Predator cannibalism can shift prey community composition toward dominance by small prey species

Cannibalism among predators is a key intraspecific interaction affecting their density and foraging behavior, eventually modifying the strength of predation on heterospecific prey. Interestingly, previous studies showed that cannibalism among predators can increase or reduce predation on heterospecific prey; however, we know less about the factors that lead to these outcomes. Using a simple pond community consisting of Hynobius retardatus salamander larvae and their associated prey, I report empirical evidence that cannibalism among predators can increase predation on large heterospecific prey but reduce that on small heterospecific prey. In a field‐enclosure experiment in which I manipulated the occurrence of salamander cannibalism, I found that salamander cannibalism increased predation on frog tadpoles but reduced that on aquatic insects simultaneously. The contrasting effects are most likely to be explained by prey body size. In the study system, frog tadpoles were too large for non‐cannibal salamanders to consume, while aquatic insects were within the non‐cannibals’ consumable prey size range. However, when cannibalism occurred, a few individuals that succeeded in cannibalizing reached large enough size to consume frog tadpoles. Consequently, although cannibalism among salamanders reduced their density, salamander cannibalism increased predation on large prey frog tadpoles. Meanwhile, salamander cannibalism reduced predation on small prey aquatic insects probably because of a density reduction of non‐cannibals primarily consuming aquatic insects. Body size is often correlated with various ecological traits, for instance, diet width, consumption, and excretion rates, and is thus considered a good indicator of species’ effects on ecosystem function. All this considered, cannibalism among predators could eventually affect ecosystem function by shifting the size composition of the prey community.     

Comparative Behavioural Studies of Larval and Adult Stages of the Phytoseiids (Acari: Mesostigmata) Typhlodromus Athiasae and Neoseiulus Californicus

We compared the behaviours of the indigenous Typhlodromus athiasae Porath and Swirski and the exotic Neoseiulus californicus (McGregor) (= Amblyseius chilenensis Dosse) relative to their persistence in apple orchards in Israel. We studied (1) larval feeding, walking, intraspecific interactions (cannibalism, touch-avoidance responses and/or touching with palps and tarsi) and tendency to aggregate (when resting), (2) predation and cannibalism on phytoseiid eggs by young females and (3) the effects of starvation for 10 days on young females relative to ambulation speed, longevity, fecundity, progeny survival and sex ratio. Larvae of T. athiasae were almost inert, did not feed and hardly walked or interacted whereas larvae of N. californicus fed, walked and interacted, mainly by touching with palps and tarsi. No cannibalism in the larval stage was observed for either species. The presence of prey increased the larval walking and intraspecific interactions of N. californicus but not of T. athiasae. Egg predation by adult females of both species was substantially higher than cannibalism, implying that both are capable of distinguishing their eggs from those of other species. Soaking eggs for 30 min in deionized water increased cannibalism in both species. During the 10 days of starvation, the ambulation speed of adult female N. californicus ranged from 1.8 to 10.1 times that of T. athiasae. The 50% lethal time value (LT₅₀) of T. athiasae (6.0 days) was significantly lower than that of N. californicus (10.4 days). None of the starved T. athiasae recuperated following the reintroduction of prey, whereas 75% of N. californicus did and oviposited after 2 days. These traits should enable N. californicus to persist when prey is scarce; however the selective predation of N. californicus eggs by T. athiasae could prevent establishment of N. californicus. The degree of specialization of these two predators is discussed.     

Effect of cannibalism on predation,oviposition and longevity of the predacious mite,Agistemus exsertus Gonzalez (Acari: Stigmaeidae)

