Costs and benefits of a cleaning mutualism

Juvenile Atlantic salmon or parr ( Salmo salar L.) maintain station at certain locations in flowing stream water. This position choice is assumed to involve the maximization of energy intake, based upon food availability which is usually directly related to water flow rate. Conversely energy expenditure, including station holding behaviour, foraging and defending preferred sites, is inversely related to water flow rate. Adaptations of parr to life in fast flows implies that station holding is energetically inexpensive at water speeds up to the maximum sustained holding speed, which is fish specific, thus the most important energetic consideration for parr is the ability to maximize food intake. Ten groups of three parr were each observed for 60 min within an artificial stream tank over a heterogeneous substratum. Individual position choice and behaviour were recorded continuously. For each location chosen by the parr the potential upstream line‐of‐sight (LOS), defined as the maximum distance upstream that the water surface would be visible, was calculated. At those sites where foraging behaviour was observed, the mean upstream potential LOS was significantly greater than at sites where other behaviours were observed and at 400 randomly generated sites within the tank. When foraging, parr usually take food from the stream drift and there is a significant time expenditure on food location, identification and catching. Results presented here would seem to confirm that to maximize time available to make these decisions, a fish would be expected to maximize the distance over which it can observe potential food particles.     

Effects of location of food delivery and social status on foraging-site selection by juvenile Atlantic salmon

Synopsis Experiments were conducted to investigate the response of juvenile Atlantic salmon,Salmo salar, to changes in the location at which food entered a stream tank. Experience with the location of food input into the system significantly influenced foraging-site selection by the dominant fish. Dominant fish changed location of their foraging site in response to a change in the location of food input, and occupied, aggressively defending, sites just downstream of the location of food introduction. In contrast, subordinate fish occupied foraging sites at the downstream end of the stream tank, regardless of location of food input. As a result of both site selection and social status, dominant fish captured significantly more prey than subordinates. Our results support the contention that salmonids choose foraging sites to maximize foraging opportunities. Our results also provide a possible explanation for the use of atypical foraging sites by individual fish within their home range over the course of a single day, as observed in a number of salmonid species in the field.     

Interactions between riparian shading and food supply: a seasonal comparison of effects on time budgets, space use and growth in Atlantic salmon Salmo salar

This study examines seasonal (winter v. summer) differences in space-time budgets, food intake and growth of Atlantic salmon Salmo salar parr in a controlled, large-scale stream environment, to examine the direction and magnitude of shifts in behaviour patterns as influenced by the availability of overhead cover and food supply. Salmo salar parr tested in the presence of overhead cover were significantly more nocturnal and occupied more peripheral positions than those tested in the absence of overhead cover. This increase in nocturnal activity was driven primarily by increased activity at night, accompanied by a reduction in daytime activity during winter. The presence of overhead cover had no effect on rates of food intake or growth for a given food supply in a given season. Growth rates were significantly higher for fish subjected to a high food supply than those subjected to a low food supply. Food supply did not affect the extent to which S. salar parr were nocturnal. These results were consistent between winter and summer. The use of riparian shading as a management technique to mitigate the effects of warming allows the adoption of more risk-averse foraging behaviour and may be particularly beneficial in circumstances where it serves also to increase the availability of food.     

Social Status and Feeding in Atlantic Salmon Salmo salar Parr: The Effect of Visual Exposure to a Dominant

Groups of four Atlantic salmon parr were established and the dominant individual identified. Feeding responses were then observed in the remaining (subordinate) fish housed in one of three conditions: in the same compartment as the dominant fish (full exposure), separated from it by a transparent partition (visual exposure) or behind an opaque partition (isolated). Subordinate parr in the full exposure condition responded to and ate fewer food pellets than did the dominant fish. However, subordinate parr in the visual exposure condition and in the isolation condition fed actively, indicating that neither visual intimidation nor long-term stress-induced reduction of appetite were responsible for the reduced food intake of subordinate parr in direct contact with dominants. Dominant parr and subordinates in the visual exposure condition, but not subordinates in the exposed condition, moved further from their feeding station when attacking food items that were contested by a competitor compared with the distance moved for uncontested items. The presence of a competitor therefore forced both dominant and visually exposed subordinates to pay higher costs while foraging.     

Why do Foraging Stream Salmonids Move During Summer?

