Why do ladybirds (Coleoptera: Coccinellidae) cannibalize?

Cannibalism of eggs by larvae ofAdalia bipunctata, an aphidophagous species of ladybirds, is important for survival when aphids are scarce. Ladybirds survive longer by eating eggs of their own species rather than aphids. Since it costs less, in terms of larval growth, to eat eggs rather than aphids, cannibalism has a strong advantage under conditions of prey scarcity.     

龟纹瓢虫的行为学观察

通过以洋槐蚜Aphis robiniae Macchiati为食物对龟纹瓢虫Propylea japonica(Thunberg)进行室内饲养和观察,记述了龟纹瓢虫幼虫的孵化、蜕皮行为,成虫的羽化、捕食、交配、产卵行为,并对幼虫成虫的自残行为进行了初步观察。结果表明幼虫孵化、蜕皮,成虫羽化后均有取食自己蜕皮时产生的残皮的行为;龟纹瓢虫成虫捕食行为分为搜寻、捕捉、清洁、静止排泄4个步骤;交配一般由雄虫发起,雌虫背负成虫进行交配。龟纹瓢虫每产一粒卵平均耗时3.75秒,两粒卵之间产卵时间间隔约31.75秒,每次产卵量平均12枚;龟纹瓢虫存在自残行为,成虫在食物不足时会取食幼虫和虫卵,高龄幼虫在饥饿的状态下也取食低龄幼虫和卵,且同龄幼虫之间也会出现相互咬食的现象。     

The use of volatile cues in recognition of kin eggs by predatory mites

1. Several animal species are known to distinguish between their own eggs and eggs of unrelated conspecifics. However, the cues involved in this discrimination are often unknown. These cues were studied using the predatory mite Gynaeseius liturivorus Ehara.
2. Adult females of these predatory mites oviposit in clusters and avoid oviposition close to eggs laid by other females, resulting in reduced cannibalism between offspring. Because predatory mites are blind, it was tested whether volatiles of eggs were used as a cue for egg recognition.
3. Adult female predatory mites were offered volatile cues of their own eggs and of unrelated conspecific eggs, and females were prevented from contacting the eggs. Predatory mites oviposited closer to their own eggs than to unrelated eggs. This preference was observed even when one own and one unrelated egg were offered as a volatile source.
4. These results suggest that adult female predatory mites can determine kinship using volatiles released from the eggs.

     

Egg cannibalism by larvae and adults of the milkweed leaf beetle (Labidomera clivicollis, Coleoptera: Chrysomelidae)

Abstract.

• 1 Egg cannibalism is a form of infanticide that has been implicated in the evolution of guarding of eggs and immatures in some species of insects. The milkweed leaf beetle, Labidomera clivicollis (Coleoptera: Chrysomelidae), exhibits three types of egg cannibalism in the field: siblicide, cannibalism of eggs by older larvae from earlier hatching egg clutches, and cannibalism of eggs by adult females. Unlike their close relative, L.suterella (Choe, 1989), female L.clivicollis do not guard their eggs or immatures. They move slowly about a patch of milkweeds laying multiple egg clutches.
• 2 First instar and older larvae cannibalized eggs in two geographically separated study populations (subspecies) in Austin, Texas, and Bridgeport, New York, U.S.A. Although adult females of both populations cannibalized eggs in the laboratory, only the Texas subspecies, L.c.rogersii, exhibited such cannibalism in the field.
• 3 In the field, correlates of siblicide varied both temporally (within subspecies) and spatially (between subspecies) in terms of whether they were statistically significant, but trends were all in the same direction. Group size was positively correlated with hatching success and siblicide, but negatively correlated with other types of predation. Siblicide was also positively correlated with egg density in a laboratory study of the Texas subspecies, L.c.rogersii.
• 4 In the laboratory, an average of 15–17% of L.c.clivicollis eggs never developed embryos. Although these were almost always cannibalized, some viable eggs were also eaten and there is no evidence that females increased the proportion of infertile eggs they laid to increase siblicide.
• 5 Field data and laboratory experiments showed that adult female L. c.rogersii cannibalized eggs while males rarely did. Females preferentially ate the eggs of other females over their own eggs in an experiment that removed spatial cues.
• 6 Although the selective context of cannibalism is not demonstrated here, I suggest that females may increase siblicide by increasing egg density and may cannibalize eggs to protect their own eggs from being eaten by second and third instar larvae produced by other females.

