOPHYTA) ASSEMBLAGE INHABITING THE HIGH‐INTERTIDAL SANDS OF NYE BEACH,OREGON1

Comparatively little is known about the vertical migration of the microphytobenthic community forming visible patches on high‐energy beaches. We collected surface and cored samples to evaluate the timing and extent of downward migration of a multispecies Euglena assemblage inhabiting Nye Beach, Oregon. Euglena density at the surface was highly variable and was not correlated with the time of low tide or instantaneous irradiance measurements; however, triplicate cores collected at low and high tides revealed a tidal rhythm in mean depth. On average, 95% of the assemblage occurred within 1 cm of the surface during low tide, but 54% of the assemblage was collected between 1 and 8 cm below the surface during high tide. A midday shading experiment revealed that short‐term changes in irradiance levels altered the Euglena density at the sediment surface by inducing vertical migration. This response to short‐term fluctuations in light may explain the weak correlation between cell density at the surface and time of day. The high‐intertidal location of these patches prevented the removal of nonmigrating cells by daily high tides, which increased the variability in surface samples and obscured the tidal migration rhythm detected in the core samples. Due in part to the semidiurnal nature of Oregon tides, this study provides in situ confirmation of past mesocosm research indicating that sediment disturbance during daily submersed periods is an important process in maintaining the quasi‐tidal rhythm in the appearance and disappearance of Euglena spp. from the surface of beaches and intertidal sandflats.     

VERTICAL MIGRATION OF EUGLENA SP. ON THE SAND BANKS OF A NORTH CAROLINA PIEDMONT STREAM

On summer days, visible patches of green appeared on the aerially-exposed sand banks of a small tributary of Mays Lake in Burlington, North Carolina. These green patches became visible approximately 3 hours after sunrise and disappeared 2 hours prior to sunset. Microscopic examination of sediment samples revealed that the appearance of color was due to the upward migration of Euglena sp. that attained densities of 2200 cells per square millimeter around solar noon. Triplicate sediment cores revealed the presence of live Euglena cells as deep as 6 cm below the sediment surface. During the day, over 80% of the population was located between the surface and a depth of 2 mm. During the night, 75% of the population was found between 2 and 6 cm below the surface. Artificially darkening the sediment surface during the day stimulated downward migration of the population. Controlled experiments with neutral density filters indicated that the number of cells on the sediment surface was directly related to incident irradiance. The results of these experiments on Euglena sp. living on the banks of a freshwater stream in central North Carolina will be contrasted with those of previous experiments conducted on Euglena proxima living on the intertidal sand flats of the Newport River estuary in coastal North Carolina.     

EFFECT OF LIGHT ON VERTICAL MIGRATION AND PHOTOSYNTHESIS OF EUGLENA PROXIMA (EUGLENOPHYTA)

During daytime low tides, sediment-inhabiting populations of Euglena proxima Dangeard migrate to the surface of intertidal sand flats to form large monospecific green patches near the Duke University Marine Laboratory in Beaufort, North Carolina. The effect of incident irradiance on the upward migration of E. proxima was studied by shading the sediment surface in the field with neutral density filters to 2, 20, 55, and 100% incident irradiance. The results of these in situ experiments revealed that E. proxima 's upward migration, as measured by population density or chl a concentration, was stimulated by decreasing incident irradiance: population densities and chl a concentrations on the sediment surface at 2% incident irradiance were greater statistically than at 100% full sunlight. In situ measurements of photosynthesis were performed by incubating 3-mL aliquots of a cell suspension in scintillation vials containing ¹⁴C inorganic carbon beneath neutral density filters. These experiments revealed that E. proxima exhibited photoinhibition at 100% full sunlight, but this depression in photosynthetic performance at high light was absent when the cells were incubated in vials containing acid-cleaned sediment. This amelioration of photoinhibition in vials containing sediment is most likely due to the downward migration of the population away from photoinhibitory light levels. These results cannot be interpreted as strictly photic effects because the temperature of the vials varied with the light treatment. The results of this study suggest that the vertical migration behavior of E. proxima and its photosynthetic physiology are intimately linked. Vertical migration provides a mechanism to deal with short-term changes in irradiance and avoid photoinhibitory light levels.     

