Feeding rhythmicity in a migratory copepod (Pseudodiaptomus hessei (Mrázek))

SUMMARY. This paper describes a day/night feeding rhythm in the copepod Pseudodiaptomus hessei in subtropical Lake Sibaya (27°20' S, 32°40' E) and examines the origin of this rhythmicity on the basis of field and laboratory observations.
Under natural conditions, adult and late copepodite stages (C_IV-C_V) of this copepod were predominantly benthic during daylight and migrated at dusk into pelagic waters in which they remained until dawn. Pronounced changes in feeding activity (based on the proportion of animals which contained food in the gut) were associated with this transition from a benthic to pelagic distribution; feeding activity was low by day and high at night. The younger copepodite stages (C_I-C_III) were less strongly benthic during daylight and showed correspondingly less pronounced diel changes in feeding activity.
Laboratory experiments involving concurrent light/dark and continuous dark regimes showed that batches of adult and C_IV—C_V stages fed throughout 24 h, although faecal pellet production increased with time under both light regimes. The laboratory observations are interpreted as negating the possibility that diel differences observed in the field arose through an endogenous feeding rhythm. The possibility that food resources in near-bottom waters was inadequate to support reasonable feeding activity is rejected on several counts. It is concluded that the feeding rhythm in P. hessei arises from an inferred infaunal distribution during daylight which would preclude filter-feeding activity. The circumstantial evidence supporting this suggestion is given and the suggestion is evaluated.     

Diel activity of the pink shrimp Penaeus duorarum

Twelve juvenile pink shrimp, Penaeus duorarum Burkenroad, were tested individually for 3-day periods in electronic shuttleboxes to determine their diel patterns of locomotor activity, in relation to a natural summer photoperiod. Nocturnal activity was twice that exhibited during the daytime; however, a bimodal pattern was evident with crepuscular peaks occurring at dawn and dusk. The dusk peak was more pronounced, with activity increasing markedly before sunset (indicative of an endogenous circadian rhythm component), and continuing during the initial hours of darkness, gradually declining later during the night with a secondary peak at dawn falling off to minimal activity during daylight. Crepuscular activity (mean of dawn and dusk) was twice the nocturnal average.     

The influence of light intensity on vertical distribution inRanatra linearis L. (Heteroptera: Nepidae)

The vertical distribution of adultRanatra linearis L. (Heteroptera: Nepidae) showed diel variations: subjects were significantly nearer the water surface during the night than during the day. This distribution was not influenced by the depth of water available. Changes in vertical distribution coincided with the natural changes of light intensity, the animals rising to the surface at dusk and descending at dawn. Variations of vertical distribution were followed during post-embryonic development. Young larvae remained very close to the water surface and also avoided the darker parts of the aquarium, in a vertical as well as in a horizontal light gradient. However, as these larvae grew older, they tended to move away from the water surface towards darker zones. The change was gradual, but especially noticeable after the third larval moult. Different biological factors possibly involved in inducing this change are discussed.     

The diel activity pattern of mountain hare (Lepus timidus) on managed heather moorland in Scotland

The research presented in this paper provides an insight into the behavioral ecology of mountain hares on heather moorland in the Lammermuir Hills of southeast Scotland. We examine the seasonal and diel activity patterns using camera traps over a period of 12 months. The rate of camera detections was calculated for the different divisions of the 24‐hr cycle (daylight, dusk, night, and dawn). During autumn and winter (October–February), the activity pattern was crepuscular with greater activity at dusk than at dawn. Daylight activity was relatively low, and there was a regular pattern of small peaks of activity during the night. In spring and summer (March–September), peaks of crepuscular activity remained evident but daylight activity was much more prevalent than during autumn and winter, and night activity was lower. We discuss the problematic definition of twilight and present an explanation for seasonal changes in the pattern of diel activity that is linked to the reproductive cycle of the mountain hare.     