     

Suitability of exotic and native lady beetle eggs (Coleoptera: Coccinellidae) for development of lady beetle larvae

Predation upon lady beetle (Coleoptera: Coccinellidae) eggs in the field is most often instances of egg cannibalism by larvae or adults while the majority of the remaining predation events upon coccinellid eggs is done by other species of Coccinellidae. Thus the recent introduction and establishment of Harmonia axyridis in the US could negatively affect native species of Coccinellidae via egg predation. However, little is known regarding the suitability of interspecific coccinellid eggs as a food source for larval development. In this study, it was found that native first or third instar Coleomegilla maculata and Olla v-nigrum larvae were incapable of surviving to the adult stage when provided solely exotic H. axyridis eggs. In stark contrast, H. axyridis larvae survived equally well when cannibalizing eggs or eating eggs of either native species. When C. maculata and O. v-nigrum were grouped as ‘native’ and compared with the exotic H. axyridis, more native eggs were attacked than exotic eggs and a higher percentage of eggs was attacked by H. axyridis larvae. Native and exotic larvae attacked a similar percentage of native eggs but native larvae attacked significantly fewer exotic eggs than did exotic larvae. These data suggest that H. axyridis may prey upon the eggs of these native species, when encountered in the field, compared with the likelihood of the native species preying upon H. axyridis eggs. Therefore, eggs of the native species C. maculata and O. v-nigrum will continue to be subjected to cannibalism and also to possible predation by other native species and the exotic H. axyridis.     

THE GENETICS AND EVOLUTION OF CANNIBALISM IN FLOUR BEETLES (GENUS TRIBOLIUM)

Cannibalism plays a major role in population regulation in Tribolium confusum, accounting for up to tenfold differences in population size between different genetic strains. I characterized the within- and between-strain genetic variation for cannibalism using standard quantitative-genetic methods. The four laboratory strains studied have similar birth and death rates but differ in their strain-specific cannibalistic tendencies. The cannibalism rates of the strains were stable for more than 60 generations of laboratory husbandry. I found considerable genetic variation for cannibalism within each strain. A genetic analysis of the between-strain differences in each of three types of cannibalism (larvae eating eggs, adults eating eggs, and adults eating pupae) showed that all three cannibalism pathways are autosomally inherited and exhibit minor degrees of dominance. Adult cannibalism of eggs and larval cannibalism of eggs appear to be genetically correlated. The differences between the “high” and “low” cannibalism strains appear to be polygenic for two kinds of cannibalism, larvae eating eggs and adults eating pupae. However, strain differences in adult cannibalism of eggs may be due to only two loci. The stability of the between-strain differences for more than 60 generations, the additive nature of inheritance, and the demonstration of considerable within-strain genetic variation suggest that cannibalism may be selectively neutral or under stabilizing selection with many adaptive peaks.     

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.     

Laboratory study of cannibalism and interspecific predation in ladybirds

Abstract.

• 1 In the absence of aphids, adult females of Adalia bipunctata (L.) showed a greater reluctance to eat eggs than males.
• 2 Eggs and young larvae were more vulnerable to cannibalism than older larvae and starved larvae were more vulnerable than well-fed larvae.
• 3 Both egg and larval cannibalism is inversely related to the abundance of aphids.
• 4 Eggs are a better food, in terms of larval growth and survival, than aphids.
• 5 In the absence of aphids interspecific predation occurred, but not equally, between the coccinellids A.bipunctata, A.decempunctata (L.), Coccinella septempunctata L. and C.undecempunctata L.
• 6 Larvae and adults of A. bipunctata and C.septempunctata were reluctant to eat conspecific eggs painted with a water extract of the other species' eggs and larvae of C. septempunctata were more likely to die after eating a few eggs of A.bipunctata than vice versa.
• 7 These results indicate that cannibalism occurs mainly when aphid prey is scarce and is adaptive in that it improves the chances of survival, and coccinellids, to varying degrees, are defended against interspecific predation.

