Indirect effects of fish on foraging behaviour and leaf processing by the isopod Lirceus fontinalis

1. We examined whether the isopod, Lirceus fontinalis, an important facultative shredder in low-order streams in the eastern United States, responded to chemical cues of the skin mucus of five fish species varying in relatedness and feeding habits, and if fish-induced alterations in the foraging activity of isopods indirectly affected leaf processing. 2. In the laboratory, isopods generally responded to the presence of fish mucus by reducing their activity. Isopods were most responsive in water containing the skin mucus of predatory sunfish (Lepomis) and least responsive in water treated with mucus from algivorous stonerollers (Campostoma anomalum). 3. Rates of leaf disc breakdown by isopods in the laboratory were significantly reduced when isopods were exposed to chemical cues of confined green sunfish (L. cyanellus). 4. The rate of leaf processing in a fishless reach of a headwater stream was four to five times greater by isopods 7—10mmTL (total length) than those ≤ 5mm TL. Rates of leaf processing by isopods in low-order streams may be affected directly by the size structure of the isopod population and indirectly by the presence of fish.     

Effect of predatory fish presence on habitat use and diel movement of the stream amphipod, Gammarus minus

SUMMARY.

• 1 We examined how the presence of predatory fish affected macro—and microhabitat use and movement patterns of the amphipod, Gammarus minus Say, in a second order stream.
• 2 Among macrohabitats, amphipods were consistently more abundant in runs than in pools. Densities in pools were low regardless of fish presence, whereas low densities in runs were correlated with fish presence. Siltation and the lack of coarse substrate particles probably accounted for the scarcity of G. minus in pools. Among microhabitats, G. minus was more abundant in leaf litter and gravel than in silt/sand substrates. Fish presence did not affect microhabitat use in the field. Laboratory manipulations showed, however, that predation rates by green sunfish (Lepomis cyanellus Rafinesque) were significantly lower on amphipods in leaf litter than in other microhabitats.
• 3 Most movement by G. minus in runs occurred at night and was directed downstream. In the laboratory, amphipods significantly reduced swimming activity at night in response to water conditioned with green sunfish. In the field, however, amphipods were only slightly less active in runs at night when exposed to chemical cues of confined green sunfish. The distribution of amphipods in streams is influenced by the presence of fish, but chemical cues alone may not be important in triggering predator avoidance behaviours in nature.

     

Behavioural correlations across situations and the evolution of antipredator behaviour in a sunfish-salamander system

Behavioural correlations are at the heart of understanding how conflicting demands shape the evolution of ecologically important behaviours. Many studies have focused on the effects of negative behavioural correlations generated by time budget conflicts within situations. We examined an alternative possibility that involves positive behavioural correlations due to behavioural carryovers across situations. Specifically, we examined the role of behavioural carryovers in governing antipredator responses of streamside salamander larvae, Ambystoma barbouri, to predatory green sunfish, Lepomis cyanellus. Earlier work showed that these larvae suffer heavy sunfish predation due to high larval exposure to fish (high proportion of time spent out of refuge). Earlier work also showed that paradoxically, despite selection pressure from fish, these larvae show higher exposure in the presence of fish (poorer antipredator behaviour) than a sister species that inhabits fishless, ephemeral ponds. The standard time budget trade-off between feeding and antipredator behaviour does not appear to explain the observed antipredator behaviours. Instead, the present study shows that the relatively large proportion of time that larvae spend out of refuge (exposed) in fish pools in the daytime can be explained in part by behavioural correlations across situations. Specifically, larvae showed positive correlations among individuals in their daytime exposure in fish pools, nighttime exposure in fish pools, and exposure in fishless pools. The benefits of high exposure in fishless conditions (associated with high feeding and developmental rates) and high exposure in fish pools at night (necessary to drift out of fish pools) have apparently overridden the predation cost of being exposed in fish pools in the day. Behavioural correlations across situations might often result in ecologically important behaviours that appear maladaptive in an isolated context. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Experimental studies on direct and indirect interactions in a three trophic-level stream system

