四种鲤科鱼类对捕食胁迫行为响应的种间比较

为了比较早期捕食胁迫经历和当前环境中存在的捕食者对鱼类行为的影响, 并考查这些影响是否存在种间差异, 研究分别考查了测定环境(有、无捕食者存在)对有、无捕食胁迫经历的鳊(Parabramis pekinensis)、草鱼(Ctenopharyngodon idellus)、鲫(Carassius auratus)和中华倒刺鲃(Spinibarbus sinensis)等4种鲤科鱼类探索性、活跃性和勇敢性的影响。结果发现: 早期捕食胁迫经历与当前环境条件对鱼类行为产生截然不同的影响, 且存在较大的种间差异。无捕食胁迫经历的鳊、草鱼和中华倒刺鲃均会对陌生的捕食者乌鳢(Channa argus)做出行为响应, 提示这3种鱼可能对陌生捕食者具有一定的识别能力, 但这种识别与猎物鱼通过捕食胁迫经历获得的识别仍具有一定差距; 具有捕食胁迫经历的鳊和中华倒刺鲃在空白环境中未表现出反捕食行为, 可能是节约能量的一种策略。总体而言, 草鱼对捕食胁迫经历和测定环境处理反应更为敏感, 而中华倒刺鲃的反应则相对保守。但当周围环境中存在捕食者时, 4种鲤科鱼类均会通过维持较高运动状态的方式来应对捕食者。维持这种应激状态可能对猎物鱼保持与捕食者的距离, 并随时保持警惕较为关键。     

Kinship reinforces cooperative predator inspection in a cichlid fish

Kin selection theory predicts that cooperation is facilitated between genetic relatives, as by cooperating with kin an individual might increase its inclusive fitness. Although numerous theoretical papers support Hamilton's inclusive fitness theory, experimental evidence is still underrepresented, in particular in noncooperative breeders. Cooperative predator inspection is one of the most intriguing antipredator strategies, as it implies high costs on inspectors. During an inspection event, one or more individuals leave the safety of a group and approach a potential predator to gather information about the current predation risk. We investigated the effect of genetic relatedness on cooperative predator inspection in juveniles of the cichlid fish Pelvicachromis taeniatus, a species in which juveniles live in shoals under natural conditions. We show that relatedness significantly influenced predator inspection behaviour with kin dyads being significantly more cooperative. Thus, our results indicate a higher disposition for cooperative antipredator behaviour among kin as predicted by kin selection theory.     

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...     

Who dares, learns: chemical inspection behaviour and acquired predator recognition in a characin fish

Individuals that dare approach predators (predator inspection behaviour) may benefit by acquiring information regarding the potential threat of predation. Although information acquisition based on visual cues has been demonstrated for fish, it is unknown whether fish will inspect predators on the basis of chemical cues or whether such inspection behaviour results in information acquisition. Here, we first ascertained whether predator inspection behaviour can be mediated by chemical cues from predators by exposing groups of predator-naive glowlight tetras (Hemigrammus erythrozonus) to the chemical cues of a potential fish predator (convict cichlid Cichlasoma nigrofasciatum) that had been fed either tetras (which possess an alarm pheromone) or swordtails (Xiphophorus helleri, which lack Ostariophysan alarm pheromones). Tetras showed a significant increase in antipredator behaviour when exposed to the tetra-diet cue, but not when exposed to the swordtail-diet cue. Chemically mediated predator inspection behaviour was also affected. Both the latency to inspect and the minimum approach distance to the predator significantly increased, and the mean number of inspectors per predator inspection visit significantly decreased when tetras were exposed to the tetra-diet versus the swordtail-diet chemical cues. We then examined a potential benefit associated with chemically mediated predator inspection behaviour. Only tetras that were initially exposed to the tetra-diet cue and that had inspected the predator acquired the visual recognition of a convict cichlid as a predation threat. Our results thus demonstrate that (1) predator inspection behaviour in the glowlight tetra can be initiated by chemical cues, (2) chemically mediated inspection behaviour is affected by the presence of alarm pheromone, and (3) inspectors benefit by acquiring the recognition of novel predators. Copyright 1999 The Association for the Study of Animal Behaviour.     

