Modelling Size-Dependent Cannibalism in Barramundi Lates calcarifer: Cannibalistic Polyphenism and Its Implication to Aquaculture

This study quantified size-dependent cannibalism in barramundi Lates calcarifer through coupling a range of prey-predator pairs in a different range of fish sizes. Predictive models were developed using morphological traits with the alterative assumption of cannibalistic polyphenism. Predictive models were validated with the data from trials where cannibals were challenged with progressing increments of prey sizes. The experimental observations showed that cannibals of 25–131 mm total length could ingest the conspecific prey of 78–72% cannibal length. In the validation test, all predictive models underestimate the maximum ingestible prey size for cannibals of a similar size range. However, the model based on the maximal mouth width at opening closely matched the empirical observations, suggesting a certain degree of phenotypic plasticity of mouth size among cannibalistic individuals. Mouth size showed allometric growth comparing with body depth, resulting in a decreasing trend on the maximum size of ingestible prey as cannibals grow larger, which in parts explains why cannibalism in barramundi is frequently observed in the early developmental stage. Any barramundi has the potential to become a cannibal when the initial prey size was <50% of the cannibal body length, but fish could never become a cannibal when prey were >58% of their size, suggesting that 50% of size difference can be the threshold to initiate intracohort cannibalism in a barramundi population. Cannibalistic polyphenism was likely to occur in barramundi that had a cannibalistic history. An experienced cannibal would have a greater ability to stretch its mouth size to capture a much larger prey than the models predict. The awareness of cannibalistic polyphenism has important application in fish farming management to reduce cannibalism.     

Analysis of sibling cannibalism among pike,Esox lucius,juveniles reared under semi-natural conditions

Synopsis Sibling cannibalism in pike, Esox lucius, larvae and juveniles living in outdoor rearing ponds was studied using stomach contents analysis. For the two initial densities tested (6 and 18 larvae m^–2, equivalent to 12 and 36 larvae m^–3), cannibalism was non-existent during the larval period (13 to 35 mm total length) and was observed only during the juvenile stages. Initial density of larvae influenced both the date of first detection of cannibalistic individuals and the rate of development of cannibalism in the population. At initial stocking densities of 18 larvae m^–2 (36 larvae m^–3), cannibalism was observed from 21 days after the start of exogenous feeding (mean total length: 60 mm) onwards. At a mean total length of 100 mm and for initial stocking densities of 6 and 18 larvae m^–2, (12 and 36 larvae m^–3), the average proportions of cannibals in the populations of juveniles were 7.8% and 41.3% and the cannibals accounted for 15.5% and 65.9% of the total pike biomass, respectively. In stomachs of cannibals, young pike were the dominant prey in terms of weight. Dry weights of invertebrate-prey were lower in cannibals than in non-cannibals of similar size. Cannibalism among pike juveniles was characterized by the prey being swallowed whole and head first in the vast majority of cases. There was a strong positive correlation between predator and prey size and the mouth size of a cannibal was found to be an important constraint determining maximum victim size. The overall mean ratio of pike prey length to pike cannibal length was 66.2% and the average ratio of prey head depth to predator mouth width amounted to 87.6%. Prey size selection could be demonstrated for several length-groups of cannibals. These results are compared with the characteristics of early cannibalism in other fish species.     

Predation risk suppresses the positive feedback between size structure and cannibalism