Cannibalism (intraspecific predation) on conspecific eggs was investigated in the predatory stigmaeid mite, Agistemus exsertus Gonzalez in the absence of eggs of Tetranychus urticae Koch (no-choice tests) and presence of three densities of prey eggs simultaneously (choice tests) in the laboratory. Data show that cannibalism occurs in immatures and adult females of the predator, which successfully developed and reproduced on conspecific eggs as an alternative prey in the absence of prey eggs. In no-choice tests, cannibalism rate on conspecific eggs by A. exsertus stages was significantly lower than the predation rate on T. urticae eggs. The predatory mite exhibited a marked decline in oviposition rate when preyed on conspecific eggs compared with feeding on prey eggs. The developmental duration and longevity of A. exsertus females were significantly longer 1.9 and 1.7 times, respectively, when fed on conspecific eggs than feeding on T. urticae eggs. The propensity of the predator towards cannibalism depends on the prey density, when T. urticae eggs and conspecific eggs are present simultaneously. Provision of increased densities of prey eggs significantly decreased cannibalism and predation by A. exsertus stages, which fed generally less on conspecific eggs than on T. urticae eggs in choice tests. The oviposition rate of the predator increased significantly, as the egg density of the prey increased. The developmental period and longevity of A. exsertus females showed significantly gradual shortness with increasing egg density of the prey.     

Threat of predation alters aggressive interactions among spotted salamander (Ambystoma maculatum) larvae

Intraspecific aggression represents a major source of mortality for many animals and is often experienced alongside the threat of predation. The presence of predators can strongly influence ecological systems both directly by consuming prey and indirectly by altering prey behavior or habitat use. As such, the threat of attack by higher level predators may strongly influence agonistic interactions among conspecifics via nonconsumptive (e.g., behaviorally mediated) predator effects. We sought to investigate these interactions experimentally using larval salamanders (Ambystoma maculatum) as prey and dragonfly nymphs (Anax junius) as predators. Specifically, we quantified salamander behavioral responses to perceived predation risk (PPR) from dragonfly nymphs and determined the degree to which PPR influenced intraspecific aggression (i.e., intraspecific biting and cannibalism) among prey. This included examining the effects of predator exposure on the magnitude of intraspecific biting (i.e., extent of tail damage) and the resulting change in performance (i.e., burst swim speed). Salamander larvae responded to PPR by reducing activity and feeding, but did not increase refuge use. Predator exposure did not significantly influence overall survival; however, the pattern of survival differed among treatments. Larvae exposed to PPR experienced less tail damage from conspecifics, and maximum burst swim speed declined as tail damage became more extensive. Thus, escape ability was more strongly compromised by intraspecific aggression occurring in the absence of predation risk. We conclude that multitrophic indirect effects may importantly modulate intraspecific aggression and should be considered when evaluating the effects of intraspecific competition.     

A Test for Male Parental Care in a Fundulid, the Bluefin Killifish, Lucania goodei

Under field conditions, breeding male bluefin killifish, Lucania goodei, have been observed aggressively defending territories from other breeding males, non-breeding females, and minnows (mainly Notropis harperi). We performed an aquarium experiment to test whether male aggression serves to protect newly deposited eggs from predation. We allowed a male and a female to spawn in a yarn mop, removed the female, and exposed the eggs to one of four treatments (spawning male present, two minnows present, spawning mal+two minnows present, no adult fish present). Mops were censused daily for seven days. Egg predation rates were highest in the male+minnows and male only treatments. Egg predation rates in the male+minnows treatment did not differ from the predicted predation rate (sum of male only and minnows only treatments). Hence, there is no evidence for male parental care in L. goodei. In addition, we compared the egg predation rates (filial cannibalism) between males of 3 different color morphs and found no evidence for differential egg cannibalism.     

Adult cannibalism in an oligophagous herbivore,the Colorado potato beetle

Cannibalism, or intraspecific predation, can play a major role in changing individual fitness and population processes. In insects, cannibalism frequently occurs across life stages, with cannibals consuming a smaller or more vulnerable stage. Predation of adult insects on one another is considered to be uncommon. We investigated adult cannibalism in the Colorado potato beetle, Leptinotarsa decemlineata (Say), which is an oligophagous herbivore specializing on plants in family Solanaceae, and an important agricultural pest. Under laboratory conditions, starvation and crowding encouraged teneral adults to feed upon each other, which reduced their weight loss during the period of starvation. However, pupae were attacked and consumed before adults. Injured beetles had a higher probability of being cannibalized than intact beetles. Males were more frequently attacked than females, but that appeared to be a function of their smaller size rather than other gender‐specific traits. Cannibalizing eggs at a larval stage did not affect beetle propensity to cannibalize adults at an adult stage. When given a choice between conspecific adults and mealworms, the beetles preferred to eat conspecifics. Cannibalistic behavior, including adult cannibalism, could be important for population persistence in this species.     