Summary We used a complete block design to experimentally study direct and indirect interactions in a three trophic-level freshwater system consisting of a top predator, the green sunfish, Lepomis cyanellus, an intermediate predator, small-mouthed salamander larvae, Ambystoma barbouri, and prey, hatchling isopods, Lirceus fontinalis. This system occurs naturally in small stream pools in central Kentucky; experiments were done in laboratory pools. Salamander larvae ate isopods and thus had a direct, negative effect on isopod survival. Accordingly, isopods responded to the presence of salamander larvae by increasing their tendency to bury themselves in the sand substrate. Fish ate salamanders and thus had a direct, negative effect on salamander survival. Salamanders responded to fish presence by increasing their time spent under plexiglass plates that simulate refuge rocks. The overall effect of fish on isopods depended on the presence of salamanders. In the absence of salamanders, fish predation on isopods had a direct, negative effect on isopod survival; isopods thus responded to the presence of fish by burying themselves in the sand. With salamanders present, fish had a positive overall effect on isopod survival; i.e., direct, negative effects of fish on isopods were outweighed by indirect, positive effects. Indirect positive effects of fish on isopods came through a reduction in salamander predation rates on isopods in the presence of fish. The mechanism involved both a decrease in the number of salamanders (a trophic-linkage indirect effect; cf. Miller and Kerfoot 1987) and a reduction in the feeding rate of individual salamanders on isopods (a behavioral indirect effect). The decrease in individual salamander feeding rates on isopods was due to reductions in both salamander activity and in spatial overlap between salamanders and isopods in the presence of fish. The latter effect reflected the fact that salamanders and isopods used different refuges from fish; salamanders went under refuge plates, whereas isopods primarily buried themselves in sand. Estimates of the relative importance of various direct and indirect effects of sunfish on isopods suggested that positive, behavioral indirect effects were of roughly the same magnitude as direct, negative effects, both of which were more important than were trophic-linkage indirect effects. Contrary to expectations, the presence of isopods did not affect the refuge use or survival of salamanders in the presence of fish.     

Effects of Predation Risk on Habitat Selection by Water Column Fish, Benthic Fish and Crayfish in Stream Pools

Predation risk can affect habitat selection by water column stream fish and crayfish, but little is known regarding effects of predation risk on habitat selection by benthic fish or assemblages of fish and crayfish. I used comparative studies and manipulative field experiments to determine whether, (1) habitat selection by stream fish and crayfish is affected by predation risk, and (2) benthic fish, water column fish, and crayfish differ in their habitat selection and response to predation risk. Snorkeling was used to observe fish and crayfish in, (1) unmanipulated stream pools with and without large smallmouth bass predators (Micropterus dolomieui >200 mm total length, TL) and (2) manipulated stream pools before and after addition of a single large smallmouth bass, to determine if prey size and presence of large fish predators affected habitat selection. Observations of microhabitat use were compared with microhabitat availability to determine microhabitat selection. Small fish (60–100 mm TL, except darters that were 30–100 mm TL) and crayfish (40–100 mm rostrum to telson length; TL) had significantly reduced densities in pools with large bass, whereas densities of large fish and crayfish (> 100 mm TL) did not differ significantly between pools with and without large bass. Small orangethroat darters (Etheostoma spectabile), northern crayfish (Orconectes virilis), and creek chubs (Semotilus atromaculatus) showed significantly greater densities in pools without large bass. The presence of large smallmouth bass did not significantly affect depths selected by fish and crayfish, except minnows, which were found significantly more often at medium depths when bass were present. Small minnows and large and small crayfish showed the greatest response to additions of bass to stream pools by moving away from bass locations and into shallow water. Small darters and sunfish showed an intermediate response, whereas large minnows showed no significant response to bass additions. Response to predation risk was dependent on prey size and species, with preferred prey, crayfish and small minnows, showing the greatest response. Small benthic fish, such as darters, are intermediate between small water column fish and crayfish and large water column fish in their risk of predation from large smallmouth bass.     