1.?Cannibalism can play a prominent role in the structuring and dynamics of ecological communities. Previous studies have emphasized the importance of size structure and density of cannibalistic species in shaping short- and long-term cannibalism dynamics, but our understanding of how predators influence cannibalism dynamics is limited. This is despite widespread evidence that many prey species exhibit behavioural and morphological adaptations in response to predation risk. 2.?This study examined how the presence and absence of predation risk from larval dragonflies Aeshna nigroflava affected cannibalism dynamics in its prey larval salamanders Hynobius retardatus. 3.?We found that feedback dynamics between size structure and cannibalism depended on whether dragonfly predation risk was present. In the absence of dragonfly risk cues, a positive feedback between salamander size structure and cannibalism through time occurred because most of the replicates in this treatment contained at least one salamander larvae having an enlarged gape (i.e. cannibal). In contrast, this feedback and the emergence of cannibalism were rarely observed in the presence of the dragonfly risk cues. Once salamander size divergence occurred, experimental reversals of the presence or absence of dragonfly risk cues did not alter existing cannibalism dynamics as the experiment progressed. Thus, the effects of risk on the mechanisms driving cannibalism dynamics likely operated during the early developmental period of the salamander larvae. 4.?The effects of dragonfly predation risk on behavioural aspects of cannibalistic interactions among hatchlings may prohibit the initiation of dynamics between size structure and cannibalism. Our predation trials clearly showed that encounter rates among hatchlings and biting and ingestion rates of prospective prey by prospective cannibals were significantly lower in the presence vs. absence of dragonfly predation risk even though the size asymmetry between cannibals and victims was similar in both risk treatments. These results suggest that dragonfly risk cues first suppress cannibalism among hatchlings and then prevent size variation from increasing through time. 5.?We suggest that the positive feedback dynamics between size structure and cannibalism and their modification by predation risk may also operate in other systems to shape the population dynamics of cannibalistic prey species as well as overall community dynamics.     

Bistability in a size-structured population model of cannibalistic fish--a continuation study

By numerical continuation of equilibria, we study a size-structured model for the dynamics of a cannibalistic fish population and its alternative resource. Because we model the cannibalistic interaction as dependent on the ratio of cannibal length and victim length, a cannibal experiences a size distribution of potential victims which depends on its own body size. We show how equilibria of the resulting infinite-dimensional dynamical system can be traced with an existing method for numerical continuation for physiologically structured population models. With this approach we found that cannibalism can induce bistability associated with a fold (or, saddle-node) bifurcation. The two stable states can be qualified as 'stunted' and 'piscivorous', respectively. We identify a new ecological mechanism for bistability, in which the energy gain from cannibalism plays a crucial role: Whereas in the stunted population state cannibals consume their victims, on average, while they are very small and yield little energy, in the piscivorous state cannibals consume their victims not before they have become much bigger, which results in a much higher mean yield of cannibalism. We refer to this mechanism as the 'Hansel and Gretel' effect. It is not related to any individual 'choice' or 'strategy', but depends purely on a difference in population size distribution. We argue that studying dynamics of size-structured population models with this new approach of equilibrium continuation extends the insight that can be gleaned from numerical simulations of the model dynamics.     

Pathogens as a factor limiting the spread of cannibalism in tiger salamanders

Summary Intraspecific predation is taxonomically widespread, but few species routinely prey on conspecifics. This is surprising as conspecifics could be a valuable resource for animals limited by food. A potential cost of cannibalism that has been largely unexplored is that it may enhance the risk of acquiring debilitating pathogens or toxins from conspecifics. We examined how pathogens affect variation in the incidence of cannibalism in tiger salamander larvae (Ambystoma tigrinum nebulosum), which occur as two environmentally-induced morphs, typicals and cannibals. Salamanders from one population were more likely than those in another to develop into cannibals, even when reared under identical conditions. Variation in the propensity to become a cannibal may be caused by variation in pathogen density. In the population with cannibals at low frequency, bacterial blooms in late summer correlated with massive die-offs of salamanders. The frequency of cannibals correlated significantly negatively with bacterial density in ten different natural lakes. In the laboratory, cannibals exposed to a diseased conspecific always preyed on the sick animal. As a result, cannibals wre more likely to acquire and die from disease than were typicals that were similarly exposed, or cannibals that were exposed to healthy conspecifics. Since conspecifics often share lethal pathogens, enhanced risk of disease may explain why cannibalism is generally infrequent. Pathogens may constrain not only the tendency to be behaviorally cannibalistic, but also the propensity to develop specialized cannibal morphologies.     

