The pay‐offs of maternal care increase as offspring develop,favouring extended provisioning in an egg‐feeding frog

Offspring quantity and quality are components of parental fitness that cannot be maximized simultaneously. When the benefits of investing in offspring quality decline, parents are expected to shift investment towards offspring quantity (other reproductive opportunities). Even when mothers retain complete control of resource allocation, offspring control whether to allocate investment to growth or development towards independence, and this shared control may generate parent–offspring conflict over the duration of care. We examined these predictions by, in a captive colony, experimentally removing tadpoles of the strawberry poison frog (Oophaga pumilio) from the mothers that provision them with trophic eggs throughout development. Tadpoles removed from their mothers were no less likely to survive to nutritional independence (i.e. through metamorphosis) than were those that remained with their mothers, but these offspring were smaller at metamorphosis and were less likely to survive to reach adult size, even though they were fed ad libitum. Tadpoles that remained with their mothers developed more slowly than those not receiving care, a pattern that might suggest that offspring extracted more care than was in mothers’ best interests. However, the fitness returns of providing care increased with offspring development, suggesting that mothers would be best off continuing care until tadpoles initiated metamorphosis. Although the benefits of parental investment in offspring quality are often thought to asymptote at high levels, driving parent–offspring conflict over weaning, this assumption may not hold over natural ranges of investment, with selection on both parents and offspring favouring extended durations of parental care.     

Territorial behavior of the Korean wood-feeding cockroach, <Emphasis Type="Italic">Cryptocercus kyebangensis</Emphasis>

We investigated the territorial behavior of Cryptocercus kyebangensis from the viewpoint of offspring protection. Aggressive behavior against adult intruders occurred more frequently in resident adults with their young than those without their young. Resident adults without offspring more frequently attacked intruders of the same sex than those of the opposite sex, whereas those with offspring attacked regardless of the sex of adult intruders. In paired residents, cooperative territorial behavior occurred in a higher proportion in pairs with offspring than those without offspring. Responses of residents to nymphal intruders depended on the head capsule size of the intruders. Resident adults rarely attacked intruders smaller than their own offspring but frequently attacked larger intruders. Resident nymphs frequently attacked intruders smaller than themselves, whereas they escaped or showed jerking behavior when they encountered larger intruders. Electronic Publication     

Are species differences in maternal effects arising from maternal care adaptive?

Parental care benefits offspring through maternal effects influencing their development, growth and survival. However, although parental care in general is likely the result of adaptive evolution, it does not follow that specific differences in the maternal effects that arise from care are also adaptive. Here, we used an interspecific cross‐fostering design in the burying beetle species Nicrophorus orbicollis and N. vespilloides, both of which have elaborate parental care involving direct feeding of regurgitated food to offspring, to test whether maternal effects are optimized within a species and therefore adaptive. Using a full‐factorial design, we first demonstrated that N. orbicollis care for offspring longer regardless of recipient species. We then examined offspring development and mass in offspring reared by hetero‐ or conspecific parents. As expected, there were species‐specific direct effects independent of the maternal effects, as N. orbicollis larvae were larger and took longer to develop than N. vespilloides regardless of caregiver. We also found significant differences in maternal effects: N. vespilloides maternal care caused more rapid development of offspring of either species. Contrary to expectations if maternal effects were species‐specific, there were no significant interactions between caretaker and recipient species for either development time or mass, suggesting that these maternal effects are general rather than optimized within species. We suggest that rather than coadaptation between parents and offspring performance, the species differences in maternal effects may be correlated with direct effects, and that their evolution is driven by selection on those direct effects.     