Contrasting diel activity and feeding patterns of four instars of Rhyacophila dorsalis (Trichoptera)

1. Ontogenetic shifts in prey choice and predator behaviour can affect food‐web structure. Therefore, it is important to establish if the diet and feeding activity differ between life‐stages of the same species. This hypothesis was tested for second, third, fourth and fifth larval instars of Rhyacophila dorsalis by comparing their diel activity and feeding patterns. Second to fifth instars collected from two streams were used either for gut analyses or for observations of their activity and feeding patterns in three stream tanks. Food was provided in excess; being organisms living in bryophytes on top of a large stone in each tank, augmented by different‐sized larvae of Ephemeroptera, Simuliidae and Chironomidae. As few first instars for gut analyses were found in the field, the diet of first instars reared in the laboratory was also studied. 2. Larvae for gut analyses were taken 1 h before dusk or dawn (n = 50 larvae per instar for each day or night sample). First and second instars fed on the smaller food items with no significant day‐night differences in diet. Gut contents indicated a progressive trend from feeding chiefly at night in third instars to almost exclusively at night in fifth instars. Fourth and fifth instars fed on the larger food items, whilst the diet of the third instar larvae overlapped with that of both the earlier and later instars. 3. Diel activity patterns of single larvae differed between instars but not within each instar (n = 20 larvae per instar). Second instars were active throughout the 24 h, with peaks at dusk, around midnight, dawn and around midday. A similar pattern was shown by third instars but the peak of activity at midday was less than the other three peaks. Prey were captured only during these peaks for both instars. Fourth and fifth instars were most active, and fed only, at night. They used an ambush strategy to capture more active prey at dusk and dawn (e.g. Baetis, Gammarus), and a searching strategy to capture more sedentary prey during the night (e.g. chironomids, simuliids). These experiments provided support for the hypothesis under test. If competition and/or interference occur between instars, then it could be reduced between earlier and later instars because of differences in their diet and diel pattern of feeding activity.     

Diel vertical movements of zooplankton in lake La Cruz (Cuenca, Spain)

The study of diel vertical movements of zooplankton in a smallmeromictic karstic lake (lake La Cruz), presenting highly stratifiedwaters, was performed using two different methodologies: (i)samples were taken along the vertical profile and were comparedat different hours in a diel cycle; (ii) some plankton trapswere located at different depths, covering different periodsof time, to catch organisms going upwards and downwards. Themain subject of this study has been the vertical movements affectingrotifers since they were dominant in the zooplankton of thislake, but we have also included the results obtained for otherzooplankton groups. The results indicate an almost general movementin epilimnetic waters corresponding to the ‘normal’pattern of migration (ascent at dusk and descent at dawn); inmetalimnetic and hypolimnetic waters, vertical movements weregreatly reduced and were more independent of the diel cycle.The results also confirm that the dense populations at theselevels of the oxic–anoxic boundary are made up of activeanimals. Among the causes accounting for our results, the cyclesof activity (mainly feeding) connected with the diel cycle couldbe important in explaining vertical movements of filter-feederrotifers. In larger zooplankters, visual predation might bethe main reason for such behavior.     

Extensive diel fish migrations in a deep ultraoligotrophic lake of Patagonia Argentina

Andean Patagonian lakes are ultraoligotrophic, highly transparent water bodies with very low densities of organisms at the different levels of their trophic food webs. Little is known about spatial distribution and habitat use by fish in these lakes. Hydroacoustic and active sampling techniques were used to study the distribution, composition, and displacement of organisms in an Andean lake throughout the diel cycle on two different moon phase days. Sound scattering layers (SSLs) were found both in the near-shore and pelagic habitats at different times of the day. These SSLs, formed by galaxiid larvae and adults, underwent displacement during the time periods of dawn and dusk, giving rise to the redistribution of organisms in the different habitats. The organisms show high sensitivity to light intensity, displaying different behaviors depending on moon phases. Extensive diel vertical migrations show that the deep pelagic habitats provide diurnal refuge for native galaxiids in Andean Patagonian lakes. Due to the high densities of galaxiid larvae and adults, it is likely that diel migrations generate an active flow of energy and matter between habitats, which could have a profound influence on whole lake dynamics.     

DIEL PATTERNS IN LIGHT ABSORPTION AND ABSORPTION EFFICIENCY FACTORS OF ISOCHRYSIS GALBANA (PRYMNESIOPHYCEAE)1

The chl a specific absorption coefficients [a* (λ), m²·mg chl a ^{? 1}] were examined in chemostat culture of the Prymnesiophyceae Isochrysis galbana (Parke) under a 12:12‐h light:dark cycle at low light (75 μmol photons·m ^{? 2}·s ^{? 1}) and high light (500 μmol photons· m ^{? 2}·s ^{? 1}) conditions. Other associated measurements such as pigment composition, cell density, and diameter as the measure of cell size were also made at the two light regimes every 2 h for 2 days to confirm the periodicity. A distinct diel variability was observed for the a* (λ) with maxima near dawn and minima near dusk. The magnitude of diel variation in a* (440) was 15% at low light and 22% at high light. Pronounced diel patterns were observed for cell size with minima near dawn and maxima near dusk. The magnitude of diel variation in cell size was 9.3% at low light and 21% at high light. The absorption efficiency factors [Q _a (440)] were determined by reconstruction using intracellular concentrations of pigments and cell size. The Q _a (440) also showed a distinct diel variability, with minima near dawn and maxima near dusk. The diel variation in a* (λ) and Q _a (λ) was primarily caused by changes in cell size due to growth, although there was some influence from diel variations in the intracellular pigment concentrations. The results presented here indicated that diel variation in a* (λ) was an important component of the optical characterization of phytoplankton.     