Intra‐cohort cannibalism and size bimodality: a balance between hatching synchrony and resource feedbacks

Cannibalistic interactions generally depend on the size relationship between cannibals and victims. In many populations, a large enough size variation to allow for cannibalism may not only develop among age‐cohorts but also within cohorts. We studied the implications of variation in hatching period length and initial cohort size for the emergence of cannibalism and bimodal size distributions within animal cohorts using a physiologically structured population model. We found that the development of size bimodality was critically dependent on hatching period length, victim density and the presence of a feedback via shared resources. Cannibals only gained enough energy from cannibalism to accelerate in growth when victim density was high relative to cannibal density at the onset of cannibalism. Furthermore, we found that the opportunity for early hatchers to initially feed on an unexploited resource increases the likelihood both for cannibalism to occur and size bimodality to develop. Once cannibals accelerated in growth relative to victims size bimodality, reduced victim numbers and relaxed resource competition resulted. Thus, in addition to that cannibals profited from cannibalism through energy extraction, their potential victims also benefited as the resource recovered due to cannibal thinning. To ensure recruitment success, it can be critical that a few individuals can accelerate in growth and reach a size large enough to escape size‐dependent predation and winter starvation. Hence, within‐cohort cannibalism may be a potentially important mechanism to explain recruitment variation especially for cannibalistic species in temperate climates with strong seasonality. However, the scope for size bimodality to develop as a result of cannibalism may be limited by low victim densities and size and food‐dependent growth rates.     

Onset and development of cannibalistic behaviour in early life stages of yellowtail

The onset and development of cannibalistic behaviour were observed in early life stages of yellowtail Seriola quinqueradiata . Cannibalistic behaviour was divided into four actions, i.e aim, chase, nip and ingestion. The frequency of chase was used as an index of cannibalistic behaviour because it always appeared in every sequence of cannibalism, although a sequence of cannibalistic behaviour sometimes stopped before nip or ingestion. No cannibalistic behaviour was observed during the larval phase until day 22 after hatching (when fish were 9.6mm T.L.) either in a rearing pond or in experimental tanks. The onset of cannibalistic behaviour was observed on day 23, coinciding with metamorphosis from the larval to juvenile phase, and it developed until day 39, with a tentative decrease between day 33 and day 36. This inverted peak corresponded roughly to the development of schooling behaviour after day 33, which was determined by distance to the nearest neighbour. In the rearing pond, suffocation of a cannibal by its prey, appeared from day 23. Field observation of juvenile yellowtails aggregating around floating seaweeds showed that cannibalism occurred in three out of 10 schools, in which six cannibals were found among 194 fish. Cannibalistic behaviour in early life stages of yellowtail may occur as a final phase of inter-individual interference and may have a role for size selection of a School member.     

Eat or be eaten: prevalence and impact of egg cannibalism on two-spot ladybirds, Adalia bipunctata

Ladybirds commonly engage in cannibalistic behaviour. Egg cannibalism by first instars is considered advantageous to the cannibal, because it not only results in direct metabolic gain but also a reduction in potential competitors. In this study, we quantified the effect of cannibalism on the development rate and survival of Adalia bipunctata (L.) (Coleoptera: Coccinellidae) larvae through development to the adult stage. We also assessed the synchrony of egg‐hatching in relation to laying order and compared the proportion of eggs cannibalized in egg batches laid as clusters or linearly. Larvae that had consumed a conspecific egg after hatching reached the adult stage 1.65 days earlier than those larvae that had not. Larval and pupal mortality was lower for cannibals compared to non‐cannibals; only 46% of non‐cannibalistic individuals reached the adult stage whereas 81% of cannibals pupated successfully. Egg cannibalism is undoubtedly advantageous to A. bipunctata larvae both in terms of faster development and increased survival. There is a positive correlation between laying and hatching order for eggs laid linearly or in a cluster. There was no significant difference in the proportion of eggs hatching in clusters or in a line (80 and 77%, respectively). The remaining eggs were either cannibalized or did not hatch. The ecological implications of these results are discussed with particular reference to trophic egg plasticity.     

