Responses of larval dragonflies to conspecific and heterospecific predator cues

Abstract.  1. In cannibalistic populations, smaller individuals are subject to predation by larger conspecifics, and small individuals commonly alter their behaviour in response to cannibals. Little is known, however, about the underlying cues that trigger such responses and how the behavioural responses to conspecific cannibals differ from heterospecific predators.
2. This study tests which cues are used for the detection of conspecific predators in the larva of the dragonfly Plathemis lydia and how the behavioural response to cannibals differed from the response to heterospecific predators.
3. Individuals were exposed to chemical cues, visual cues, and a combination of both cues from conspecifics as well as no predator and heterospecific predator controls during which their activity and feeding rates were observed.
4. Individuals increased their activity, spatial movement and feeding behaviour in response to either visual or chemical cues from conspecific predators, which was opposite to responses displayed with cues from heterospecific predators. Interestingly, the responses to visual and chemical cues from conspecifics combined were weaker than to either cue in isolation and similar to the no cue control.
5. The results clearly indicate that individuals are able to use chemical and visual cues to detect even very subtle differences in phenotype of conspecific predators.
6. The opposite response in behaviour when exposed to conspecific cannibals vs. heterospecific predators suggests that the presence of cannibals will increase the mortality risk of small individuals due to heterospecific predation. This risk-enhancement is likely to have important consequences for the dynamics of predator–prey interactions.     

The effects of stimulus type on consistency of responses to conflicting stimuli in Siamese fighting fish

Individuals must often decide among a range of behavioral responses in a given situation and individuals will likely vary from one another in the choices they make. A rarely explored example of this is when both a male and a female are present simultaneously and a male must decide to fight or court. Consistent individual differences have been found in decision-making in Siamese fighting fish; however, dummies were used to explore this question and it is possible that individuals respond differently in this situation when faced with live stimuli as a result of feedback. Therefore, this study investigated whether males behave similarly to dummy and live stimuli by exposing them to male and female conspecifics both simultaneously and separately. Subjects were tested first with dummies and then in the same situations using live conspecifics. It was hypothesized that males would respond in the same manner to both the dummy and live fish. The results indicated that individuals behaved similarly to the dummy fish and the live fish in corresponding conditions. In addition, repeatability levels suggest that individuals behaved similarly across trials of a given treatment type while differing from one another in their responses. This study demonstrates that males are consistent in their responses even when receiving feedback from conspecifics and supports the continued use of dummy stimuli as an effective tool.     

Consistency in boldness expression varies with ecological context in a jumping spider

Animals can adjust their behaviours depending on ecological context (i.e., behavioural plasticity), and an individual's response to a given context may also vary from occasion to occasion (intra‐individual variability). Recognizing the roles of both behavioural plasticity and intra‐individual variability is important in understanding how behavioural diversity is maintained within populations. However, how the ecological context itself influences the individual behavioural response and intra‐individual variability (e.g., how variable an individual is in their behavioural expression) remains largely unexplored. Here, we examine boldness expression (the duration of startle response) in a specialised spider‐eating jumping spider, Portia labiata, across three contexts following a mild disturbance: presence of a conspecific intruder (most dangerous), environmental change but no conspecific intruder, and no conspecific intruder or environmental change (safest). We found that context does not significantly influence the average boldness expression at the population level. However, each individual responded to each context differently, and the repeatability of boldness expression—the proportion of behavioural variation attributable to the between ‐individual level—is context‐dependent. We also found that in the presence of a conspecific intruder, spiders behave less predictably than in the environmental change context, but not differently from the safest context. These findings may suggest that the presence of conspecifics influences behavioural consistency in individuals, but that this may occur without influencing the population average behaviour.     

Frustrative reward omission increases aggressive behaviour of inferior fighters

Animals use aggressive behaviour to gain access to resources, and individuals adjust their behaviour relative to resource value and own resource holding potential (RHP). Normally, smaller individuals have inferior fighting abilities compared with larger conspecifics. Affective and cognitive processes can alter contest dynamics, but the interaction between such effects and that of differing RHPs has not been adjudged. We investigated effects of omission of expected reward (OER) on competing individuals with contrasting RHPs. Small and large rainbow trout (Oncorhynchus mykiss) were conditioned to associate a light with reward. Thereafter, the reward was omitted for half of the fish prior to a contest between individuals possessing a 36–40% difference in RHP. Small control individuals displayed submissive behaviour and virtually no aggression. By contrast, small OER individuals were more aggressive, and two out of 11 became socially dominant. Increased aggression in small OER individuals was accompanied by increased serotonin levels in the dorsomedial pallium (proposed amygdala homologue), but no changes in limbic dopamine neurochemistry were observed in OER-exposed individuals. The behavioural and physiological response to OER in fish indicates that frustration is an evolutionarily conserved affective state. Moreover, our results indicate that aggressive motivation to reward unpredictability affects low RHP individuals strongest.     

