Filial cannibalism in an assassin bug

Filial cannibalism has been described in many fish species with male parental care, and has typically been explained as a response to high energetic costs of brood defence and decreased feeding opportunities during the period of care. We investigated filial cannibalism in an insect, the assassin bug Rhinocoris tristis. In this species, males guard eggs of a number of females, cannibalizing some of their offspring within the brood. We monitored guarding males in both the field and the laboratory. Males typically ate eggs around the periphery of the brood, which were those most likely to have been parasitized by wasps. However, cannibalism persisted in the laboratory in the absence of parasites, and the number of cannibalized eggs was related to the length of care and overall brood size, suggesting that males use eggs as an alternative source of food. This conclusion was further supported by the fact that males in the field did not lose weight while guarding, despite being unable to forage efficiently while caring. Males were also observed to adopt broods, but in a laboratory experiment did not eat more eggs from adopted than from their own broods.     

Filial Cannibalism in the Biparental Fish Cichlasoma nigrofasciatum (Pisces: Cichlidae) in Response to Early Brood Reductions

Filial cannibalism (eating one's own offspring) may enhance a parent's lifetime reproductive success if the costs associated with this behaviour are outweighed by its benefits. An organism might limit its parental care to relatively high quality offspring and eat the others. Or, an organism capable of repeated breeding in the same season might eat the survivors of a brood that was reduced by predators, and breed again. In this study, we manipulated brood size of convict cichlids by removing 0 % (control), 33 % (ER 33) or 66 % (ER 66) of the eggs spawned. The results show that smaller broods are more likely to be cannibalized by their parent(s) than are larger broods. Furthermore, pairs with reduced broods were preparing to respawn; female gonad weights were significantly higher in the brood-reduction groups than in the control group. In a second experiment, we show that the behaviour of the parents differed significantly between experimental groups; the ER 66 group performed less parental care than the control group. Moreover, females invested more parental effort than males. The results suggest that filial cannibalism may be a tactic used by parental convict cichlids to enhance their lifetime reproductive success.     

Seasonal pattern of filial cannibalism by Apogon doederleini mouthbrooding males

In temperate waters of southern Japan, mouthbrooding males of the cardinal fish Apogon doederleini sometimes ate their entire brood within a day of spawning. In spite of annual variation in age structure and length of the breeding season, however, the annual cannibalism rate (broods cannibalized to total broods spawned) was nearly constant, 12–16%. Fish 1 year old frequently cannibalized early broods of the season, especially the first brood, whereas the cannibalism rate by 2 year-old fish did not vary within the season. In contrast, the cannibalism by older males increased late in the season. This tendency was attributed to their different allocation patterns to growth and reproduction. For 1 year-old fish, which can enhance future reproductive success by growth and cannibalizing on early broods instead of mouthbrooding (at relatively low temperatures), this may be a tactic for investing energy in growth. For fish 3 years, which have more breeding cycles and grow little, cannibalizing late broods may be energetic compensation for their poor somatic condition. No seasonal variation of cannibalism by 2 year-old fish can be explained by their intermediate nature of growth and reproduction. Multiple mate availability is one factor facilitating the cannibalism by 2 and 3 + year-old fish whose operational sex ratio is female-biased.     

Parental fish change their cannibalistic behaviour in response to the cost-to-benefit ratio of parental care

Partial filial cannibalism, the act of cannibalizing some offspring, has been explained as a response to the high energetic cost of care. I tested this hypothesis by manipulating the cost-to-benefit ratio of care in the scissortail sergeant, Abudefduf sexfasciatus, a tropical damselfish with male care. Background egg mortality was lower than the incidence of cannibalism, confirming that males did not just dispose of dead eggs. Investment in the current brood affected future investment, because males forced to skip a brood cycle put more effort into courtship during the following cycle and obtained larger broods than did unmanipulated males. Any factor influencing the cost-to-benefit ratio of parental care should also affect the incidence of cannibalism. I reduced the cost of care by supplementary feeding and reduced the benefit of care by simulating a decrease in paternity certainty through simulated intrusions by non-nesting males. Supplementary feeding significantly reduced partial filial cannibalism by parental males, a result compatible with the hypothesis that eggs are consumed to cover the energetic costs of parental care. Cannibalism decreased regardless of whether males were fed with conspecific eggs or crabmeat. Cannibalism was only reduced but not fully eliminated by supplementary feeding, and residual levels of cannibalism after feeding were similar to the background rate of egg mortality. Simulated intrusions by non-nesting males led to an increase in filial cannibalism and a decrease in parental effort.     