Offspring of older parents are smaller—but no less bilaterally symmetrical—than offspring of younger parents in the aquatic plant Lemna turionifera

Offspring quality decreases with parental age in many taxa, with offspring of older parents exhibiting reduced life span, reproductive capacity, and fitness, compared to offspring of younger parents. These “parental age effects,” whose consequences arise in the next generation, can be considered as manifestations of parental senescence, in addition to the more familiar age‐related declines in parent‐generation survival and reproduction. Parental age effects are important because they may have feedback effects on the evolution of demographic trajectories and longevity. In addition to altering the timing of offspring life‐history milestones, parental age effects can also have a negative impact on offspring size, with offspring of older parents being smaller than offspring of younger parents. Here, we consider the effects of advancing parental age on a different aspect of offspring morphology, body symmetry. In this study, we followed all 403 offspring of 30 parents of a bilaterally symmetrical, clonally reproducing aquatic plant species, Lemna turionifera, to test the hypothesis that successive offspring become less symmetrical as their parent ages, using the “Continuous Symmetry Measure” as an index. Although successive offspring of aging parents older than one week became smaller and smaller, we found scant evidence for any reduction in bilateral symmetry.     

Parental condition affects early life-history of a coral reef fish

Parents can exert a range of non-genetic effects on the growth and survival of their offspring. In particular, parents may modify the size or condition of their offspring depending on the amount of energy they have available for reproduction. In this study, the body condition of breeding pairs of the coral reef fish Acanthochromis polyacanthus was experimentally manipulated to test the effects of parental condition on reproductive output and offspring life history traits. Parents in good condition commenced breeding earlier, had higher reproductive output, and their eggs exhibited increased survival during embryogenesis, compared to parents in poorer condition. Increased reproductive output was attained through more reproductive bouts over the breeding season that contained both a greater number and an increased size of eggs. The offspring from parents in good condition were larger at hatching, with larger yolk reserves and increased survival on endogenous reserves. Larger size is expected to provide benefits to offspring through reduced susceptibility to size-selective mortality. The range of offspring characteristics modified by parental condition could result in a greater proportion of offspring from good condition parents recruiting to the population.     

Foster rather than biological parental telomere length predicts offspring survival and telomere length in king penguins

Because telomere length and dynamics relate to individual growth, reproductive investment and survival, telomeres have emerged as possible markers of individual quality. Here, we tested the hypothesis that, in species with parental care, parental telomere length can be a marker of parental quality that predicts offspring phenotype and survival. In king penguins (Aptenodytes patagonicus), we experimentally swapped the single egg of 66 breeding pairs just after egg laying to disentangle the contribution of prelaying parental quality (e.g., genetics, investment in the egg) and/or postlaying parental quality (e.g., incubation, postnatal feeding rate) on offspring growth, telomere length and survival. Parental quality was estimated through the joint effects of biological and foster parent telomere length on offspring traits, both soon after hatching (day 10) and at the end of the prewinter growth period (day 105). We expected that offspring traits would be mostly related to the telomere lengths (i.e., quality) of biological parents at day 10 and to the telomere lengths of foster parents at day 105. Results show that chick survival up to 10 days was negatively related to biological fathers’ telomere length, whereas survival up to 105 days was positively related to foster fathers’ telomere lengths. Chick growth was not related to either biological or foster parents’ telomere length. Chick telomere length was positively related to foster mothers’ telomere length at both 10 and 105 days. Overall, our study shows that, in a species with biparental care, parents’ telomere length is foremost a proxy of postlaying parental care quality, supporting the “telomere – parental quality hypothesis.”     

Functional aspects of prolonged parental care in Bewick's swans

In Bewick's swans prolonged parental care is important in protecting offspring from feeding competition with other individuals in winter flocks. When separated from their parents, cygnets were less successful in aggressive encounters, were frequently displaced and spent less time feeding than when close to their parents. parents protected their offspring by intervening in encounters on their behalf. They were able to do so because of their high dominance rank within the flock. Differences in dominance rank among families were correlated with differences in dominance of cygnets both with and away from their parents, and preliminary evidence suggested that cygnets may assume their parents' dominance, the effect continuing into the second winter. Parental care involved measurable costs to both parents at the beginning of the winter.     