We hypothesize that foraging stream salmonids move during summer because (1) they monitor habitat conditions at a reach scale (100s of m), and (2) dominant fish move when conditions in their present foraging location become sub-optimal relative to conditions at other locations in the reach. To test these ideas, we quantified temporal variation in foraging habitat quality between late spring and early fall in a reach of a small Rocky Mountain brook charr, Salvelinus fontinalis, stream, predicted optimal-foraging fish distributions within the reach, and experimentally manipulated access to foraging sites and measured fish responses. Our results show that high-quality foraging sites were located at certain places in the reach during one period, but at different places during others, consistent with the hypothesis that fish movement is required if dominant fish are to occupy high-quality foraging sites throughout summer. The optimal foraging model was able to predict foraging locations within study pools, but not the exact location of individual fish within the pools or the reach. However, empirical evidence suggests that fish were distributed in order to maximize energy intake at the reach scale. Finally, dominant fish excluded from their preferred foraging location either left the pools (three of six cases), or began to occupy focal points of the next largest fish which, in turn, exited the pool (two of six cases). If habitat selection was occurring only within habitat units, then large fish, when excluded from their preferred locations, would select the next best locations within the pool. Taken together, these results suggest that charr use summertime movements to both monitor habitat conditions at a large spatial scale, and to gain access to optimal foraging locations even as conditions change temporally.     

Turbidity amplifies the non‐lethal effects of predation and affects the foraging success of characid fish shoals

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Short-term cues used by foraging trout in a California stream

Stream salmonids choose foraging locations to maximize the energy benefit of foraging within the constraints of size-mediated dominance hierarchies and predation risk. But, because stream habitats are temporally variable, fish must use a search process to monitor changing habitat conditions as a means of locating potentially-better foraging locations. I explored the cues used by the cutthroat trout, Oncorhynchus clarki clarki, when searching for food at the pool scale by artificially increasing prey availability at different locations by using special feeders and by manipulating pool velocities. Behavior of individually marked fish was monitored from stream bank platforms under unmanipulated control conditions and under seven experimental sets of conditions involving different combinations of feeder location and velocity manipulation. Under natural conditions fish elected to forage in the deepest (>50 cm), fastest (0.10–0.25 m s⁻¹) locations and within 1 m of structure cover, but would readily move to shallower (<30 cm) water away from cover if velocities were manipulated to be highest there. Although fish did not locate feeders unless they were placed in high-velocity areas, when high velocity was provided fish would move into very shallow water (<20 cm) if prey were delivered there. Responses of individual trout to manipulations indicated that water velocity was the main physical cue used by fish to decide where to forage, and that fish could also learn about new food sources by observing conspecifics. Overall, results indicated fish were not “perfect searchers” that could quickly locate new food resources over short time scales, even when the new resources were within a few meters of the fish’s normal foraging location. When given the correct cues, however, fish could detect new food sources and defend them against subordinate fish. Movement of new fish into and out of the study pools during the ten-day observation period was common, consistent with the idea that trout used movement as a means of exploring and learning about habitat conditions at the reach scale.     

Feeding ecology of wintering terns in Guinea-Bissau

We studied the feeding ecology of Little Terns Sterna albifrons , Sandwich Terns S. sandvicensis and Royal Terns S. maxima in the Archipélago dos Bijagós (11°40'N, 15°45'W) in Guinea-Bissau (West Africa) during the winter of 1992/1993. More than 95% of all prey taken by these terns were roundfish, ranging in weight from 0.3 to 40 g. Birds usually fed alone, but sometimes they were observed feeding in mixed-species flocks consisting of 15–200 individuals. Capture rate ( n fish per hour foraging) in these flocks was higher than that of solitary birds. However, smaller fish were caught by birds foraging in flocks, so food intake rate (g/h) did not differ between solitary and flock-feeding birds. The relationships between foraging behaviour of the three tern species and abiotic factors, such as time, tide and water clarity, have been investigated. Capture rate of Royal Terns increased with water clarity. For Little Terns and Sandwich Terns, food intake rate was lower in the most turbid waters compared to clearer waters. There was very little foraging activity during high tide. For Little Terns and Royal Terns, food intake rate was about twice as high during receding and low tides as during an incoming tide. Food intake rate averaged 8 g/h in Little Terns, 60 g/h in Sandwich Terns and 45 g/h in Royal Terns. With a rough model, we estimate the maximum rate of daily energy expenditure of terns wintering in the tropics at 3 × BMR (defined as energy expenditure of inactive bird at thermoneutrality in a post-absorptive state during the resting phase of the daily cycle). From an energetic viewpoint, wintering Sandwich Terns in Guinea-Bissau seem to have an easy living.     

