Foraging rhythm of great crested grebes Podiceps cristatus adjusted to diel variations in the vertical distribution of their prey Osmerus eperlanus in a shallow eutrophic lake in The Netherlands

Summary Great crested grebes Podiceps cristatus only foraged for an hour or two during dawn and again during dusk on Lake IJsselmeer in August-September. During this time of the year the adult birds are in wing moult and temporarily unable to fly. The food of grebes consisted almost exclusively of smelt Osmerus eperlanus, the most numberous pelagic fish. Simultaneous sonar registrations and trawl net fishing showed that smelt moved to the water surface during the twilight periods. During day and night they were concentrated near the bottom. We argue that grebes have the best foraging opportunities during twilight when much of their prey is near the surface, where light intensities allow the fish to be detected and captured. When the smelt are in the upper water layers the distance to the covered to get the prey (i.e. diving time and cost) is also least.     

Piscivore-Prey Fish Interactions: Mechanisms behind Diurnal Patterns in Prey Selectivity in Brown and Clear Water

Environmental change may affect predator-prey interactions in lakes through deterioration of visual conditions affecting foraging success of visually oriented predators. Environmental change in lakes includes an increase in humic matter causing browner water and reduced visibility, affecting the behavioural performance of both piscivores and prey. We studied diurnal patterns of prey selection in piscivorous pikeperch (Sander lucioperca) in both field and laboratory investigations. In the field we estimated prey selectivity and prey availability during day and night in a clear and a brown water lake. Further, prey selectivity during day and night conditions was studied in the laboratory where we manipulated optical conditions (humic matter content) of the water. Here, we also studied the behaviours of piscivores and prey, focusing on foraging-cycle stages such as number of interests and attacks by the pikeperch as well as the escape distance of the prey fish species. Analyses of gut contents from the field study showed that pikeperch selected perch (Perca fluviatilis) over roach (Rutilus rutilus) prey in both lakes during the day, but changed selectivity towards roach in both lakes at night. These results were corroborated in the selectivity experiments along a brown-water gradient in day and night light conditions. However, a change in selectivity from perch to roach was observed when the optical condition was heavily degraded, from either brown-stained water or light intensity. At longer visual ranges, roach initiated escape at distances greater than pikeperch attack distances, whereas perch stayed inactive making pikeperch approach and attack at the closest range possible. Roach anti-predatory behaviour decreased in deteriorated visual conditions, altering selectivity patterns. Our results highlight the importance of investigating both predator and prey responses to visibility conditions in order to understand the effects of degrading optical conditions on piscivore-prey interaction strength and thereby ecosystem responses to brownification of waters.     

Day–night changes in the spatial distribution and habitat preferences of freshwater shrimps, Gammarus pulex, in a stony stream

1. As many invertebrates are nocturnal, their spatial distribution and habitat preferences may change from day to night. Both aspects are examined for Gammarus pulex by testing the hypotheses: (i) a power function was a suitable model for the spatial distribution of the shrimps in both day and night; (ii) diurnal and nocturnal spatial distributions were significantly different; (iii) diurnal and nocturnal habitat preferences were significantly different. Five different life‐stages were treated separately. To ensure that the conclusions were consistent, large samples were taken near midday and midnight in April, June and November over 4 years at two sites about 3 km apart in a stony stream: downstream (n = 30) and upstream (n = 50). 2. The first and second hypotheses were supported at both sites. A power function, relating spatial variance (s²) to mean (m), was an excellent fit in all analyses (P < 0.001, r² > 0.91), i.e. the spatial variance was density‐dependent. All five life‐stages were aggregated in the day. At night, the degree of aggregation increased for juveniles at higher densities but decreased for juveniles at lower densities, increased for immature females and males, but decreased slightly for mature females and especially mature males, the latter being close to a random distribution. There were no significant differences between sites, in spite of the lower numbers at the downstream site. 3. The third hypothesis was tested at only the upstream site and supported by comparisons between shrimp densities and 13 physical variables (distance from bank, water depth, water velocity, ten particle size‐classes), and three non‐physical variables (dry weights of bryophytes, leaf material, organic detritus). During the day, densities were strongly related to particle sizes with the following preferences: 0.5–8 mm for juveniles, 8–256 mm for the other life‐stages with a weaker relationship for males. There were no significant positive relationships with the other variables, apart from bryophytes for immature shrimps and adults. At night, densities were unrelated to particle size; juveniles and immature shrimps preferred low water velocities near the banks, often where leaf material and organic detritus accumulated, females often preferred medium water velocities slightly away from the banks, and males showed no habitat preferences. 4. Day samples do not provide a complete picture of habitat preferences and probably identify refuge habitats. Day–night changes in spatial distribution and habitat preferences are an essential part of the behavioural dynamics of the shrimps and should be investigated in other species.     

