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.     

Male mating behaviour reduces the risk of sexual cannibalism in an Australian praying mantid

In cases where sexual cannibalism represents a sexual conflict, we should expect to find male traits that reduce the risk of cannibalism. In fact, a wide variety of such traits have been proposed, including elaborate courtship displays, cautious approach behaviours, and opportunistic mating whilst a female is feeding. However, there is very little direct evidence that these behaviours actually reduce the risk of sexual cannibalism for males, and the evidence that does exist comes mainly from spider studies. In this study of Pseudomantis albofimbriata praying mantids we found evidence of various male behaviours that reduced the risk of sexual cannibalism. Most males attempted to avoid detection entirely and others moved in a slow and stealthy manner. We also observed five cases of opportunistic mating, but saw no evidence of any male courtship routine. It seems that mounting from the rear of a female or slowly approaching her from the front, allows many males to go unnoticed and, therefore, substantially reduces the risk of sexual cannibalism in this mating system. Interestingly, we could not identify any female traits that males use to assess the risk of sexual cannibalism. It may be that P. albofimbriata males use alternative means to assess these risks. The presence of these behavioural strategies by males suggests a net cost of being consumed, and that sexual cannibalism is likely to be a female strategy in this system.     

Sexual cannibalism in scorpions: fact or fiction?

Females of several scorpion species are currently deemed to be sexual cannibals, but this reputation is, however, largely built on anecdotal evidence from the older literature. Theoretical models predict that given the low rates of female-male encounter, males should allow themselves to be cannibalized after sperm transfer. The present paper examines the information available for a number of species to determine (a) whether sexual cannibalism is as widespread as previously suggested, and (b) whether it is likely to involve male self-sacrifice. Information on three scorpions currently regarded as sexual cannibals is presented: the bothriurids Bothriurus bonariensis (C.L. Koch) and Urophonius jheringii Pocock, and the buthid Leiurus quinquestriatus (Hemprich & Ehrenberg). No post-mating cannibalism was ever observed in these species, nor in another 19 representatives of four families. In L. quinquestriatus only two cases of cannibalism of the male by the female occurred, both in the beginning of courtship and after long periods of food deprivation. We suggest that if sexual cannibalism occurs in scorpions, it is most likely to be of an 'economic' type, where females perceive males as prey rather than mates, and not a 'self-sacrificial', post-mating cannibalism enhancing the fitness of the offspring of the male.     

The strange occurrence of male cannibalism on adult females in snakes

Cannibalism is widespread in the animal kingdom. The most common, well-known cases are found in some arthropod systems where the larger females occasionally consume the smaller males before, during, or after mating (i.e., sexual cannibalism). The killing and consumption of females by males are, on the other hand, seemingly much rarer and theoretically more difficult to explain, at least among adults, because the consumption of females would result in the loss of potential mates for males. Here, we use observations submitted to a community science reporting system to describe three instances of adult males feeding on adult females in the Montpellier snake, Malpolon monspessulanus, in the south of France. We discuss our observations in the theoretical framework of cannibalism to provide potential explanations for these observations and briefly review empirical findings of cannibalism in vertebrates, which confirms that this sex-reversed form of sexual cannibalism is rare in nature.     

Spawning behaviour and acoustic communication in Atlantic cod

Cannibalism of small numbers of offspring by a parent has been proposed as an adaptive parental strategy, by providing energy to support parental care. There are few empirical studies, however, to support this hypothesis. The beaugregory damselfish, Stegastes leucostictus , is a marine teleost that does not actively ventilate its eggs by fanning them. Partial cannibalism is common in this species, but in field studies was found to be unrelated to ration level. Filial cannibalism differed from predation in the pattern of egg eating; filial cannibalism was characterised by a random pattern of egg loss from a clutch rather than an aggregated distribution. Embryos developed quicker and had higher survival rates when they were at low densities and in nest sites where oxygen levels were high, and experimental reduction of oxygen levels increased rates of filial cannibalism. Here I present a hypothesis for filial cannibalism in the beaugregory damselfish; males cannibalise egg clutches in order to reduce clutch density and improve oxygen supply to the remaining embryos. I use a model of filial cannibalism to demonstrate how oxygen mediated cannibalism may be adaptive, and discuss the evolution of filial cannibalism in the beaugregory damselfish and other teleosts.     

