Shoaling behaviour of sticklebacks infected with the microsporidian parasite, <Emphasis Type="Italic">Glugea anomala</Emphasis>

We compared the shoaling behaviour of three-spined sticklebacks, Gasterosteus aculeatus, infected with the microsporidian, Glugea anomala, to that of non-infected conspecifics. Infected fish lost significantly more weight than non-infected fish during a period of food deprivation, suggesting a metabolic cost to parasitism. In binary shoal choice tests, non-infected test fish showed an association preference for a shoal of non-infected over a shoal of infected conspecifics; infected test fish displayed no preference. Infected fish, however, showed a higher overall tendency to shoal than non-parasitised fish. Furthermore, infected fish occupied front positions within a mixed school. We consider the behavioural differences between infected and uninfected fish in the context of their potential benefits to the fish hosts and the parasites.     

Is there always an influence of shoal size on predator hunting success?

Theoretical and empirical studies predict that there should be a decrease in hunting success of predators with increasing prey group size. Most of these studies investigated situations in which predator and prey were in full view of each other before, during and after an attack. In this study, single rock bass Ambloplites rupestris were given an opportunity to launch surprise attacks at shoals of creek chub Semotilus atromaculatus that ranged in size from two to 13 fish. There was no significant influence of either shoal size or attack distance on predator success rate and no significant relationship between attack distance and shoal size. Furthermore, it was found that the leading fish of a shoal was attacked significantly more often than fish in other shoal positions, indicating that predation risk was not shared equally among shoal members. Also, leading fish in larger shoals (eight to 13 fish) were not more likely to survive a predator attack than ones in small shoals (two to seven fish).The consequences of these results are discussed in the general context of antipredator benefits of grouping.     

Crimson Spotted Rainbowfish (Melanotaenia duboulayi) Change Their Spatial Position according to Nutritional Requirement

Decision making in moving animal groups has been shown to be disproportionately influenced by individuals at the front of groups. Therefore, an explanation of state-dependent positioning of individuals within animal groups may provide a mechanism for group movement decisions. Nutritional state is dynamic and can differ between members of the same group. It is also known to drive animal movement decisions. Therefore, we assayed 6 groups of 8 rainbowfish foraging in a flow tank. Half of the fish had been starved for 24h and half had been fed 1h prior to experimental start. Groups were assayed again one week later but individuals were allocated to the opposite nutritional treatment. During the assay the positions of individually identified fish were recorded as were the number of food items they each ate and the position within the group they acquired them from. Food-deprived fish were more often found towards the front of the shoal; the mean weighted positional score of food-deprived fish was significantly larger than that of well-fed fish. Individuals were not consistent in their position within a shoal between treatments. There was a significant positive correlation between mean weighted positional score and number of food items acquired which displays an obvious benefit to front positions. These results suggest that positional preferences are based on nutritional state and provide a mechanism for state-dependent influence on group decision-making as well as increasing our understanding of what factors are important for group functioning.     

Influence of black spot disease on shoaling behaviour in female western mosquitofish, <Emphasis Type="Italic">Gambusia affinis</Emphasis> (Poeciliidae,Teleostei)

Parasites can fundamentally alter the cost–benefit ratio of living in a group, e.g. if infected individuals increase the predation risk of shoal mates. Here, the effect of an infection with a trematode, Uvulifer sp. (Diplostomatidae) on the shoaling behaviour of female western mosquitofish, Gambusia affinis, was investigated. The parasite examined causes a direct phenotypical change of the host by forming black spots on its body surface. When given a choice between a stimulus shoal and no shoal, we found shoaling tendencies to be significantly reduced in infected focal fish. In another experiment, we tested for association preferences relative to the infection status of the stimulus fish. Given the choice between an infected and a healthy stimulus fish, both infected and healthy focal fish preferred to associate with non-infected stimulus fish. Our results suggest that (1) the cost–benefit ratio of shoaling might be different for infected and non-infected individuals. Infected fish may be more affected by competition for food within a shoal. (2) Associating with infected conspecifics appears to be costly for female mosquitofish, maybe due to increased predation risk.     

Parasitism, oddity and the mechanism of shoal choice

When choosing between shoals differing in Schistocephalus solidus infection status, uninfected test sticklebacks Gasterosteus aculeatus showed a preference for joining uninfected conspecifics when shoal sizes were equal, but reversed this preference when the relative size of the infected shoal was increased by a factor of 3. When given a choice between a shoal composed of size-matched minnows Phoxinus phoxinus and a shoal composed of the same number of all uninfected or all S. solidus -infected sticklebacks, test fish always preferred the sticklebacks, regardless of their infection status, over the minnow shoal. These observations suggest that species, parasite status and shoal size are all of importance when fish decide which shoal to join.     