A survey of intraspecific predation within the class Mammalia

The occurrence of cannibalism in mammals was studied during a general review of cannibalism (Polis, 1981). In total, there were 146 references documenting intraspecific predation in 75 species of mammals distributed between seven orders. Of these references, 138 refer to cannibalism whereas eight studies refer to intraspecific killing unaccompanied by cannibalism. Of the papers that specified the identity of both the predator and the prey, approximately 80% referred to infanticide (including cannibalism). It appears that infanticide often functions as part of a reproductive strategy. Juveniles and adults are occasional intraspecific prey for some species of Insectivora, Chiroptera, Primates, Lagomorpha, Rodentia, Carnivora, and Artio-dactyla. Cannibalism was best documented for the carnivores, rodents and primates. In general, cannibalism often occurs during normal predatory activities and is a function of low food availability, environmental stress, and a high density of conspecifics.     

Density-dependent predation and implications for stock enhancement with Japanese flounder

In laboratory predation trials kinsen-gani crab Matuta lunaris rarely preyed on hatchery-reared (HR) Japanese flounder Paralichthys olivaceus , even at extremely elevated prey densities ( c . 90 fish m⁻²), but fed readily on sacrificed Japanese founder. This suggests that under similar conditions in the wild predator-prey interactions between kinsen-gani and juvenile Japanese flounder are likely to be weak. Larger Japanese flounder readily cannibalized smaller conspecifics, suggesting that within-release group cannibalism may contribute considerably to post-release mortality of HR Japanese flounder when sufficient size variation exists within groups of released fish. Kinsen-gani presence had no effect on within-release group cannibalism rates of large Japanese flounder on small Japanese flounder. Within-release group cannibalism rates were significantly affected by densities of smaller (prey) fish, implying that the number of fish released (and thus post-release densities) in stock enhancement programmes may significantly affect post-release mortality rates. The probability of within-release group cannibalism increased rapidly as the cannibal-to-prey size ratio exceeded 1.6, suggesting that minimizing size variation of released HR individuals will reduce post-release cannibalism rates. These results have implications for Japanese flounder stock enhancement release strategies, and may be applicable to other finfish stock enhancement programmes.     

Cannibalism as a selective force on offspring size in fish

Cannibalism may cause considerable mortality on juvenile fish and it has been hypothesised that it may exercise selection on offspring size in that larger offspring may enjoy a size refuge. For this to be evolutionarily advantageous the survival of individual offspring must compensate for the reduced fecundity implied by larger offspring size. We develop a model which combines standard assumptions of size‐dependent mortality with adult cannibalism to investigate the potential for cannibalism to act as selective force on offspring size. We find that for this potential to be realised, the mortality due to cannibalism must exceed a threshold value that is a decreasing function of non‐cannibalistic predation intensity, cannibalized size range width and the average cannibalized size. If cannibalism exceeds this threshold, the model predicts evolution of offspring size towards refuges above or below cannibalized size range depending on initial offspring size. Cannibalistic mortality cannot be so great that the population is non‐viable, however, the range of parameter values describing cannibalistic intensity allowed within these boundaries is wide. On this basis, we suggest that cannibalism is a potential mechanism for offspring size selection.     

Effect of emigration on cannibalism and intraguild predation in aphidophagous ladybirds

Abstract.  1. The incidence and timing of emigration, cannibalism, and intraguild predation of larvae of three aphidophagous ladybirds (Coleoptera: Coccinellidae), Harmonia axyridis Pallas, Coccinella septempunctata brucki Mulsant, and Propylea japonica Mulsant, relative to the presence of prey was determined in the laboratory in single- and mixed-species populations.
2. In single-species populations, 80% of the larvae of C. s. brucki emigrated prior to the extinction of the aphid population and no larvae were lost due to cannibalism; however > 80% of the larvae of the other two species were still present when the aphid became extinct and the losses due to cannibalism for H. axyridis and P. japonica were 25% and 14% respectively. Finally, 28% of the P. japonica larvae completed their development, whereas no larvae of the other two species became adult.
3. In mixed-species populations, mortality of P. japonica attributable to cannibalism or intraguild predation increased greatly to 60%, whereas that of the other two species remained about the same. Consequently, survival of H. axyridis larvae improved and survival of P. japonica worsened; however the survival of C. s. brucki larvae was not affected by the other two species. Early emigration by C. s. brucki larvae may have enabled them to escape intraguild predation by H. axyridis in this system.     

