Ibanex, C. M. and F. Keyl. 2010. Cannibalism in cephalopods. Review In Fish Biology and Fisheries 20: 123-136.

Cannibalism refers to the action of consuming a member of the same species and is common in many taxa. This paper reviews the available literature on cannibalism in cephalopods. All species of the class Cephalopoda are predators and cannibalism is common in most species whose diet has been studied. Cannibalism in cephalopods is density-dependent due to their aggressive predatory and in case of the octopuses territorial nature. It also depends upon local and temporal food availability and of the reproductive season. Cannibalistic behaviour is positively related to the size of both cannibal and victim. It can affect population dynamics of cephalopods in periods of low food availability and/or high population abundance. Cephalopods are generally restricted in their ability to store energy. It is thus assumed that cannibalism is part of a population energy storage strategy enabling cephalopod populations to react to favourable and adverse environmental conditions by increasing and reducing their number. Finally, we propose five orientation points for future research on cannibalism in cephalopods. Richardson, M. L. Mitchell, R. F. Reagel, P. F. and L. M. Hawks. 2010. Causes and Consequences of Cannibalism in Noncarnivorous Insects. Annual Review in Entomology 55: 39-53. We review the primary literature to document the incidence of cannibalism among insects that typically are not carnivorous. Most of the cannibalistic species were coleopterans and lepidopterans, and the cannibals often were juveniles that aggregate or that overlap in phenology with the egg stage. Cannibalism can be adaptive by improving growth rate, survivorship, vigor, longevity, and fecundity. It also can play an important role in regulating population density and suppressing population outbreaks, stabilizing host plantinsect relationships, and reducing parasitism rates. Cannibalism often was favored by density-dependent factors for herbivores that feed in concealed feeding situations (such as stem borers, leafminers), but also by density-independent factors (such as high ambient temperature) for herbivores that feed in exposed feeding situations. Banerji, A. and P. J. Morin. 2009. Phenotypic plasticity, intraguild predation and anti-cannibal defences in an enigmatic polymorphic ciliate. Functional Ecology 23: 427-434. 1. Inducible trophic polymorphisms are greatly underappreciated forms of phenotypic plasticity that allow organisms to respond dynamically to the environmental variation by enabling them to change the trophic level upon which they feed. Although inducible trophic polymorphisms occur in a diverse array of organisms, their costs, benefits and their consequences for long-term population and community dynamics are poorly understood. 2. We studied the inducible trophic polymorphism of the freshwater hymenostome ciliate Tetrahymena vorax, whose isogenic populations can contain three distinct morphs: pyriform, bacterivorous microstomes; larger, carnivorous macrostomes; and elongate, 'tailed' microstomes. We tested whether (i) the tailed microstome constitutes an inducible defence against macrostomes and (ii) the transformation of microstomes into macrostomes is size-dependent. We also describe the dynamics of the three morphs in the presence and absence of an intraguild prey (Colpidium) across a gradient of growth medium concentrations to infer potential trade-offs in the success of different morphs at different productivity levels. 3. Macrostomes do not discriminate between pyriform microstomes and readily consumed heterospecific prey (Colpidium). Tailed microstomes display greatly reduced susceptibility to consumption by macrostomes as compared with undefended, pyriform microstomes. Morph dynamics are consistent with the hypothesis that tailed microstomes function as an inducible defence against cannibalism; tailed microstomes and macrostomes appear simultaneously, in both the presence and absence of Colpidium. At low productivity, T. vorax achieves higher rates of growth when feeding on

Colpidium instead of on bacteria. At higher productivity, this pattern is reversed, with growth rates maximized in the absence of Colpidium. 