Consequences of temperature and temperature variability on swimming activity,group structure,and predation of endangered delta smelt

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Predators Exacerbate Competitive Interactions and Dominance Hierarchies between Two Coral Reef Fishes

Predation and competition are critical processes influencing the ecology of organisms, and can play an integral role in shaping coral reef fish communities. This study compared the relative and interacting effects of competition and predation on two competing species of coral reef fish, Pomacentrus amboinensis and P. moluccensis (Pomacentridae), using a multifactorial experiment. Fish were subjected to the sight and smell of a known predator (Pseudochromis fuscus), the presence of the heterospecific competitor (i.e., P. amboinensis vs. P. moluccensis), or a combination of the two for a period of 19 days. The sub-lethal effects of predator/competitor treatments were compared with controls; a combination of otolith microstructure analysis and observations were used to determine otolith growth patterns and behaviour. We predicted that the stress of competition and/or predation would result in strong sub-lethal impacts, and act synergistically on growth and behavioural patterns. We found strong evidence to support this prediction, but only for P. amboinensis, which suffered reductions in growth in both predator and competitor treatments, with the largest reductions occurring when subjected to both predation and competition concurrently. There was strong evidence of asymmetrical competition between the two damselfish species, with P. moluccensis as the dominant competitor, displaying strong aggressive behaviour towards P. amboinensis. Growth reductions for P. amboinensis in predator/competitor treatments appeared to come about primarily due to increases in shelter seeking behaviour, which significantly reduced the foraging rates of individuals compared with controls. These data highlight the importance of predator/competitor synergisms in influencing key behaviours and demographic parameters for juvenile coral reef fishes.     

Interpopulational differences in the use of the Tit-For-Tat strategy during predator inspection in the guppy,Poecilia reticulata

Summary Prior work (Dugatkin, 1991a) has argued that guppies, originating from an area of high predation pressure, employ the Tit-For-Tat strategy during predator inspection visits. Guppies from the high predation site show all three characteristics associated with Tit-For-Tat; nice, retaliatory and forgiving behaviour. Since predation is the major selective force favouring Tit-For-Tat during predator inspection, we predicted fish from an area of low predation would not display the Tit-For-Tat strategy. Our results confirm this prediction; males display retaliatory, but not nice and forgiving behaviour, while females display none of the three characteristics. Thus as predation pressure changes, so too does the conditional nature of the cooperative strategy used during predator inspection.     

Antipredator Behaviour and Suppressed Aggression by Convict Cichlids in response to Injury-released Chemical Cues of Conspecifics but not to those of an Allopatric Heterospecific

In aquatic environments, chemical cues serve as an important source of information for the detection of predation risk. Here, we investigate the response of convict cichlids, Cichlasoma nigrofasciatum, to injury-released chemical cues. We exposed pairs of juvenile convict cichlids first to dechlorinated tap water (control), then later to one of two test stimuli: 1. chemical cues from injured convict cichlids; or 2. chemical cues from injured mosquito fish, Gambusia affinis. Gambusia are allopatric and phylogenetically unrelated to convict cichlids. Gambusia skin was used to control for a general response to injured fish. In response to conspecific cues, convict cichlids significantly increased time spent near the bottom of test aquaria and time under a shelter object. In response to Gambusia skin, convict cichlids tended to increase time spent near the tank bottom but did not increase use of the shelter object. There was a trade-off between antipredator and agonistic behaviours. In response to convict cichlid cues, there was a significant reduction in the frequency of approaches and bites. Gambusia skin extract had no significant effect on aggressive behaviour. These data suggest a species-specific antipredator response to conspecific alarm pheromones in a New World cichlid fish and demonstrate a trade-off between predator avoidance and intraspecific aggression. Further, the presence of an alarm response in this model species sets the stage for the use of chemical cues as a research tool to manipulate predation risk in studies of the interaction between predation risk and reproductive behaviour.     