Role of surface chemicals in egg cannibalism and intraguild predation by neonates of two aphidophagous ladybirds, Propylea dissecta and Coccinella transversalis

Abstract:  The role of surface chemicals in egg cannibalism and intraguild predation by neonates of two aphidophagous ladybirds, Propylea dissecta and Coccinella transversalis were examined. Neonates of both species prefer to eat non-sibling conspecific eggs than heterospecific eggs, with higher preference in P. dissecta . Surface chemicals appear to play a major role for the preference of conspecific eggs, as the ladybird behaviour was reversed when these chemicals were interchanged. The surface chemicals present on the eggs possibly act as feeding stimulants to conspecific neonates but not for heterospecific neonates. Egg clustering enhances the effect of surface chemical and stimulate non-sibling egg cannibalism and appears to reduce intraguild predation. Egg clustering is advantageous to ladybirds, as it can stimulate non-sibling cannibalism by neonates. It appears that aggregation of chemicals present on the conspecific egg surface possibly attracts the hungry conspecific neonate, providing the first meal for its survival, while protecting against heterospecific predators. Neonates of both ladybirds were reluctant to eat heterospecific eggs.     

Genetic control of geographic variation for cannibalism in the southwestern corn borer,Diatraea grandiosella

The southwestern corn borer,Diatraea grandiosella Dyar, is a subtropical insect whose range has expanded northward within this century. Geographic variation exists between populations at the extremes of the range for a suite of morphological and behavioral characters, including cannibalism. Laboratory colonies established from a Missouri population (37°N lat.) with a high cannibalism rate and a Mexican population (19° N lat.) with a low cannibalism rate were used to examine the genetic basis of cannibalism by means of a controlled breeding design. When larvae were held in pairs on artificial diet in 30 ml plastic cups at 30° C 16L:8D, the highest incidence of cannibalism in both Missouri and Mexican populations was found in larvae 15 to 18 days-of-age (5th and 6th instars). Under these conditions, cannibalism was expressed in 33% of the pairs formed from Missouri larvae and 11% of the pairs formed from Mexican larvae. First and second generation crosses between Missouri and Mexican populations showed an intermediate level of cannibalism, whereas backcrosses to the parental generations showed a regression to the parental phenotypes. These results indicate that cannibalism is under additive genetic control. Larvae from the Missouri parental population were more likely to consume their victim (i.e., cannibalize) once intraspecific killing had occurred than were larvae from the Mexican parental population or from any of the crosses, and female larvae were cannibals more often than were males. Under the conditions employed, no advantage was detected for cannibalism over instraspecific killing for larval weight gain or growth rate.     

Optimal reproductive phenology under size‐dependent cannibalism

Intra‐cohort cannibalism is an example of a size‐mediated priority effect. If early life stages cannibalize slightly smaller individuals, then parents face a trade‐off between breeding at the best time for larval growth or development and predation risk from offspring born earlier. This game‐theoretic situation among parents may drive adaptive reproductive phenology toward earlier breeding. However, it is not straightforward to quantify how cannibalism affects seasonal egg fitness or to distinguish emergent breeding phenology from alternative adaptive drivers. Here, we devise an age‐structured game‐theoretic mathematical model to find evolutionary stable breeding phenologies. We predict how size‐dependent cannibalism acting on eggs, larvae, or both changes emergent breeding phenology and find that breeding under inter‐cohort cannibalism occurs earlier than the optimal match to environmental conditions. We show that emergent breeding phenology patterns at the level of the population are sensitive to the ontogeny of cannibalism, that is, which life stage is subject to cannibalism. This suggests that the nature of cannibalism among early life stages is a potential driver of the diversity of reproductive phenologies seen across taxa and may be a contributing factor in situations where breeding occurs earlier than expected from environmental conditions.     