Spatial dynamics and expanded vertical niche of blue sharks in oceanographic fronts reveal habitat targets for conservation

Dramatic population declines among species of pelagic shark as a result of overfishing have been reported, with some species now at a fraction of their historical biomass. Advanced telemetry techniques enable tracking of spatial dynamics and behaviour, providing fundamental information on habitat preferences of threatened species to aid conservation. We tracked movements of the highest pelagic fisheries by-catch species, the blue shark Prionace glauca, in the North-east Atlantic using pop-off satellite-linked archival tags to determine the degree of space use linked to habitat and to examine vertical niche. Overall, blue sharks moved south-west of tagging sites (English Channel; southern Portugal), exhibiting pronounced site fidelity correlated with localized productive frontal areas, with estimated space-use patterns being significantly different from that of random walks. Tracked female sharks displayed behavioural variability in diel depth preferences, both within and between individuals. Diel depth use ranged from normal DVM (nDVM; dawn descent, dusk ascent), to reverse DVM (rDVM; dawn ascent, dusk descent), to behavioural patterns where no diel differences were apparent. Results showed that blue sharks occupy some of the most productive marine zones for extended periods and structure diel activity patterns across multiple spatio-temporal scales in response to particular habitat types. In so doing, sharks occupied an extraordinarily broad vertical depth range for their size (1.0-2.0 m fork length), from the surface into the bathypelagic realm (max. dive depth, 1160 m). The space-use patterns of blue sharks indicated they spend much of the time in areas where pelagic longlining activities are often highest, and in depth zones where these fisheries particularly target other species, which could account for the rapid declines recently reported for blue sharks in many parts of the world's oceans. Our results provide habitat targets for blue shark conservation that may also be relevant to other pelagic species.     

Vertical migrations of herring,Clupea harengus,larvae in relation to light and prey distribution

Synopsis The influence of light and prey abundance on the vertical distribution of herring larvae was evaluated by three investigations made under calm weather conditions in the North Sea off the Scottish coast. The investigations took place at different time after hatching and the vertical distributions of three size groups of larvae (mean sizes 8,15 and 19 mm) were related to time of day and the vertical distribution of copepods. No migratory behaviour of copepods was observed but their vertical distribution differed between investigations. In the investigation on intermediate sized larvae, copepod density peaked at the pycnocline (40 m). Larvae concentrated at this depth at noon. At dawn and dusk larvae migrated towards the surface and the vertical distributions fluctuated semidielly. In the two other investigations, copepods were homogeneously distributed in the water column and after migration towards the surface at dawn larvae stayed in the upper water column during the day. The observations suggest that the daytime vertical distribution of larvae in calm weather is mainly determined by feeding conditions: the larvae move to depths were light is sufficient for feeding, and refinement within that zone is made according to a compromise between optimal light conditions for feeding and optimal prey densities.     

Relaxed circadian rhythm in zooplankton along a latitudinal gradient

To test whether aquatic invertebrates are able to adjust their diel migratory cycle to different day length and presence of predators, we performed standardized enclosure experiments in shallow lakes at four different latitudes from southern Spain, with strong night–day cycles, to Finland where daylight is almost continuous during summer. We show here that nearly continuous daylight at high latitudes causes a relaxation in diel migratory behaviour in zooplankton irrespective of predation risk. At lower latitudes, however, similar conditions lead to pronounced diel rhythms in migration. Hence, zooplankton may show local behavioural adaptations in their circadian rhythm. They are also able to make risk assessments as to whether diel migration is beneficial or not, manifested in a lack of diel migration at near constant daylight, irrespective of predator presence. Our results provide an additional explanation to previous knowledge regarding diel migrations among aquatic invertebrates by showing that both physical (light) and biological (predation) factors may affect the migratory behaviour.     

