Species segregation during predator evasion in cyprinid fish shoals

SUMMARY. 1. When a predator attacks a mixed-species group, individual prey arc faced with a conflict between retaining the advantages of large group size and the disadvantages of remaining among aliens. Experiments were designed to show how this conflict is resolved in fish shoals.
2. Simulated attacks with a model predator were made upon mixed and single species shoals of three cyprinid species under semi-natural conditions in a fluvarium.
3. Measurements of elective group size, neighbour identity and distance, and encounters within and between species revealed behaviour which sorted fish into species at the onset of predator attack.
4. A control experiment demonstrated the same effect for fish of matched size.
5. The value to the individual of segregative behaviour in response to predator threat is discussed.     

The social organization of free-ranging fish shoals

The phenotypic composition of social groups can affect the foraging efficiency and predation risk of their individual members. In shoaling fishes, active choice of shoal companions on the basis of phenotypic characters such as body length has been reported. In this study, we investigated the phenotypic composition of free-ranging, multi-species fish shoals in a temperate freshwater lake. We collected 21 entire shoals, each consisting of up to four species and comprising 3622 fish in total. Shoals were strongly assorted by species, body length, parasite prevalence and parasite load. Using a mark-recapture procedure involving 785 banded killifish ( Fundulus diaphanus ), we found no evidence for shoal fidelity or site fidelity, as individuals from different shoals mixed extensively after just 24 h and ranged all over the study site irrespective of initial capture locations. Killifish showed strong changes in shoal-size choice over 24 h, and no evidence was found that shoals might break up into sub-units of individuals that are more phenotypically assorted than their original shoals. The mechanisms by which assortative groups may arise and the consequences of low group fidelity for the evolution of cooperative behaviour are discussed.     

Positioning behaviour in fish shoals: a cost–benefit analysis

Shoal position can have a strong influence on individual fitness. Individuals in front positions of shoals were observed to have higher feeding rates than individuals elsewhere. Manipulation of nutritional state showed that hungry individuals had a stronger preference for front positions and that the duration of food-deprivation was positively correlated with the degree of the position preference. On the other hand, front positions (like other peripheral positions) probably incur costs in terms of increased predation risks. Experiments with Schreckstoff showed that frightened individuals seek the central part of the shoal. This suggests that individuals rotate their shoal positions according to the tradeoff between energy intake and predation risk.     

Familiarity affects social network structure and discovery of prey patch locations in foraging stickleback shoals

Numerous factors affect the fine-scale social structure of animal groups, but it is unclear how important such factors are in determining how individuals encounter resources. Familiarity affects shoal choice and structure in many social fishes. Here, we show that familiarity between shoal members of sticklebacks (Gasterosteus aculeatus) affects both fine-scale social organization and the discovery of resources. Social network analysis revealed that sticklebacks remained closer to familiar than to unfamiliar individuals within the same shoal. Network-based diffusion analysis revealed that there was a strong untransmitted social effect on patch discovery, with individuals tending to discover a task sooner if a familiar individual from their group had previously done so than if an unfamiliar fish had done so. However, in contrast to the effect of familiarity, the frequency with which individuals had previously associated with one another had no effect upon the likelihood of prey patch discovery. This may have been due to the influence of fish on one another''s movements; the effect of familiarity on discovery of an empty ‘control’ patch was as strong as for discovery of an actual prey patch. Our results demonstrate that factors affecting fine-scale social interactions can also influence how individuals encounter and exploit resources.     

Maintaining fish diversity in Thailand: variations in foraging behavior

We examined foraging adaptations and diet among dominant fishes within two species-rich riffle habitats in Thailand headwater rivers during the early and late portions of the dry season. All species exhibited diel temporal variations in feeding activity. Some species fed principally during daylight, while others fed mainly during darkness. Feeding patterns within species were generally spatially and temporally consistent. All seven fishes exploited the same benthic prey, but their foraging activities were related to spatially distinct water velocities. Diets overlapped with Ephemeroptera and Trichoptera the quantitatively most important prey. Ostracods, while not a large contributor to diet biomass, were numerically important in the diet of Homaloptera smithi. Trophic diversification through diel temporal feeding patterns and microhabitat separation by water velocity are major tactics in maintaining high diversity in riffle assemblages in Thai headwater rivers.     