Effects of food availability on temporal activity patterns and growth of Atlantic salmon

1. Patterns of sheltering and activity are of fundamental importance in the ecology of animals and in determining interactions among predators and prey. Balancing decreased mortality risk when sheltering with increased feeding rate when exposed is believed to be a key determinant of diel patterns of sheltering in many animals. 2. Despite lower foraging efficiency at night than during the day, Atlantic salmon Salmo salar parr are nocturnal during winter and at low summer temperatures. Nocturnal activity also occurs at warm water temperatures during summer, but little is known about the functional significance of this behaviour. 3. This study aimed to determine: (1) the preferred activity and shelter pattern of Atlantic salmon parr during warm summer months, and (2) their response to variations in food availability when balancing growth rate (G) and mortality risk (M), as expressed through time out of shelter. We differentiated among four potential responses to reduced food availability: (1) no response; (2) G decreases but M remains constant; (3) G remains constant but M increases; and (4) G decreases and M increases. 4. Time exposed from shelter was inversely related to food availability. Fish subject to high food availability were significantly less active during the day than those with restricted rations. However, food availability had no significant effect on the extent to which fish were active at night. There was no evidence of variation in growth rate with food availability. 5. Salmon were predominantly nocturnal at high ration levels, consistent with their previously reported behaviour during winter. Rather than switching to diurnal behaviour at high temperatures per se, as previously was supposed, it appears that the fish are diurnal only to the extent needed to sustain a growth rate, and this extent depends on food availability. 6. Atlantic salmon parr modulate the amount of time they are active rather than growth when responding to variations in food availability over an order of magnitude.     

Influence of sexual maturity on feeding, growth and energy stores of wild Atlantic salmon parr

Maturing Salmo salar parr had significantly lower short-term energy stores (indicated by hepato-somatic index) in August, and significantly lower energy stores and growth rate (indicated by RNA : DNA) in September than immature parr captured from the same stream sites on the same date. There were no significant differences in gut fullness or protein concentrations, suggesting that up to early September the main energetic consequences of maturation were a reduction in allocations to growth and short-term energy storage, but not a mobilization of long-term stored energy in the form of proteins. These are the first observations of relative food intake and energy storage for maturing parr under natural conditions, and also the first to assess growth effects on wild fish before completion of the maturation process.     

Effects of prey abundance on the foraging behaviour, diving efficiency and time allocation of breeding Guillemots Uria aalge

The foraging behaviour of Guillemots Uria aalge at sea was compared between 2 years of radically different food abundance. Radio telemetry was used to determine foraging locations and diving patterns. In the poor compared with the good food year, foraging trips were much longer, the birds foraged more than six times further from their breeding sites, they spent over five times as much time diving when at sea and their estimated energy expenditure was twice as great. Time spent foraging in the poor food year was at the expense of time spent sitting at the colony. The duration of a foraging trip was a poor indicator of distance travelled but a good indicator of the amount of time spent diving. Mean dive durations, surface pause durations and interbout periods did not differ between years, but individuals made more than four times as many dives per diving bout in the poor food year. Surface pause lengths did not vary with water depth in either year. In the poor food year, birds made shorter surface pauses for a dive of a gi^ven duration than in the good food year, possibly accepting a lactic acid debt in order to maximize searching time, The duration of the interbout period was positively related to the number of dives in the previous bout, and dives tended to get shorter in long diving sequences, suggesting possible exhaustion effects. These data demonstrate that breeding Guillemots have the capacity to adjust their foraging behaviour and time budgets in response to changes in food abundance, but this flexibility was not sufficient to compensate fully for the very low food abundance experienced by birds in this study.     

The Importance of Egg Size and Social Effects for Behaviour of Arctic Charr Juveniles

Early behaviour can determine food intake and growth rate with important consequences for life history and survival in fishes. Egg size is known to affect growth rate of young Arctic charr but its influence on the development of behaviour is poorly documented. It is believed that egg size influence on growth and potentially on the behaviour of young fish decreases over time, minimized by the effects of social factors. Shortly after first feeding, we examined differences in mobility and foraging of Arctic charr in relation to egg size and social environment. The behaviour of juveniles from small and large eggs was compared five times over the course of development and in three different experimental settings: long‐term isolation (isolation before hatching), short‐term isolation vs. group rearing and mixed size group vs. homogeneous size groups. Egg size affected foraging behaviour and mobility of fish: fish coming from large eggs were more mobile and foraged more than fish coming from small eggs. Social environment affected foraging behaviour, mobility and space use: fish in a group were more mobile, foraged more and responded faster to food delivery than isolated fish. The interaction of egg size and social effects was seen primarily in foraging activities but did not affect mobility or space use. Large fish in groups foraged more than the three other groups: large fish in isolation, small fish in groups and small fish in isolation. Agonistic behaviour was rarely observed and there was no significant effect of group composition on agonistic behaviour. We discuss the importance of egg size and social effects at early stages of development with a focus on the evolutionary ecology of Arctic charr.     