Evidence for gravitactic behaviour in benthic diatoms

Vertical migration by diatoms is a well-known phenomenon, occurring in intertidal and subtidal benthic biofilms. It is partially endogenously driven, as cell movements can be observed in the absence of external stimuli such as light, temperature or water cover. Although vertical migration of diatoms under constant conditions has often been attributed to geotactic orientation, this hypothesis has never been experimentally demonstrated. Our study tested the gravitactic nature of the vertical migratory behaviour of benthic diatoms in sedimentary biofilms, using an experimental setup designed to distinguish gravitaxis from surface-oriented cell movements. The hourly variation of surface diatom biomass during migratory cycles was compared in homogenized sediment samples kept facing upwards (surface-oriented and gravity stimuli coinciding; controls) and facing sideways or downwards (surface-oriented and gravity stimuli not coinciding). During the experiments, sediment samples were kept in complete darkness in custom-made, sealed measuring chambers designed to avoid any contact with atmospheric air and the formation of physico-chemical gradients near the surface. Microalgal biomass was monitored non-intrusively using PAM fluorometry, by measuring dark-level fluorescence, F_o. The results showed a clear effect of sample orientation in relation to the gravitational stimulus. In the controls, a biphasic pattern in surface biomass was observed, with the formation of a clear biomass peak (three- to six-fold increase) followed by a slower decrease. In contrast, in samples facing sideways or downwards, surface biomass also varied but to a much lesser extent (typically < two-fold). These results strongly suggest that, in the absence of light, upward vertical migration of benthic diatoms is mostly guided by negative gravitaxis, supporting the often hypothesized capacity of these cells to sense and use gravity to move vertically within the sediment.     

Persistent,vertical-migration rhythms in benthic microflora

The diurnal migratory rhythm of the epipelic diatom association of freshwater streams has been investigated in laboratory studies. Movement of the diatom flora up to the surface of the sediment and down beneath the surface occurs once every 24 hours reaching a peak of cell numbers at the surface at approximately the same time each day. This migratory movement persists in the laboratory for at least eleven days under alternating light/dark conditions. It is also expressed in continuous darkness but the number of cells migrating is less than under conditions of alternate illumination and darkening. In continuous illumination the rhythm is disturbed.

All the common diatom species behave in approximately the same way as the population.

From the data, the existence of three separate rhythms of motility, phototaxis and geotaxis have been deduced. During the first half of the light cycle the cells increase their motility and become positively phototactic and during the latter half of this cycle they lose their positive phototaxy, acquire a positive geotaxy and finally mobility decreases.

These observations are compared with previous studies.     

EFFECT OF SUBSURFACE NUTRIENT SUPPLIES ON THE VERTICAL MIGRATION OF EUGLENA PROXIMA (EUGLENOPHYTA)1

Euglena proxima Dangeard inhabits intertidal sand flats and displays a tidal rhythm in vertical migration. During daytime low tides when the sand flat is emersed, millions of cells are visible on the sediment surface, but the population remains below the surface at all other times. An earlier study demonstrated that the extent of downward migration of E. proxima is reinforced by the presence of a subsurface layer of black sediment. The present study was designed to test the hypothesis that the higher availability of inorganic nutrients or organic substrates in or above the black layer is responsible for the enhancement of downward migration in E. proxima. This hypothesis was tested experimentally by manipulating the bottom water in 24 mesocosm containers in a tidal tank. Six replicates of each of the following nutrient treatments were tested: seawater control; deep porewater collected from 70 cm below the sediment surface; seawater enriched with ammonium, nitrate, and phosphate; and seawater enriched with acetate, glucose, and the preceding inorganic nutrients. Multivariate analysis of variance revealed that the chl a biomass and chl a‐weighted mean depth of the population at high tide were significantly greater for replicates receiving inorganic nutrients. There was no difference between those receiving only inorganic nutrients and those enriched with inorganic nutrients, acetate, and glucose. These findings represent the first experimental evidence that subsurface nutrients are an important resource that reinforces the maintenance of vertical migration behavior in benthic microalgae.     