Initiative, personality and leadership in pairs of foraging fish

Studies of coordinated movement have found that, in many animal species, bolder individuals are more likely to initiate movement and shyer individuals to follow. Here, we show that in pairs of foraging stickleback fish, leadership is not merely a passive consequence of temperamental differences. Instead, the act of initiating a joint foraging trip out of cover itself brings about a change in the role that an individual plays throughout the subsequent trip, and success in recruiting a partner affects an individual's tendency to initiate the next trip. On each joint trip, whichever fish took the initiative in leading out of cover gains greater influence over its partner's behaviour, which persists even after several changes in position (i.e. termination attempts and re-joining). During any given trip, the initiator is less responsive to its partner's movements than during trips initiated by the partner. An individual's personality had an important effect on its response to failure to recruit a partner: while bold fish were unaffected by failures to initiate a joint trip, shy individuals were less likely to attempt another initiation after a failure. This difference provides a positive feedback mechanism that can partially stabilise social roles within the pair, but it is not strong enough to prevent occasional swaps, with individuals dynamically adjusting their responses to one another as they exchange roles.     

Effects of bottom trawling on fish foraging and feeding

The effects of bottom trawling on benthic invertebrates include reductions of biomass, diversity and body size. These changes may negatively affect prey availability for demersal fishes, potentially leading to reduced food intake, body condition and yield of fishes in chronically trawled areas. Here, the effect of trawling on the prey availability and diet of two commercially important flatfish species, plaice (Pleuronectes platessa) and dab (Limanda limanda), was investigated over a trawling intensity gradient in the Irish Sea. Previous work in this area has shown that trawling negatively affects the condition of plaice but not of dab. This study showed that reductions in local prey availability did not result in reduced feeding of fish. As trawling frequency increased, both fish and prey biomass declined, such that the ratio of fish to prey remained unchanged. Consequently, even at frequently trawled sites with low prey biomass, both plaice and dab maintained constant levels of stomach fullness and gut energy contents. However, dietary shifts in plaice towards energy-poor prey items were evident when prey species were analysed individually. This, together with a potential decrease in foraging efficiency due to low prey densities, was seen as the most plausible cause for the reduced body condition observed. Understanding the relationship between trawling, benthic impacts, fish foraging and resultant body condition is an important step in designing successful mitigation measures for future management strategies in bottom trawl fisheries.     

Glacial clay affects foraging performance in a Patagonian fish and cladoceran

Climate change is altering temperatures and precipitation patterns all over the world. In Patagonia, Argentina, predicted increase in precipitation together with rapidly melting glaciers increase the surface runoff, and thereby the transport of suspended solids to recipient lakes. Suspended solids affect the visual conditions in the water which in turn restricts visual foraging. The native fish Aplochiton zebra Jenyns, and its filter-feeding cladoceran prey, Daphnia commutata Ekman, were subjected to foraging experiments at three turbidity levels. A. zebra foraging rate was substantially reduced at naturally occurring turbidity levels and the filtering rate of D. commutata was reduced at the highest turbidity level. This indicates that Daphnia may be partly released from predation from A. zebra at the same time as it can maintain relatively high feeding rates as turbidity increases. Lower foraging rates at the same time as the metabolic demand increases, through increased temperatures, may result in larger effects on A. zebra than could be expected from increases in turbidity or temperature alone. Turbidity may, as an indirect effect of climate change, decrease planktivore foraging rates and thereby alter the interaction strength between trophic levels.     

Opsin switch reveals function of the ultraviolet cone in fish foraging

Although several studies have shown that ultraviolet (UV) wavelengths are important in naturally occurring, visually guided behaviours of vertebrates, the function of the UV cone in such behaviours is unknown. Here, I used thyroid hormone to transform the UV cones of young rainbow trout into blue cones, a phenomenon that occurs naturally as the animal grows, to test whether the resulting loss of UV sensitivity affected the animal''s foraging performance on Daphnia magna, a prey zooplankton. The distances and angles at which prey were located (variables that are known indicators of foraging performance) were significantly reduced for UV knock-out fish compared with controls. Optical measurements and photon-catch calculations revealed that the contrast of Daphnia was greater when perceived by the visual system of control versus that of thyroid-hormone-treated fish, demonstrating that the UV cone enhanced the foraging performance of young rainbow trout. Because most juvenile fishes have UV cones and feed on zooplankton, this finding has wide implications for understanding the visual ecology of fishes. The enhanced target contrast provided by UV cones could be used by other vertebrates in various behaviours, including foraging, mate selection and communication.     