Cannibalism in pike fry, Esox lucius L.: some experiments with fry densities

Pike fry were kept in 0.054-m³ tanks at densities of 50, 100 and 150 (277, 555, 833 fry m⁻²) for seven weeks during which the development of social and feeding behaviour was observed. Zooplankton, macro-invertebrates and perch fry were provided sequentially as food; the pike fry were allowed to feed ad libitum.
Zooplanktivorous fry stopped growing at 22 mm whilst, in the presence of abundant suitable alternative prey, 1–4% turned cannibalistic at 5 weeks of age; cannibals subsequently grew rapidly (mean 1.88 mm day⁻¹). Cannibalism ensued in all tanks when the ratio of predator size: prey size was c .2:1.
Fry tended to space-out evenly in the tanks with no overt aggression or territoriality. Behaviour was typified by remaining still for long periods, particularly subsequent to the onset of cannibalism. Cannibals were attracted by fry movements which often initiated attacks.
Daily per capita mortality rates showed no crowding effects before cannibalism but significant density-dependent mortality due to cannibals. Consumption rates of cannibals varied between 0.63 and 6.0 fry per cannibal per day. Cannibals accounted for 54–96% of daily mortality in the experimental tanks. These results are discussed in relation to proposed mechanisms of pike population density regulation.     

Preference for Cannibalism and Ontogenetic Constraints in Competitive Ability of Piscivorous Top Predators

Occurrence of cannibalism and inferior competitive ability of predators compared to their prey have been suggested to promote coexistence in size-structured intraguild predation (IGP) systems. The intrinsic size-structure of fish provides the necessary prerequisites to test whether the above mechanisms are general features of species interactions in fish communities where IGP is common. We first experimentally tested whether Arctic char (Salvelinus alpinus) were more efficient as a cannibal than as an interspecific predator on the prey fish ninespine stickleback (Pungitius pungitius) and whether ninespine stickleback were a more efficient competitor on the shared zooplankton prey than its predator, Arctic char. Secondly, we performed a literature survey to evaluate if piscivores in general are more efficient as cannibals than as interspecific predators and whether piscivores are inferior competitors on shared resources compared to their prey fish species. Both controlled pool experiments and outdoor pond experiments showed that char imposed a higher mortality on YOY char than on ninespine sticklebacks, suggesting that piscivorous char is a more efficient cannibal than interspecific predator. Estimates of size dependent attack rates on zooplankton further showed a consistently higher attack rate of ninespine sticklebacks compared to similar sized char on zooplankton, suggesting that ninespine stickleback is a more efficient competitor than char on zooplankton resources. The literature survey showed that piscivorous top consumers generally selected conspecifics over interspecific prey, and that prey species are competitively superior compared to juvenile piscivorous species in the zooplankton niche. We suggest that the observed selectivity for cannibal prey over interspecific prey and the competitive advantage of prey species over juvenile piscivores are common features in fish communities and that the observed selectivity for cannibalism over interspecific prey has the potential to mediate coexistence in size structured intraguild predation systems.     

Precopulatory Sexual Cannibalism Causes Increase Egg Case Production,Hatching Success,and Female Attractiveness to Males

Precopulatory sexual cannibalism is an extreme form of sexual conflict that can entail significant costs to the cannibalized individual and a variety of costs and benefits to the cannibal itself. Characterizing these costs and benefits is fundamental to our understanding of how this behavior evolves. Using the spider Agelenopsis pennsylvanica, we tested the reproductive consequences of precopulatory sexual cannibalism by staging cannibalization events and comparing the performance of experimental cannibals against natural cannibals (i.e., those that cannibalized on their own) and non‐cannibals. We found two performance benefits associated with precopulatory sexual cannibalism: first, experimental cannibals were more likely to produce egg cases than non‐cannibals, and second, egg cases from experimental cannibals and natural cannibals were significantly more likely to hatch than those produced by non‐cannibals. We then tested whether males were more likely to approach the webs of experimental cannibals vs. non‐cannibalistic control females. Our data demonstrate that sexual cannibalism increases female attractiveness to males. Although this result seems counterintuitive, in fact, rates of precopulatory sexual cannibalism were much lower in females that had already cannibalized their first male: 38% of sexually naïve females engaged in precopulatory sexual cannibalism, whereas only 5% of females engaged in cannibalism a second time. Thus, males that approach cannibals receive two benefits: they are less likely to be cannibalized precopula, and they have the possibility of mating with females that have a higher probability of producing viable egg cases. Taken together, our data suggest that precopulatory sexual cannibalism affords females numerous benefits and may have a hand in shaping male mate choice decisions.     