Burrowing abilities and swash behavior of three crabs, Emerita analoga Stimpson, Blepharipoda occidentalis Randall, and Lepidopa californica Efford (Anomura, Hippoidea), of exposed sandy beaches

To investigate factors related to the distribution of intertidal species, and specific predictions of the swash exclusion hypothesis for exposed sandy beaches, we compared the burrowing abilities and swash behavior of three species of anomuran crabs in the superfamily Hippoidea (Emerita analoga, Blepharipoda occidentalis and Lepidopa californica) which commonly inhabit the intertidal and shallow subtidal zones of beaches along the California coast. Burrowing times in the laboratory increased significantly with crab size for all species in five sediment grain sizes ranging from fine sand to gravel (0.15 to 3.24 mm). For each species, burrowing times differed significantly among sand grain sizes, ranging from 0.3 to 21.5 s. Burrowing times for the hippid crab, E. analoga, were relatively constant across sediment types, while those of the albuneid crabs, B. occidentalis and L. californica, were rapid in fine to medium sands, and much slower in coarser sediments. Our results indicate that E. analoga is a substrate generalist while L. californica and B. occidentalis are substrate sensitive. Pre-burrowing times and behavior, distance moved, and burrowing times differed among the species in the swash zone. Combined times of preburrowing and burrowing were shorter than the swash period (6 s) for most E. analoga individuals. Fifty percent of the individuals of L. californica reached the substrate and burrowed in the swash period, while no individuals of B. occidentalis burrowed in that time. Pre-burrowing behavior and time may be valuable in explaining spatial and temporal patterns in the distribution of hippoid crabs on California beaches. Our results support predictions of the swash exclusion hypothesis concerning the burrowing and locomotory abilities of sandy beach macrofauna. The substrate generalist characteristics, and unique orientation and swimming abilities of the hippid crab, E. analoga, in intertidal swash may help explain the success of this species and its congeners, and have important implications for understanding patterns of macrofauna community structure on exposed sandy beaches in California and other regions.     

Influence of light quality and gassing on the vertical migration of diatoms inhabiting the Wadden Sea

Diatoms inhabiting the Wadden Sea show a rhythmic migration pattern, which is superimposed by the tidal rhythm. In addition to light intensity, light quality has a pronounced influence on the upward-directed migration, thus giving some information on the nature of the relevant photoreceptors. Maximum diatom migration occurred when sediment surfaces were illuminated with blue light. The cell densities in blue light exceeded those of white light control experiments 1.8-fold. Furthermore, we registered a minor peak in the red light region, which reached approximately 60% of the white light controls. Cryptochrome and/or phototropin may thus be involved and act as photoreceptors for the vertical migration pattern. Flushing sediment surfaces of freshly mixed Wadden Sea sediments with air, O₂, CO₂ or N₂ did not show a significant influence of O₂ on the upward migration. The disappearance of diatoms from sediment surfaces which were flushed with CO₂ is most probably caused by the acidification of the sediment bed.Communicated by E. Rachor     

Hesionides riegerorum n. sp., a New Interstitial Freshwater Polychaete from the United States

Hesionides riegerorum n. sp. is a new freshwater polychaete species. It was found in considerable numbers in the interstitial habitat of sandy banks of the Chowan River in North Carolina, USA. There are no general differences in the organization and behavior compared to the marine species of this genus. The new species, however, is unequivocally distinguished from them by slight quantitative differences and more striking qualitative features. The localities not far from the coast point to an immigration path for this freshwater polychaete that has been shown also for most of the other marine interstitial immigrants: directly from sandy intertidal beaches along brackish estuaries into the limnopsammal.     