Parasites and parental care in male upland bullies (Eleotridae)

Parental care of eggs by male upland bullies Gobiomorphus breviceps was studied in the field and in the laboratory, with special emphasis on the effect of infection by trematode cysts. In the field, the proportion of old eggs in the nest affected the time taken by males to return to their nest after being chased away, but only in one year of the study. No effects of male size, male parasite load, number of eggs per nest or nest characteristics were observed. In the laboratory, heavily-infected males spent more time fanning their eggs than lightly-infected ones. Smaller males also tended to spend more time fanning than large males. Male size, parasite load and number of eggs per nest had no measurable effect on the aggressiveness of guarding males towards intruders, weight loss in males or egg incubation time. Parental investments did not vary as a function of age of the eggs. All males cannibalized some of their eggs, and there appeared to be a threshold brood size below which males consumed their entire brood. Overall, results were not consistent with those of studies on other fish species, and indicate that rules of investment in parental care may be different for fish inhabiting harsh, unpredictable environments.     

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.

     

Hurry-up and hatch: selective filial cannibalism of slower developing eggs

Filial cannibalism (the consumption of one's own offspring) is thought to represent an adaptive strategy in many animals. However, little is known about the details of which offspring are consumed when a parent cannibalizes. Here, we examined patterns of within-brood filial cannibalism in the sand goby (Pomatoschistus minutus). Males spawned sequentially with two females, and we asked whether males cannibalized selectively with regard to egg size or the order in which eggs were received. Males preferentially consumed the larger eggs of the second female they spawned with. Because larger eggs took longer to hatch, and because female 2's eggs were up to 1 day behind those of female 1, such preferential cannibalism might allow males to decrease the time spent caring for the current brood and re-enter the mating pool sooner. More work is needed to understand the fitness consequences of such selective cannibalism.     

Is the small clutch size of a Corsican blue tit population optimal?

In an attempt to test predictions of the optimisation hypothesis of life history traits in birds, we estimated fitness consequences of brood size manipulations. Experiments were carried out over a period of 4 years in a Mediterranean population of blue tits Parus caeruleus which is confronted with a particular set of environmental constraints. Effects of brood size manipulation were investigated in relation to year-to-year variation in environmental conditions, especially caterpillar abundance. There was a strong variation in the effects of brood size manipulation depending on year. Most effects were on offspring quality (fledging mass, tarsus length). The absolute number of recruits did not significantly differ among categories (reduced, control, enlarged broods) but varied considerably among years. Females recruited from enlarged broods were of lower quality, started to breed later and laid fewer eggs than those recruited from control and reduced broods. Neither parental survival nor reproductive performances of adults in year n + 1 was affected by brood size manipulation in year n. Thus there was no evidence for a cost of reproduction in this population. Since the number of recruits did not depend on brood size manipulation (recruitment rates were higher in reduced broods), but recruits from reduced broods were of better quality compared with other groups, we conclude that adults lay a clutch that is larger than that which is predicted by the optimisation hypothesis. Producing more young could incur some penalties because offspring from large broods are of lower quality and less likely to recruit in the population. Two possible reasons why decision rules in this population seem to be suboptimal are discussed. Received: 10 March 1998 / Accepted: 1 July 1998     

Survival for specific life intervals of smallmouth bass, Micropterus dolomieu, during parental care