Algal Turbidity Reduces Risk Assessment Ability of the Three‐Spined Stickleback

Recent anthropogenic increases in algal turbidity in aquatic habitats have been suggested to affect the ability of fish to assess predation risk. We investigated the response of feeding three‐spined sticklebacks (Gasterosteus aculeatus) exposed to a sudden appearance of an avian predator (the silhouette of common tern, Sterna hirundo), under clear and turbid water conditions. As stickleback use social cues to aid in predator avoidance, we also tested whether turbidity affected social information use by manipulating group size. We found that in turbid water, a smaller proportion of fish would escape from the feeding area, that the distance escaped was shorter and that a smaller proportion of fish fled into shelter. Larger group size was associated with longer escape distance and greater shelter use. However, there was no effect of group size on the proportion of fish that escaped the arena. The effect of group size was similar for turbid and clear water. Our finding that the fish showed a weaker antipredator response suggests that turbidity impedes their risk assessment capability. However, the sticklebacks were still able to benefit of the social facilitation provided by being in a group. This suggests that algal turbidity has detrimental effects on the ability of sticklebacks to assess predation risk from avian predators in shallow water. An implication is that in shallow water fish may be more vulnerable to avian predation under turbid conditions.     

Behavioural variation between piscivore and insectivore rainbow trout Oncorhynchus mykiss

A proactive-reactive continuum integrating multiple (i.e., 3+) dimensions of animal behaviour has been reported as a major axis of behavioural differentiation, but its stability along a biological hierarchy from individuals to populations remains speculative. Piscivore and insectivore rainbow trout (Oncorhynchus mykiss) represent closely related ecotypes with strong ecological divergence driven by selection for a large-bodied piscivorous lifestyle with fast juvenile growth vs. selection for smaller adult body size and lower growth associated with an insectivorous diet. To evaluate whether differences in behaviour between ecotypes are consistent with a proactive-reactive axis and consistent along a biological hierarchy, the authors examined variation in emergence time from a shelter, exploration, activity and predator inspection among individuals, populations and ecotypes of juvenile piscivore and insectivore rainbow trout O. mykiss. As expected, the faster-growing piscivore ecotype was more proactive (i.e., shorter emergence time, exploration and predator inspection) than the more reactive insectivore ecotype. This behavioural contrast was partly maintained across populations, although activity differences were most pronounced among populations, rather than emergence time. Insectivore fry showed substantial variation in behavioural expression among individuals within populations; by contrast, piscivores showed highly similar proactive behaviours with significantly lower inter-individual variation in behavioural expression, suggesting intense selection on behaviour supporting their faster growth. This work suggests that piscivore and insectivore O. mykiss broadly differ in behaviour along a proactive vs. reactive continuum, and highlights the greater multidimensionality of behavioural expression within the insectivore ecotype. Contrasting behaviours between ecotypes may result from differential selection for slow vs. fast juvenile growth and associated metabolism, and may contribute to adult trophic specialization.     

Escape tactics by a neotropical montane lizard: a comparison of flight responses against natural and nonnatural predators

Prey must judge the level of threat related to a given predator attack to employ the most efficient escape strategy at a low cost. In this context, the capacity of prey to correctly assess the threat from predators might optimise their decision on the strategy employed to flee. We examined the escape behaviour of the montane lizard Tropidurus montanus in response to different types of predatory stimuli. Lizards stimulated by a model of a natural predator ran shorter distances straight towards shelter and spent less time in flight. On the other hand, when exposed to a model of a nonnatural predator, T. montanus performed longer flights in distance and time, running across open and exposed substrates in the view of predators. These results showed that the lizard T. montanus made different escape decisions according to the type of predatory stimulus.     