     

The Use of Conspecific Chemical Cues for Cannibal Avoidance in California Newts (Taricha torosa)

Adult California newts, Taricha torosa (Caudata: Salamandridae), prey on both conspecific egg masses and larvae. We used both laboratory and field experiments to determine if newt larvae use conspecific chemical cues to avoid adult intraspecific predators. Larvae showed significant hiding responses when exposed to adult cues in both the field and the laboratory. Larvae did not exhibit significant hiding responses to chemical cues from invertebrate predators.     

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.     

Context-Dependent Plastic Response during Egg-Laying in a Widespread Newt Species

Previous research on predator-induced phenotypic plasticity mostly focused on responses in morphology, developmental time and/or behaviour during early life stages, but the potential significance of anticipatory parental responses has been investigated less often. In this study I examined behavioural and maternal responses of gravid female smooth newts, Lissotriton vulgaris, in the presence of chemical cues originating from invertebrate predators, Acilius sulcatus water beetles and Aeshna cyanea dragonfly larvae. More specifically, I tested the extent of oviposition preference, plasticity in egg-wrapping behaviour and plasticity in egg size when females had the possibility to lay eggs at oviposition sites with and without predator cues during overnight trials. I found that individuals did not avoid laying eggs in the environment with predator cues; however, individuals that deposited eggs into both environments adjusted the size of the laid eggs to the perceived environment. Females deposited larger eggs earlier in the season but egg size decreased with time in the absence of predator cues, whereas individuals laid eggs of average size throughout the investigated reproductive period when such cues were present. Also, egg size was found to be positively related to hatching success. Individuals did not adjust their wrapping behaviour to the presence of predator cues, but females differed in the extent of egg-wrapping between ponds. Females’ body mass and tail depth were also different between ponds, whereas their body size was positively associated with egg size. According to these results, female smooth newts have the potential to exhibit activational plasticity and invest differently into eggs depending on temporal and environmental factors. Such an anticipatory response may contribute to the success of this caudate species under a wide range of predator regimes at its natural breeding habitats.     

Inter- and intraspecific predation on immatures by adult females in Euseius finlandicus, Typhlodromus pyri and Kampimo dromus aberrans (Acari: Phytoseiidae)

In a series of experiments, the interspecific predation and cannibalism on immatures by the adult females of Euseius finlandicus, Kampimodromus aberrans and Typhlodromus pyri were examined under laboratory conditions. The three species showed differing tendencies to prey on each other's motile immature stages. Euseius finlandicus females consumed more larvae and protonymphs than the females of T. pyri and K. aberrans. In cages without free water E. finlandicus ate a range of 6.51 larvae or 5.31 protonymphs of T. pyri and 5.27 larvae or 5.95 protonymphs of K. aberrans per female per day. Kampimodromus aberrans and T. pyri females exhibited a greater tendency to prey on heterospecifics than on conspecifics. When feeding on phytoseiid immatures and without free water, T. pyri females survived longer and laid more eggs than the females of E. finlandicus and K. aberrans. Adult females of E. finlandicus and T. pyri having free water and preying on heterospecific protonymphs were able to maintain egg laying during the whole experimental period of 12 days. The females of all three species had difficulties in piercing phytoseiid eggs, and the number of sucked eggs per female per day was low. Kampimodromus aberrans females ate 0.48 eggs of T. pyri daily, which was the highest recorded number. The great tendency to interspecific predation on motile immature phytoseiids by the females of E. finlandicus is discussed with regard to the dominance of this species on deciduous trees and bushes in Austria.     

Indiscriminate oophagy by ant larvae: an explanation for brood serial organization?