Contrasting Parental Tasks Influence Parental Roles for Paired and Single Biparental Cichlid Fish

Biparental care of young occurs when both parents provide some sort of care for offspring and can include a wide variety of behaviors, yet often studies focus on single aspects of parental care when trying to determine how each parent contributes. Here, we presented the biparental convict cichlid fish (Amatitlania nigrofasciata) with two conflicting parental behaviors: retrieval of non‐swimming offspring that have been displaced from the nest and defense against a conspecific intruder, a potential brood predator. We examined single males, single females, and pairs of parents to determine how males and females each contributed to overall parental care. Traditionally in this species, parents exhibit a division of labor (in which each parent contributes to all parental activities), but also exhibit a division of roles (in which males tend to favor the role of defense and females tend to favor more direct care, spending more time with offspring). We hypothesized that single parents would compensate for their absent mate, but that males and females would still favor preferred roles. Additionally, we hypothesized that there would be an asymmetrical expansion of roles, with females being more flexible. Our results show that the preferred roles of both parents were evident even when parents were without their mates and that males and females differed in their compensatory levels, at least when compared to the behaviors of the intact pair. Contrary to our prediction, females seem unable to fully compensate for the defensive behaviors usually exhibited by males, while males shifted completely to retrieve displaced offspring.     

Strain‐specific effects of parental age on offspring in Drosophila melanogaster

Maternal age is generally known to be negatively correlated with the lifespan of offspring in several animal models including yeast, rotifers, flies, and possibly in humans. However, several reports have shown positive effects of parental age on offspring lifespan. Thus, there was a need to investigate further the inconsistent results on the effect of parental age on lifespan. In this study, the effects of parental age on offspring fitness and lifespan were examined by using Drosophila melanogaster. The lifespan of offspring from old parents was significantly increased compared with that of the young counterparts in the Canton‐S (CS) strain but not in other D. melanogaster strains, such as Oregon‐R (OR) and w¹¹¹⁸. To find out why the lifespan is increased in the offspring from old parents in CS flies, fitness components that could modulate lifespan were examined in CS flies. Egg weight and body weight were reduced by parental aging and the offspring of old fathers or old mothers developed faster than that of the young. In addition, the offspring of old parents had increased resistance to oxidative and heat shock stresses. However, reproductive capacity, mating preference, and food intake were unaffected by parental aging. These results indicate that parental aging in CS strain D. melanogaster has beneficial effects on the lifespan and fitness of offspring. The presence of strain‐specific manner effects suggests that genetic background might be a significant factor in the parental age effect.     

Costly parenting: physiological condition over time and season in males of the giant waterbug Abedus dilatatus

Parental care patterns increase offspring fitness but may drive energetic costs to parents. The costs associated with parental care can change over time, decreasing the condition of parents that experience prolonged parental care. Thus, males can modulate parental effort based in the relative fitness cost/benefit pay‐offs under different stages and environmental conditions. The present study assesses the condition of parental males of Abedus dilatatus Say by measuring their lipid, glycogen and carbohydrate contents. We compare the condition in parental males that have experienced recent and prolonged care and that were also collected in the summer and winter. Waterbug males provide parental care via carrying and ventilating eggs on their back. Winter males are smaller and carry fewer eggs compared with summer males. Males with recent care and carrying more eggs present a more lipid content. However, at the end of care, males carrying more eggs present less lipids than males with smaller egg‐pads. Additionally, we find that males collected in the summer present more carbohydrates than males in the winter. Moreover, larger males with prolonged care present less carbohydrates than smaller males, in contrast to males with recent care where there is a positive relationship between size and carbohydrate content. Our results suggest that parental care in A. dilatatus may be a sexually‐selected trait, as has been found in related species, and further experiments could test this idea. This is the first study to provide evidence of physiological costs related to exclusive paternal care in arthropods.     