Balancing the dilution and oddity effects: decisions depend on body size

Background

Grouping behaviour, common across the animal kingdom, is known to reduce an individual''s risk of predation; particularly through dilution of individual risk and predator confusion (predator inability to single out an individual for attack). Theory predicts greater risk of predation to individuals more conspicuous to predators by difference in appearance from the group (the ‘oddity’ effect). Thus, animals should choose group mates close in appearance to themselves (eg. similar size), whilst also choosing a large group.

Methodology and Principal Findings

We used the Trinidadian guppy (Poecilia reticulata), a well known model species of group-living freshwater fish, in a series of binary choice trials investigating the outcome of conflict between preferences for large and phenotypically matched groups along a predation risk gradient. We found body-size dependent differences in the resultant social decisions. Large fish preferred shoaling with size-matched individuals, while small fish demonstrated no preference. There was a trend towards reduced preferences for the matched shoal under increased predation risk. Small fish were more active than large fish, moving between shoals more frequently. Activity levels increased as predation risk decreased. We found no effect of unmatched shoal size on preferences or activity.

Conclusions and Significance

Our results suggest that predation risk and individual body size act together to influence shoaling decisions. Oddity was more important for large than small fish, reducing in importance at higher predation risks. Dilution was potentially of limited importance at these shoal sizes. Activity levels may relate to how much sampling of each shoal was needed by the test fish during decision making. Predation pressure may select for better decision makers to survive to larger size, or that older, larger fish have learned to make shoaling decisions more efficiently, and this, combined with their size relative to shoal-mates, and attractiveness as prey items influences shoaling decisions.     

Consistency of leadership in shoals of mosquitofish (Gambusia holbrooki) in novel and in familiar environments

In social animal groups, an individual's spatial position is a major determinant of both predation risk and foraging rewards. Additionally, the occupation of positions in the front of moving groups is generally assumed to correlate with the initiation of group movements. However, whether some individuals are predisposed to consistently occupy certain positions and, in some instances, to consistently lead groups over time is as yet unresolved in many species. Using the mosquitofish (Gambusia holbrooki), we examined the consistency of individuals' spatial positions within a moving group over successive trials. We found that certain individuals consistently occupied front positions in moving groups and also that it was typically these individuals that initiated group decisions. The number of individuals involved in leading the group varied according to the amount of information held by group members, with a greater number of changes in leadership in a novel compared to a relatively familiar environment. Finally, our results show that the occupation of lead positions in moving groups was not explained by characteristics such as dominance, size or sex, suggesting that certain individuals are predisposed to leadership roles. This suggests that being a leader or a follower may to some extent be an intrinsic property of the individual.     

The effect of shoal size on patterns of body colour segregation in mollies

Individual white and black mollies Poecilia latipinna spent significantly more time near the larger group when given the choice between two shoals of similar colouration to themselves. When given the choice between a large and a small shoal of dissimilar colouration to themselves, black test fish spent significantly more time with the larger shoal while white test fish showed no preference for either group. Both white and black mollies chose the smaller of two shoals when given the choice between a large dissimilarly coloured shoal and a small similarly coloured shoal. The results indicate that mollies actively discriminate between shoals on the basis of both body colour and shoal size. However, body colour segregation appears to have a stronger influence on shoal choice.     

Optimal group positioning after a predator attack: the influence of speed, sex, and satiation within mobile whirligig swarms

The position of animals within fish shoals, bird flocks, andinsect swarms is related to individual differences in hunger,body size, and defenses. These differences relate to the waythat animals balance multiple selection pressures includingfood-distribution and predator-attack patterns. However, therole of drafting/slipstreaming (taking advantage of the vorticesof those in front of you) and sex on the position of individualswithin a polarized group has not been well studied. For example,although hungry fish have been found to prefer positions towardthe front of a shoal on average, the mitigating factors of sex,recent predator exposure, and drafting have not been factoredinto this response. We conducted a controlled laboratory experimentwith individually marked whirligig beetles (Coleoptera: Gyrinidae)where sex and feeding level were controlled and the positionof beetles in a polarized group (in a flow tank) was analyzedat 2 different water speeds after exposure to a simulated predator.It was predicted that males and females would balance foragingand predator avoidance needs differently, as suggested by sexualsegregation theory and that males might be likely to occupyfront positions because of greater energetic needs. We foundthat in slow water males were more likely to occupy front positions,whereas in fast water females did, suggesting a different trade-offbetween the sexes in the need to forage versus save energy (draft).Additionally, we found that in slow water it was the hungrymales that came to the group's front, whereas hungry femaleswere more likely to move back. These are some of the first observationsof the positional complexity with which individuals in congregationsdisplay, and several adaptive and nonadaptive explanations forthe observed patterns are suggested.     