A comparative study of cannibalism and predation in seven species of flour beetle

Abstract 1. The present study quantified egg and pupal cannibalism, and interspecific predation on eggs and pupae, by larvae and adults of seven species of flour beetle (Tribolium spp.) under laboratory conditions: T. anaphe, T. brevicornis, T. castaneum, T. confusum, T. destructor, T. freemani, and T. madens. 2. Variation among species in cannibalism and predation propensities did not reflect taxonomic affinities within the genus, indicating that these behaviours were shaped by ecology at species level. 3. Within species, larvae and adults displayed different propensities for cannibalism and predation, leading to the conclusion that these behaviours evolve independently in the two life stages. 4. All species behaved as intraguild predators to some degree, especially in the adult stage. 5. Three general patterns of cannibalism and predation were described by principal component mapping and cluster analysis. 6. The first group comprised three cosmopolitan pest species that were more voracious as adults than as larvae: T. castaneum, T. confusum, and T. destructor. It is proposed that stored product environments select for high adult voracity because the costs associated with emigration from such rare, but resource‐rich, habitats intensifies interference competition among adults. 7. The second group consisted of species that inhabit natural environments and that were more voracious as larvae: T. anaphe, T. freemani, and T. madens. Habitats for these species are probably numerous, but generally poor in quality, a situation that intensifies larval competition, while favouring earlier adult emigration. 8. The largest species, T. brevicornis, demonstrated inconsistent voracity between life stages and was the only species with chemically defended pupae. 9. It is proposed that consumption of eggs provides primarily nutritional benefits, whereas consumption of pupae has a more important role in interference competition.     

Male opportunistic mating increases with intensity of female sexual cannibalism in 3 web-building spiders

Sexual conflict is common in animals, and female sexual cannibalism represents an extreme form of sexual conflict. Males in many species have evolved a variety of strategies to circumvent or decrease the risk of female sexual cannibalism. Opportunistic mating, by which a male mates with a female when she is disturbed or when she is feeding or undertaking moulting, is one of such kinds of strategies, and widely occurs in many animals, especially in spiders. However, whether the occurrence of male opportunistic mating depends on the intensity of female sexual cannibalism remains largely unexplored. We predicted a positive correlation between them. In this study, we tested this prediction by performing a series of mating trials in the laboratory using 3 species of web-building spiders with different intensities of female sexual cannibalism: Nephila pilipes, Nephilengys malabarensis, and Parasteatoda tepidariorum. We found that the occurrence of male opportunistic mating was positively, though not statistically significantly, correlated with the intensity of female sexual cannibalism, thus supporting our hypothesis. All together, we provide evidence that male opportunistic mating may have evolved to respond to the selection pressure posed by female sexual cannibalism.     

Underwater sound emission as part of an antipredator mechanism in Ceratophrys cranwelli tadpoles

We report the emission of underwater sounds in the tadpoles of a second member of the family Ceratophryidae, Ceratophrys cranwelli. These tadpoles produce a short metallic‐like sound, which consists of short trains of pulses at Gosner stages 25, 28, and 37. Experiment I was designed to record underwater sounds and their characteristics. Experiment II was designed to test: (i) if at higher densities (total number of tadpoles/L) but fixed predator–prey proportions C. cranwelli larvae are cannibalistic, (ii) if cannibalism increases at higher proportions of predators at a fixed density, and (iii) if tadpoles display a mechanism of intraspecific recognition that may diminish the frequency of cannibalism. Each treatment combines larvae of C. cranwelli (predator) with those of Rhinella arenarum (prey). The number of live and dead individuals was recorded during 72 h, and the following variables were calculated: time to eat the first and second prey, time without eating, time to eat a congener, and number of events of cannibalism. The results indicate that relative predator–prey availability affects the frequency of predation between conspecifics. We consider that an antipredator mechanism exists in C. cranwelli tadpoles and that the underwater sound is part of it.     