4. The reduced consumption rate of tailed microstomes by cannibalistic macrostomes, together with the simultaneous induction of tailed microstomes and macrostomes, suggests that both morphs comprise a coordinated adaptive response to the presence of intraguild prey. Hawley, T. J. 2009. The Ecological Significance and Incidence of Intraguild Predation and Cannibalism among Anurans in Ephemeral Tropical Pools. Copeia 4: 748-757. Ephemeral pools are traditionally considered to be tadpole refuges, with few or no predators. While it is recognized that some tadpoles are predaceous, little attention has been given to understanding how tadpoles shape age and size structures of populations and the composition of tadpole assemblages. I examined the incidence of intraguild predation (IGP) among three species that co-occur In ephemeral pools In the Osa Peninsula, Costa Rica. I found a high Incidence of IGP among the three species, with IG predators differing in their voracity for prey eggs and IG prey differing In their vulnerability to IGP. I also examined the factors that influence the Intensity of cannibalism and tadpole performance In a model species, Smilisca phaeota. I manipulated predator and prey size and developmental stage, tadpole density, food quantity, and tadpole relatedness in microcosm experiments. Cannibalism in S. phaeota was dependent on tadpole predator and prey size and developmental stage, with medium and large tadpoles causing almost complete mortality of embryos, but little mortality of mobile hatchlings. In contrast, predation by small tadpoles was restricted primarily to the embryo developmental stage. Tadpole mortality was high In low plant food quantity treatments, suggesting a high Incidence of cannibalism. Other measures of tadpole performance, including survival to metamorphosis, metamorph mass, and larval period, were influenced by tadpole density and plant food quantity. Additions of conspecific prey eggs and degree of tadpole relatedness did not affect tadpole performance. My results suggest that tadpoles are capable of playing a regulatory role, Influencing the relative abundance and composition of species in pools through differential IGP and cannibalism on eggs and hatchlings. The Intensity of IGP In ephemeral pools Is likely Influenced by the morphology of IG predators, reproductive mode and fecundity of IG prey, and priority effects. My study supports the view that we need to reconsider the position tadpoles occupy within the aquatic food webs, with greater appreciation for their role as predators. Osamu, K. Trussell, G. C., Nishimura, K. and T. Ohgushi. 2009. Inducible defenses in prey intensify predator cannibalism. Ecology 90: 3150-3158. Trophic cascades are often a potent force in ecological communities, but abiotic and biotic heterogeneity can diffuse their influence. For example, inducible defenses in many species create variation in prey edibility, and size-structured interactions, such as cannibalism, can shift predator diets away from heterospecific prey. Although both factors diffuse cascade strength by adding heterogeneity to trophic interactions, the consequences of their interaction remain poorly understood. We show that inducible defenses in tadpole prey greatly intensify cannibalism in predatory larval salamanders. The likelihood of cannibalism was also strongly influenced by asymmetries in salamander size that appear to be most important in the presence of defended prey. Hence, variation in prey edibility and the size structure of the predator may synergistically affect predatorprey population dynamics by reducing prey mortality and increasing predator mortality via cannibalism. We also suggest that the indirect effects of prey defenses may shape the evolution of predator traits that determine diet breadth and how trophic dynamics unfold in natural systems.

Moller, H., L. S. Lee, B. B. Paterson, and D. Mann. 2008. Cannibalism contributes significantly to the diet of cultured sand crabs, Portunus pelagicus (L.): A dual stable isotope study. Journal of Experimental Marine Biology and Ecology 361: 75-82. The significance of cannibalism in the diet of juvenile pond-cultured blue swimmer crabs (Portunus pelagicus (L.)) was investigated using dual stable isotope analysis of carbon and nitrogen. In a laboratory feeding experiment, delta N-15 demonstrated a constant trophic shift (Delta delta N-15 approximate to+1.6 parts per thousand.), and therefore seemed to be a reliable indicator for assessing trophic position for P. pelagicus. This agrees with previously reported trends. Difference in growth rate did not seem to influence delta N-15 values. In contrast, delta C-13 did not display consistent shifts between trophic levels (range of Delta delta C-13: +1 to +1.7%.). The results from the pond experiment showed that larger individuals had a more enriched delta N-15 than smaller individuals, which, when compared to the results from the laboratory experiment, indicates that larger individuals were at a higher trophic level. This is most likely due to cannibalism prevailing in the pond rather than a direct result of faster growth rate. Cannibalistic behaviour might further increase growth, resulting in the observed positive correlation between size and delta N-15. Rudolf, V. H. W. and J. Armstrong. 2008. Emergent impacts of cannibalism and size refuges in prey on intraguild predation systems. Oecologia 157: 675-686. 