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.     

Diel vertical migration and gut pigment rhythm of Paracalanus parvus,P. crassirostris,Acartia erythraea and Eucalanus subcrassus (Copepoda,Calanoida) in Tolo Harbour,Hong Kong

Diel vertical migration and gut pigment rhythm of calanoid copepods were studied in the highly eutrophic waters of Tolo Harbour, Hong Kong in summer between July 30 and August l, 1991 and in winter between January 24 and 25, 1992. Paracalanus parvus and P. crassirostris showed neither diel vertical migration nor diel gut pigment rhythm during the summer study. Acartia erythraea and Eucalanus subcrassus exhibited prominent diel vertical migration and diel gut pigment rhythm. Diel changes in gut pigment content of surface dwelling individuals and increase of gut pigment content before the onset of nocturnal upward migration suggest that gut pigment rhythm in E. subcrassus was not the result of animals moving in and out of a food-rich surface layer. During the winter study, the harbour was affected by a bloom of Noctiluca scintillans. P. parvus and P. crassirostris resided in deeper waters presumably to avoid the dense populations of N. scintillans at the surface. P. crassirostris remained non-migratory during the winter study. There was a nocturnal increase in the number of P. parvus in the surface waters, although no diel change in the mean depth of the population was observed. Both Paracalanus species showed diel gut pigment rhythm. The presence of diel gut pigment rhythm in the non-migratory P. crassirostris during the winter study suggested the presence of an independent feeding rhythm.     

The distribution and diel vertical migration of Pseudodiaptomus hessei (Mrázek) (Calanoida: Copepoda) in a subtropical lake in southern Africa

The vertical distribution and diel migratory behaviour of Pseudodiaptomus hessei in a freshwater lake is described. All stages showed a pronounced diel migration. During daylight naupliar and copepodite instars were almost exclusively benthic in shallow areas (<10m). Copepodite V and adult stages were predominantly benthic even at 40 m. The diurnal vertical distribution pattern implies age-related differences in photosensitivity and a possible depth-regulatory mechanism, based on the existence of differential photosensitivity, is offered to account for day-depth control. Nocturnal vertical distribution, studied approximately fortnightly during 1970–71, showed no clear seasonal variation. Variations in pattern, most obvious in adult instars, correlated with prevalent lunar conditions. Nocturnal or midnight sinking was evident, particularly in post-naupliar stages. Dusk rise and dawn descent were performed at very low light intensities and paralleled changes in light penetration. Dawn descent was less rapid than the dusk rise. An ephemeral dawn rise was sometimes observed but was not a consistent feature of all stages. An endogenous rhythm of locomotor activity was recorded under laboratory conditions. Activity was low during daylight, increased sharply at dusk and then decreased through the night to reach daytime levels at dawn. The stimulus provided by changes in light intensity and/or endogenous activity changes can account for the dusk and dawn movements of P. hessei but a definitive identification of the direct migratory stimulus is not possible. Attention is called to a closer examination of the role of endogenous activity rhythms in vertical migratory behaviour. The possible adaptive value of vertical migration to P. hessei is examined briefly.     