Growing large in a low grade environment: size dependent foraging gain and niche shifts to cannibalism in Arctic char

Arctic char ( Salvelinus alpinus ) living in allopatry in lakes often show a distinct size dimorphism where the large char prey upon small char. We studied the relationship between size scaling of energy intake, total metabolic costs and levels of storage energy related to the niche shift from invertebrate feeding to cannibalism in an alpine Arctic char population. Gill-net samples consisted of two distinct size modes, small invertebrate feeding char (92–172 mm total length) and large cannibalistic char (265–606 mm total length). Food consumption was estimated for a one year period using a radioactive tracer (¹³⁷Cs). The energy intake followed a hump-shaped relationship with fish size for invertebrate feeding char and increased monotonically with size for cannibalistic char. Total metabolic costs increased exponentially with size for both groups. Char feeding in the invertebrate niche had a maximum net energy gain at 140 mm total length with an upper size limit for positive net energy gain at 180 mm. For cannibals the relationship between size and net energy gain formed a hump shaped function with maximum net energy gain at 400 mm and a lower size limit for positive net energy gain at 230 mm. The size at maturation for both small and large char was close to the size for maximum energy gain in their respective niches. Immature individuals close to the upper size limit for positive net energy gain in the invertebrate niche had low levels of storage lipids but tended to have high mass specific growth rates. This may imply a reallocation from reversible (stored) energy to structural (length) growth as a strategy to transform into the size-range where the cannibalistic niche successfully could be exploited.     

Sibling cannibalism in Koi carp, Cypvinus carpio L., larvae and juveniles reared under controlled conditions

The potential impact of sibling cannibalism on the size and population structure of captive populations of Koi carp, Cyprinus carpio L., which were fed only dry foods, was determined. The rate of cannibalism was found to be positively density-dependent, with the highest rate being recorded at initial stocking densities of 40 larvae 1 (33.3% of the fish had been eaten by cannibals by day 52). Two types of cannibalism could be distinguished: type I, where the head of the prey was rejected once the caudal region of the prey had been progressively ingested; type II, where the prey were caught either head-first or tail-first and were ingested completely. The first type was limited to the period from the ninth to the 15th day of rearing, and the second type started on the 18th day. Measurement of the mouth, head and tail allowed the calculation of predicted cannibal -prey relationships. The consequences and implications of coeval sibling cannibalism in culture systems are discussed.     

Maternal cannibalistic preferences affect offspring preferences and development in Menochilus sexmaculatus Fabricius (Coleoptera: Coccinellidae): A transgenerational investigation

Cannibalism has been widely reported across taxa. However, the heritability and expression of cannibalistic traits have been least explored. The variation in the expression of cannibalism is likely to exist amongst the population affecting the propensity of cannibalism. Thus, to know whether the mother has any role in the transgenerational transmission of this trait in a ladybird beetle, Menochilus sexmaculatus, we studied the interaction between maternal and offspring prey preferences and its effect on, development duration and body weight of offspring over generations. An insignificant effect of maternal dietary history on offspring prey preference was observed across generations except for the non-cannibalistic adults who significantly preferred aphids over eggs. The long-term detrimental effect of cannibalism was found in cannibals with increased developmental duration and decreased body weight of offspring over generations. In conclusion, the results show that maternal diet did not affect the offspring preferences in M. sexmaculatus but cannibalism had a profound generational effect on the cannibalistic propensity, development and body weight of offspring across generations shows that larval dietary history and nutritional composition of prey contribute to the expression of cannibalistic behaviour across generations.     