An analysis of a nonlinear stage-structured cannibalism model with application to the Northeast Arctic cod stock

A two-dimensional stage-structured population model with nonlinear cannibalism terms is studied. We show that there is a large parameter interval where the nontrivial equilibrium of the model is the only stable attractor, but that there also exist parameter intervals where we find quasiperiodic, periodic and chaotic dynamics. Moreover, in the interplay between increasing the fecundity and increasing the cannibalism pressure, the former turns out to be a destabilizing effect while the latter tends to act in a stabilizing fashion. Finally, we have applied the model to the North Atlantic cod stock using ICES biomass estimates. Our main conclusion from this study is that the combined effect of recruitment and cannibalism may not serve as an explanation of the observed fluctuations in the cod stock.     

Adaptive filial cannibalism in the beaugregory damselfish

Cannibalism of small numbers of offspring by a parent has been proposed as an adaptive parental strategy, by providing energy to support parental care. There are few empirical studies, however, to support this hypothesis. The beaugregory damselfish, Stegastes leucostictus, is a marine teleost that does not actively ventilate its eggs by fanning them. Partial cannibalism is common in this species, but in field studies was found to be unrelated to ration level. Filial cannibalism differed from predation in the pattern of egg eating; filial cannibalism was characterised by a random pattern of egg loss from a clutch rather than an aggregated distribution. Embryos developed quicker and had higher survival rates when they were at low densities and in nest sites where oxygen levels were high, and experimental reduction of oxygen levels increased rates of filial cannibalism. Here I present a hypothesis for filial cannibalism in the beaugregory damselfish; males cannibalise egg clutches in order to reduce clutch density and improve oxygen supply to the remaining embryos. I use a model of filial cannibalism to demonstrate how oxygen mediated cannibalism may be adaptive, and discuss the evolution of filial cannibalism in the beaugregory damselfish and other teleosts.     

Avoidance of post-coital cannibalism in the brachyurid paddle crab Ovalipes catharus

Sexual cannibalism was examined experimentally in the New Zealand paddle crab Ovalipes catharus, where it is the female that risks cannibalism during and after sexual activity. In this species copulation only occurs when the female is soft-shelled after moulting and she is most vulnerable to cannibalism. Male paddle crabs protect and copulate with the females until the females are no longer sexually receptive. Males appear able to identify a female with whom they have recently copulated. After a brief separation, significantly fewer females were cannibalized in cases where a female was returned to her original sexual partner than in those that had exchanged partners. The asymmetry, where females gain some protection during moulting but males receive no such advantage, may, at least partially, explain the skewed sex ratios which are sometimes found in samples of O. catharus from nature.     

Life history responses depend on timing of cannibalism in a damselfly

1. Cannibalism has often been suggested as an important mechanism to reach the necessary developmental stage and size before a critical time horizon is reached, but this role has been largely unexplored. We studied effects of cannibalism on the life history of the damselfly Lestes viridis under combinations of a time constraint (by manipulating the perceived time available in the growth season) and a biotic constraint (density). 2. Larvae had a faster development and growth rate when reared at high time stress (late photoperiod). They also had a higher growth rate and mass at emergence when cannibalism occurred (density 2 and 4). Cannibalism occurred earlier at higher density. Accelerated life history responses (faster development and growth rate) and a higher mass at emergence were dependent upon the timing of cannibalism. Responses were more pronounced or only present if cannibalism occurred early in the larval period. 3. Our data suggest that cannibalism may not only act as a lifeboat mechanism by enabling cannibals to survive detrimental ecological conditions, but may also act as a compensatory mechanism to keep life history variables near‐optimal at life history transitions, even under sub‐optimal conditions.     

Does New Guinea cannibalism have nutritional value?