NONDESTRUCTIVE TRACING OF MIGRATORY RHYTHMS OF INTERTIDAL BENTHIC MICROALGAE USING IN VIVO CHLOROPHYLL A FLUORESCENCE1,2

In vivo chlorophyll (Chl) a fluorescence was measured in undisturbed intertidal sediments with the purpose of tracing the vertical migratory rhythms of benthic microalgae. A pulse amplitude fluorometer, an instrument which does not require physical contact with the sample, was used, thus allowing successive measurements to be taken on the same sample without causing any type of disturbance to the sediment structure. The basis of the method is the possibility to detect changes in the Chl a concentration near the sediment surface caused by the vertical movement of the microalgae. This requires the verification of two conditions: the possibility to follow changes in the sediment Chl a content from fluorescence intensity, and a sediment photic depth smaller than the vertical distances covered by the moving microalgae. Both conditions were experimentally verified in intertidal muddy sediments of the Tagus estuary, Portugal. In vivo fluorescence was shown to vary linearly with the sediment Chl a content, and the sediment photic depth was estimated to reach 0.27 mm, a value clearly smaller than the reported depths for microalgal migrations. Sediment samples kept under in situ conditions exhibited large hourly Variations (over 400%) in the Chl a fluorescence intensity, which were closely synchronized with the daytime periods of emersion. The rhythmic fluctuations in Chl a fluorescence were confirmed further to represent microalgal migration by (1) its endogenous nature (fluorescence continued to follow diurnal and tidal cycles after removal of environmental stimuli), (2) its dependence on the vertical distribution of the microalgal population within the sediment (vertically homogenized samples failed to display fluorescence variations), and (3) the lack of significant temperature and light effects on the fluorescence emission under in situ conditions (tested in three species representative of the main groups found in the studied microphytobenthic communities—the diatom Phaeodactylum tricornutum (Böhlin), the cyanobacterium Spirulina maxima (Setch. et Gard.), and the euglenophyte Euglena granulata (Klebs) Lemm.). The results obtained indicate that, in spite of the potential concurrent effects of factors other than the Chl a concentration on the fluorescence intensity, in vivo Chl a fluorescence can be used to trace nondestructively the migratory behavior of benthic microalgae.     

Behavioural versus physiological photoprotection in epipelic and epipsammic benthic diatoms

Benthic diatoms are dominant primary producers in intertidal marine sediments, which are characterized by widely fluctuating and often extreme light conditions. To cope with sudden increases in light intensity, benthic diatoms display both behavioural and physiological photoprotection mechanisms. Behavioural photoprotection is restricted to raphid pennate diatoms, which possess a raphe system that enables motility and hence positioning in sediment light gradients (e.g. via vertical migration into the sediment). The main physiological photoprotection mechanism is to dissipate excess light energy as heat, measured as Non-Photochemical Quenching (NPQ) of chlorophyll fluorescence. A trade-off between vertical migration and physiological photoprotection (NPQ) in benthic diatoms has been hypothesized before, but this has never been formally tested. We exposed five epipelic diatom species (which move in between sediment particles) and four epipsammic diatom species (which live in close association with individual sand grains) to high light conditions, and characterized both NPQ and the relative magnitude of the migratory response to high light. Our results reveal the absence of a significant downward migratory response in an araphid diatom, but also in several raphid epipsammic diatoms, while all epipelic species showed a significant migratory response upon high light exposure. In all epipsammic species the upregulation of NPQ was rapid and pronounced; NPQ relaxation in low light conditions, however, occurred faster in the araphid diatom, compared with the raphid epipsammic species. In contrast, all epipelic species lacked a strong and flexible NPQ response and showed higher susceptibility to photodamage when not able to migrate. While overall our results support the vertical migration-NPQ trade-off, the lack of strong relationships between the capacity for vertical migration and NPQ within the epipsammic and epipelic groups suggests that other factors as well, such as cell size, substrate type and photoacclimation, may influence photoprotective strategies.     

A change in the pattern of vertical migration of Chaoborus flavicans after the introduction of trout

The pattern of diel vertical migration exhibited by Chaoborusflavicans in Lake Lenore, Washington has changed over the pastdecade. Formerly Chaoborus larvae and pupae were present inthe water column during the day. Since 1982, a strong patternof diel vertical migration has been evident in this population.Third and fourth instar larvae and pupae reside in the sedimentsduring the day and then ascend to the surface waters at night.The change in migratory activity of C. flavicans was coincidentwith the introduction of cutthroat trout (Salmo clarki henshawi)into Lake Lenore in 1979. Predation by trout on fourth instarlarvae and pupae of C. flavicans was probably responsible forthe observed changes in migratory behavior.     