Paternity and egg cannibalism in the ringlegged earwig Euborellia annulipes (Dermaptera: Anisolabididae)

Cannibalism is a common occurrence in nature, and many cannibals prey on relatively small and defenseless life stages, such as eggs or young juveniles. Such behavior provides many benefits to the cannibal, but cannibalistic individuals also face risks, including the cost of decreasing their inclusive fitness by eating close relatives such as siblings or offspring. This risk can be mitigated if cannibals can recognize and avoid preying on kin. Here, we tested whether male ringlegged earwigs Euborellia annulipes avoid cannibalizing eggs that they had sired. In this species, females care for their own eggs, but males provide no care and frequently prey upon eggs. We found that when presented with an unguarded clutch of eggs, male earwigs nearly always cannibalized some eggs, but that the proportion of eggs eaten was smaller if the male had sired the clutch. This suggests that males can distinguish between their own offspring and unrelated offspring and that they avoid harming their kin.     

Does Female Personality Determine Mate Choice Through Sexual Cannibalism?

Animal personalities (e.g. consistent across‐context behavioural differences between individuals) can lead to differences in mate choice. However, evidence for this link remains limited. Pre‐mating sexual cannibalism can be a behavioural syndrome (i.e. a suboptimal personality) in which adaptive female aggression towards heterospecific prey spills over on non‐adaptive aggression towards courting males, independently of the female mating or feeding status (i.e. the ‘aggressive spillover hypothesis’, ASH). On the other hand, sexual cannibalism can also be a form of mate choice by which females selectively kill or mate with males depending on the male phenotype. We introduce the hypothesis that the most aggressive females in the population will not only attack males more frequently, but will be less likely to impose sexual selection on males through sexual cannibalism. Assuming that in a field common garden experiment in which females were fed ad libitum the rate of weight gain by a female may reflect her voracity or aggressiveness, we show that in the cannibalistic burrowing wolf spider Lycosa hispanica (formerly L. tarantula), voracity towards heterospecific prey predicts a female's tendency towards sexual cannibalism. Unmated females with higher weight gains were more cannibalistic and attacked males regardless of the male phenotype. On the other hand, females that were less voracious tended to be less cannibalistic, and when they did kill a male, they were selective, killing males in poorer condition and mating with those in better condition. Our results demonstrate that females with different phenotypes (growth rates) differently imposed selection on male condition, tentatively supporting the hypothesis that female aggression levels can spill over on sexual selection through sexual cannibalism.     

The adaptive significance of polymorphism in the rotifer Asplanchna. Humps in males and females

Summary Experiments show that humps or body wall outgrowths protect males and cruciform females of Asplanchna sieboldi from predation by conspecific, cannibal females. Humped prey were eaten by large cannibals to a significantly lesser extent than similarly-sized or even larger, non-humped prey.Since males and male-producing females, which are typically cruciform, co-occur with the largest, most cannibalistic morphotypes, their protection by humps may greatly increase the efficiency of sexual reproduction.An analysis of female diets and the occurrence of humped males and females in the different species of Asplanchna shows that these humped forms are found only in those species in which the females can eat comparatively large prey. This relationship provides strong indirect evidence for the theory that humps in both males and polymorphic females specifically evolved as a defense mechanism against predation by conspecific, cannibal females.Supported by National Science Foundation Grant GB-32182.     

Emergent impacts of cannibalism and size refuges in prey on intraguild predation systems