This paper examines the question of the nutritional value of cannibalism. Although other authors have concluded that the practice does not have such value, we argue that this cannot properly be determined except in the context of the total subsistence economy and local human ecology. The paper also presents a format for the empirical investigation of foodgetting and new ethnographic information about New Guinea cannibalism. Our major conclusion is that this practice does have nutritional value for certain human groups, specifically tropical peoples living at lowmedium population densities and exploiting a diverse range of animal foods.     

Egg cannibalism by Coleomegilla maculata lengi neonates: preference even in the presence of essential prey

Abstract 1. Egg cannibalism among coccinellids has been reported widely, however reasons why this type of behaviour is observed so frequently have been neglected. This experiment was undertaken to clarify whether cannibalistic behaviour is advantageous to Coleomegilla maculata lengi Thimberlake neonates and to understand the reasons for high levels of egg cannibalism.
2. Benefits gained by neonate cannibals were ascertained by comparing survival, developmental time, and second-instar weight of C. maculata larvae that were allowed to cannibalise conspecific eggs or not. Preference and behaviour tests were also conducted to assess the reasons for high levels of egg cannibalism.
3. Cannibal neonates grew faster and were heavier than non-cannibals. The developmental time of neonates was influenced more by prey vulnerability than by prey quality.
4. In choice tests, where three different proportions of conspecific eggs and aphids were offered (33–67, 50–50, and 67–33%), C. maculata neonates always consumed significantly more eggs. Manly's preference indexes indicated that neonates showed a consistent preference for conspecific eggs.
5. Seventy-five per cent of neonates observed went directly towards eggs and 90% of the first prey consumed by neonates were an egg. When aphids were painted with extract of crushed eggs and eggs with crushed aphids to determine whether neonates found eggs by chemical cues, neonates preferred aphids painted with egg extract to eggs painted with aphid extract.
6. It was concluded that C. maculata neonates benefited from cannibalistic behaviour. Moreover, egg cannibalism is not related only to frequency of encounter; chemical cues are also involved in egg searching.     

Territoriality mediates atypical size‐symmetric cannibalism in the Amblypygi Phrynus longipes

Cannibalism can have important demographic and ecological effects on populations. Typically, cannibalism is size‐structured, where larger individuals eat smaller conspecifics. Initial cursory observations of the whip spider, Phrynus longipes, however, suggested that cannibalism might not be size‐structured in this species, perhaps because cannibalism is often a by‐product of territory contests. We staged paired interactions and recorded latency to escalate to physical aggression or cannibalize to understand the dynamics of cannibalism and resource contests. We employed a multimodel comparative approach to tease apart the contest characteristics that best predicted cannibalism during behavioral trials between P. longipes opponents. We found that, while armament size symmetry predicted escalation of contests, cannibalism was best predicted by body mass asymmetry. Further, cannibalism was most likely to occur among individuals similar in armament, but dissimilar in body mass. This suggests a discrepancy between phenotypes that may have evolved to communicate resource holding potential (e.g., armaments which benefit individuals if dishonest), and body mass as a cue of resource holding potential.     

Cases of maternal cannibalism in wild bonobos (<Emphasis Type="Italic">Pan paniscus</Emphasis>) from two different field sites,Wamba and Kokolopori,Democratic Republic of the Congo

Maternal cannibalism, whereby a mother consumes her own offspring, occurs in various animal taxa and is commonly explained by nutritional stress or environmental pressures. It is rare in nonhuman primates and is considered an aberrant behavior only observed under high-stress conditions. It was therefore surprising when, in the first reported case of cannibalism in wild bonobos, a mother consumed part of the dead infant at LuiKotale. Here we report two more cases of maternal cannibalism by wild bonobos at two different study sites, Wamba and Kokolopori. The dead infants’ mothers participated in the cannibalism in both cases. At Kokolopori, although the mother did consume part of the carcass, it was held and shared by another dominant female. At Wamba, the mother was a dominant female within the community and was the primary consumer of the carcass. In both cases, cannibalism resembled other meat-eating events, with the dominant female controlling meat consumption. Infanticide was not observed in either case, but its occurrence could not be ruled out. Although rare, the occurrence of maternal cannibalism at three different study sites suggests that this may represent part of the behavioral repertoire of bonobos, rather than an aberrant behavior.     