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.     

Effects of Ultraviolet Radiation on Diel Vertical Migration of Crustacean Zooplankton: An in situ Mesocosm Experiment

The objective of this study was to expand the spatial scale of previous experiments on the effects of ultraviolet radiation (UVR) on diel vertical migration (DVM) by freshwater zooplankton. We conducted an in situ mesocosm experiment in highly UVR transparent Lake Giles, Pennsylvania, in which we imposed two treatments: ambient UVR and UVR-shielded. Mesocosms (3440 L, 0.74 m diameter, 8 m deep) were large enough to include a spatial refuge from UVR and permit relatively large-scale DVM. Daphnia catawba adopted a significantly deeper distribution during the day in the ambient UVR treatment compared to the UVR-shielded treatment, but effects of UVR were absent at night. In contrast, DVM by Leptodiaptomus minutus was unaffected by the UVR treatment. In both treatments, Leptodiaptomus minutus were most abundant at the bottom of the mesocosms during the day and exhibited a more uniform distribution across depths at night. These results suggest that UVR, along with temperature, algal resources, and predators, may affect zooplankton DVM in aquatic ecosystems.     

Sleeping cluster patterns and retiring behaviors during winter in a free-ranging band of the Sichuan snub-nosed monkey

Little information is available on the sleeping cluster pattern and retiring behavior of Sichuan snub-nosed monkeys (Rhinopithecus roxellana). Here, we provide observational data on a provisioned free-ranging band in the Qinling Mountains, central China. The results suggest that winter night activity of R. roxellana is a compromise between antipredator and thermoregulatory strategies and an adaptation to ecological conditions of their temperate habitat. Monkeys retired between 1804 and 1858 h in winter. In support of the antipredation hypothesis, all monkeys slept in trees at night, whereas 18.8% of individuals slept on the ground during the day. Also, the study band was more spatially cohesive at night than in daytime, with shorter distances between one-male units. Keeping warm is critical for survival in freezing temperatures. Monkeys often slept in the lower stratum of the tree canopy, avoiding the upper canopy where it is cold and windy. They formed larger sleeping clusters at night than in daytime. The most common types of night-sleeping clusters were adult females and juveniles, followed by adult females with other adult females. These accounted for 60.2% of the total records. The frequency of female–male clusters is two times greater, and that of adult male–juvenile clusters was four times less at night than during the day. The variations in composition of sleeping clusters suggest affiliative patterns at night-sleeping sites differ from those during the day.     

Habitat overlap of enemies: temporal patterns and the role of spatial complexity

Environmental heterogeneity can promote coexistence of conflicting species by providing spatial or temporal refuges from strong interactions (e.g., intraguild predation, competition). However, in many systems, refuge availability and effectiveness may change through time and space because of variability in habitat use by either species. Here I consider how the intensity of intraguild predation risk varies from day to night for aquatic insects that use both vegetated and open water habitats. Large (1,265 l) and small (42 l) mesocosms were used to test the hypothesis that Buenoa would choose an open-water habitat that minimized predation by the ambush predator Notonecta during the day, but that at night Buenoa would safely use both vegetated and open water. Regardless of container size, Notonecta remained in vegetated water during the day and exploited both habitats at night, despite exhibiting greatest instantaneous predation rates in open water during the day. In contrast, Buenoa maintained an even distribution throughout the mesocosms during day and night, even though habitat-specific predation risks were fivefold lower in open waters than in vegetation during the day and habitat-specific predation risk would have been reduced threefold by fully exploiting open waters. Thus, temporal heterogeneity was both beneficial and detrimental to Buenoa; darkness of night reduced predation, but spatial refuges also disappeared. Together, these patterns suggest that while environmental heterogeneity can dampen intense biotic interactions, enemies do not select habitats solely on the basis of conflict avoidance. Instead, it appears that habitat-specific variation in other biotic (e.g., visual predators) or physical factors (e.g., UV radiation) may also mediate species interactions by influencing habitat selection.     

