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.     

The evolution of filial cannibalism and female mate choice strategies as resolutions to sexual conflict in fishes

Filial cannibalism (the consumption of one's own viable offspring) is common among fish with paternal care. In this study, I use a computer simulation to study simultaneous evolution of male filial cannibalism and female mate choice. Under certain conditions, selection on parental males favors filial cannibalism. When filial cannibalism increases a male's probability to raise the current brood successfully, filial cannibalism also benefits the female. However, when egg eating is a male investment into future reproduction, a conflict between female and male interests emerges. Here I investigate how female discrimination against filial cannibals affects evolution of filial cannibalism and how different female choice criteria perform against filial cannibalism. The introduction of discriminating females makes the fixation of filial cannibalism less likely. I introduced three different female choice criteria: (1) females who could discern a male's genotype, that is, whether the male was going to eat eggs as an investment in future reproductive events; (2) energy-choosing females that preferred to mate with males who had enough energy reserves to live through the current brood cycle without consuming eggs; and (3) females that preferred to mate with already mated males, that is, males with eggs in their nest. Genotype choice never coexisted with filial cannibals at fixation and filial cannibals were unable to invade a population with genotype-choosing females. Energy choice was successful only when males had high energy reserves and were less dependent on filial cannibalism as an alternative energy source. The egg choosers frequently coexisted with the cannibals at fixation. When the female strategies were entered simultaneously, the most frequent outcome for low mate sampling costs was that both the cannibals and the egg choice was fixed and all other strategies went extinct. These results suggest that sexual conflicts may not always evolve toward a resolution of the conflict, but sometimes the stable state retains the conflict. In the present case, this was because the egg-preference strategy had a higher fitness than the other female strategies. The outcome of this simulation is similar to empirical findings. In fish with paternal care, male filial cannibalism and female preference for mates with eggs commonly co-occur.     

Parents benefit from eating offspring: density-dependent egg survivorship compensates for filial cannibalism

Abstract Why should animals knowingly consume their own young? It is difficult to imagine many circumstances in which eating one's own young (i.e., filial cannibalism) actually increases an individual's fitness; however, filial cannibalism commonly co‐occurs with parental care in fishes. The evolutionary significance of filial cannibalism remains unclear. The most commonly accepted explanation is that filial cannibalism is a mechanism by which caring males gain energy or nutrients that they reinvest into future reproduction, thereby increasing net reproductive success. There is mixed support for this hypothesis and, at best, it can only explain filial cannibalism in some species. A recent alternative hypothesis suggests that filial cannibalism improves the survivorship of remaining eggs by increasing oxygen availability, and thus increases current reproductive success. This theory has received little attention as of yet. We evaluated the hypothesis of oxygen‐mediated filial cannibalism in the sand goby by examining the effect of oxygen and egg density on the occurrence of filial cannibalism, evaluating the effects of partial clutch cannibalism on the survivorship of remaining eggs, and comparing potential costs and benefits of filial cannibalism related to the net number of eggs surviving. Indeed, we found that oxygen level and egg density affected the occurrence of cannibalism and that simulated partial clutch cannibalism improved survivorship of the remaining eggs. Additionally, because increased egg survivorship, stemming from partial egg removal, compensated for the cost of cannibalism (i.e., number of eggs removed) at a range of cannibalism levels, filial cannibalism potentially results in no net losses in reproductive success. However, oxygen did not affect egg survivorship. Thus, we suggest a more general hypothesis of filial cannibalism mediated by density‐dependent egg survivorship.     

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.     

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.     

Sex-biased egg cannibalism in spawning walleye pollock: the role of reproductive behavior

We studied inadvertent egg cannibalism in spawning stocks of walleye pollock, Theragra chalcogramma, in the Gulf of Alaska and the Bering Sea between 1986 and 1996. Male pollock had on average 3 times more eggs in their stomachs than females. For both sexes adult fish of average body length had more eggs in their stomach than did smaller or larger fish. When maturity of fish was taken into account, actively spawning males had the highest numbers of eggs in their stomachs. We found weak evidence for a diel variation; the number of eggs per stomach was high in both sexes during the day and lower during the night. We suggest two possible explanations for this phenomenon. Sex-biased egg cannibalism may reflect the differential time spent in layers of high egg densities. Hydroacoustic and trawl catch data from both areas suggest that males aggregate deeper than females. Spawning takes place in the deeper layers of the fish aggregation, so males spend more time in high egg densities. Alternatively, males may be more active than females and increased gill ventilation and/or drinking rates may be responsible for the differences in egg cannibalism.     

Do the risks of egg cannibalism and desiccation influence the choice of oviposition sites in the land snail Arianta arbustorum?