We tested whether daily mortality rates (DMR) of smallmouth bass offspring were influenced by life interval, offspring density and growth, parental male attributes, and selected mortality factors during parental care in a regulated Virginia stream. Mortality averaged 9.5% per day (range 5.2–13.9%) and 94.1% total (range 80.9–99.5%) from egg deposition to the juvenile period (29–36 d) for individual broods. Offspring losses were primarily attributed to fungus (Saprolegnia parasitica) infection of eggs and to American eel, Anguilla rostrata, predation. DMR were significantly higher for the interval from swim-up of larvae to metamorphosis relative to earlier and later intervals. There was no significant autocorrelation of DMR among life intervals for individual broods, indicating that relative mortality rates were inconsistent among broods through time. DMR were also uncorrelated with the number of offspring per brood, offspring growth rates, and parental male attributes, except during egg and embryo intervals. Daily egg mortality was negatively related to male size and positively related to the number of eggs per nest, suggesting that density-dependent egg mortality may have been partially offset in nests of larger males. Larger males received more eggs, tended to maintain larger broods throughout parental care, and contributed a high proportion of the total number of juveniles reared. This revised version was published online in August 2006 with corrections to the Cover Date.     

Filial cannibalism in Tilapia mariae

In 152 isolated pairs of Tilapia mariae the parents ate 41% of their broods before these reached the frees-wimming stage. The cannibalized broods were usually eaten at the egg-stage. Most brood-eating was done by the males, as concluded from direct observations and from variation between males (but not between females) in the tendency of eat broods. Cannibalistic behaviour was more common when: 1. the relative weight of female (female weight to male weight) was small (< .4) 2. the relative weight was large (> .7) but not too large (> .85) 3. the female was large (> 70 g) 4. the pair spawned at certain times of the year (outside breeding peaks) 5. the spawn was exceptionally small (less than 100 eggs). In normally sized spawns neither the number of eggs nor the size of the eggs or the spawn could be related to cannibalism. The probability of cannibalism was not altered in the second spawnings of the pairs after an experience of a first unsuccessful spawning. Many of these changes in frequency of cannibalism can be related to the males' expected reproductive success. Thus they possibly reflect an underlying natural adaptive variation which partly has been modified by the aquarium milieu.     

Interaction between parental care and sibling competition: parents enhance offspring growth and exacerbate sibling competition

Species with elaborate parental care often also show intense sibling competition over resources provided by parents, suggesting joint evolution of these two traits. Despite this, the evolution of elaborate parental care and the evolution of intense sibling competition are often studied separately. Here, we examine the interaction between parental food provisioning and sibling competition for resources through the joint manipulation of the presence or absence of parents and brood size in a species with facultative parental care: the burying beetle Nicrophorus vespilloides. The effect of the interaction between the presence or absence of parents and brood size was strong; brood size had a strong effect on growth when parents provided care, but no effect when parents were absent. As in previous studies, offspring grew faster when parents were present than when parents were absent, and offspring grew faster in smaller broods than in larger broods. Our behavioral observations showed that brood size had a negative effect on both the amount of time parents spent providing resources to individual offspring and the offspring's effectiveness of begging, confirming that the level of sibling competition increased with brood size. Furthermore, offspring in larger broods shifted more from begging toward self-feeding as they grew older compared to offspring in small broods. Our study provides novel insights into the joint evolution of parental care and sibling competition, and the evolution of offspring begging signals. We discuss the implications of our results in light of recent theoretical work on the evolution of parental care, sibling competition, and offspring begging signals.     

Twofold cost of reproduction: an increase in parental effort leads to higher malarial parasitaemia and to a decrease in resistance to oxidative stress

Parental effort is usually associated with high metabolism that could lead to an increase in the production of reactive oxidative species giving rise to oxidative stress. Since many antioxidants involved in the resistance to oxidative stress can also enhance immune function, an increase in parental effort may diminish the level of antioxidants otherwise involved in parasite resistance. In the present study, we performed brood size manipulation in a population of great tits (Parus major) to create different levels of parental effort. We measured resistance to oxidative stress and used a newly developed quantitative PCR assay to quantify malarial parasitaemia. We found that males with an enlarged brood had significantly higher level of malarial parasites and lower red blood cell resistance to free radicals than males rearing control and reduced broods. Brood size manipulation did not affect female parasitaemia, although females with an enlarged brood had lower red blood cell resistance than females with control and reduced broods. However, for both sexes, there was no relationship between the level of parasitaemia and resistance to oxidative stress, suggesting a twofold cost of reproduction. Our results thus suggest the presence of two proximate and independent mechanisms for the well-documented trade-off between current reproductive effort and parental survival.     