Diel vertical migrations of invertebrate predators (Leptodora kindtii ,Thermocyclops taihokuensis, andMesocyclops sp .) in a shallow, eutrophic lake

We studied vertical distribution patterns of three invertebrate predators – Leptodora kindtii, Mesocyclopssp., and Thermocyclops taihokuensis– in a shallow eutrophic lake, Lake Suwa , Japan. From June to October in 2000 and 2001, we collected samples in the lake center in order to examine the vertical distribution patterns and the densities of the predators in the water column during the day (0900) and at night (2330). We also examined phototactic behavior of Leptodora in the laboratory. The three invertebrate predators showed clear migration patterns. Leptodora and Thermocyclops displayed a typical migration, avoiding the surface and maintaining a high abundance in deeper water during the day, and being distributed uniformly during the night. Mesocyclops, on the other hand, showed no clear vertical distribution pattern in the water column. However, Mesocyclops showed higher densities in the water column during the night than during the day. It suggests that they stayed just above the bottom during the day and migrated upward during the night. Leptodora also showed such a density difference between day and night. In the laboratory, Leptodora showed strong negative phototactic behavior. The observed density changes between day and night in Leptodora and Mesocyclops suggests the possible underestimation of their population density by usual sampling methods, and thus the impact of predation on populations of prey zooplankton species may also be underestimated in shallow water bodies.     

Caridina nilotica in Lake Victoria: abundance, biomass, and diel vertical migration

Caridina nilotica (Decapoda: Atyidae) in offshore waters of Lake Victoria were investigated with both day and night sampling over a period of two years. Offshore populations are mainly planktonic rather than benthic, and the animals exhibit diel vertical migrations into near-surface waters at night. These changes in diel abundance as well as the size-frequency distribution of the migrating shrimp suggest that the migratory behavior is in response to visual planktivory, because only the very smallest individuals (2–4 mm) remain in surface waters during the day. During October 1992, abundances were estimated both by vertical net sampling and by underwater video transect methods. Concordance was established between abundances estimated by the two methods. Only about 9% (night) to 14% (day) of the Caridina population appeared to be epibenthic. We suggest that the behavior of the animal is consistent with the hypothesis that it is not a strict detritivore as previously reported; rather it may engage in facultative planktivory, especially at night.     

Inshore and offshore diel migrations in European benthopelagic mysids, genera Gastrosaccus, Anchialina and Haplostylus (Crustacea, Mysidacea)

The nocturnal migratory activity of the six most abundant Gastrosaccinaeliving off the European coast, namely Gastrosaccus sanctus,Gastrosaccus spinifrr, Anchialina agilis, Haplostylus lobatus,Haplostylus lobatus var. armata and Haplostylus norrnani. wasstudied. Both Gastrosaccus perform horizontal and vertical migrationsnear sandy beaches. These migrations involve the whole population,which exhibits a superficial or hyponeustonic distribution inthe water column. Anchialina and Haplostylus perform verticalmigrations from inshore to slope waters. The timing of pelagicphases, as well as the distribution in the water column, dependon age. sex, depth and clarity. Male A.agilis and H.lobatusshow intense migratory activity leading to a hyponeustonic distributionin mid-shelf and slope waters a few hours after dusk. Femalesand juveniles show continuous pelagic phases and a uniform distributionfrom dusk to dawn. Haplosiylus lobatus var. armata and H.normanipopulations exhibit the same behaviour, consisting of overnightmigrations and a uniform to deep nocturnal distnbution. In shallowwaters, male Anchialina and Haplostylus no longer exhibit hyponeustonicbehaviour and their nocturnal distribution becomes more andmore deep as clarity increases. Above the slope, animals thatmigrate from the continental shelf during the night sometimesadopt permanent pelagic behaviour above more than 200 or 300m in depth. but upward nocturnal migration persists and leadsto superficial stratification, albeit delayed. Slow morningdescents to the bottom can occur even further from the 500 misobath.