Diving patterns of a Ross seal (Ommatophoca rossii ) near the eastern coast of the Antarctic Peninsula

In January 1987 we documented the diving patterns of a female Ross seal (Ommatophoca rossii) in the marginal pack-ice zone near the eastern coast of the Antarctic Peninsula for 2 days using a microprocessor-based time-depth recorder. The seal hauled out during the day and dived continually when in the water at night. Dives averaged 110 m deep and 6.4 min long; the deepest dive was 212 m and the longest 9.8 min. Dives were deepest near twilight and shallowest at night; this pattern suggests that the seal's prey, presumably mid-water squid and fish, may have been making vertical migrations or changing predator-avoidance behavior in response to diel light patterns. The dives of this Ross seal were substantially deeper, on average, than those of crabeater seals (Lobodon carcinophagus), which forage in the same areas on Antarctic krill (Euphausia superba). Received: 15 August 1996 / Accepted: 22 February 1997     

Diel activity patterns of the fish community in a temperate stream

The diel activity patterns of fishes in a temperate New Brunswick stream were studied during the summer over 5 years. Young‐of‐the year Atlantic salmon Salmo salar and blacknose dace Rhinichthys atratulus were more active during the day than at night, whereas lake chub Couesius plumbeus, brook trout Salvelinus fontinalis and adult white suckers Catostomus commersonii were more active at night than during the day. Because fishes were as likely to be nocturnal as diurnal, the data suggest that more night‐time sampling is needed to provide an unbiased view of fish community structure in temperate streams.     

Laboratory-simulated naturally-decreasing day lengths, twilight and aphid photoperiodism

Abstract. Day-length changes, as well as periods of twilight, that occur in the course of each natural day-night cycle, were recreated for two chosen latitudes in a computer-controlled 'natural-day-length simulator'.The photoperiodic responses of two aphid species, Aphis fabae and Megoura viciae , were examined in conditions which mimicked late summer to autumn at two simulated latidudes, 51.5N (e.g.Ascot, Southern Britain) and 60°N (e.g.southern tip of Shetland Islands, Northern Britain), with temperatures between 16°C ('night') and 18°C ('day').The responses under simulated natural photoperiodic conditions were similar to those observed under conventional experimental conditions of squarewave light-dark cycles (with abrupt lights-on and lights-off and constant light intensities during the light phase): both aphid species responded to civil twilight as light, and the critical day lengths (including civil twilight) for the induction of sexual morphs by the two aphid species observed in the simulator were the same as those found in squarewave light-dark cycles.
Autumn field experiments (51.5°N) with the same clones of A. fabae and M.viciae revealed much longer critical day lengths for gynopara and male induction in A. fabae compared with those in the laboratory, but the same critical day length for ovipara induction in M.viciue. Minimal night temperatures in the field were on average 6°C, whereas maximal day temperatures declined from around 30°C in early September to 12°C at the end of October; it seems that the critical day lengths in A.fabae are temperature dependent, whereas the findings for M.viciae confirm that the critical day length is temperature compensated.     

First Year Ontogenetic Diet Patterns in Two Coexisting <Emphasis Type="Italic">Sander</Emphasis> Species, <Emphasis Type="Italic">S. lucioperca</Emphasis> and <Emphasis Type="Italic">S. volgensis</Emphasis> in Lake Balaton

First year ontogenetic diet patterns and size-dependent diet partitioning of two coexisting piscivorous percids, pikeperch and Volga pikeperch were analysed in Lake Balaton. Diet patterns of the two pikeperches followed a common trend, catching larger preys at larger sizes, but in details they were different. Until 20–30 mm size age-0 pikeperch fed on planktonic microcrustaceans and were found only in the pelagic zone. After this size, from the end of May, a part of the stock shifted habitat and moved to the littoral zone to feed on Limnomysis benedeni. Majority of the stock stayed in the pelagic zone and shifted to Leptodora kindtii. In the pelagic group early piscivory, mainly cannibalism, occurred only sporadically. However, the few individuals that reached 100 mm size during the first year were typically piscivorous. Volga pikeperch stayed in the pelagic zone during the first year and shifted from planktonic microcrustaceans to benthic macroinvertebrates. General patterns of the first year diet ontogeny were similar in the different areas of Lake Balaton in both pikeperches with only some differences in the secondary food resources, especially in the share of chironomids. Chironomids were eaten at highest ratio by the two fish species in the Keszthely basin in accordance with their abundance pattern. Maximum prey sizes increased significantly with predator size in both pikeperch and Volga pikeperch. However, pikeperch ate much larger prey than did Volga pikeperch at the same size although they have a similar mouth gape to length ratio. These two, closely related pikeperches, although occur in the same habitat and have similar patterns of diet ontogeny, effectively partition food resources in Lake Balaton during the first growing season, except in the very beginning of their ontogeny at L = 5–10 mm size. It appears that age-0 Volga pikeperch is not a significant competitor for the economically highly rated pikeperch in Lake Balaton. On the other hand, it is age-0 pikeperch that might affect Volga pikeperch during its early ontogeny, in the planktivorous stage.     