Sex‐Specific Parental Care in Response to Predation Risk in the European Roller,Coracias garrulus

Sublethal effects of predation constitute an important part of predation effects, which may modulate prey population and community dynamics. In birds, the risk of nest predation may cause a reduction in parental activity in the care of offspring to reduce the chance of being detected by predators. In addition, parents may modify their parental food allocation preferences within the brood in response to predation risk. Our aim in this study was to evaluate the effects of risk of nest predation on parental care and within‐nest food allocation in the European Roller (Coracias garrulus), an asynchronously hatching bird. We manipulated brood predation risk by placing a snake model near the nests that simulates the most common nest predator in the Mediterranean region. Our results show that males but not females increased their provisioning rate when they were exposed to the model and that despite this, nestlings’ body mass decreased in response to this temporary increase in predation risk. We did not find evidence that parents changed their food allocation strategy towards senior or junior nestlings in their nests in response to predation risk. These results show that the European roller modifies parental care in response to their perception of predation risk in the nest and a sex‐specific sensitivity to the threat, which suggests a different perception of offspring reproductive value by parents. Finally, our results show that changes in parental behaviour in response to nest predation risk might have consequences for nestling fitness prospects.     

PARENTAL EFFECTS IN PLANTAGO LANCEOLATA L. I.: A GROWTH CHAMBER EXPERIMENT TO EXAMINE PRE- AND POSTZYGOTIC TEMPERATURE EFFECTS

In spite of the potential evolutionary importance of parental effects, many aspects of these effects remain inadequately explained. This paper explores both their causes and potential consequences for the evolution of life-history traits in plants. In a growth chamber experiment, I manipulated the pre- and postzygotic temperatures of both parents of controlled crosses of Plantago lanceolata. All offspring traits were affected by parental temperature. On average, low parental temperature increased seed weight, reduced germination and offspring growth rate, and accelerated onset of reproduction by 7%, 50%, 5%, and 47%, respectively, when compared to the effects of high parental temperature. Both pre- and postzygotic parental temperatures (i.e., prior to fertilization vs. during fertilization and seed set, respectively) influenced offspring traits but not always in the same direction. In all cases, however, the postzygotic effect was stronger. The prezygotic effects were more often transmitted paternally than maternally. Growth and onset of reproduction were influenced both directly by parental temperature as well as indirectly via the effects of parental temperature on seed weight and germination. Significant interactions between parental genotypes and prezygotic temperature treatment (G × E interactions) show that genotypes differ in their intergenerational responses to temperature with respect to germination and growth. The data suggest that temperature is involved in both genetically based and environmentally induced parental effects and that parental temperature may accelerate the rate of evolutionary change in flowering time in natural populations of P. lanceolata. The environmentally induced temperature effects, as mediated through G × (prezygotic) E interactions are not likely to affect the rate or direction of evolutionary change in the traits examined because postzygotic temperature effects greatly exceed prezygotic effects.     

Biparental care is predominant and beneficial to parents in the burying beetle Nicrophorus orbicollis (Coleoptera: Silphidae)

Parenting strategies can be flexible within a species and may have varying fitness effects. Understanding this flexibility and its fitness consequences is important for understanding why parenting strategies evolve. In the present study, we investigate the fitness consequences of flexible parenting in the burying beetle Nicrophorus orbicollis, a species known for its advanced provisioning behaviour of regurgitated vertebrate carrion to offspring by both sexes. We show that, even when a parent is freely allowed to abandon the carcass at any point in time, biparental post‐hatching care is the most common pattern of care adopted in N. orbicollis. Furthermore, two parents together raised more offspring than single parents of either sex, showing that the presence of the male can directly influence parental fitness even in the absence of competitors. This contrasts with studies in other species of burying beetle, where biparental families do not differ in offspring number. This may explain why biparental care is more common in N. orbicollis than in other burying beetles. We suggest how the fitness benefits of two parents may play a role in the evolution and maintenance of flexible biparental care in N. orbicollis.     

Delayed dispersal in the Zaisan mole vole (Ellobius tancrei): helping or extended parental investment?