The influence of light on the diel vertical migration of young-of-the-year burbot Lota lota in Lake Constance

The diel vertical distribution of young-of-the-year (YOY) burbot Lota lota in the pelagic zone of Lake Constance was compared to light intensity at the surface and to the light intensity at their mean depth. Lota lota larvae inhabited the pelagic zone of Lake Constance from the beginning of May until the end of August. From early June, after the stratification of the water column, fish performed diel vertical migrations (DVM) between the hypolimnion and epilimnion. The amplitude of DVM increased constantly during the summer and reached 70 m by the end of August. Lota lota started their ascent to the surface after sunset and descended into the hypolimnion after sunrise. As the YOY fish grew from May to August, they experienced decreasing diel maximum light intensities: in May and early June L. lota spent the day at light intensities >40 W m⁻², but they never experienced light intensities >0·1 W m⁻² after the end of June. From this time, L. lota experienced the brightest light intensities during dusk and dawn, suggesting feeding opportunities at crepuscular hours. The present study implies, that YOY L. lota in the pelagic zone of Lake Constance increased their DVM amplitude during the summer to counteract a perceived predation risk related to body size and pigmentation.     

What goes down must come up: symmetry in light-induced migration behaviour of Daphnia

During a short period of the year, Daphnia may perform a phenotypically induced diel vertical migration. For this to happen, light-induced swimming reactions must be enhanced both at dawn and at dusk. Enhanced swimming in response to light intensity increase can be elicited by fish-associated kairomone in the laboratory, if food is sufficiently available. However, during the light change at dusk the Daphnia are still in the hypolimnion, where no fish kairomone is present and both temperature and food availability is low. Still, what goes down must come up. This raises questions about how Daphnia tunes its light-induced swimming behaviour to prevailing conditions such that a normal diel vertical migration can be performed. We investigated the symmetry in behavioural mechanism underlying these diel vertical migrations in the hybrid Daphnia galeata × hyalina (Cladocera; Crustacea), with special interest for the environmental cues that are known to affect swimming in response to light increase. That is, we tested whether fish- associated kairomone, food availability, and temperature affected both swimming in response to light intensity increase and decrease similarly. We quantified swimming behaviour during a sequentially increased rate of light change. Vertical displacement velocity was measured and proved to be linearly related to the rate of the light change. The slope (PC) of the function depends on the value of the factors kairomone concentration, food availability, and temperature. The changes of the PC with kairomone concentration and with temperature were similar both at light intensity increases and decreases. The PC also increased with food concentration, although during light increases in a different way from during light intensity decreases. Low food availability inhibited swimming in response to light intensity increase, but enhanced swimming in response to light intensity decrease. Hence, ascent from the deep water layers with low food concentration at dusk is facilitated. These causal relations are part of a proximate decision-making mechanism which may help the individual Daphnia to tune migration to predation intensity and food availability.     

Daily activity patterns of coyotes (Canis latrans) in a tropical deciduous forest of western Mexico

We studied the activity patterns of the coyote (Canis latrans) in a tropical deciduous forest in the Mexican Pacific coast over 3 years. Fifteen coyotes (six females, nine males) were fitted with radio-collars equipped with activity sensors to determine the influence of seasonality (dry vs. wet), gender (males vs. females) and diel intervals (dusk, night, dawn, and day) on activity patterns. We found differences in activity patterns between diel intervals, but the only pair of diel intervals that showed significant differences was dawn (more active) vs. day (less active). We found no differences due to sex or season on any of the four studied diel intervals. Coyote activity patterns in this tropical forest could be responding to prey availability, human avoidance or thermoregulation.     

Preliminary observations on predator/prey relationships between chaetognaths and copepods in the Java sea

There is evidence that copepods undertake vertical migrations in shallow waters off the north coast of Java. They aggregate at the surface of the sea at dawn and dusk but sink to deeper levels in the middle of the day. Chaetognaths feed on copepods and tend to occur in areas which are rich in their prey. They do not however follow them on vertical migrations during daylight.     

Food selectivity and diel vertical distribution of Chaoborus edulis (Diptera, Chaoboridae) in Lake Malawi

1. Diel diet and vertical distribution patterns of the larval instars of Chaoborus edulis were studied in deep water near the central part of Lake Malawi, Africa.
2. First instar larvae contained very little food in their crops and probably depended on reserves from the egg. Second, third and fourth instars fed on zooplankton and were size-selective in their feeding. The mean size of prey eaten by the three instars was significantly different from each other, with larger instars feeding on larger prey. Smallest available prey was selected against and the upper size of prey was probably constrained by larval gape. Nauplii were not found in any of several thousand larvae examined. Phytoplankton did not form a significant part of the diet.
3. There was a progressive and related increase in diel periodicity in feeding and vertical migrations of successive instar stages. Fourth instars migrated particularly large distances. Such migrations removed them from their zooplankton food supply but avoided predators. A refuge from predators is probably found in or near the permanent zero oxygen boundary, at depths greater than 200 m.     