Challenging the Aggressive Spillover Hypothesis: Is Pre‐Copulatory Sexual Cannibalism a Part of a Behavioural Syndrome?

Pre‐copulatory cannibalism – females devouring males during courtship – may bring no benefit to either sex. The ‘aggressive spillover hypothesis’ (ASH) posits that pre‐copulatory cannibalism represents a spillover of female aggressiveness from the juvenile foraging context, when aggressiveness is advantageous, to the adult mating context, when aggressiveness may be non‐adaptive or maladaptive. The ASH suggests that individuals exhibit limited plasticity in aggressive behaviours because they are genetically canalised for indiscriminate aggressiveness towards prey and conspecifics, including males. Hence, a tendency to employ pre‐copulatory cannibalism is a part of the female aggression syndrome, an assertion generally accepted in the personality field. We here re‐evaluate the previous findings in the light of personality criteria, which we propose for ASH validation: between‐individual differences, repeatability and heritability in tendency for pre‐copulatory attacks (and pre‐copulatory cannibalism) and voracity towards prey, and their correlation. To re‐evaluate ASH and to allow for additional or alternative explanations, we ask whether pre‐copulatory cannibalism depends on female hunger, mating status, size and/or male quality. Finally, we ask whether cannibalistic females have a reduced reproductive success as predicted by the ASH. While repeatability and heritability in voracity towards prey and its correlation with the tendency to engage in pre‐copulatory cannibalism were found in certain systems, we lack any evidence for repeatability and heritability in pre‐copulatory cannibalistic attempts and for its maladaptiveness. Rather than only resorting to the ASH, foraging and mate choice hypotheses may also explain pre‐copulatory cannibalism. We suggest clarifying the use of the terms sexual cannibalism (effect) and female aggressiveness or tendency to attack and devour males (cause), and argue that male strategies to avoid cannibalism should be considered. We propose testing the ASH as the explanation for pre‐copulatory cannibalism in those cases where female tendency to devour males correlates with actual pre‐copulatory cannibalism and when all the above criteria are fulfilled. Finally, we propose future directions for studying the ASH.     

You eat what you are: personality‐dependent filial cannibalism in a fish with paternal care

Many animal parents invest heavily to ensure offspring survival, yet some eventually consume some or all of their very own young. This so‐called filial cannibalism is known from a wide range of taxa, but its adaptive benefit remains largely unclear. The extent to which parents cannibalize their broods varies substantially not only between species, but also between individuals, indicating that intrinsic behavioral differences, or animal personalities, might constitute a relevant proximate trigger for filial cannibalism. Using a marine fish with extensive paternal care, the common goby (Pomatoschistus microps), we investigated the influence of animal personality on filial cannibalism by assessing (1) behavioral consistency across a breeding and a nonbreeding context; (2) correlations between different breeding (egg fanning; filial cannibalism) and nonbreeding (activity) behaviors, and, in a separate experiment; (3) whether previously established personality scores affect filial cannibalism levels. We found consistent individual differences in activity across contexts. Partial filial cannibalism was independent of egg fanning but correlated strongly with activity, where active males cannibalized more eggs than less active males. This pattern was strong initially but vanished as the breeding season progressed. The incidence of whole clutch filial cannibalism increased with activity and clutch size. Our findings indicate that filial cannibalism cannot generally be adjusted independently of male personality and is thus phenotypically less plastic than typically assumed. The present work stresses the multidimensional interaction between animal personality, individual plasticity and the environment in shaping filial cannibalism.     