Diet and feeding habits of Economidichthys pygmaeus (Perciformes: Gobiidae) in Lake Pamvotis,NW Greece

The purpose of the study was to define the diet composition, feeding preferences, seasonal, size and sex related changes in diet, feeding strategy and diel cycle of Economidichthys pygmaeus. Important features of the overall feeding patterns of the goby include (i) feeding activity, (ii) bottom habits and (iii) consumption of food items to identify the dietary breadth. Stomach contents of 533 specimens, 13.42–48.65 mm total length (TL), collected by hand net (2 mm mesh size) from October 2006 to September 2007 were analyzed. The percentage of 76 empty stomachs (14.26%) did not vary significantly with season. The food composition suggested that the goby is a carnivorous fish. Prey items identified in the stomachs belonged to four major groups: Crustaceans, Insects, Mollusca and Plants. Copepods were the most important (%IRI = 57.51), especially in fish larger than 23 mm TL. Chironomids (%IRI = 43.23) constituted the main prey for fish <23 mm TL. Diet composition showed little seasonal variation. Copepoda dominated the diet in autumn and winter (43.4%; 82.4%) and were replaced by Chironomidae larvae in spring (39.7%) and summer (47.7%). During the reproductive season, large males show a narrow food spectrum as opposed to large females, reflecting the different activity pattern exhibited by males and females during the breeding season, since the former supports parental care. According to the modified Costello graphical method, specialist individuals form the goby population that feeds on two preferential prey types (e.g. Copepods, Chironomids). Nevertheless, they consume some occasional prey (e.g. Cladocera, Insects, Gastropods). The species foraged chiefly at night and early morning, during lower light intensity. The highest feeding activity recorded at night (02.00 hours) and during early morning (08.00 hours) could be related to the cryptic behaviour displayed by the species.     

Fishing spiders, green sunfish, and a stream-dwelling water strider: male-female conflict and prey responses to single versus multiple predator environments

Many studies have experimentally addressed the effects of a particular predator species on prey behavior. In nature, however, prey frequently face multiple species of predators that often vary in their predatory mode and in their level of predation risk. Relatively few studies have considered prey responses under these complex conditions. In Kentucky, the stream-dwelling water strider (Aquariusremigis) coexists with many potentially dangerous predators, two of which are the green sunfish (Lepomiscyanellus) and the fishing spider (Dolomedesvittatus). Green sunfish occupy stream pools and attack water striders from below. In contrast, fishing spiders hunt along stream shorelines where they perch on overhanging vegetation or rocks and attack water striders near shore. We compared how A. remigis individuals respond to these two very different predators in pools with one or both predators. The presence of sunfish in pools had strong effects on male water strider behavior, including increased use of three types of refuge from sunfish (riffles, climbing out of the water, sitting on the water but at the edges of pools), decreased activity and a decreased number of aggressive males on the water. Spiders also influenced water strider behavior; male water striders avoided spiders by shifting away from the edges of pools. Comparisons of the effects of the two predator species showed that in general, antipredator responses by male water striders were stronger in pools with fish alone than in those with spiders alone. In the presence of both predators, male water strider behavior (microhabitat use and activity) was generally similar to behavior in the presence of fish alone. In contrast, female water striders showed no significant response to the presence of sunfish, and little response to the presence of spiders. This lack of response could be because females spent much of their time in refuges even in the absence of predators (apparently hiding from harassment by males). Both spiders and fish caused decreases in water strider mating activity. The presence of fish reduced both the number of matings per pool (mating frequency), and mean mating durations. Spiders induced a decrease in mean mating duration, but not in mating frequency. The largest reductions in mating activity occurred in pools with both predators present. Pools with either spiders or fish alone suffered 15–20% water strider mortality during our experiment (versus no mortality in predator-free pools). Extant theory suggests that when prey face conflicting microhabitat responses to two predators (as in this study), the predators should have facilitative effects on predation rates (i.e., prey that avoid one predator are often killed by the other and vice versa). Mortality rates in pools with both predators present, however, were not significantly different from that predicted by a null model of multiple predator effects. The lack of predator facilitation can be explained by the compensatory reductions in water strider activity and mating activity in the presence of both predators. Received: 26 August 1996 / Accepted: 12 June 1998     

Nocturnal increases in the use of near-surface water by pond animals

We assessed diel animal habitat use in three shallow ponds, using unbaited funnel traps, a large column sampler, and sweep net collections in the upper stratum (0–0.3 m) of littoral and open habitats. In all three ponds, more animals were caught at night than during the day, indicating that use of near-surface waters was greatest at night, particularly in the fishless ponds. All methods yielded similar patterns. Our results demonstrate that nocturnal observations of pond animals are necessary to describe their ecology, even in fishless ponds where diel differences in habitat use or behavior might not be anticipated.     