The highly social subterranean voles of genus Ellobius (mole vole) represent a unique model to compare with both social bathyergids and surface-dwelling voles. Unlike most arvicolines, mole voles display the prolonged obligatory delay of dispersal. In subterranean rodents, this delay may be associated with benefits of cooperation (weaned offspring help their parents), an extended parental investment (parents care for weaned offspring), or both. To identify the role offspring aged <3 months play in mole vole families, we estimated their contribution to important group activities. We found that juveniles contributed very little to daily living activities up to the age of approximately 2 months. The older offspring carried objects at least as frequently as breeders, but they engaged in gnawing obstacles less frequently than their fathers. Male breeders and non-breeders engaged in gnawing more than the respective categories of females. Although young mole voles clearly did not surpass their parents in performing the more energetically costly activities, they gained body mass by 26 g (130 %) between days 30 and 90, often surpassing their parents in weight. Based on our results, in E. tancrei an extremely prolonged infancy is followed by intensive building of body reserves. This ontogenetic trajectory appears to be distinct from the patterns described for social bathyergids and for most voles.     

Are offspring begging levels exaggerated beyond the parental optimum? Evidence from a bidirectional selection experiment

Parental care involves elaborate behavioural interactions between parents and their offspring, with offspring stimulating their parents via begging to provision resources. Thus, begging has direct fitness benefits as it enhances offspring growth and survival. It is nevertheless subject to a complex evolutionary trajectory, because begging may serve as a means for the offspring to manipulate parents in the context of evolutionary conflicts of interest. Furthermore, it has been hypothesized that begging is coadapted and potentially genetically correlated with parental care traits as a result of social selection. Further experiments on the causal processes that shape the evolution of begging are therefore essential. We applied bidirectional artificial selection on begging behaviour, using canaries (Serinus canaria) as a model species. We measured the response to selection, the consequences for offspring development, changes in parental care traits, here the rate of parental provisioning, as well as the effects on reproductive success. After three generations of selection, offspring differed in begging behaviour according to our artificial selection regime: nestlings of the high begging line begged significantly more than nestlings of the low begging line. Intriguingly, begging less benefitted the nestlings, as reflected by on average significantly higher growth rates, and increased reproductive success in terms of a higher number of fledglings in the low selected line. Begging could thus represent an exaggerated trait, possibly because parent–offspring conflict enhanced the selection on begging. We did not find evidence that we co‐selected on parental provisioning, which may be due to the lack of power, but may also suggest that the evolution of begging is probably not constrained by a genetic correlation between parental provisioning and offspring begging.     

The role of transgenerational effects in adaptation of clonal offspring of white clover (<Emphasis Type="Italic">Trifolium repens</Emphasis>) to drought and herbivory

Environmentally induced transgenerational effects can increase success of offspring and thereby be adaptive if offspring experience conditions similar to the parental environment. The ecological and evolutionary significance of these effects in plants have been considered overwhelmingly in the context of sexual generations. We investigated whether drought stress and jasmonic acid, a key hormone involved in induction of plant defenses against herbivores, applied in the parental generation, trigger transgenerational effects in clonal offspring of Trifolium repens and whether these effects are adaptive. We found that drought stress experienced by parents significantly affected phenotypes of offspring ramets. Offspring ramets were bigger if they were produced in the parental water regime (control/drought). Repeated application of jasmonic acid to parents increased the subsequent growth of offspring ramets produced by stolons after they were disconnected from the parental clone. However, these offspring ramets experienced similar herbivory by the generalist Spodoptera littoralis caterpillar as did control offspring ramets, indicating that this jasmonic acid application in the parental generation did not result in a transgenerational effect comprising increased herbivory resistance. We conclude that, overall, environmental interaction in the parental generation can trigger transgenerational effects in clonal plants and some of these effects can be adaptive. Moreover, transgenerational effects in clonal plants that significantly influence their growth and behavior can ultimately affect the evolutionary trajectories of clonal populations.     