The relationship between foraging and shoal position in a mixed shoal of roach (Rutilus rutilus) and chub (Leuciscus cephalus): a field study

Feeding rates of mixed shoals of juvenile roach and chub were observed in a shallow stream near Cambridge (UK). Roach at the front of the shoal had significantly higher feeding rates than roach at the back and than chub in either front or back positions. Position in the shoal also had a significant effect on the kind of food consumed, with front roach feeding more on plankton and back roach more on bottom food. Altogether 36 fish from the stream were caught and marked. Half of these were deprived of food and the other half well-fed for 3 days in captivity. After release 36% of them joined their old shoal again. Individuals from the starved group occupied front positions significantly more often than well-fed fish, but after 2 days this difference disappeared.     

Cross-shoal variability in the feeding habits of migrating Atlantic cod (Gadus morhua)

Prey intake and selection were related to within-shoal position for Atlantic cod (Gadus morhua) engaged in annual migration across the Newfoundland shelf in the northwest Atlantic. Comparisons made among fish occupying five regions from the front to rear of a large (>10 km across) migrating shoal indicated that leading fish, or scouts, were larger, ate more food by weight, and had a more varied diet than did fish at other positions. Also, scouts consumed more preferred prey types (fish and pelagic invertebrates) than did fish at other positions. In contrast, trailing fish consumed few fish prey but a larger proportion of benthic invertebrates. Our results are the first to document systematic heterogeneous feeding success among members of a free-ranging and migrating fish shoal in the open ocean.     

The effect of sex and shoal size on shoaling behaviour in Danio rerio

Male and female zebra fish Danio rerio were given choices of shoals that differed in sex and size. Male zebra fish preferred to associate with female shoals over male shoals, but had no preference when given a choice between a mixed‐sex shoal and either a male or female shoal. Female zebra fish showed no significant preference when given a choice between male and females shoals, nor between mixed‐sex shoals and either male or female shoals. When given choices between shoals of differing size, females preferred to associate with the larger shoal, whether or not they were composed of males or females. Males, however, had no preference for larger shoals over smaller shoals, whether or not they were composed of males or females. These results showed that male zebra fish were capable of distinguishing between males and females solely on the basis of visual cues. Furthermore, these results demonstrated a significant difference between the shoaling choices of male and female zebra fish, which may indicate a difference in the function of shoaling for the two sexes.     

The Effect of Hunger on Shoal Choice in Golden Shiners (Pisces: Cyprinidae,Notemigonus crysoleucas)

When an animal has a choice of joining one group over another, its decision may depend on its relative vulnerabilities to predation and starvation. For example, a well-fed animal may choose a large group of individuals with body size matching its own because this gives good protection against predators, but a hungry animal may prefer smaller groups made up of smaller individuals because this decreases food competition. To test this idea, a choice between various shoals was given to golden shiners, Notemigonus crysoleucas, that were either well fed or deprived of food for 48 h. In a choice of 10 vs. 3 shoalmates, both well-fed and hungry shiners spent more time near the shoal of 10. In a choice of 20 vs. 3 shoalmates, both well-fed and hungry shiners again preferred the larger shoal, but in one replicate this preference was significantly weaker in the hungry fish. This reduced preference did not appear to be an artefact of increased mobility by hungry fish searching for food. In a choice between shoals of small vs. large conspecifics, small well-fed shiners, small hungry shiners, and large well-fed shiners preferred shoalmates with body size matching their own, but large hungry shiners preferred smaller individuals. These results are consistent with the hypothesis that hungry fish sacrifice safety from predation in their shoaling behaviour (by avoiding larger groups to a certain extent and by risking the oddity effect) so as to decrease food competition.     