Summary Last instar larvae ofLasius niger under standard laboratory conditions and abundant food supply feed on conspecific eggs and, if forced to a choice, show a significant statistical preference for trophic versus fertilized eggs (approx. 60% of the trials observed). On the other hand, they are unable to discriminate between kin and non-kin eggs, both fertilized and trophic. Fertilized eggs killed by freezing and trophic eggs handled in the same way are also selected in a random manner.Last instar larvae ofMessor semirufus regularly failed to discriminate in a significant way between trophic and fertilized and between kin and non-kin eggs.Substantial experimental evidence confirms that larvae of both these species — as already suggested in the literature for other ants-rely on trophic eggs as an essential component of their diet in order to be able to develop. Additionally,L. niger last instar larvae appear to be able to attack and pierce the egg chorion with their own mandibles, while this capacity is at least very reduced or very rare inM. semirufus. First and second instar larvae of both species never succeeded in piercing the egg membrane alone.These findings imply that the right (i.e. trophic) eggs should be presented to the larvae by the workers (i.e. worker discrimination should be assumed) and, at least in the case of youngLasius larvae, and probably for all stages ofMessor larvae, the egg membrane must be pierced by the workers in order to allow the larvae to feed.The following evolutionary sequence is suggested to explain the origin of trophic eggs in ants: 1) larval oophagy (obligatory at least for the species founding new colonies in an independent claustral manner), 2) generalized facultative or obligatory larval oophagy for larvae of all colonial stages, 3) production of trophic eggs by the queen(s) and/or workers in order to avoid cannibalism of nestmates.Lack of discrimination or weak discrimination capacity between trophic and fertilized eggs and the consequent larval cannibalism in form of oophagy, as demonstrated in this paper, is suggested to explain age segregation among eggs and larvae from workers, a widespread phenomenon in ants which must have been selected to avoid the oophagy of viable eggs.     

Chrysoperla rufilabris (Neuroptera: Chrysopidae) females do not avoid ovipositing in the presence of conspecific eggs

Previous experiments have demonstrated that green lacewing (Neuroptera: Chrysopidae) adults could be attracted to field crops using artificial honeydew. To be effective as a biological control method, such a technique would require that the increase in female abundance translate in an increase egg deposition. An experiment was conducted to evaluate whether the honeydew-feeding females of the green lacewing Chrysoperla rufilabris (Burmeister) avoid laying eggs in the presence of conspecific eggs. The potential risk associated with oviposition in a site already occupied by conspecific eggs was also studied. The preference of C. rufilabris larvae for kin and non-kin eggs and the susceptibility of C. rufilabris eggs to cannibalism relative to their age was determined. The results demonstrate that females are not reluctant to oviposit in the presence of conspecific eggs. Larvae show no preference for kin or non-kin eggs, and lacewing eggs become less susceptible to cannibalism as they age. This indicates that the risk of egg cannibalism by neonate in the field may be low. The results are discussed from ecological and biological control points of view.     

Effect of intraguild predation on the survival and development of three species of aphidophagous ladybirds: consequences for invasive species

Abstract 1 Survival and development of hatchling larvae of three aphidophagous ladybirds (Coleoptera: Coccinellidae), Harmonia axyridis Pallas, Coccinella septempunctata brucki Mulsant and Adalia bipunctata Linnaeus, when fed their own and the other species eggs were recorded. 2 In all three species, the larvae survived when fed conspecific eggs. 3 The percentage of larvae of H. axyridis that survived decreased to 35% and 85% when fed eggs of A. bipunctata and C. s. brucki, respectively. All the larvae of A. bipunctata and C. s. brucki died after eating eggs of H. axyridis. None of the larvae of C. s. brucki died after eating eggs of A. bipunctata, whereas 46% of those of A. bipunctata died after eating eggs of C. s. brucki. 4 In general, larvae were reluctant to eat the eggs of other species. However, larvae of C. s. brucki showed less reluctance than H. axyridis to eat the eggs of A. bipunctata. 5 The consequence of this for invasive species of ladybird is discussed.     

Differentiated egg size of the cannibalistic salamander <Emphasis Type="Italic">Hynobius retardatus</Emphasis>