Intergenerational effects of inbreeding in Nicrophorus vespilloides: offspring suffer fitness costs when either they or their parents are inbred

Inbreeding depression is the reduction in fitness caused by mating between related individuals. Inbreeding is expected to cause a reduction in offspring fitness when the offspring themselves are inbred, but outbred individuals may also suffer a reduction in fitness when they depend on care from inbred parents. At present, little is known about the significance of such intergenerational effects of inbreeding. Here, we report two experiments on the burying beetle Nicrophorus vespilloides, an insect with elaborate parental care, in which we investigated inbreeding depression in offspring when either the offspring themselves or their parents were inbred. We found substantial inbreeding depression when offspring were inbred, including reductions in hatching success of inbred eggs and survival of inbred offspring. We also found substantial inbreeding depression when parents were inbred, including reductions in hatching success of eggs produced by inbred parents and survival of outbred offspring that received care from inbred parents. Our results suggest that intergenerational effects of inbreeding can have substantial fitness costs to offspring, and that future studies need to incorporate such costs to obtain accurate estimates of inbreeding depression.     

The Effect of Female Condition on Maternal Care in the European Earwig

Parental care typically enhances offspring fitness at costs for tending parents. Asymmetries in genetic relatedness entail potential conflicts between parents and offspring over the duration and the amount of care. To understand how these conflicts are resolved evolutionarily, it is important to understand how individual condition affects offspring and parental behaviour and whether parents or offspring make active choices in their interactions. Condition effects on offspring have been broadly studied, but the effect of parental condition on parent–offspring interactions is less well understood, in particular in species where care is facultative and offspring have the option to beg for food from the parents or to self‐forage. In this study, we carried out two experiments in the European earwig Forficula auricularia, a system where females provide facultative care, in which we manipulated female condition (through a high‐food and low‐food treatment) and the degree by which mothers and offspring could make active choices. In a first experiment, where female mobility was limited, female condition had no significant effect on the rate of offspring self‐foraging, which increased with nymph age. In a second experiment, nymph access to food was limited and females in poor nutritional condition provided food to significantly fewer nymphs than high condition females. In both experiments, offspring attendance remained at a constantly high level and was independent of female condition even after experimental separation of females and offspring. Our results show that earwig nymphs do not use cues of female condition to adjust rates of self‐foraging, that females control food provisioning depending on their own condition, and that females and nymphs share control over offspring attendance, a form of care not influenced by female condition.     

Control of parental investment changes plastically over time with residual reproductive value

Evolutionary conflict between parents and offspring over parental resource investment is a significant selective force on the traits of both parents and offspring. Empirical studies have shown that for some species, the amount of parental investment is controlled by the parents, whereas in other species, it is controlled by the offspring. The main difference between these two strategies is the residual reproductive value of the parents or opportunities for future reproduction. Therefore, this could explain the patterns of control of parental investment at the species level. However, the residual reproductive value of the parents will change during their lifetime; therefore, parental influence on the amount of investment can be expected to change plastically. Here, we investigated control of parental investment when parents were young and had a high residual reproductive value, compared to when they were old and had a low residual reproductive value using a cross‐fostering experiment in the burying beetle Nicrophorus quadripunctatus. We found that parents exert greater control over parental investment when they are young, but parental control is weakened as the parents age. Our results demonstrate that control of parental investment is not fixed, but changes plastically during the parent's lifetime.     

Adjustment of parental expenditure in sympatric <Emphasis Type="Italic">Onthophagus</Emphasis> beetles (Coleoptera: Scarabaeidae)

Many organisms adjust their parental expenditure to offspring in response to resource quality. However, the mechanisms underlying the adjustment in parental expenditure are not well understood. We examined the adjustments in parental expenditure and subsequent offspring performance in two sympatric, closely related dung beetles, Onthophagus ater and O. fodiens, that were provided either monkey, deer, horse, or cow dung. The egg contained within each dung brood mass provisioned by the parent beetles develops to adulthood underground. Thus, the size of the brood mass roughly represents the amount of parental expenditure. The brood mass size differed between the two species and among the four dung types. Results of offspring performance suggested that O. ater parents optimally adjusted the brood mass size in response to dung quality, whereas O. fodiens parents did not. We hypothesized that brood mass size in O. ater may increase with prolonged egg maturation caused by the lower nutrition level of cow dung. In addition, our complex results may be explained in part by the specific threshold concept of dung quality (i.e., water content and nutritional level).