Free Amino Acids Mediate Association Preferences in Fish

Chemical information is used in a variety of contexts including habitat recognition, foraging and predator avoidance, and a plethora of studies have shown that an individual's distinct chemical profile can mediate interactions with conspecifics. Interestingly, recent work has revealed that diet alone is sufficient to change the chemical profile of individuals, and with it, the way other individuals perceive and interact with them. Free amino acids are known to be utilised across species in a range of contexts, including during predator–prey and host–parasite interactions, and have been widely postulated to underpin diet‐mediated social interactions, especially in fish. However, so far no empirical evidence has been presented to support this suggestion. Using an established behavioural assay in three‐spined sticklebacks (Gasterosteus aculeatus), we aimed to assess association preferences in groups of fish fed experimental diets which differed by a single free amino acid. Our results demonstrate that free amino acids alone are sufficient to mediate interindividual association preferences, raising the possibility that such a mechanism may be widespread among aquatic animals.     

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.     

Lateralization of Aggression during Reproduction in Male Siamese Fighting Fish

A preference for the left‐eye use during aggressive interactions has been widely reported in the literature, even though in some cases the direction of lateralization varies among individuals within populations. Laterality of aggression in male Siamese fighting fish has been described in a number of studies, yet very little is known about lateralization of aggression during reproduction and/or parental care in fish. Here, we aimed to investigate the relationship between the different reproductive phases and lateralization in eye use during aggressive interactions in males of Siamese fighting fish. Lateralization in eye use is influenced during the early reproductive state, before and after the bubble nest construction stages. We found that nest‐holding males preferentially used the right eye before and after bubble nest construction independent of the sex of the intruder. During the later reproductive phases, aggressiveness increased whereby the direction of lateralization rather than the degree was influenced supporting the hypothesis that reproductive state influences behavioral consistency in Siamese fighting fish.     

Cannibalism of parasitoid‐attacked conspecifics in a non‐carnivorous caterpillar

Cannibalism, the killing and consumption of conspecifics, can even occur in insect species typically considered to be non‐carnivorous. Of particular interest is the cannibalism of parasitoid‐attacked conspecifics, which could reduce parasitism levels in subsequent generations for that conspecific population. This study reports on the occurrence and some of the consequences of cannibalism in parasitoid‐attacked obliquebanded leafroller, Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae). We show that larvae of C. rosaceana, which is considered to be an herbivorous caterpillar species, did not prey upon live conspecifics, but readily consumed conspecifics attacked by Habrobracon gelechiae Ashmead (Hymenoptera: Braconidae). Further examination found that C. rosaceana larvae feeding on parasitoid‐attacked conspecifics, since their fourth instar, suffered a higher mortality and reduction in body size than those fed on plant material only. The cannibalism of attacked conspecifics did not appear to offer any nutrient benefits for the cannibal. To our best knowledge, this is the first empirical example of the occurrence and some of the consequences of cannibalism by a non‐carnivorous insect on its parasitoid‐attacked conspecifics. We discuss the adaptive significance of such cannibalism on parasitoid‐attacked conspecifics with respect to a trans‐generational fitness gain for the population through the killing of the parasitoids, thereby reducing parasitism in subsequent generations.     

The winner and loser effect: integrating multiple experiences

An important question in state-dependent behaviour is how multiple influences on state are integrated to determine current behaviour. Aggressive behaviour is known to be affected by a prior contest experience. Nevertheless, whether and how multiple prior fighting experiences are integrated into a fighting decision remain unexplored. In this study, individuals of Rivulus marmoratus (Cyprinodontidae), a hermaphroditic fish, were given different combinations of two prior fighting experiences to investigate: (1) the effect of penultimate experiences on the probability of winning a subsequent contest; (2) the relative effect of a recent win and loss; and (3) whether the effect of a winning experience was as short lived as observed in other species. Penultimate and recent fighting experiences were given to the test fish approximately 48 and 24 h prior to the dyadic contests, respectively. From the results of the five types of contests staged, we conclude that: (1) penultimate fighting experiences had a significant effect on the probability of winning a subsequent contest; (2) a more recent experience had a more pronounced effect than an earlier experience, which suggested that the effect of a fighting experience would decay and/or the effect of a recent experience would interfere with the effect of an earlier experience; (3) no asymmetric effect between a winning experience and a losing experience was detected; and (4) the effect of both a winning and a losing experience lasted for at least 48 h in R. marmoratus which was the maximum time tested in these experiments. The possible reasons for the differences in results among studies of experience effects on contest outcomes are discussed. Copyright 1999 The Association for the Study of Animal Behaviour.     