Size-segregative behaviour in minnow shoals

Shoals composed of equal numbers of two size-classes of European minnows were observed undisturbed, feeding and after threat from a pike in a large arena tank.
The time/frequency budget and analysed sequences of behaviour of the two size-classes were very similar. Irrespective of size, for standard behaviour measures, fish in the shoal behaved similarly under the same external influences, including predator threat.
In contrast, however, the distribution of the two size-classes provided evidence of size segregation within the shoal. This was brought about by individual minnows making shoaling responses preferentially to their own size-class. After exposure to the predator, shoaling responses changed and differed between small and large minnows.
The outcomes of contests at foraging patches were governed primarily by fish size and information asymmetry rather than by occupation of a feeding site.
The experiment shows that asymmetrical pay-offs in foraging and in response to predator threat are the probable reasons for size-segregation behaviours. This conclusion supports the views of earlier workers that mechanical sorting by swimming speed is not an important factor in size segregation in shoals.     

Temporal variation in foraging group structure of a size-structured stream fish community

Temporal variation in foraging group structure of a fish assemblage was examined in a flood-prone stream in southern Hokkaido, Japan. Foraging behaviour was observed underwater for four species which inhabit the water column: ayu, Plecoglossus altivelis, white-spotted charr, Salvelinus leucomaenis, masu salmon, Oncorhynchus masou, and Japanese dace, Tribolodon hakonensis, with each species being categorized into five size classes (species-size group; SSG). Based on foraging behaviour, each SSG of the fish assemblage was classified into one of four foraging groups: algae grazers, drift foragers, benthos-drift foragers, and omnivores, defined as SSG exhibiting similar foraging behaviour. All size classes of ayu, and of charr and salmon were categorized as algae grazers and drift foragers, respectively, throughout the study period. In contrast, size classes of dace were categorized as drift foragers, benthos-drift foragers, or omnivores with the same size classes often assigned to different foraging groups from month to month. Digestive tract contents of the fishes in the four foraging groups reflected their observed foraging behaviour, and foraging groups were therefore regarded as representing trophic groups. Abundance and membership of each foraging group varied in accordance with changes in abundance of SSG due to their growth, immigration, emigration, and/or mortality. Moreover, due to numerical dominance within the assemblage, plasticity in foraging behaviour of small- and medium-sized dace also played a key role in determining variability in the foraging group structure. Relative frequencies of two types of foraging behaviour, algae nipping and benthos foraging, of the small-sized dace were significantly correlated with the level of each resource, whereas no significant relationship was detected between foraging frequencies of the medium-sized dace and either resource. Fluctuations in foraging group structure within this assemblage occurred through niche shifts of some component members and by changes in SSG composition.     

Body length variation within multi-species fish shoals: the effects of shoal size and number of species

Theory predicts that selection should favour phenotypic homogeneity in fish shoals, and field studies have indeed confirmed that variation in body length within fish shoals is significantly lower than expected from a random distribution of fish among shoals. We investigated the extent to which variation in fish body length within shoals is determined by the shoal mean of body length, the number of species in a shoal, and the overall shoal size. We collected 34 fish shoals, ranging in size from 6 to 776 individuals, from the littoral zone of a Canadian lake. Shoals consisted of up to four different species, with multi-species shoals being larger and more frequent than single-species ones. The strongest determinant of body length variation within shoals was the shoal mean of body length, followed by the number of fish species in a shoal; i.e. multi-species shoals were less size-assorted than single-species ones. A more detailed analysis showed that the higher body length variation observed in multi-species shoals was due to increased body length variation both within and between component species. Shoal size had no significant effect on body length variation within shoals. Potential explanations of the positive relationship between body length variation and the number of species in a shoal are suggested. The implications of the above results for the evolution of multi-species shoals are discussed. Received: 6 May 1997 / Accepted: 14 October 1997     

Experimental test of the influence of the size of shoals and density of fish on parasite infections

There is an increasing interest in the relationship between adult coral reef fishes and their parasites; however, there is almost no information concerning post larvae (i.e. newly recruited) fish and their parasites. Newly recruited and juvenile fish are highly sensitive to their environment and they have a very low survival rate. Fish grouping behavior has been studied in several ways and many studies have shown that increasing group size might be a way to share a cost linked to parasitism, via a dilution effect. To my knowledge, this hypothesis has never been tested for juveniles of fish. The potential influence of school size and density of a juvenile coral reef fish, Haemulon flavolineatum (Haemulidae), on the success of a monogenean fish parasite, Haliotrema sp., was studied. First, the fish acquired their monogeneans when they were more than 40 mm long, size corresponding also to a change in their foraging behavior. Second, there was a positive effect of fish density on the success of the establishment of the parasite on the host population. Finally, although the parasite success was not directly related to fish group size, the parasite abundance was higher for intermediate shoal size. These results are discussed in terms of the consequence parasitism may have on a fish's shoaling and to a larger extent on the fish population.