Shoaling behaviour in a surface-dwelling and a cave-dwelling population of a barb <Emphasis Type="Italic">Garra barreimiae</Emphasis> (Cyprinidae,Teleostei)

We studied shoaling behaviour in a species of fish (Garra barreimiae) from Oman. We compared two populations (a surface-dwelling and a cave-dwelling population) with different theoretical costs and benefits of shoaling. We measured the tendency to associate with a shoal of conspecifics. The stimulus shoal was confined to (1) clear Plexiglas cylinders in light, (2) wire-mesh cylinders in light, or (3) wire-mesh cylinders in darkness. The surface form exhibited a strong preference for the stimulus shoal during the experiments in light, but also in darkness, when only non-visual cues from the shoal could be perceived. The cave form did not show a preference when solely visual cues were available (Plexiglas cylinder). When non-visual cues from the shoal could be perceived (wire-mesh), the cave form did show a preference to associate with the shoal, but the shoaling tendency was considerably weaker than in the surface form. The shoaling tendency has probably been genetically reduced in the cave form.     

Overriding the oddity effect in mixed-species aggregations: group choice by armored and nonarmored prey

Because "odd" individuals often suffer disproportionately highrates of predation, solitary individuals should join groupswhose members are most similar to themselves in appearance.We examined group-choice decisions by individuals in armoredand nonarmored species and predicted that either (1) the oddityeffect would result in preference for conspecific groups forsolitary individuals of both species, or (2) individuals inthe armored species would prefer to associate with groups containingindividuals of the more vulnerable species. Armored brook sticklebacks(Culaea inconstans) and nonarmored fathead minnows (Pimephalespromelas) have the same predators and often occur together instreams. In mixed-species shoals, yellow perch (Perca flavescens)attacked minnows earlier and more often than sticklebacks. Wetested whether solitary minnows and sticklebacks preferred toassociate with conspecific or heterospecific shoals under conditionsof both low and high predation risk. When predation risk washigh, minnows preferred to associate with conspecifics overheterospecifics, as predicted by the oddity effect. In contrast,sticklebacks preferentially associated with groups of minnowsover groups of conspecifics when predation risk was high. Whenpredation risk was low, solitary individuals of both speciespreferentially associated with conspecific over heterospecificshoals. Stickleback shoal choices under low-risk conditionsmay have been influenced by interspecific competition for food.In feeding experiments, minnows were more efficient foragersthan sticklebacks, so it should benefit sticklebacks to avoidminnows unless predation risk is high. Therefore, for armoredprey, the benefits of associating with more vulnerable preyappear to override the costs of both the oddity effect and foodcompetition when predation risk is high.     

Shoal Choice in the Banded Killifish (Fundulus diaphanus,Teleostei, Cyprinodontidae): Effects of Predation Risk,Fish Size,Species Composition and Size of Shoals

Banded killifish (Fundulus diaphanus) were presented individually with a choice of shoaling with either of two stimulus shoals which differed in shoal size, species composition, and fish body size, before and after a simulated avian predator attack. When threatened, test fish preferred to shoal with the larger of two conspecific shoals, but only if members of both stimulus shoals were of the same size class as the test fish. Otherwise, they preferred to shoal with similarly sized fish irrespective of shoal size; threat of predation increased the magnitude of this preference. Furthermore, test fish preferred a shoal of similarly sized shiners (Notemigonus crysoleucas) over larger killifish, when shoal sizes were identical. This indicates that body size plays a key role in shoal choice, overriding the effects of shoal size and species preference. Notwithstanding the above, shoal choice was affected by predator threat only when differences between shoal size or body size of stimulus fish were large.     

Group Choice as a Function of Group Size Differences and Assessment Time in Fish: The Influence of Species Vulnerability to Predation

The shoal-choice behaviour of two species of fish that differ in their vulnerability to predation was compared. Individuals of threespine stickleback, Gasterosteus aculeatus, and creek chub, Semotilus atromaculatus, were presented with a simultaneous choice of two equidistant stimulus shoals of conspecifics that differed in membership size (5 vs. 6 fish, 5 vs. 7, 5 vs. 8, 5 vs. 9 and 5 vs. 10). Test fish were allowed to view the stimulus shoals from a standard distance for either 10–20 or 120–150s before being frightened with a stimulus from an overhead light and released to join either shoal. We observed which shoal (the smaller or the larger one) the test fish approached. Preference for the larger stimulus shoal generally increased with increasing shoal size difference and with the duration of the assessment period, and was more pronounced in chub (the more vulnerable of the two species). For the short assessment period, chub showed a significantly stronger preference for the larger stimulus shoal than sticklebacks, whereas there was no significant difference between species for the long assessment period. Furthermore, chub responded more readily to small differences in shoal size (of 1–3 fish) than sticklebacks, for both short and long assessment periods. The above results are consistent with the hypothesis that chub, as the more vulnerable of the two species (in terms of predation), should be able to identify the larger of two shoals more quickly and should be more sensitive to small differences in shoal size than sticklebacks.     