Do competing males cooperate? Familiarity and its effect on cooperation during predator inspection in male three‐spined sticklebacks (Gasterosteus aculeatus)

We investigated the trade‐off between conflict and cooperation, using predator inspection behaviour in sticklebacks as a model system. Male three‐spined sticklebacks ( Gasterosteus aculeatus ) compete for territories during the breeding season and it has been demonstrated that the level of aggression between territorial neighbours declines with time, a phenomenon known as the dear enemy effect. In this experiment we examine whether this increase in familiarity between territorial neighbours can facilitate an increase in cooperation during predator inspection events. This was analyzed using male sticklebacks from four pond populations, two with and two without predatory rainbow trout ( Oncorhyncus mykiss ). Males were each exposed to five different treatments: (1) the presentation of a live rainbow trout when alone; (2) with a newly revealed territorial neighbour; (3) with this neighbour after two days of familiarization; (4) with another unfamiliar neighbour (termed a 'floater'); and (5) a second solitary trial (to provide controls at the start and end of the experiment). As predicted, fish from predator‐sympatric populations showed higher levels of predator inspection and lower rates of misdirected territorial aggression towards the predator throughout. However, familiarity between neighbouring males did not facilitate an increase in predator inspection behaviour. Instead, predator inspection behaviour decreased throughout treatments involving the presence of any sort of neighbouring male. Familiarity between neighbours did not influence their ability to cooperatively inspect, but only the nature of any aggressive territorial behaviours, all of which detracted equally from individual inspection effort.     

Do Larval Damselflies make Adaptive Choices When Exposed to Both Parasites and Predators?

The importance of multiple enemies from different trophic levels on investment in defence by prey has, with some exceptions, received little attention. Some defences may make the victim more susceptible to other enemies; this latter situation applies to predators and parasites of larval damselflies. Baker and Smith [Oecologia 109 (1997) 622) showed that larval damselflies were as active in the presence of both mites and fish as they were when only mites were present, an apparently maladaptive behaviour that results in higher fish predation. In this paper, we further examine this maladaptive behavioural response to multiple enemies (fish predators and mite parasites) and test whether their defence responses are a result of the order in which they experience the parasite or predator, and/or if behavioural ‘personalities’ exist, such that some individuals show anti‐predator behaviours and other show anti‐parasite behaviours. Order of experience did not affect the four main behaviours (groom, crawl, turn and swim) exhibited when larval damselflies were simultaneously exposed to fish and mites. Grooming levels increased in response to mites, decreased in response to fish and when exposed to both mites and fish were similar to when they were exposed to mites alone. Duration of the other three behaviours was lower in the presence of both mites and fish. The crawling ‘personalities’ were evident. The apparently maladaptive response of high grooming levels in the presence of mites and fish is not a result of order of experience or ‘personalities’. It may be a result of relatively high encounter rates with mite parasites, compared with the encounter rates with fish. Lower encounter rates can result in diminishing investment in defence against an enemy.     

The carabid <Emphasis Type="Italic">Pterostichus melanarius</Emphasis> uses chemical cues for opportunistic predation and saprophagy but not for finding healthy prey

The sentinel prey method can quantify predation pressure in various habitats. Real prey is assumed to more realistically mimic the predator experience but the predator can rarely be identified. Artificial prey made of plasticine may lack real chemical cues, but provides information about predator identity. However, the relationship between predation pressure registered by artificial versus real prey is not clear. We tested the relative attractiveness of artificial caterpillars, and intact, wounded, or dead larvae of the cabbage moth (Mamestra brassicae) for the carabid predator Pterostichus melanarius Illiger (Coleoptera: Carabidae). P. melanarius adults were attracted to dead caterpillars more than to live or wounded ones. Coating artificial caterpillars with caterpillar haemolymph increased their attractiveness. However, predators were not attracted more to healthy, real caterpillars than to “untreated” artificial ones. We conclude that using artificial caterpillars does not underestimate predation pressure by this carabid on healthy caterpillars.     