Diel activity and thermoregulatory behavior of a fully aquatic frog: Xenopus laevis

Ten adult Xenopus laevis were tested individually for 48-hr periods, following an initial 24-hr introductory period, in electronic shuttleboxes which allowed them to control water temperatures without operant conditioning. Locomotor activity was recorded via photocell-monitored light beams. The frogs were nocturnal, being nearly twice as active at night as during the day. The mean preferred temperature was 22.4°C, with no significant difference between night (22.5°C) and day (22.3°C), although the modal preferendum shifted from 24°C by day to 22°C at night, with a corresponding change in skewness. The range of voluntarily occupied temperatures was 14–32°C by day and 14–29°C at night. The median thermal preferendum was 22°C both day and night.     

Migratory behaviour of adult pikeperch (Stizostedion lucioperca) in a lowland river

The behaviour of radio-tagged adult pikeperch (Stizostedion lucioperca (L.)) from two areas in the Danish River Gudenaa were recorded prior to, and during the spawning period. Eight of 13 tagged fish in the lower reaches of the river were located throughout the whole study. Five of these fish moved upstream to various sites in the river prior to spawning, which occurred from late April to June. The three remaining fish moved to the fjord. These movements were interpreted as a spawning migration, and it is suggested that the lower reaches of the River Gudenaa constitute an over-wintering area for pikeperch, which use different spawning areas. Ten pikeperch caught just downstream of an impassable hydropower plant were radio-tagged and translocated upstream of the dam to a reservoir. Within a week, half of the fish moved to a lake situated more than 30 km upstream the reservoir. This behaviour is hypothesised to be a homing response. The study reveals that the pikeperch is a highly mobile species with a complicated migration pattern, even in relatively small river systems.     

Autumnal shift from diurnal to nocturnal peaking feeding activity of Rutilus rutilus in boreal lake littoral zones

The possibility that the feeding activity of roach Rutilus rutilus shows an autumnal shift towards hours of lower light intensity in shallow and illuminated habitats in temperate watercourses as a consequence of relatively increasing daytime predation risk was tested in a littoral zone of a large boreal lake using trapping experiments. Autumnal feeding activity of R. rutilus first showed a striking shift from day to twilight when water temperature fell from 20 to 10° C, and a low but steady night activity prevailed in late autumn, when temperature had fallen to 5° C and the length of the night had increased. The size of the captured littoral R. rutilus also increased throughout the autumn. Both shifts may be linked to increasing daytime predation risk and decreasing food demand and availability.     

Horizontal distribution of pelagic zooplankton in relation to predation gradients

An investigation was conducted examining the horizontal and vertical distribution of zooplankton in Lake Miramar, a southern California reservoir. Daphnia and Mesocyclops populations were most abundant offshore and in deeper water during the day but appeared to move toward shore and upward at night. The results of inshore zooplankton sampling provided no evidence chat the diel horizontal migration pattern was a result of sampler avoidance by zooplankton. Inshore-offshore differences in Daphnia and Mesocyclops abundance and diel migrations were reduced during winter and early spring. Rotifer zooplankters exhibited less seasonal variation in their horizontal distributions than did the large crustacean zooplankters at all times of the year. It is hypothesized that the spatial distribution of zooplankton is related to predation gradients in Lake Miramar. The dominant planktivore in the reservoir, young-of-the-year Micropterus salmoides. was abundant from late May through December and much less so from January to early May. They were largely restricted to the littoral zone and this produced horizontal gradients of planktivory which varied in strength seasonally and from day lo night. It appears that crustacean zooplankton in Lake Miramar avoid areas with abundant planktivores during the day but migrate into these areas at night when the intensity of planktivory is reduced. Rotifers exhibit less horizontal heterogeneity and no significant diel migrations, which is attributed to the reduced risk of predation that rotifers experience relative to crustacean zooplankters. A graphical model is proposed to integrate our understanding of diel vertical and horizontal migrations of zooplankton. In this model, gradients of predation are completely vertical in offshore areas and strongly horizontal in near shore areas. Gradients of food availability are roughly similar to those of predation intensity. Plankiers respond to these gradients by migrating in a path parallel to gradients of predation at dawn and parallel to gradients of food availability after dark.     