Intraseasonal effects of clutch manipulation on parental provisioning and residual reproductive value of eastern phoebes (Sayornis phoebe)

Summary First clutches of double-brooded eastern phoebes Sayornis phoebe were manipulated (up two eggs, down 2 eggs or no change) to test for intraseasonal reproductive tradeoffs and to test whether size of first brood influenced food delivery rates to nestlings and nestling quality in second broods.Considering all nests from both broods, rate of feeding nestlings increased linearly with brood size but nestling mass per nest decreased with increasing brood size. High nestling weights in small broods may have resulted from parents delivering better quality food, but we did not test this.Among treatment groups in first broods, nestlings from decreased broods weighed more than those in control or increased broods. Treatment did not influence the likelihood that second nests would be attempted after successful first nests nor did it alter the interval between nests. Nestlings of parents that renested weighed more than those of parents that did not, regardless of treatment, suggesting that post-fledging care may preclude renesting. Mass of individual females did not change between broods, regardless of brood size. Clutch sizes of second attempts were not affected by manipulations of first broods but increasing first broods reduced the number of nestlings parents were able to raise to day 11 in their second broods. However, manipulation of first broods did not affect mean nestling mass per nest of nestlings that survived to day 11.In phoebes, parents of small first broods are able to raise nestlings in better condition. We predict that in harsh years, parents of small first broods would be more likely to renest. Parents of enlarged first broods sacrificed quality of offspring in second broods, which seems a reasonable strategy if nestlings from second broods have lower reproductive value.     

The effect of partial brood loss on male desertion in a cichlid fish: an experimental test

There is little experimental evidence testing whether currentbrood size and past brood mortality influence mate desertion.In the cichlid Aequidens coeruleopunctatus both parents initiallydefend offspring. In a field study, all experimental broods,irrespective of initial brood size (222.9 ± 60.4, mean± SD), were manipulated to a size of 100 fry. Neitherthe duration nor investment of females in parental care differed between control and brood reduced pairs, even though care seemedcostly. On average, females lost 5.1 ± 4.8% of initialweight while guarding a brood until independence. In contrast,males with experimentally reduced broods guarded fry for significantlyfewer days before deserting their mate than did males fromcontrol pairs with natural-sized broods (20.5 ± 7.5 vs. 14.2 ± 6.2 days). In at least 20% of cases (n = 9/45),the deserting male immediately mated with another female. Maleswith experimentally reduced broods also spent less time guardingfry before deserting and attacked fewer brood predators thandid males with control broods. For broods manipulated to have100 fry, there was a significant negative relationship betweenthe days until male desertion and the proportion of the initialbrood removed. This indicates that male assessment of the futuresuccess of the current brood (hence its reproductive value)is based on past mortality and/or that there is variation amongmales in the expected size of future broods. Both current broodsize and brood size relative to initial brood size are thereforepredictors of male, but not female, parental behavior and matedesertion. Female care may be unaffected by brood reductiondue to limited breeding opportunities and partial compensationfor reduced male care.     

Adjusting parental investment to changing environmental conditions: the effect of food ration on parental behaviour of the convict cichlid, Cichlasoma nigrofasciatum

Animals that care for their offspring may vary the amount of care provided for a particular brood in relation to environmental conditions. Food availablity is one factor that may affect the costs and benefits associated with parental investment. The convict cichlid, Cichlasoma nigrofasciatum, is a small, substrate-spawning cichlid from Central America. Both male and female provide parental care for eggs and fry. Paris were kept at one of three ration levels, high, medium or low. Time spent in parental fanning by females was positively related to ration. Males spent less time fanning than females and their parental behaviour varied considerably between individuals. Males on the high ration spent slightly more time fanning than those on the lower rations. The number of eggs produced per spawning and the post-spawning weight of both males and females were significantly and positively related to ration. Foraging frequency of both males and females was inversely related to ration. There was no significant effect of ration on the frequency of mouthing eggs and young or on intra-pair aggression. Eggs of low-ration fish hatched earlier than those of medium- and high-ration fish but there was no significant difference in the number of days that the young survived. These results suggest that the allocation of time and effort between parental and maintenance activities differs in relation to food supply. Parents may provide more care for the large brood produced when food is plentiful but place more emphasis on their own survival when food is short and broods are small.     