Interactions between Males of the Lizard Liolaemus tenuis: Roles of Familiarity and Memory

For territorial organisms, recognition of familiar individuals can reduce the frequency and intensity of aggressive encounters (‘dear enemy’ phenomenon), stabilize social systems, and reduce the cost of territory maintenance. Here, we investigated the behavioural events displayed during contests between familiar and unfamiliar individuals in the lizard Liolaemus tenuis (Liolaemidae), a species in which males are territorial. The behaviours recorded were attack, warning, evasion, and submission, and the latencies to the first aggressive (attack or warning) behaviour. Additionally, we assessed the ability of individuals to remember a familiar conspecific after a period without social interaction. Individual males reduced and delayed aggressive behaviour directed towards socially familiar individuals compared with unfamiliar ones. These results suggest that males distinguished between familiar and unfamiliar conspecific males and are in agreement with the ‘dear enemy’ phenomenon. Other behaviours were similar in the contests between familiar and unfamiliar individuals. Recognition of familiar conspecifics was lost after 20 d without social interactions. This may be relevant for interactions with floater males or with neighbours that lose their territory and subsequently attempt to fight for their ex‐neighbour's territory.     

Smell, learn and live: The role of chemical alarm cues in predator learning during early life history in a marine fish

The speed with which individuals can learn to identify and react appropriately to predation threats when transitioning to new life history stages and habitats will influence their survival. This study investigated the role of chemical alarm cues in both anti-predator responses and predator identification during a transitional period in a newly settled coral reef damselfish, Pomacentrus amboinensis. Individuals were tested for changes in seven behavioural traits in response to conspecific and heterospecific skin extracts. Additionally, we tested whether fish could learn to associate a previously novel chemical cue (i.e. simulated predator scent) with danger, after previously being exposed to a paired cue combining the conspecific skin extract with the novel scent. Fish exposed to conspecific skin extracts were found to significantly decreased their feeding rate whilst those exposed to heterospecific and control cues showed no change. Individuals were also able to associate a previously novel scent with danger after only a single previous exposure to the paired conspecific skin extract/novel scent cue. Our results indicate that chemical alarm cues play a large role in both threat detection and learned predator recognition during the early post-settlement period in coral reef fishes.     

Infection with a trematode parasite differentially alters competitive interactions and antipredator behaviour in native and invasive crayfish

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Who's Watching Me: Female Siamese Fighting Fish Alter Their Interactions in Response to an Audience

It is well established that interactions between conspecifics are often influenced by the presence of passive bystanders. Individuals have been found to alter their behavior in a variety of contexts, from foraging to aggression, based on the presence, sex, or identity of an audience. This audience effect may influence not only the nature of a signaling event but also the evolution of signal structure as signals may have to convey information across a distance. Additionally, audience individuals may use information obtained by watching in later encounters with these individuals, which may act as a selection pressure on communication. Communication networks in Siamese fighting fish, Betta splendens, are particularly well studied, with audience effects influencing both male–male interactions and male–female interactions. However, the effects of an audience on female–female interactions have not been examined in this species or any other. This study examined the interactions of pairs of females in three different audience conditions (male, female, and no audience). The results suggest that female–female interactions are affected by the presence of an audience as interactant‐directed gill flaring, the most commonly performed behavior, was performed more with an audience present. Additionally, the sex of the audience seemed to be influential, reflected by a difference in the frequency of interactant‐directed behaviors when a female vs. a male audience was present. This study is one of the first to demonstrate that females modify their behavior as a result of being watched and stresses the importance of examining audience effects in a variety of social contexts.     

Asymmetrical Resource Ownership Increases Owners’ and Reduces Non‐Owners’ Motivation to Fight in the Mangrove Rivulus,Kryptolebias marmoratus

Resource ownership often increases an individual's aggressiveness and its probability of defeating a competitor. Individuals contesting resource owners could therefore incur higher costs, making individuals reluctant to compete with owners. We tested the hypothesis that animals use asymmetry in resource ownership as a cue for contest costs and adjust contest decisions accordingly. Using a mangrove rivulus fish (Kryptolebias marmoratus), we staged (1) contests with a randomly assigned asymmetry in resource ownership (one fish was provided with a shelter) and (2) contests in which neither fish had a shelter. Owners that were in their shelters at the contest start showed a greater tendency to fight and won more contests than their intruder opponents; those outside the shelter at the start did not. Compared with fish in contests with no shelters at stake, shelter owners had a higher tendency to fight whether or not they were in their shelters at the start; intruders, however, had a lower tendency to fight only against owners that were inside the shelter at the start. These results demonstrate (1) that ownership status influences both owners’ and intruders’ contest decisions (and in opposite directions), producing a detectable ownership advantage and (2) that intruders required confirmation of their opponents’ ownership status before retreating without challenging them. Ownership status per se is therefore important to the fish's contest decisions.     