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.     

Diel infestation dynamics of gnathiid isopod larvae parasitic on Caribbean reef fish

Infestation dynamics of parasitic gnathiid isopods on Caribbean reefs were studied throughout the 24-h diel cycle. Gnathiid infestation on caged longfin damselfish (Stegastes diencaeus) peaked strongly at dawn, remained low during the remainder of the day, and increased again at night until about midnight. Gnathiids were less abundant during the pre-dawn period. Peak loads on fish retrieved at dawn were the highest reported in any study thus far. The dawn peak consisted almost exclusively of individuals from the smallest size class, whereas nocturnal activity consisted almost exclusively of individuals of the largest size class. Because of the high rates of infestation at night and dawn, and the high variation in parasite loads on fish collected during that time, reduction of parasite infestation may play an important role in the selection of nocturnal and crepuscular shelter holes and settlement sites by reef fishes.     

Diel variations in the assemblage structure and foraging ecology of larval and 0+ year juvenile fishes in a man‐made floodplain waterbody

This study investigated diel variations in zooplankton composition and abundance, and the species composition, density, size structure, feeding activity, diet composition and prey selection of larval and 0+ year juvenile fishes in the littoral of a man‐made floodplain waterbody over five 24 h periods within a 57 day period. There was a significant difference in the species composition of diurnal and nocturnal catches, with most species consistently peaking in abundance either during daylight or at night, reflecting their main activity period. There were no consistent diel patterns in assemblage structure or the abundance of some species, however, most likely, respectively, due to the phenology of fish hatching and ontogenetic shifts in diel behaviour or habitat use. There were few clear diel patterns in the diet composition or prey selection of larval and 0+ year juvenile roach Rutilus rutilus and perch Perca fluviatilis, with most taxa consistently selected or avoided irrespective of the time of day or night, and no obvious shift between planktonic and benthic food sources, but dietary overlap suggested that interspecific interactions were probably strongest at night. It is essential that sampling programmes account for the diel ecology of the target species, as diurnal surveys alone could produce inaccurate assessments of resource use. The relative lack of consistent diel patterns in this study suggests that multiple 24 h surveys are required in late spring and early summer to provide accurate assessments of 0+ year fish assemblage structure and foraging ecology.     

Predator-dependent diel migration by <Emphasis Type="Italic">Halocaridina rubra</Emphasis> shrimp (Malacostraca: Atyidae) in Hawaiian anchialine pools

Diel migration is a common predator avoidance mechanism commonly found in temperate water bodies and increasingly in tropical systems. Previous research with only single day and night samples suggested that the endemic shrimp, Halocaridina rubra, may exhibit diel migration in Hawaiian anchialine pools to avoid predation by introduced mosquito fish, Gambusia affinis, and perhaps reverse migration to avoid the predatory invasive Tahitian prawn, Macrobrachium lar. To examine this phenomenon in greater detail, we conducted a diel study of H. rubra relative abundance and size at 2-h intervals in three anchialine pools that varied in predation regime on the Kona-Kohala Coast of Hawai‘i Island. We found two distinct patterns of diel migration. In two pools dominated by visually feeding G. affinis, the abundance of H. rubra present on the pool bottom or swimming in the water column was very low during the day, increased markedly at sunset and remained high until dawn. In contrast, in a pool dominated by the nocturnal predator M. lar, H. rubra density was significantly lower during the night than during the day (i.e., a pattern opposite to that of shrimp in pools containing fish). In addition, we observed that the mean body size of the shrimp populations varied among pools depending upon predator type and abundance, but did not vary between day and night in any pools. Our results are consistent with the hypothesis that H. rubra diel migratory behavior and size distributions are influenced by predation regime and suggest that diel migration may be a flexible strategy for predator avoidance in tropical pools where it may be a significant adaptive response of endemic species to introduced predators.     