Larvae of the salamander, Hynobius retardatus, are carnivorous, and even though there are two morphs, a typical morph and a broad-headed or “cannibal” morph, both are cannibalistic. They also sometimes eat other large prey, for example larvae of the frog, Rana pirica. In natural habitats, use of both conspecific and R. pirica larvae as food may contribute more strongly to high survival and substantially to fitness when larval densities are higher, because early-stage H. retardatus larvae sometimes experience scarcity of their typical prey. In cannibalistic oviparous amphibians, larger individuals that developed from larger eggs can more efficiently catch and consume larger prey and thus their survival may be better than that of smaller individuals developed from smaller eggs. Populations might therefore diverge in respect of egg size in response to variation in the density of conspecific and R. pirica larvae in natural ponds, with eggs being larger when larval density is higher. I examined how variance in hatchling size correlated with the incidence of cannibalism, and whether increasing larval density in natural ponds correlated with increasing egg size. Variance in initial larval body size facilitated cannibalism, and egg size increased as larval density in the ponds increased. In ponds with high larval density, where cannibalism and large prey consumption is a critical factor in offspring fitness, the production of fewer clutches with larger eggs, and thus of fewer and larger offspring, results in greater maternal fitness. Variation among the mean egg size in populations is likely to represent a shift in optimum egg size across larval density gradients.     

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.     

Water mold infection but not paternity induces selective filial cannibalism in a goby

Many animals heavily invest in parental care but still reject at least some of their offspring. Although seemingly paradoxical, selection can favor parents to neglect offspring of particularly low reproductive value, for example, because of small survival chances. We here assess whether filial cannibalism (FC), where parents routinely eat some of their own young, is selective in response to individual offspring reproductive value. We performed two independent laboratory experiments in the common goby (Pomatoschistus microps) to test whether caring fathers preferentially cannibalize eggs of a given infection history and paternity. While males did not discriminate kin from nonkin eggs, they consumed significantly more eggs previously exposed to water mold compared to uninfected eggs. Our findings clearly show that parents differentiate between eggs based on differences in egg condition, and thus complement the prevailing view that FC arises for energetic reasons. By preventing the spread of microbial infections, the removal of molded eggs can constitute an important component of parental care and may represent a key driver of selective FC in a wide array of parental fish.     

Cannibalizing Harmonia axyridis (Coleoptera: Coccinellidae) larvae use endogenous cues to avoid eating relatives

Cannibalism is widespread among many different organisms, and can have both negative and positive fitness consequences. Avoiding eating relatives can minimize negative fitness consequences of cannibalism. Such avoidance requires kin discrimination, but evidence for this ability among cannibals is limited with little data that address the cues used in such discrimination. We examined whether larvae of the ladybird beetle Harmonia axyridis avoid eating their relatives. We further manipulated environmental factors to create individuals that had similar or dissimilar rearing environments to begin to test for endogenous versus exogenous recognition cues. In our experiments, third-instar larvae were much less likely to cannibalize if they were interacting with a relative. Larvae that did cannibalize kin required more encounters, and significantly delayed cannibalism, compared to larvae cannibalizing unrelated individuals. Acquired cues were less important. Even though the different rearing environment resulted in significantly different phenotypic effects, similarity or dissimilarity of rearing environment had no effect on cannibalism, and there was no interaction between environments and relatedness in cannibalism. We suggest that H. axyridis has a well-developed kin discrimination system, and that kin recognition in this ladybird beetle is based on endogenous rather than exogenous cues. We also argue that these cues reflect either direct or indirect genetic effects on larval phenotypes.     

Nest and Egg Recognition by Salamanders in the Genus Desmognathus: A Comprehensive Re‐Examination

We conducted a comparative investigation of nest relocation and egg recognition by four species of streamside salamanders indigenous to eastern North America: Seepage Salamander (Desmognathus aeneus), Santeetlah Salamander (Desmognathus santeetlah), Ocoee Salamander (Desmognathus ocoee), and Northern Dusky Salamander (Desmognathus fuscus). Females of all four species were able to relocate their nest sites following displacement of 1 m. Upon return to a nest site following natural displacement, females must be able to recognize their eggs and, in some instances, choose between their own clutch and the unattended clutch of a conspecific. In two‐choice behavioral tests, female salamanders of all four species moved randomly within the test chamber in the absence of eggs, but preferred their own eggs to a filter paper blank. One species, D. fuscus, exhibited a similar attraction to conspecific eggs suggesting that the presence of eggs, regardless of their origin, may stimulate maternal care in physiologically primed females. We found that egg discrimination is not as infallible as previously reported. All four species spent more time with their own eggs than they did with the eggs of a conspecific female; however, this trend was significant for D. ocoee only. Interspecific variation in egg discrimination is attributed to differential selection associated with the life history of individual species.