Redistribution of juvenile salmonid fishes after localized catastrophic depletion

Juvenile Atlantic salmon and brown trout were depleted at three sites ( c . 108–380 m²) of a natural stream during the summer months of 1991 and 1992. Local population changes and movements of fish marked in sections adjacent to each depleted area were monitored thereafter. There was very little movement of marked salmon parr into the central regions of the depleted areas following the immediate post-marking period. Upstream movement by young-of-the-year fish from high density sections in mid-late summer was noted for trout but not salmon. Unmarked 1-year-old salmon parr immigrated into depleted areas in June 1992, and the pattern of recolonization was consistent with migration upstream from the adjoining river. It is concluded that resident salmon were very strongly site-attached and resource tracking was of no functional significance as a compensatory mortality mechanism. The occurrence of a long distance migratory component in the population during early-mid summer indicates that this, rather than local resource tracking, constitutes a potential compensatory mechanism.     

Feeding motivation and response to predation risk in Atlantic salmon parr adopting different life history strategies

Feeding intensity was measured before (baseline level) and after (disturbed level) brief exposure to a potential predator in groups of 0+ Atlantic salmon, Salmo salar. parr destined to follow either a fast-growth, early-smolting life history (upper modal group or UMG fish) or a slow-growth, late-smolting life history (lower modal group or LMG fish). Feeding intensity decreased following exposure to the predator. While the absolute decrease in feeding intensity is constant regardless of baseline level, the proportionate decrease is negatively related to pre-presentation feeding intensity. Parr that are strongly motivated to feed thus maintain a higher food intake and incur greater risks when foraging in the presence of a predator. No differences in baseline or disturbed feeding intensity of UMG and LMG parr were found up to September of their first year. Previous work has shown that baseline feeding intensities decrease in LMG fish after September but increase in UMG fish. In the present study, these differences were reflected in greater disturbed feeding intensities in UMG fish, but fish destined to follow different life history patterns do not differ in risk-taking, once these differences in baseline feeding motivation have been taken into account.     

Axes of fear for stream fish: water depth and distance to cover

To better understand habitat-specific predation risk for stream fish, we used an approach that assumes animals trade off food for safety and accurately assess risk such that predation risk can be measured as a foraging cost: animals demand greater harvest rates to occupy riskier locations. We measured the foraging cost of predation risk for juvenile salmonids within enclosures in a natural stream at locations that varied in water depth and distance to cover. Measurements relied on a food delivery apparatus and direct observations that allowed estimation of “giving-up” harvest rates – food delivery rates at which animals left the feeding apparatus. Juvenile steelhead about 120 mm fork length exhibited sharp increases in giving-up harvest rate with decreasing water depth and refused to use the feeding device even when offered extreme food delivery rates in water ≤20 cm deep. Giving-up harvest rates were less affected by the distance to cover. Assuming the gradients we observed in giving-up harvest rates reflect predation risk, the results of this study can be applied to spatially explicit models of stream fish populations that incorporate risk into both habitat selection and mortality due to predation.     

Intraspecific variation in competitive ability and food intake in salmonids: consequences for energy budgets and growth rates

The interactions between dominance status, feeding rate and growth in rainbow trout, Salmo gairdneri Richardson, were analyzed using published data on experimental populations. There was a positive correlation between metabolic expenditure and food intake in both dominant and subordinate fish, but dominants obtained a greater intake for a given expenditure than did subordinates. Subordinates that adopted a high–return/high–cost foraging strategy actually expended more energy than they acquired, whereas those that minimized energy expenditure obtained a net energy gain. This led to the surprising finding that the growth rate of subordinates was negatively correlated with food intake.     