Laboratory tests were conducted to examine whether egg-laying Arianta arbustorum prevents egg cannibalism in its offspring by the choice of oviposition sites. Arianta arbustorum did not react to the presence of conspecific eggs when starting to oviposit, thus exposing its eggs to the risk of between-batch cannibalism. Snails preferred to oviposit in soil of higher moisture content even at high air humidity, reducing the risks of later egg desiccation and within-batch cannibalism. Both the types of oviposition sites chosen and their spatial distribution, as observed in 11 natural populations of A. arbustorum near Uppsala, Sweden, suggest that abiotic factors, such as a permanently moist microhabitat, may exert a stronger selection pressure on the oviposition-site choice than does between-batch egg cannibalism. As a result, the dispersion of egg batches observed in the field mainly reflects the spatial heterogeneity of habitats.     

When to care for, abandon, or eat your offspring: the evolution of parental care and filial cannibalism

Parental care and filial cannibalism (the consumption of one's own offspring) co-occur in many animals. While parental care typically increases offspring survival, filial cannibalism involves the killing of one's young. Using an evolutionary ecology approach, we evaluate the importance of a range of factors on the evolution of parental care and filial cannibalism. Parental care, no care/total abandonment, and filial cannibalism evolved and often coexisted over a range of parameter space. While no single benefit was essential for the evolution of filial cannibalism, benefits associated with adult or offspring survival and/or reproduction facilitated the evolution of cannibalism. Our model highlights the plausibility of a range of alternative hypotheses. Specifically, the evolution of filial cannibalism was enhanced if (1) parents could selectively cannibalize lower-quality offspring, (2) filial cannibalism increased egg maturation rate, (3) energetic benefits of eggs existed, or (4) cannibalism increased a parent's reproductive rate (e.g., through mate attractiveness). Density-dependent egg survivorship alone did not favor the evolution of cannibalism. However, when egg survival was density dependent, filial cannibalism invaded more often when the density dependence was relatively more intense. Our results suggest that population-level resource competition potentially plays an important role in the evolution of both parental care and filial cannibalism.     

Cannibalism and intraguild predation of eggs within a diverse predator assemblage

Greater biodiversity among aphid predators sometimes leads to greater predator reproductive success. This could occur if cannibalism of predator eggs is consistently stronger than intraguild predation, such that diversity dilutes cannibalism risk when total predator densities remain constant across diversity levels. We compared the frequency of cannibalism versus intraguild predation by adult predators of four species [the lady beetles Coccinella septempunctata L. and Hippodamia convergens Guerin-Meneville, and the predatory bugs Geocoris bullatus (Say) and Nabis alternatus Parshley] on the eggs of three predator species (all of these predators but Nabis). For both coccinellid species, egg predation averaged across all intraguild predators was less frequent than cannibalism. In contrast, Geocoris eggs were generally more likely to be consumed by intraguild predators than by conspecifics. Closer inspection of the data revealed that Geocoris consistently consumed fewer eggs than the other species, regardless of egg species. Indeed, for lady beetle eggs it was relatively infrequent egg predation by Geocoris that brought down the average across all heterospecific predators, masking the fact that adults of the two lady beetles were no more likely to act as egg cannibals than as intraguild predators. Nabis ate eggs of the two beetles at approximately equal rates, but rarely ate Geocoris eggs. Female predators generally consumed more eggs than did males, but this did not alter any of the patterns described above. Altogether, our results suggest that species-specific differences in egg predation rates determined the relative intensity of egg intraguild-predation versus cannibalism, rather than any more general trend for egg cannibalism to always exceed intraguild predation.     

Sibling cannibalism in aphidophagous ladybirds: its impact on sex-dependent development and body weight

Abstract:  The effects of sibling egg cannibalism on sex-dependent development and adult body weight were determined by rearing simultaneously two groups of sibling larvae of ladybirds, viz. Propylea dissecta and Coccinella transversalis individually. The first group included cannibals (i.e. neonates, which took a sibling egg as the first meal and later fed on aphids, Aphis craccivora ) and the other included non-cannibals (reared exclusively on aphids). The cannibal larvae developed faster with heavier adults than non-cannibals. This expedited development and nutritional advantage (increased body weight) was greater in first instars indicating maximum benefit of sibling cannibalism to them. Sibling cannibalism was relatively more advantageous to male than to female ladybirds. Laboratory data predict that the larger size of adults in the field could possibly be the result of sibling egg cannibalism at the neonate level.     