Recent Physical Encounters Affect Chemically Mediated Retreat-Site Selection in a Gecko

Chemoreception is well documented in a range of animals but the effects of recent behavioural interactions on an animal's response to subsequent chemical information are poorly understood. Using a series of laboratory experiments with a rock‐dwelling gecko (Oedura lesueurii), we examined whether latest social experience affects how animals used chemoreception to mediate microhabitat selection. Males of this species defend retreat‐sites and the outcomes of physical duels are maintained during second physical contests staged up to 1 wk after first contests. Retreat‐site selection experiments showed that past social experience affects the behavioural responses of adult male geckos to chemical cues from conspecific male lizards. When offered a choice between retreat‐sites labelled with chemicals from an unknown male vs. the opponent from a recent contest, geckos that won their previous fight selected retreat‐sites at random. In contrast, under the same conditions, geckos that lost their previous fight significantly preferred retreat‐sites scented with an unknown male conspecific over those scented with the winning opponent. During these experiments males that lost their previous fight were significantly more vigilant, and spent greater time near the retreat‐site of the unknown male, compared with males that won their previous fight. These results support the notion that recent physical social encounters can affect the way that animals respond to chemical information from potential competitors.     

The Influence of Social Environment: Behavior of Unrelated Adults Affects Future Juvenile Behaviors

Juveniles' behaviors are often influenced by the behaviors of conspecifics. Most experimental studies of the influence of conspecific behavior vary the social environment by the presence or absence of conspecifics or investigate the impact of the outcome of social encounters (winner/loser effects) but less frequently expose individuals to variation in behavioral phenotypes present in the social environment. Based on previous work showing that juveniles of the salamander Plethodon cinereus are likely to interact frequently with non‐parental adults, I hypothesized that territorial adults in the social environment alter the future behaviors of juveniles. I measured the intracohort social behaviors of juvenile salamanders collected from two geographic areas, Michigan (MI) and Virginia (VA), before and after housing with ostensibly territorial (VA) or non‐territorial (MI) adults. There were overall effects of adult territoriality and aggression on the behavior of juveniles. However, juveniles from populations in MI were especially susceptible to behavioral modification. Compared with behaviors prior to being housed with adults, MI juveniles increased investigatory and escape behaviors in juvenile–juvenile interactions after being housed with adults that displayed territorial behaviors and decreased investigatory and escape behaviors after being housed with non‐territorial adults. This study shows that not only is a specific behavior, territoriality of adult salamanders, a social environment that modifies future juvenile behaviors, but the effects of social environment may differ between populations.     

Cross-species familiarity in shoaling fishes

Preferential association with familiar shoal mates confers a number of potentially important benefits to individuals, including improved anti-predator effects and the reduction of aggression in competitive interactions. Until now, however, familiarity has been demonstrated purely between conspecifics. Here, we present evidence that familiarity preferences can override natural preferences for conspecifics. Individual focal fishes (chub, Leuciscus cephalus) were given a choice of two stimulus shoals of the same size composed of conspecifics or of heterospecifics (minnows, Phoxinus phoxinus) in a flow tank. A series of four treatments was carried out to investigate the effects of familiarity, induced by a 15 day association between the focal fish and the stimulus fishes, on the choices made by the focal fish. Focal fishes showed a significant preference for conspecifics over heterospecifics when both stimulus shoals were composed of non-familiar individuals. Focal fishes also showed a significant preference for stimulus shoals composed of familiar fishes over stimulus shoals composed of non-familiar fishes when both shoals were conspecific and when both shoals were heterospecific. Finally, the preference of focal fishes for conspecifics disappeared when the alternative, a shoal of heterospecifics, was composed of familiar individuals. The importance of this work is discussed in the context of species interactions in free-ranging shoals.     