Piscivory and cannibalism in Arctic charr

Piscivory and cannibalism in Arctic charr, Salvelinus alpinus , were studied in three lakes in northern Norway: Guolasjavri, which contains only charr, Takvatn, where Arctic charr coexist with three-spined sticklebacks, Gasterosteus aculeatus and brown trout, Salmo trutta , and Stuorajavri, where whitefish, Coregonus lavarelun dominate a fish community containing six species. The prevalence of piscivory in the Arctic charr populations generally increased with increasing predator size. In all three lakes, many charr larger than 20 cm were piscivorous, but the extent of piscivory and cannibalism varied. The greatest prevalence of cannibalism was found in Guolasjavri, where 27% of charr greater than 20 cm in length had fed upon smaller conspeciflcs. In Takvatn, 5% of charr larger than 20 cm were cannibalistic, and an additional 9% had eaten three-spined sticklebacks. In Stuorajavri, up to 74% of the charr greater than 20 cm had eaten whitefish but cannibalism was not recorded. The possible role of cannibalism in population regulation within Arctic charr populations is considered.     

Evolution of cannibalism: referring to costs of cannibalism

A rational explanation for cannibalism is that it would be favored under conditions of crowding of conspecific individuals and/or low availability of alternative prey with the fear of starvation, so as to maximize individual fitness. Cannibalism has, however, not evolved and is not maintained by a simple individual optimization, while it has evolved and is maintained as a game among population members. We analysed the attainable state of an evolutionary cannibalism game within a framework that reflects the minimum essence of cause-effect in the cannibalism phenomenon. Cannibalism is predator-prey interaction among conspecifics. Immediate direct payoffs (survival in the interaction among conspecifics) and indirect payoffs (growth results in potential productivity and survival against the threat of starvation) would be included. No morphological specialization and no size priority of cannibalism individuals are assumed as conservative situations in which we analyse the possibility of cannibalism. Cannibalism would be possible under the conservative condition, if initially the wild population's cannibalism rate is not sufficiently lower than a threshold value. Crowding and/or low availability of alternative prey with the fear of starvation facilitates cannibalism evolution. Energy gain from conspecific prey would be attenuated by costs of counterattacks by conspecific victims and by challenge cost of its own. Discounting net intake energy required in the arms race for cannibalism challenge result in a relative disadvantage of having a high cannibalism rate and makes an evolutionary equilibrium of low cannibalism rate, even when potential profitability of conspecific prey is high.     

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.     

No Evidence for Kin Discrimination in Cannibalistic Tree-Hole Mosquitoes (Diptera: Culicidae)

Kin selection theory suggests that cannibalism is more likely to spread and be maintained if cannibalism of close relatives can be preferentially avoided. One important group of insects in which kin discrimination might be expected to evolve is cannibalistic tree-hole mosquitoes. Larvae of these species develop in small, ephemeral water bodies, where they regularly encounter both relatives and nonrelatives. In this study we compared the degree of sib cannibalism with the degree of nonsib cannibalism in two ecologically distinct mosquito species: an anautogenous (blood-feeding) detritivore Trichoprosopon digitatum (Culicinae) and an autogenous predator Toxorhynchites moctezuma (Toxorhynchitinae). Despite a wealth of literature documenting kin-biased discrimination in other insects, neither of these tree-hold mosquito species preferentially consumed nonrelatives. In the case of Tr. digitatum, the size-dependent nature of larval cannibalism may preclude additional selection for preferential consumption of nonrelatives, but in the autogenous Tx. moctezuma the direct nutritive and indirect competitive benefits of indiscriminate cannibalism may outweigh the immediate costs of consuming a relative.     

Cannibalism in prehistoric Europe

The key argument for the identification of prehistoric cannibalism is provided by analysis of close similarities in the treatment of human and animal remains. Such analysis requires precise data on depositional context, meticulous excavation records, detailed bone modification studies, a relatively large sample of human and animal postcranial bones, and data on local mortuary practices. With the exception of Fontbrégoua Cave, these necessary conditions are lacking at all Stone Age European sites where it has been hypothesized that cannibalism occurred. The alternative hypothesis of secondary burial practices has been proposed informally for some sites and, in a more formal and detailed way, for Krapina and Fontbrégoua. However, this hypothesis does not have a higher probability, is not justified by current data, and uses ethnographic analogies to prop up interpretations of materials for which contextual data are missing or have been neglected. At Fontbrégoua, cannibalism remains the simplest and most plausible explanation of the evidence; at Krapina and other sites the available evidence is insufficient to prove either secondary burial or cannibalism.