Games among cannibals: competition to cannibalize and parent-offspring conflict lead to increased sibling cannibalism

Sibling cannibalism occurs in many species, yet understanding of sibling cannibalism as an adaptation currently lags behind understanding of other antagonistic interactions among siblings. Observed sibling cannibalism phenotypes likely reflect the interaction between competitive games among siblings and parent-offspring conflict. Using a game-theoretic approach, we derive optimal offspring cannibalism behaviour and parental modifiers that limit or facilitate cannibalism. The results are compared to contemporary frequency-independent analysis. With the addition of game interactions among siblings or parent-offspring co-evolution, our model predicts increased cannibalism (compared to the frequency-independent prediction), as offspring compete to eat siblings. When infertile eggs are present--strengthening competition--offspring risk eating viable siblings in order to gain access to infertile eggs, intensifying parent-offspring conflict. We use the results to make new predictions about the occurrence of sibling cannibalism. Additionally, we demonstrate the utility of trophic egg laying as a maternal mechanism to promote egg eating.     

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.     

Sibling and non-sibling cannibalism by larvae of a lady beetleHarmonia axyridis Pallas (Coleoptera: Coccinellidae) in the field

Sibling (cannibalism among siblings) and non-sibling cannibalism (cannibalism among non-siblings) were studied in a natural population of a lady beetle, Harmonia axyridisPallas . Of all the eggs laid (n=2269), 24.76% (n=562) were killed by sibling cannibalism and 36.10% (n=819) were killed by non-sibling cannibalism. Sibling cannibalism occurred constantly and intensively in most egg batches throughout the entire oviposition period. On the other hand, non-sibling cannibalism was more intense in the middle and late oviposition periods, and when the egg batches were close to an aphid colony. This may be due to the high density of H. axyridis larvae relative to aphid density in the middle and late oviposition periods and also the larvae searching intensively near an aphid colony.     

Egg cannibalism in Ascia monuste in the field; opportunistic, preferential and very frequent

Newly hatched caterpillars of the specialist herbivorous species Ascia monuste (Lepidoptera, Pieridae) feed on egg chorion before ingesting plant material. They also ingest their exoskeletons and perform egg cannibalism in both the field and the laboratory. Females lay eggs on plants which already have conspecifics (eggs and/or caterpillars) leading to the observation of cannibalism in the field. Two favorable conditions are required for such an occurrence: (1) eggs laid close together on the same leaf, and (2) the finding by older caterpillars of eggs on the same plant. Oviposition preference experiments in cages showed that females do not avoid laying on plants with eggs. However, they do avoid laying eggs on plants predated by caterpillars. All instars perform cannibalism in the field and, in addition to this, the ingested eggs are healthy and cannibalism occurs in the presence of abundant food. The ingestion of eggs from their own clutch was more intense when the clutch was large. Contact with eggs is also important for cannibalism. Caterpillars ingested healthy eggs during three successive days. According to the results, it is possible to state that egg cannibalism in A. monuste is opportunistic, preferential and highly frequent and that there is a tolerance to a diet based on animal protein in this species. Since A. monuste obtains nutrients from both animal and vegetable resources, it is probably better to consider it as opportunistic omnivorous.     

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.     

Cannibalism and protective behavior of eggs in Arctic charr (Salvelinus alpinus)

From video recordings of spawning events, we quantified protective and cannibalistic behavior of Arctic charr occurring immediately after spawning. The number of fish cannibalizing on stray eggs was examined regarding (a) whether more than one male shed milt during the spawning event, that is, whether sperm competition occurred, (b) whether the sperm competition included few or many males, that is, the intensity of sperm competition, and (c) the density of fish at the spawning site. Response behavior toward egg cannibalism was also examined among females and dominant males in order to determine any parental investment toward protecting the eggs after spawning. Cannibalistic behavior was seen in almost 50% of the spawnings, and the multiple spawning events showed the highest numbers of fish cannibalizing on eggs. Both the number of males releasing milt and the number of fish approaching the spawning site were positively correlated with egg cannibalism. Sperm competition was, however, not a prerequisite for egg cannibalism. Although we also observed partial filial cannibalism, protective behavior of eggs was seen both among dominant males and females, suggesting that charr actually conduct parental care.     

Environmental influences on fecundity, egg viability and egg cannibalism in the scale insect predator, Chilocorus nigritus

The effects of temperature on age-specific fecundity, egg viability and egg cannibalism were examined in the coccidophagous coccinellid, Chilocorus nigritus being fed on the cyanophyllum scale, Abgrallaspis cyanophylli. There were no significant differences in lifetime fecundity or oviposition rate at constant temperatures in the range of 20 to 30°C although there was a trend for both parameters to increase with increasing temperatures. Total fecundity ranged from five to 1890 eggs per female whilst the oviposition rate varied from 0.2–12.1 eggs day^-1. Egg viability was significantly greater at 26 and 30°C (63 and 74%, respectively) than at lower temperatures (18 to 30%) whilst egg cannibalism was highest at 24 and 26°C (being the mid range of the temperatures tested). Mean pre-oviposition periods varied from 54 days at 20°C to 8.2 days at 30°C. The effect of relative humidities in the range of 33% to 75% on oviposition rate, egg viability and egg cannibalism was measured and found to have no significant effect on either parameter.     