Ontogeny of Diel Pattern of Stream-Margin Habitat Use by Emerging Brown Trout, Salmo Trutta, in Experimental Channels: Influence of Food and Predator Presence

Age-0 brown trout, Salmo trutta, inhabit shallow and slow-flowing habitats where they can easily maintain stationary swimming positions. However, recent results have shown that they use deeper and faster habitats during daylight than at night, suggesting the occurrence of a nocturnal movement toward stream-margin habitats. Experiments were conducted to describe precisely when this diel pattern of habitat use appears during ontogeny. In two indoor channels, free-embryo brown trout were deposited under the gravel. When emerging, alevins were free to choose between margin (2 cm deep, 0-2 cm s^-1) or deep habitat (12 cm, 2-4 cm s^-1), or to leave the channel (upstream or downstream). During the week of emergence, upstream and downstream catches, fish habitat use (deep habitat or margin), and fish behavior (resting or swimming) were measured by direct observations and trap counts. Three treatments were performed: (1) fish artificially fed on drifting invertebrates, (2) fish exposed to predators (bullhead, Cottus gobio), and (3) control channels (no food, no predator). In control and food channels, a diel pattern of habitat use was observed 1-2 days after the emergence started. Most fish rested in the margin at night, whereas they moved towards the deep habitat during daylight to hold stationary swimming positions. In the presence of bullhead, most trout were cryptic, and visible fish stood in the margin during both daylight and at night. The importance of predation risk and foraging behavior on the ontogeny of the diel pattern of habitat use is discussed. Results support the direct development without larva from free-embryo via alevin in brown trout.     

THE HABITATS AND HABITAT PREFERENCES OF CLARIAS GARIEPINUS (PISCES: CLARIIDAE) IN A CLEAR COASTAL LAKE (LAKE SIBAYA,SOUTH AFRICA)

SUMMARY

The habitat preferences of the sharptooth catfish, Clarias gariepinus, in a clear coastal lake, Lake Sibaya, are described. Larval and small juvenile catfish inhabit flotsam and the rootstocks of plants in very shallow marginal areas. Larger juveniles inhabit densely-vegetated marginal pools, and may venture into open areas at night. Adult catfish (over 350 mm TL) occupy offshore areas, where their density is highest in terrace and sheltered bay habitats, and least in slope and profundal habitats. Adult catfish undergo a diel migration into shallow water at night and into deeper water during the day which is more marked at low lake levels. The methods used to capture catfish, and the main habitats in Lake Sibaya, are also described.     

Diel variation in diet composition of a riverine fish community

Although diel food habit studies have been undertaken on a number of individual species, few studies have examined diel variation in the diets of fish communities. We examined the diel diet variation and feeding periodicity of a fish community in the Juniata River, Pennsylvania. Nine species, totalling 1,098 fish, were collected at 4-h intervals over a 24-h period in October 1989, in numbers sufficient to describe their diel variation in diet composition. Diel variation in diet composition was evident in all species, as no single prey taxon was dominant in the diet of any species during any 4-h interval. Ephemeropterans were the most important prey taxa for four species of centrarchids, whereas chironomids were the main prey of banded killifish, mimic shiners, and spotfin shiners. Algae was the major component in the diet of spottail shiners, whereas bluntnose minnows contained mostly detritus. Feeding activities of rock bass, redbreast sunfish, and pumpkinseed occurred at low levels throughout the day; peak feeding occurred from 2000 to 0400 hours. Food consumption of smallmouth bass increased throughout the day with peak consumption occurring at 2000 hours. Non-centrarchids fed little during daylight hours and showed peak activity at 2000–2400 h. Construction of a 24-hour diet from six 4-h interval estimates and feeding periodicity data provided a comprehensive representation of the diel feeding ecology of all species collected.     