Many organisms undergo ontogenetic niche shifts due to considerable changes in size during their development. These ontogenetic shifts can alter the trophic position of individuals, the type and strength of ecological interactions across species, and allow for cannibalism within species. In this study we ask if and how the interaction of a size refuge and cannibalism in the prey alters the dynamics of intraguild predation (IGP) systems. By manipulating the composition of large cannibalistic (Aeshna umbrosa) and predatory (Anax junius) dragonfly larvae in mesocosms we show that the interaction of cannibals and predators was non-linear and increased the survival of prey. The structure of the final resource community shared by prey and predator differed between small and large dragonfly treatments but not within size classes across species. In general, the small prey stage showed similar shifts in microhabitat use and refuge use when exposed to either conspecific cannibals or predators, while large cannibals showed no clear anti-predator response. However, further behavioral experiments revealed that specific behavioral components, such as distances between individuals or number of movements, differed when individuals were exposed to either cannibals or predators. This indicates that individuals discriminated between conspecific or heterospecific predators. Furthermore, in similar experiments large cannibals and predators showed different behaviors when exposed to conspecifics rather than to each other. These changes in behavior are consistent with the observed increase in prey survival. In general, the results indicate that cannibalism and ontogenetic niche shifts can result in behavior-mediated indirect interactions that reduce the impact of the predator on the mortality of its prey and alter the interactions of IGP systems. However, they also indicate that size is not the sole determinant and that we also need to account for the species identity when predicting the dynamics of communities.     

The potential for intracohort cannibalism in age-0 walleye pollock,Theragra chalcogramma,as determined under laboratory conditions

Synopsis In laboratory experiments, we tested the capability of larger age-0 walleye pollock to consume smaller members of their cohort. In separate aquaria, 81 pairs of juveniles covering a wide range of size differences (total lengths differing by 12 to 61 mm) were held and monitored over a 4 day period. Complete consumption, in which a smaller fish was swallowed whole by a larger fish, occurred 11% of the time. In 36% of the pairs, attacks by the larger fish resulted in mortality of the smaller fish. The mouth width:body depth ratio between the larger and smaller fish of a pair differed significantly depending on whether the smaller fish survived, was killed but not consumed, or was ingested whole by the larger fish. Cannibalistic individuals could consume fish close to the maximum size physically possible under gape limitation; at this size the length of the cannibal was approximately 1.7 times the length of the prey. Length-frequency distributions of age-0 pollock in field concentrations suggested that, at least in some geographical areas, potential cannibals and prey commonly co-occur. Unsuccessful predatory attacks by larger individuals may have additional detrimental effects on smaller pollock in natural populations.     

Learned Recognition of Intraspecific Predators in Larval Long-Toed Salamanders Ambystoma macrodactylum

The ability of prey to detect predators and respond accordingly is critical to their survival. The use of chemical cues by animals in predator detection has been widely documented. In many cases, predator recognition is facilitated by the release of alarm cues from conspecific victims. Alarm cues elicit anti‐predator behavior in many species, which can reduce their risk of being attacked. It has been previously demonstrated that adult long‐toed salamanders, Ambystoma macrodactylum, exhibit an alarm response to chemical cues from injured conspecifics. However, whether this response exists in the larval stage of this species and whether it is an innate or a learned condition is unknown. In the current study, we examined the alarm response of naïve (i.e. lab‐reared) larval long‐toed salamanders. We conducted a series of behavioral trials during which we quantified the level of activity and spatial avoidance of hungry and satiated focal larvae to water conditioned by an injured conspecific, a cannibal that had recently been fed a conspecific or a non‐cannibal that was recently fed a diet of Tubifex worms. Focal larvae neither reduced their activity nor spatially avoided the area of the stimulus in either treatment when satiated, and exhibited increased activity towards the cannibal stimulus when hungry. We regard this latter behavior as a feeding response. Together these results suggest that an anti‐predator response to injured conspecifics and to cannibalistic conspecifics is absent in naïve larvae. Previous studies have shown that experienced wild captured salamanders do show a response to cannibalistic conspecifics. Therefore, we conducted an additional experiment examining whether larvae can learn to exhibit anti‐predator behavior in response to cues from cannibalized conspecifics. We exposed larvae to visual, chemical and tactile cues of stimulus animals that were actively foraging on conspecifics (experienced) or a diet of Tubifex (naïve treatment). In subsequent behavioral treatments, experienced larvae significantly reduced their activity compared to naive larvae in response to chemical cues of cannibals that had recently consumed conspecifics. We suggest that this behavior is a response to alarm cues released by consumed conspecifics that may have labeled the cannibal. Furthermore, over time, interactions with cannibals may cause potential prey larvae to learn to avoid cannibals regardless of their recent diet.