Aggression and risk assessment during predatory interactions between first-instarSinea diadema (Fabricius) (Hemiptera: Reduviidae)

The behaviors exhibited by first-instarSinea diadema (F.) (Hemiptera: Reduviidae) during intraspecific interactions are described. Five distinct behaviors were characterized: posturing, striking, stalking, stridulating, and beaking. Distinct bouts of grappling were also noted. Interactions between unfed 2-day-old nymphs, unfed 3-day-old nymphs, and unfed 5-day-old nymphs were recorded and analyzed. The behavior of the nymphs changed with the duration of food deprivation. As the period of food deprivation increased from 2 to 5 days, the frequency of posturing decreased as the frequency of beaking increased. During noncannibalistic interactions, posturing or striking by one nymph was most frequently followed by posturing by the other nymph. In contrast, during cannibalistic interactions, posturing or striking was usually followed by beaking. The incidence of grappling increased with the duration of food deprivation. Moreover, grappling always preceded cannibalism. Although first instars were capable of capturing and killing conspecifics of the same age, cannibalism occurred only after 4 days of food deprivation. These results suggest that a decision-making process is involved. During interactions the nymphs can assess the vulnerability of the opponent. Whereas recently emerged nymphs usually forego the risk of attacking conspecifics, the attacks made by nymphs deprived of food are more persistent despite the danger of predation by their opponent.     

Sibling cannibalism among juvenile vundu under controlled conditions. I. Cannibalistic behaviour, prey selection and prey size selectivity

Sibling cannibalism among vundu Heterobranchus longifilis larvae started at the age of 4 days, with the prey caught tail-first then swallowed up to the head, which was eventually discarded (type I cannibalism). At 17 days old, this type of cannibalism vanished and was replaced by the ingestion of the whole prey (type II cannibalism), which could only be exerted by predators six times as heavy as their prey. Type II cannibalism consisted of a seemingly opportunistic ambush attack by a formerly passive predator towards a disorientated prey. It required no preliminary aggression or chase, or even contact with the prey, suggesting that the attack was not mediated by the tactile sense, and that cannibalism was independent of aggressive behaviour. When alternative food resources (formulated feed, live tilapia prey) were available, the intensity of cannibalism decreased but pellet-eaters or tilapia predators always achieved lower growth rates than those feeding on conspecifics, suggesting that cannibalism was the most advantageous foraging tactic. Losses to cannibalism among populations of 30-day old juvenile vundu with an initial ratio of 4% of cannibals were as high as 75·5–79·9% over 15 days. Predation peaked during the first days (up to 2·8 prey C⁻¹ day⁻¹), then vanished progressively as surviving prey grew faster than cannibals and escaped their predation. Cannibals preferred consuming the largest prey available with respect to the logistics of cannibalism (body weight ratio of 6·0). This preference for large prey was interpreted both as a foraging tactic aiming to maximize the energetic return, and as foraging strategy enabling the cannibals to exploit their prey as long as possible. Based on these data, comprehensive models of the impact of cannibalism on vundu populations were developed.     

Cohabitation of juvenile females with mature males promotes sexual cannibalism in fishing spiders

Precopulatory sexual cannibalism, or female predation of a potentialmate before mating, illustrates an extreme intersexual conflict.Unlike sexual cannibalism occurring after mating, precopulatorysexual cannibalism cannot be construed as a male strategy. Thus,research on the adaptive significance of this phenomenon hasfocused on female benefits. In the present study, I test theidea that precopulatory sexual cannibalism represents an adaptivefemale trade-off between the material costs and benefits ofmating with a male (forgoing food, securing sperm) and preyingon a male (forgoing sperm, securing food). I pay particularattention to the rarely tested prediction that precopulatorysexual cannibalism by virgin females should increase as eachfemale's expectation of future mating opportunities increases.I use the phenomenon of cohabitation between adult males andjuvenile females nearing sexual maturity as a means to manipulatefemale expectation of future mate availability. Results indicatethat feeding on a male has significant positive effects on severalmeasures of female fecundity. However, the likelihood of precopulatoryattacks was not explained by a female's recent feeding history.Finally, as predicted, juvenile female cohabitation with maturemales (expectation of future mating opportunities) heightensthe prevalence of precopulatory attacks by virgin females, suggestingthat juvenile experience can alter a female's propensity forsexual cannibalism. This is the first study to suggest thatjuvenile experience can alter a female's propensity for sexualcannibalism. This finding emphasizes the point that studiesof sexual selection and mating systems need to consider theeffects of juvenile experience on adult behavior.