Horizontal and vertical migration patterns of Phorrnidium corallyticum and Beggiatoa spp. associated with black-band disease of corals

An in situ field study of the motility patterns exhibited by Phormidium corallyticum and Beggiatoa spp. in black-band disease of corals was conducted over a 5-day period. Measurements were made at a spatial resolution of 50 m to document the horizontal migration of black-band across living coral tissue, while vertical migrations within the band were documented by observation and macrophotography of the black-band surface. It was determined that horizontal migration occurred both day and night, with the fastest movements by the front of the band during the day and the back of the band at night. Beggiatoa would rise to the band surface at night, and would often remain above the cyanobacterial population during extended periods of illumination the following day. The migration patterns are discussed in terms of motility cues and microbial physiology.     

Habitat partitioning of native and exotic Daphnia in gradients of temperature and food: mesocosm experiments

1. Invasion of tropical zooplankton into temperate lakes provides an interesting opportunity to explore habitat segregation in a thermal gradient. 2. We explored differing vertical positioning of native and exotic Daphnia (Daphnia mendotae and Daphnia lumholtzi) in a large indoor mesocosm system (Plön plankton towers) during 2 month‐long experiments. The two towers were manipulated to provide gradients of both temperature (15–29 °C) and algal food (0.05–0.58 mg C L^?1) and a day–night cycle. 3. Both juvenile and adult D. lumholtzi showed a ‘typical’ vertical migration pattern, with higher densities in the epilimnion at night than during the day. They avoided the food‐poor middle layer. In contrast, D. mendotae adults showed little tendency to migrate into the epilimnion at night, remaining in the cooler hypolimnion while juveniles migrated. The vertical distribution of D. mendotae appeared unaffected by the presence of D. lumholtzi. 4. The strong migration behaviour of D. lumholtzi in the absence of fish cues suggests that this behaviour may be a constitutive trait. Habitat partitioning of the two species is probably the result of different thermal tolerances, with D. mendotae constrained to remaining in deeper water by high temperatures in the epilimnion and the tropical D. lumholtzi able to use the warm epilimnion at night.     

Diel locomotor activity of brook charr, as determined by radiotelemetry

Locations by radiotelemetry during eight 24–h sampling periods in 1991, 1992, and 1993 were used to evaluate the diel activity patterns of adult brook charr Salvelinus fontinalis in two lakes of the Laurentian Shield (Québec, Canada). Based on the minimum distance travelled between two consecutive locations, adult brook charr were more active at dusk and at night than during the day. During daylight periods, individual fish displayed strong site selection, remaining in the same area along the shore from day to day. The fish left their inshore position at dusk, were highly mobile during the night, and returned to their focal point at sunrise, exhibiting diel homing behaviour. These results are supported by an increase in gillnet captures of adult brook charr in Lac Melchior at night. Data obtained by gillnet fishing in lakes Bondi and Simpson showed that juvenile (1+) brook charr were active throughout the diel cycle and that young-of-the-year were active mainly during the day, suggesting that there is an ontogenetic change in the diel activity of brook charr. Despite its limitations (restrictions on the number and size of tracked individuals, post-manipulation mortality, lifetime of transmitter battery), radiotelemetry allows for collection of multiple behavioural observations (distances travelled, instantaneous activity, site selection and homing) during the activity cycle of individual fish, and thus provides a finer degree of spatial resolution than alternative methods such as gillnetting.     

To what extent do environmental factors affect the long-distance nocturnal post-fledging movements of the Reed Warbler?

We studied the effects of weather and the lunar cycle on long-distance nocturnal pre-migratory flights of Reed Warblers (Acrocephalus scirpaceus). Noturnal tape luring was used to capture the birds, and the study was carried out in a habitat atypical of this species on the Courish Spit (southeastern Baltic) between1999 and 2002. A total of 443 juvenile Reed Warblers were captured during 120 nights of trapping. Based on data on the moult and body condition of the birds, it was possible to identify 163 individuals as being on post-fledging movements. More than half of the birds (54.0%) were captured at the end of night during the nautical and civil twilight period; the remaining individuals were caught during the astronomical twilight period and darkest periods of night. Many birds performed pre-migratory flights in the middle of the night, the period during which a large part of the moon was visible. We suggest that the increased visibility under the full moon may provide conditions in which Reed Warblers increase the distance or intensity of nocturnal post-fledging movements. During the entire course of the night, the birds generally preferred to fly under rainless conditions, limited cloud cover and/or in still air. The birds performed flights under winds stronger than 2 m s⁻¹ when the wind was blowing along the axis of the spit. We also suggest that the birds generally fly along the spit. We found a weak but significant relationship between the numbers of Reed Warblers captured during post-fledging movements and the individual weather parameters and their interaction. Our data suggest that endogenous stimuli rather than weather parameters or lunar cycle phase determine the decision of Reed Warblers to undertake pre-migratory long-distance flights at night.