Dying from the lesser of three evils: facilitation and non‐consumptive effects emerge in a model with multiple predators

Prey modify their behaviour to avoid predation, but dilemmas arise when predators vary in hunting style. Behaviours that successfully evade one predator sometimes facilitate exposure to another predator, forcing the prey to choose the lesser of two evils. In such cases, we need to quantify behavioural strategies in a mix of predators. We model optimal behaviour of Atlantic cod Gadus morhua larvae in a water column, and find the minimal vulnerability from three common predator groups with different hunting modes; 1) ambush predators that sit‐and‐wait for approaching fish larvae; 2) cruising invertebrates that eat larvae in their path; and 3) fish which are visually hunting predators. We use a state‐dependent model to find optimal behaviours (vertical position and swimming speed over a diel light cycle) under any given exposure to the three distinct modes of predation. We then vary abundance of each predator and quantify direct and indirect effects of predation. The nature and strength of direct and indirect effects varied with predator type and abundance. Larvae escaped about half the mortality from fish by swimming deeper to avoid light, but their activity level and cumulative predation from ambush predators increased. When ambush invertebrates dominated, it was optimal to be less active but in more lit habitats, and predation from fish increased. Against cruising predators, there was no remedy. In all cases, the shift in behaviour allowed growth to remain almost the same, while total predation were cut by one third. In early life stages with high and size‐dependent mortality rates, growth rate can be a poor measure of the importance of behavioural strategies.     

Influence of predation risk on the sheltering behaviour of the coral-dwelling damselfish, <Emphasis Type="Italic">Pomacentrus moluccensis</Emphasis>

Predation is a key ecosystem function, especially in high diversity systems such as coral reefs. Not only is predation one of the strongest top-down controls of prey population density, but it also is a strong driver of prey behaviour and function through non-lethal effects. We ask whether predation risk influences sheltering behaviour of damselfish living in mutualism with branching corals. Host corals gain multiple advantages from the mutualistic relationship which are determined by the strength of damselfish sheltering. Distance travelled by the Lemon Damselfish Pomacentrus moluccensis away from their host colony was measured here as a proxy for sheltering strength and was expected to be shortest under highest predation risk. Predation risk, defined as a function of predator abundance and activity, turbidity and habitat complexity, was quantified at four reef slope sites in Kepulauan Seribu, Indonesia. Damselfish sheltering strength was measured using stationary unmanned video cameras. Small damselfish (< 2 cm) increased their sheltering strength under high turbidity. Predator feeding activity, but not abundance, influenced damselfish sheltering strength. Contrary to our expectations, sheltering behaviour of adult damselfish decreased under high predator activity. While these observations are in line with risk-averse behaviour by juvenile P. moluccensis, they may indicate the presence of sentinel behaviour in the adults of this species. Habitat complexity seemed to be less important as a driver of damselfish behaviour. These counterintuitive results may indicate complex social behaviour and age-specific strategies for predator avoidance.     

The relative importance of food and shelter for seagrass-associated invertebrates: a latitudinal comparison of habitat choice by isopod grazers

The generality of mechanisms affecting habitat choice and grazing in seagrass meadows was evaluated in a latitudinal comparison of seagrass grazers from the temperate (60°N) Baltic Sea and the subtropical (30°N) Gulf of Mexico. Using similar habitat choice experiment set-ups in Finland and the USA, the role of food type, habitat complexity and predation hazard on habitat choice of the isopods Idotea baltica (Pallas) and Erichsonella attenuata Harger were tested. When shelter was provided by both living and artificial seagrass, epiphytic food resources on artificial vegetation were clearly preferred by both species, although Idotea was attracted to epiphyte-free seagrass when no alternative food was present. When choosing between food and shelter, both species preferred epiphytic food over shelter. However, under predation hazard of fish, Erichsonella clearly switched to the habitat offering shelter, while the presence of a predatory fish produced no preference for shelter by Idotea. Food type may be considered as an universal mechanism that partly determines the presence of grazers in seagrass habitats and is, in the absence of a predator, more important than shelter. Predation risk affected the behaviour of the grazers, but the response varied between species possibly due to varying importance of fish predation in the areas studied. Received: 16 November 1998 / Accepted: 13 February 1999     

Refuge use by fish as a function of body length-related metabolic expenditure and predation risks