Linking Behavior and Community Ecology: Interspecific Aggression Provides Evidence for Competition Between a Migrant and Resident Warbler

Intraspecific aggression is known to be an important behavior structuring bird communities, but interspecific aggression has been studied less frequently. Because of a high degree of similarity in foraging niches, I hypothesized that American redstarts (Setophaga ruticilla) and Adelaide's warblers (S. adelaidae) would interact aggressively during the non‐breeding season in southwest Puerto Rico. I used a crossover experiment to determine whether these species were aggressive to heterospecifics, presenting decoys and vocalizations of these two species, along with a control, to individuals and observing their vocal and physical responses. However, the study used only a single playback and decoy for each species, limiting the generality of the conclusions. Both species responded aggressively to decoys of conspecifics and heterospecifics, while no individual responded to the control. Responses to conspecifics were stronger than responses to heterospecifics as hypothesized, although the differences were significant only in Adelaide's warblers. The interspecific aggression observed in this study, combined with previous studies showing a high degree of overlap in space use and foraging niches and probable food limitation, strongly suggests that these species are competing for food during the non‐breeding season.     

Spatial patterns of the frog Oophaga pumilio in a plantation system are consistent with conspecific attraction

The conspecific attraction hypothesis predicts that individuals are attracted to conspecifics because conspecifics may be cues to quality habitat and/or colonists may benefit from living in aggregations. Poison frogs (Dendrobatidae) are aposematic, territorial, and visually oriented—three characteristics which make dendrobatids an appropriate model to test for conspecific attraction. In this study, we tested this hypothesis using an extensive mark‐recapture dataset of the strawberry poison frog (Oophaga pumilio) from La Selva Biological Station, Costa Rica. Data were collected from replicate populations in a relatively homogenous Theobroma cacao plantation, which provided a unique opportunity to test how conspecifics influence the spatial ecology of migrants in a controlled habitat with homogenous structure. We predicted that (1) individuals entering a population would aggregate with resident adults, (2) migrants would share sites with residents at a greater frequency than expected by chance, and (3) migrant home ranges would have shorter nearest‐neighbor distances (NND) to residents than expected by chance. The results were consistent with these three predictions: Relative to random simulations, we observed significant aggregation, home‐range overlap, and NND distribution functions in four, five, and six, respectively, of the six migrant–resident groups analyzed. Conspecific attraction may benefit migrant O. pumilio by providing cues to suitable home sites and/or increasing the potential for social interactions with conspecifics; if true, these benefits should outweigh the negative effects of other factors associated with aggregation. The observed aggregation between migrant and resident O. pumilio is consistent with conspecific attraction in dendrobatid frogs, and our study provides rare support from a field setting that conspecific attraction may be a relevant mechanism for models of anuran spatial ecology.     

Wood warblers copy settlement decisions of poor quality conspecifics: support for the tradeoff between the benefit of social information use and competition avoidance

Social information use in songbird habitat selection commonly involves a conspecific attraction strategy. Individuals copy the breeding‐site choices of conspecifics, that is, bias their own settlement decisions towards sites (tracts of spatially limited habitat with similar structure) already occupied by others. In order to be adaptive, social information use has to be discriminative. Especially the decisions of good quality individuals, i.e. measuring high at observable fitness correlates, should be copied more frequently than those of poor quality individuals. It is unknown, however, whether songbirds discriminatively use conspecific presence by evaluating the quality of information providers in habitat selection. We experimentally tested whether wood warblers Phylloscopus sibilatrix selectively copied settlement decisions of conspecifics in relation to the quality of observed individuals. We also tested whether the use of social cues was influenced by the population density at a particular site in the preceding year. We found that wood warblers selectively used intraspecific social information, but in a pattern opposite to that expected based on existing hypotheses. Wood warblers copied breeding‐site choices of poor quality conspecifics and despite temporary attraction to sites where the presence of good quality individuals was simulated, they did not ultimately settle near these individuals. Population density in the preceding year did not influence settlement patterns. We argue that when making settlement decisions, wood warblers assessed the expected level of local intraspecific competition and selectively copied breeding‐site choices of conspecifics or refused to settle, depending on competitive abilities of observed individuals. This adds a novel aspect to the patterns and processes of social information use proposed thus far, and provides support for the predicted negative effect of intraspecific competition on benefit of information. Moreover, it seems that habitat selection in wood warblers is a complex decision‐making process, in which initial decisions are adjusted after acquiring more accurate information. Synthesis Social information use in songbird habitat selection commonly involves copying the breeding‐site choices of conspecifics (so‐called conspecific attraction). To be adaptive, this strategy has to be discriminative, but almost no empirical studies have tested this assertion. Our study shows that birds may selectively use social information by copying settlement decisions of poor quality conspecifics, but avoid settling near good quality individuals, likely because of their high competitive abilities. This decision‐making pattern supports the predicted, yet not experimentally tested, tradeoff between information value and cost of competition in social information use. Our study highlights also that the use of social cues in settlement decisions may be both positively and negatively biased.