Diel shifts in the structure and function of nearshore estuarine fish communities

Day–night shifts in the nearshore fish fauna of a temperate microtidal estuary were assessed using a holistic suite of structural and functional community attributes. Mean fish species richness and diversity (taxonomic distinctness) were higher at night across all regions of the estuary and seasons, concurring with the findings of numerous comparable studies reviewed worldwide, while the diel period in which mean abundance was higher varied among seasons. Likewise, species and functional guild compositions (the latter based on feeding modes and habitat use) both differed significantly between day and night, with the extent of the diel shift again varying seasonally. Daytime fish communities were characterized by higher abundances of Atherinidae, Sillaginidae and Mugilidae, while Gobiidae were far more abundant at night. Marked shifts in size composition were also evident, with smaller fishes (<100 mm total length, L_T) being more prevalent during the day and larger fishes (≥200 mm L_T) proportionally more abundant at night. The above diel shifts were feasibly related to a range of predator–prey interactions and feeding‐related movements, namely a nocturnal decrease in top‐order avian piscivory coupled with an increase in invertebrate prey availability, resulting in changes in the presence and catchability of certain fish species in shallow estuarine waters.     

Predator avoidance costs and habituation to fish chemicals by a stream isopod

1. We examined whether long-term exposure to chemical cues of predatory longear sunfish (Lepomis megalottis) affected growth and rates of leaf processing by the isopod, Lirceus fontinalis, an important facultative shredder in low-order streams in the eastern United States, and whether isopods habituated to sunfish chemicals. 2. Long-term (24 days) exposure to fish chemicals did not affect growth or the extent of leaf processing by isopods in the laboratory. Thus, chemical cues alone may not be important in triggering long-term predator avoidance behaviours that could potentially reduce fitness or affect community processes like Litter breakdown. 3. Isopods exposed to fish chemicals for 3, 9 and 15 days were significantly more active than individuals unexposed to fish chemicals, when transferred to another environment with fish chemicals. This result, coupled with results of the growth experiment, suggests that isopods habituate to fish chemical stimuli and that time to habituation is c. 3 days or less. Because fish chemicals can mislead prey about predator presence, and hence be a non-threatening stimulus, they are probably important in eliciting only short-term antipredatory behaviours by isopods.     

A shift in the habitat use pattern of a lentic galaxiid fish: an acute behavioural response to an introduced predator

Despite potentially reducing predation mortality, behavioural responses of native species to introduced predators may still have sub-lethal impacts. In video-recorded laboratory trials, we examined the effects of introduced brown trout, Salmo trutta, on the short-term behaviour of a threatened, lake-dwelling galaxiid fish and confirmed a suspected diel pattern in habitat use by this species. We found that Galaxias auratus followed a distinct diel pattern in the use of complex habitats and open water, which was significantly altered by the presence of brown trout. In trials without the introduced predator, G. auratus used complex habitats (rocks or macrophytes) during the day, and open water during the night. In trials with brown trout present, G. auratus spent significantly less time in open water and rarely ventured out of the macrophytes. However, when given the option of using only rocky substrate or open water, which is the more common situation in the lakes to which this galaxiid is endemic, the fish reduced the amount of time they spent in the open water during the night, but still spent more time in open water than when macrophytes were available. Spending the daylight hours amongst the cover of rocks or macrophytes is most likely an adaptation to reduce the risk of predation by visual predators, and the pattern of reduced use of open water habitats in the presence of brown trout is an acute response to the close proximity of the introduced predator. The difference in the nocturnal use of macrophytes and rocks when trout are present may be related to differences in feeding opportunities or success within these habitats.     

Effects of size and early sexual maturity on downstream migration during smolting in Baltic samon (Salmo salar)

The effects of size and previous sexual maturity on downstream migration in two-summer-old Baltic salmon were studied during their normal seaward migratory period in spring. We used an automatic PIT (Passive Integrated Transponder) tag monitoring system for individual recognition of tagged fish. Size affected the probability of migration among both previously immature and previously mature males while there was no such relationship among females. Nearly all females and 63.6% of the previously immature males migrated during the smolting period, while only 24.5% of the previously mature males did. In migrating fish the individual activity level increased from almost zero in May to about 450 antenna passages per day in mid June, coinciding with a pronounced increase in ambient water temperature. During the period of increasing activity in early June the fish changed from night activity to day activity and finally activity all day and night. The fish moved downstream in schools but no specific school size dominated. This paper demonstrates the strong effect of body size and previous sexual maturity on the probability of migration.     

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

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