Theory predicts that staying in a refuge has benefits in terms of predator avoidance and costs in terms of lost feeding opportunities. In this study, we investigated how the relative importance of these costs and benefits changes with increasing body length. This is of particular interest in animals such as fish, which show continuous growth throughout their lives. Our results suggest that larger fish are subject to lower predation risks and are less affected by food deprivation than small fish, with fish decreasing their responses to food-deprivation treatments more strongly with increasing body length than to predation treatments. This may explain our observation that large fish emerged later from a refuge than small ones and spent shorter times outside the refuge. The key role of differential responses to food deprivation was further illustrated by the finding that the relative weight loss of individual fish was strongly correlated with a reduction in hiding time even in the absence of body length differences. The importance of inter-individual differences in metabolic rates for the decision-making behaviour of animals is discussed.     

捕食驯化对胭脂鱼和中华倒刺鲃游泳行为、应激和免疫功能的影响

鱼类对环境的行为和生理适应能力与其在自然界的资源变动状况密切相关, 研究选取胭脂鱼(Myxocyprinus asiaticus)和中华倒刺鲃(Spinibarbus sinensis)幼鱼为研究对象, 考察1周捕食者(乌鳢, Channa argus)驯化对2种鱼类的运动能力(最大匀加速速度)、特异(血浆IgM水平)和非特异免疫(血浆溶菌酶含量)指标和抗氧化能力(血浆超氧化物歧化酶活性)的影响, 及在有无捕食者急性暴露两种条件下的驯化和非驯化鱼群自发游泳行为(游泳速度、运动时间比和个体间距离)和应激反应(血浆皮质醇水平)。研究发现: (1)总体上中华倒刺鲃比胭脂鱼有更快的游泳速度、更为活跃的自发游泳行为、更高的血浆皮质醇和IgM水平; (2)1周捕食驯化导致两种鱼类血浆皮质醇水平、特异免疫和非特异免疫水平的上升, 并且中华倒刺鲃比起胭脂鱼表现的更加明显; (3)急性捕食者暴露导致血浆皮质醇水平上升, 个体间距离下降, 但后者仅在非驯化组有所体现。研究表明: (1)捕食驯化鱼类通过皮质醇动员特异和非特异免疫应对应激, 这些生理和行为的改变可能有利于鱼类增强避敌能力或加快非致死捕食损伤的快速恢复。这表明捕食驯化可作为潜在的增殖放流前的生态锻炼; (2)2种鱼类的行为、免疫等生理基础状态及其对外界刺激的响应程度不尽相同, 未来相似的环境变化可能对两种鱼类的资源产生不同影响。     

Nesting Success in Crimson Finches: Chance or Choice?

In avian systems, nest predation is one of the most significant influences on reproductive success. Selection for mechanisms and behaviours to minimise predation rates should be favoured. To avoid predation, breeding birds can often deter predators through active nest defence or by modifying behaviours around the nest (e.g. reducing feeding rates and vocalisations). Birds might also benefit from concealing nests or placing them in inaccessible locations. The relative importance of these strategies (behaviour vs. site selection) can be difficult to disentangle and may differ according to life history. Tropical birds are thought to experience higher rates of predation than temperate birds and invest less energy in nest defence. We monitored a population of crimson finches (Neochmia phaeton), in the Australian tropics, over two breeding seasons. We found no relationship between adult nest defence behaviour (towards a model reptile predator) and the likelihood of nest success. However, nest success was strongly related to the visibility of the nest and the structure of the vegetation. We found no evidence that adult nest building decisions were influenced by predation risk; individuals that re‐nested after a predation event did not build their nest in a more concealed location. Therefore, predator avoidance, and hence nest success, appears to be largely due to chance rather than due to the behaviour of the birds or their choice of nesting sites. To escape high predation pressures, multiple nesting attempts both within and between seasons may be necessary to increase reproductive success. Alternatively, birds may be limited in their nest‐site options; that is, high‐quality individuals dominate quality nest sites.