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. Rudolf, V.H.W. 2007. The interaction of cannibalism and omnivory: Consequences for community dynamics. Ecology 88: 2697-2705. Although cannibalism is ubiquitous in food webs and frequent in systems where a predator and its prey also share a common resource (intraguild predation, IGP), its impacts on species interactions and the dynamics and structure of communities are still poorly understood. In addition, the few existing studies on cannibalism have generally focused on cannibalism in the top-predator, ignoring that it is frequent at intermediate trophic levels. A set of structured models shows that cannibalism can completely alter the dynamics and structure of three-species IGP systems depending on the trophic position where

cannibalism occurs. Contrary to the expectations of simple models, the IG predator can exploit the resources more efficiently when it is cannibalistic, enabling the predator to persist at lower resource densities than the IG prey. Cannibalism in the IG predator can also alter the effect of enrichment, preventing predator-mediated extinction of the IG prey at high productivities predicted by simple models. Cannibalism in the IG prey can reverse the effect of top-down cascades, leading to an increase in the resource with decreasing IG predator density. These predictions are consistent with current data. Overall, cannibalism promotes the coexistence of the IG predator and IG prey. These results indicate that including cannibalism in current models can overcome the discrepancy between theory and empirical data. Thus, we need to measure and account for cannibalistic interactions to reliably predict the structure and dynamics of communities. Wise, D. H. 2006. Cannibalism,food limitation, intraspecific competition and the regulation of spider populations. Annual Review of Entomology 51; 441-465. Cannibalism among generalist predators has implications for the dynamics of terrestrial food webs. Spiders are common, ubiquitous arthropod generalist predators in most natural and managed terrestrial ecosystems. Thus, the relationship of spider cannibalism to food limitation, competition, and population regulation has direct bearing on basic ecological theory and applications such as biological control. This review first briefly treats the different types of spider cannibalism and then focuses in more depth on evidence relating cannibalism to population dynamics and food web interactions to address the following questions: Is cannibalism in spiders a foraging strategy that helps to overcome the effects of a limited supply of calories and/or nutrients? Does cannibalism in spiders reduce competition for prey? Is cannibalism a significant density-dependent factor in spider population dynamics? Does cannibalism dampen spider-initiated trophic cascades? Denno RF, Mitter MS, Langellotto GA, Gratton C, and D. L. Finke. 2004. Interactions between a hunting spider and a web-builder: consequences of intraguild predation and cannibalism for prey suppression. Ecological Entomology 29: 566-577. 1. Antagonistic interactions among invertebrate predators such as intraguild predation and cannibalism have the potential to dampen top-down impacts on shared prey at lower trophic levels. Two abundant spider predators, the large wolf spider Pardosa littoralis and the small sheet-web builder Grammonota trivitatta co-occur on the salt marshes of eastern North America where they both attack planthoppers (Prokelisia spp.), the dominant herbivores on the marsh. Experiments both in the laboratory and field were used to assess the incidence of intraguild predation and cannibalism in these spiders and elucidate how such antagonistic interactions influence planthopper suppression. 2. Functional response experiments showed that with an increase in planthopper prey density, Grammonota captured more prey but not a higher proportion of that offered. Pardosa exhibited the same response when Grammonota were offered as intraguild prey. Both functional responses were type I over the range of prey densities offered. 3. Grammonota is moderately cannibalistic, and the presence of planthopper prey reduced the incidence of cannibalism. 4. Factorial experiments in the laboratory showed that Pardosa but not Grammonota reduced planthopper prey populations when prey density was low. By contrast, at high prey densities, both Pardosa and Grammonota had significant adverse effects on planthopper populations. Moreover, there was an interactive effect such that Grammonota reduced planthopper populations relatively more when Pardosa was absent than when it was present. 5. There was direct evidence for the intraguild predation of Grammonota by Pardosa such that fewer Grammonota survived in the presence of Pardosa than when it was absent. This result occurred whether planthopper prey were abundant or not.