Propensity towards cannibalism among Hypoaspis aculeifer and H. miles, two soil-dwelling predatory mite species

In biological control programmes, the two predatory soil mites Hypoaspis aculeifer and H. miles are often applied against soil-borne pests like mushroom flies, springtails and mites. Although the mites show high consumption rates on varying prey types in Petri dish experiments and in greenhouses, their overall efficiency is sometimes limited. We hypothesized that intraspecific interactions, like cannibalism, could contribute to this decreased competence. Therefore, experiments were conducted to show the propensity of H. aculeifer and H. miles to cannibalise. Adult mites and nymphs were introduced as predators with conspecific eggs, larvae, nymphs, adult females or males as prey and the number of killed individuals was recorded. Additionally, the oviposition rate on conspecific prey was quantified and the correlation with the number of prey consumed was calculated to assess the influence of cannibalism on egg production. The results illustrate that cannibalism occurs infrequently in both Hypoaspis spp., the only exception being H. aculeifer nymphs, which cannibalised one conspecific egg per day. Moreover, cannibalism never occurred in the presence of alternative prey. Oviposition rate decreased during the experiment in both species but it was positively correlated with the cannibalism rate only for H. aculeifer. The benefit of cannibalism for populations of H. aculeifer and H. miles is discussed.     

Egg sac damage and previous egg sac production influence truncated parental investment in the wolf spider,Pardosa milvina

Life history theory predicts that iteroparous animals adaptively partition reproductive effort between current and future reproduction. When rearing costs of current offspring exceed the potential benefits, parental care should be terminated and deferred toward future reproduction. We tested two related predictions that follow from life history theory: (a) parents should be sensitive to offspring viability and withhold parental care if offspring survival probability drops and future reproductive opportunities are likely, and (b) parents should be less sensitive to offspring survival probability when future reproduction is unlikely and maximize parental care late in life. The wolf spider, Pardosa milvina, demonstrates extensive parental care; however, they may also abandon or cannibalize their egg sacs. We tested the effects of egg sac damage and production of a previous egg sac on egg sac abandonment and cannibalism decisions. Among four egg sac groups (1st egg sac intact, 1st egg sac damaged, 2nd egg sac intact, 2nd egg sac damaged), we daily monitored egg sac abandonment and cannibalism and measured differences in egg sac searching, protection, and grooming among removed and damaged egg sacs (N = 116 with 1st egg sac and 88 with 2nd egg sac). Females with first egg sacs abandoned and cannibalized damaged egg sacs significantly more compared to unmanipulated egg sacs; however, females with second egg sacs were insensitive to egg sac damage. Females also spent significantly more time protecting second egg sacs compared to first egg sacs and groomed damaged egg sacs significantly more than undamaged. These results support the general predictions of life history theory that indicate that abandonment and cannibalism should decrease with diminished future reproductive potential and that parents should be less sensitive to indicators of offspring survival probability late in life.     

The relationship between dietary specialism and availability of food and water on cannibalistic interactions among predatory mites in protected crops

The interplay between dietary specialism, the tolerance of food and water stress and level of cannibalism is likely to be important in determining the outcome of biological control using inundative releases of multiple natural enemies, such as phytoseiid mites in protected crops. The dietary specialist, Phytoseiulus persimilis, with a short immature development time (4-5 days) when plentiful food was available had a low ability to survive without food (5 days), even with access to water. The dietary generalists, Neoseiulus californicus, N. cucumeris and lphiseius degenerans, had longer immature development times (by up to 2 days) than P. persimilis. Survival ability differed amongst the generalist species when they were starved but provided with constant access to water. Both N. californicus and N. cucumeris survived the longest (8-10 days) and I. degenerans survived the shortest period (4 days). No negative intra-specific interaction between immatures was observed with P. persimilis when food was available and in the absence of food this species tended to starve rather than act cannibalistically. Both N. californicus and N. cucumeris showed a low degree of cannibalism between immatures, either when food was available, or when starved but given access to water. Even when food was available survival of I. degenerans fell by 30% in 4 days and remained at 60-70% for 3 further days; survival continued to decline rapidly when they were starved but provided with water. This indicates that immatures of I. degenerans could either feed on dead conspecifics or that they were capable of a degree of cannibalism. Adult females of P. persimilis did not feed on conspecific eggs even when deprived of food but provided with water. Adult female N. californicus and N. cucumeris did feed on conspecific eggs but at a low level (<1 egg per day), which occurred only after 48 h starvation. Although egg cannibalism occurred more consistently with adult female I. degenerans than with other mite species it was at a low level (<1 egg per day). If the tendency to cannibalism, not just of eggs but with more susceptible life stages such as larvae, is reduced when water is available freely this could be important in determining the interactions that occur under natural conditions.