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.     

Parental investment in an anadromous population of threespine sticklebacks: An experimental study

Summary Two previously published field studies of threespine sticklebacks conducted to test predictions of parental investment (PI) theory yielded different conclusions about how males invest in their young. Males of a freshwater population invested more in older/larger broods whereas environmental factors played no role in PI decisions. In contrast, males of an anadromous population adjusted levels of PI in response to environmental factors rather than brood characteristics. The current laboratory study attempts to determine possible reasons for the discrepancy between these two studies. We addressed the following questions. Do males invest more in larger/older broods? Do environmental factors affect levels of parental investment? Males were more aggressive (a measure of PI) in defending eggs than empty nests and free-swimming fry. However, aggression was similar for different-aged eggs and different-aged fry. Moreover, when we manipulated brood sizes (fry), males did not change their level of investment. Males increased their level of aggressive defence in response to a rise in water level, indicating that they can adjust their level of investment in response to changes in environmental conditions that affect brood survival.     

Cumulative effects matter: multi‐brood responses of Daphnia to hypoxia

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Maternal investment in a spider with suicidal maternal care, Stegodyphus lineatus (Araneae, Eresidae)

Providing parental care is costly for the parent, but generally beneficial for the young whose survival, growth and reproductive value can be increased. Selection should strongly favour an optimal distribution of parental resources, depending on the relationship between the costs and benefits for parents and their offspring. Parental care is characterized by trade offs in investment, for example between egg size and number of young or providing resources at the egg stage versus the post-hatching stage. Females of the spider Stegodyphus lineatus (Eresidae) produce a single small brood with small eggs and provide the young with regurgitated fluid and later, with their body contents via matriphagy. We asked whether females adjust the investment of resources differentially into eggs, regurgitation feeding and matriphagy, and how maternal investment affects the size of the young at dispersal. We followed the growth of young of broods in the lab and in the field and manipulated brood size in order to determine the pattern of resource allocation. We found that brood size was positively correlated with body mass: larger females had larger broods. Females provided 95% of their body mass to the young, allocating more resources to regurgitation than to matriphagy. Females provided regurgitated food to the young according to the brood size, providing less food when the brood was reduced. Maternal resources had a large influence on offspring mass at dispersal, which is likely to affect their future fitness. The study shows the importance of the female's body mass and her resource allocation decisions for her reproductive outcome.     

Azure‐winged Magpies Cyanopica cyanus trade off reproductive success and parental care by establishing a size hierarchy among nestlings

It is common in birds that the sizes of nestlings vary greatly when multiple young are produced in one nest. However, the methods used by parents to establish size hierarchy among nestlings and their effect on parental provisioning pattern may differ between species. In the Azure‐winged Magpie Cyanopica cyanus, we explored how and why parents controlled the sizes of nestlings. Asynchronous hatching was the main cause of size hierarchy within the brood, although the laying of larger eggs later in the laying sequence reduced this effect. Parents with asynchronous broods produced more eggs and fledged more nestlings than those with synchronous broods but their brood provisioning rates, food delivery per feeding bout and feeding efficiency did not differ. We performed a cross‐fostering experiment to synchronize some asynchronous broods. Provisioning rates of asynchronous broods were lower than those of synchronized broods, but the daily growth rates and fledging body mass of their nestlings were not different. Our findings indicate that parents of asynchronous broods can achieve higher reproductive success than those of synchronous broods based on the same parental care, and the same reproductive success as those of synchronized broods based on less parental care. It appears that parent birds can better trade off reproductive success and parental care by establishing a size hierarchy among nestlings.