Disruption of seasonality in growth hormone-transgenic coho salmon (Oncorhynchus kisutch) and the role of cholecystokinin in seasonal feeding behavior

Seasonal variation in daily food intake is a well-documented phenomenon in many organisms including wild-type coho salmon where the appetite is noticeably reduced during periods of decreased day length and low water temperature. This reduction may in part be explained by altered production of cholecystokinin (CCK) and growth hormone (GH). CCK is a hormone produced in the brain and gut that mediates a feeling of satiety and thus has an inhibitory effect on food intake and foraging behaviour. Growth hormone (GH) enhances feeding behaviour and consequently growth, but its production is reduced during winter. The objectives of this study were: first, to compare the seasonal feeding behaviour of wild and GH-transgenic coho salmon; second, to determine the behavioural effect of blocking the action of CCK (by using devazepide) on the seasonal food intake; and third, to measure CCK expression in brain and gut tissues between the two genotypes across seasons. We found that, in contrast to wild salmon, food intake in transgenic salmon was not reduced during winter indicating that seasonal control of appetite regulation has been disrupted by constitutive production of GH in transgenic animals. Blocking of CCK increased food intake in both genotypes in all seasons. The increase was stronger in wild genotypes than transgenic fish; however blocking CCK in wild-type fish in winter did not elevate appetites to levels observed in the summer. The response to devazepide was generally faster in transgenic than in wild salmon with more rapid effects observed during summer than during winter, possibly due to a higher temperature in summer. Overall, a seasonal effect on CCK mRNA levels was observed in telencephalon with levels during winter being higher compared to the summer in wild fish, but with no seasonal effect in transgenic fish. No differences in seasonal CCK expression were found in hypothalamus. Higher levels of CCK were detected in the gut of both genotypes in winter compared to summer. Thus, CCK appears to mediate food intake among seasons in both wild-type and GH-transgenic salmon, and an altered CCK regulation may be responsible at least in part for the seasonal regulation of food intake.     

Annual rhythms of swimming behaviour and seawater adaptation in young baltic salmon,Salmo salar,associated with smolting

Synopsis Sexually immature and sexually mature precocious male Baltic salmon,Salmo salar, parr from Umeälven (Ume river) were tested for rheotactic behaviour and adaptation to seawater before, during, and after the time period for smolt migration. Size of fish at the beginning of the experiment in January was on average 13.5 cm. Rheotactic behaviour was tested in annular stream tanks with photocells to measure upstream and downstream movements. Samples of fish were given a Seawater challenge test at monthly intervals in order to determine their ability to adapt to 20%. saltwater. During spring, both immature and sexually precocious parr became silvery and showed progressive development of downstream-directed movements. In early June the fish exhibited good hypoosmoregulatory ability in 20%. saltwater and swimming was predominantly downstream. During late June and early July there was a marked reversal in swimming behaviour, accompanied by a dramatic change in saltwater adaptation. The fish moved mainly upstream and showed decreasing ability to meet the seawater challenge test. This was accompanied by a loss of silvery coloration. The annual cycle of swimming behaviour and seawater adaptation is discussed in relation to the appearance of a smolt-window, i.e., a critical interval for smolt migration.     

Impact of Beaver Dams on Abundance and Distribution of Anadromous Salmonids in Two Lowland Streams in Lithuania

European beaver dams impeded movements of anadromous salmonids as it was established by fishing survey, fish tagging and redd counts in two lowland streams in Lithuania. Significant differences in abundancies of other litophilic fish species and evenness of representation by species in the community were detected upstream and downstream of the beaver dams. Sea trout parr marked with RFID tags passed through several successive beaver dams in upstream direction, but no tagged fish were detected above the uppermost dam. Increase in abundances of salmonid parr in the stream between the beaver dams and decrease below the dams were recorded in November, at the time of spawning of Atlantic salmon and sea trout, but no significant changes were detected in the sections upstream of the dams. After construction of several additional beaver dams in the downstream sections of the studied streams, abundance of Atlantic salmon parr downstream of the dams decreased considerably in comparison with that estimated before construction.     

Foraging Rules for Group Feeders: Area Copying Depends upon Food Density in Shoaling Goldfish

The decision rules governing forage area copying behaviour were investigated in shoaling fish. Shoaling goldfish were offered two equal food patches, one of which was adjacent to an equal-sized shoal feeding behind a transparent barrier. When food was low, goldfish foraged according to an area copying rule, but under high and zero food area copying disappeared. Only under high food density did equal numbers of fish feed at both sites as predicted by foraging theory. Under zero food the fish were less certain about where to forage. Precise visual cues from feeding fish were required: non-feeders did not attract area copiers. Furthermore, area copying was task-dependent since it reappeared strongly if fish were not able to forage on patches like their fellows. Control experiments eliminated an increase in group size for anti-predator advantage as an explanation. Two sequential decisions: to stay or move, and to join or leave may explain the results, which are not accommodated by simple optimality models. These decisions may be based on a comparison of current food intake with the anticipation of a higher reward by foraging socially.