Interpopulation Differences in Shoaling Behaviour in Guppies (Poecilia reticulata): Roles of Social Environment and Population Origin

In Trinidad, guppies (Poecilia reticulata) in high‐predation localities show more cohesive shoaling behaviour than those living with less dangerous predators in low‐predation sites. We evaluated the relative contributions of population origin (i.e. genetic and/or maternal effects) and social environment on the expression of shoaling by assessing the behaviour of juveniles reared in a range of social conditions. Focal individuals, offspring of guppies from populations from high‐ or low‐predation localities, were reared in a multifactorial experiment; we created four different social conditions by manipulating the source and demography of the conspecific residents with whom focal individuals interacted. We found that high‐predation fish displayed a stronger propensity to shoal than low‐predation ones. Our results also suggest a role for interactions between the source of the focal individuals, the demography of the group in which they were reared and the origin of the guppies with whom they were reared. Depending on their origin (high‐ vs. low‐ predation) and rearing density, our focal fish were more likely to shoal if they were reared with high‐predation residents. Learning from high‐predation residents, aggressive interactions with low‐predation residents and/or phenotype matching could have played a role in driving this effect of social environment. This effect of the phenotype of conspecifics on shoaling development would enhance heritable differences in shoaling propensity such that both could contribute to the well‐documented difference in shoaling behaviour of high‐ and low‐predation guppies in natural populations.     

The effect of hunger, shoal size and the presence of a predator on shoal cohesiveness in bluntnose minnows, Pimephales notatus Rafinesque

Shoals of fish vary in their degree of cohesiveness as certain conditions such as hunger, predation risk and shoal size vary: this study relates this variation to the benefits of shoaling. Shoals of bluntnose minnows, Pimephales notatus Rafinesque, consisting of different numbers of individuals were starved for 5, 24 or 72 h and allowed to forage in the presence or absence of a predator. Cohesiveness was measured as the dispersion offish within the shoal (the radius of the longest axis of the shoal corrected for the number of fish in the shoal), straggle frequency (the number of movements by individuals of at least 5 body lengths away from the shoal) and aggressive interactions between shoal members. Cohesiveness increased as shoal size and the benefits per individual increased. Cohesiveness also increased in the presence of a predator, as did the importance of shoaling as a means of reducing predation risk. Cohesivenes decreased as hunger level increased, possibly as a result of increased competition for food among shoal members.     

鱼群中饥饿个体比例和模拟捕食风险对德玛森小岩鲷群体行为的影响

研究旨在探讨同质(所有个体均正常摄食或饥饿)或异质(由不同饥饿个体比例组成的鱼群)鱼群的隐蔽所利用及觅食行为差异,以及上述行为对模拟捕食刺激的响应。实验选取喜好集群的德玛森小岩鲷(Chindongo demasoni)幼鱼为研究对象,以设置了隐蔽所臂和食物臂的六臂迷宫为竞技场,考察不同饥饿个体比例鱼群(8尾成员,分别为8F0S、7F1S、4F4S、1F7S和0F8S, F为正常投喂个体, S为饥饿个体),在自发状态及遭遇模拟捕食刺激下在迷宫不同区域的分布和成群动态。研究发现:(1)8F0S鱼群偏好隐蔽所臂;随着鱼群饥饿个体成员比例上升,鱼群在食物臂分布呈线性增加趋势, 0F8S鱼群在隐蔽所臂和食物臂的分布已不再存在差异;(2)隐蔽所臂的鱼群成群频率随饥饿鱼比例的上升呈下降趋势,但食物臂的鱼群成群频率并未随饥饿鱼比例上升而上升,且鱼群单次持续时间和总体时间占比均不受鱼群内部饥饿个体占比的影响;(3)模拟捕食刺激导致实验鱼在隐蔽所臂分布显著上升,应激状态下几乎所有集群均发生在隐蔽所臂,且该变化不受鱼群组成的影响。研究表明:(1)在陌生环境下德玛森小岩鲷的行为策略是优先避敌,其次才是营养需求...