The substrate bin—a new sampling device for studying diel vertical migratory movements on to and off lake sediments

The substrate bin, used to study diel migratory movements of the copepod Pseudodiaptomus hessei on to and off bottom interfacial areas is described. It consists essentially of a transparent Perspex box open at its upper end and supported upright within a metal framework. A lid, hinged along one face of the open end of the bin, opens and closes automatically through a simple system of levers as the apparatus is lowered on to or raised off a solid substrate. The apparatus is lowered on to the lake bed where it opens. Diel movements on to or off the shallow layer of bottom substrate enclosed within the bin can be examined by retrieving bins serially at different times. Using this apparatus in conjunction with standard net sampling techniques, clear evidence for an alternation of benthic and pelagic phases during diel cycles was obtained. Notwithstanding an apparent oversampling of the benthic component, the magnitude of which cannot be defined precisely, it is concluded that the apparatus provides a simple and reliable method of sampling.     

The thermoregulatory function of diel vertical migration for a juvenile fish,Cottus extensus

Juvenile sculpin (Cottus extensus) less than 30 mm long exhibit a diel vertical migration in the limnetic zone of Bear Lake (Utah-Idaho). Using mid-water and bottom trawls we found that these fish inhabit the bottom of the lake (5° C) during the day but migrate 30–40 m into the water column at night where they reside in the metalimnion or epilimnion at temperatures near 13–16°C. Larger fish do not migrate into the water column. Stomach analyses demonstrated that the young-of-the-year fish do not migrate into the water column to feed: from July to October their diet is 70–93% benthic ostracods and copepods, and pelagic prey are rarely consumed. Furthermore, gut fullness of the sculpin increases through the daylight period and decreases through the night, reaching minimum levels just before the dawn descent. Laboratory experiments demonstrated that the diel migration would increase digestion rate from 3%/h at profundal temperatures, to 22%/h in the warmer surface water, thus allowing the fish to empty their guts overnight and permit feeding the following day. Additionally, sculpin held in a temperature and feeding regime that mimicked that experienced by migrating fish grew 300% faster than those reared at 5° C. Given the overwhelming importance of fast growth for juvenile fishes, a post-feeding thermotaxs that increases digestion may be a common phenomenon increasing growth, and affecting the distribution and bioenergetic relationships of fish.     

Diel migration and site fidelity in a stream-dwelling cyprinid, Leuciscus leuciscus

During May and June 1995, radio-telemetry was used to record diel movements between different, clearly defined daytime and night-time habitats in dace Leuciscus leuciscus . The frequent and often rapid return of individuals to previously occupied locations at dawn and dusk suggests differential habitat suitability according to light intensity, and appears similar in some respects to roosting behaviour in birds.     

Diel and seasonal habitat and food segregation of three sympatric Coregonus lavaretus forms in a subarctic lake

Habitat use of three sympatric whitefish Coregonus lavaretus forms was determined using hydroacoustics, pelagic trawling and epibenthic gillnetting in the subarctic Lake Muddusjärvi during the day and night in June, August and September. Whitefish constituted 97% of the numerical catches and whitefish with high number of gill rakers (DR) were the most abundant whitefish form. Forms with low numbers of gill rakers used only epibenthic habitats during both the day and night in all study periods: large whitefish with low numbers of gill rakers (LSR) dwelled mainly at depths 0–10 m, whereas small whitefish with low numbers of gill rakers (SSR) used deeper (>10 m) habitats. LSR and SSR whitefish consumed mainly benthic macroinvertebrates during all study occasions. The planktivorous DR whitefish used both epibenthic and pelagic habitats, but vertical habitat selection varied both over time of day and season. In June, when light intensity was continuously high, DR whitefish did not perform diel vertical migrations. In August and September, when dark nights were distinguishable, DR whitefish ascended from the bottom to the pelagic at dusk to feed on zooplankton, and descended at dawn. DR whitefish used pelagic habitats only at the lowest light intensities during the night, which was probably related to the high predation risk from brown trout Salmo trutta .