6. Field releases of Grammonota in open plots resulted in significant but small decreases in the density of planthopper prey, both nymphs and adults. 7. Enhancing densities of Pardosa in open plots resulted in Grammonota suppression. The intraguild predation of Grammonota at this enhanced Pardosa density, however, did not preclude Pardosa from significantly reducing planthopper populations. 8. Although there was evidence that Grammonota reduced planthopper populations and that the intraguild predation of Grammonota by Pardosa occurred, the strength of these interactions was relatively weak given the low consumption rate of planthoppers by Grammonota (< 3 day(-1)) and Grammonota by Pardosa (approximate to 2 day(-1)). Thus, weak asymmetric intraguild predation among spiders on the marsh likely dampens but does not eliminate the ability of Pardosa to exert significant top-down control on planthopper populations. Wahlstrom E, Persson L, Diehl S, and p. Bystrom P. 2000. Size-dependent foraging efficiency, cannibalism and zooplankton community structure. Oecologia 123: 138-148. To examine size-dependent food web interactions in systems with cannibalism, we compared the abundances of zooplankton and phytoplankton over 2 years in four lakes with cannibalistic perch (Perca fluviatilis ) of which two also supported the top predator pike (Esox lucius). The abundance of perch 2 years and older was lower in lakes with pike than in lakes with only perch. In contrast, the abundance of small perch (young-of-the-year and I-year old) was lower in lakes with only perch suggesting that intense cannibalism reduced these size classes to low levels in lakes lacking pike. Functional response experiments with differently sized perch and zooplankton showed that the attack rate of small perch susceptible to cannibalism was much higher than that of large cannibalising perch. The optimal body size of perch with respect to attack rate was also lower for small zooplankton prey than for large zooplankton. The zooplankton communities in lakes with only perch were dominated by the relatively small species Ceriodaphnia quadrangula and Bosmina spp. and total zooplankton biomass was higher in these lakes than in lakes with both pike and perch. In contrast, the mean size of cladoceran zooplankton was largest in lakes with both pike and perch owing to a dominance of the large zooplankton species Holopedium gibberum in these lakes. We relate these patterns to (1) the low foraging efficiency of large perch on small zooplankton and (2) the low abundance of small zooplanktivorous perch (due to cannibalism) in lakes with only perch. The differences in zooplankton community structure also resulted in different seasonal dynamics of phytoplankton between lakes. Cannibalism introduces a vertical heterogeneity to food webs that causes consumer-resource dynamics that are not predictable from linear food chain models. Holt RD, and G. A. Polis. 1997. A theoretical framework for intraguild predation. American Naturalist 149: 745-764. Many important issues in community ecology revolve around the interplay of competition and predation. Species that compete may also be locked in predator-prey interactions, a mixture of competition and predation known as ''intraguild predation'' (IGP). There is growing evidence for the importance of IGP in many natural communities, yet little formal ecological theory addresses this particular blend of interactions. In this article, we explore the consequences of incorporating IGP into standard models of exploitative competition and food chains (a general resource-consumer model, a Lotka-Volterra food chain model, and Schoener's exploitative competition model). Our theoretical analyses suggest a general criterion for coexistence in IGP systems: the intermediate species (the prey in intraguild predation) should be superior at exploitative competition for the shared resource, whereas the top species (the predator) should gain significantly from its consumption of the intermediate species. Along gradients in environmental productivity, coexistence is most likely at intermediate levels of productivity. Analyses of the models reveal the potential for alternative stable states in systems with

IGP; these are particularly likely if the top predator gains little benefit from consuming the intermediate predator. We further show that IGP can lead to unstable population dynamics, even when all pairwise interactions are inherently stable and each species can increase when rare. Persistent, strong IGP raises a puzzle of species coexistence, particularly in productive environments. We conclude by comparing IGP with related community modules (i.e., food chains, exploitative competition, apparent competition) and discussing mechanisms that should foster coexistence in systems with strong IGP. Fagan WF, and G. M. Odell. 1996. Size-dependent cannibalism in praying mantids: Using biomass flux to model size-structured populations. American Naturalist 147: 230-268. Here we investigate how cannibalism, a widespread phenomenon in nature, influences the population dynamics and the differential success of emergence-timing strategies in praying mantids. Relying on an extraordinarily complete data set describing the ecophysiology of a single mantid species, we construct a size-structured model based on ''biomass flux'' to study how size-dependent cannibalism influences this species' population ecology. Further, we demonstrate how this model accurately predicts mantid developmental patterns under both laboratory and field conditions. We conclude that for mantids, and perhaps generalist ambush predators overall, descriptions of biomass flux can be satisfactorily used to model growth without explicit reliance on encounter rate terms. In particular, this autecological approach to modeling size-structured interactions provides a method for revealing how sensitively development hinges on small differences in emergence time.