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Chemical cues and signals, especially in insects, play a pivotal role in mediating interactions between individuals. Past studies have largely focused on adult semiochemicals and have neglected those of juvenile stages. Especially in the context of parental care, the larval odor might have a profound impact on parenting behavior, guiding parents in how much resources they should allocate to the different developmental stages. However, whether ontogenetic changes occur in subsocial species and whether larval-emitted scents influence parent–offspring interactions is largely unknown. Using 3 different sampling techniques, we analyzed the cuticular and VOC profile of the 3 larval instars of the burying beetle Nicrophorus vespilloides , which is known for its elaborate parental care. We found distinct differences in the cuticular and VOC profiles across the 3 larval stages. Second-instar larvae, which receive more frequent feedings from parents than the other larval stages, released greater amounts of acetophenone, methyl geranate, and octanoic acid isopropyl ester than the first and third instar. Additionally, using a newly developed bioassay with automated video tracking, we found that adding the odor of second-instar larvae to first-instar larvae increased the number of maternal feeding trips. Our results suggest that the odor produced by larvae plays an important role in mediating parent–offspring interactions. Given these findings, burying beetles might emerge as a promising candidate for identifying a potential begging pheromone. [ABSTRACT FROM AUTHOR]

The widely spread evolutionary strategy of parental care is considered an important driver of social evolution. Although offspring were long thought to primarily interact competitively, recent studies revealed the potential importance of sibling cooperation. Theories suggest that the degree of cooperation in offspring interactions depends on the degree of offspring dependence on parental care: offspring unable to forage on their own should compete more, whereas more independent juveniles may increase the degree of cooperation. In this study, we tested the occurrence of sibling cooperation in the absence of posthatching care in several burying beetle species exhibiting varying degrees of offspring dependence. To this end, we measured larval growth rate and survival in the presence and absence of prehatching care using different brood sizes. We found that sibling cooperation cannot be exclusively explained by offspring dependence on parental care. Although only species with more independent larvae cooperated when receiving prehatching care, larval cooperation occurred across species in the absence of care. The latter result suggests that sibling cooperation was already present in an early ancestor of the genus Nicrophorus. Overall, these findings give important insights into the transition from facultative to obligate family life. [ABSTRACT FROM AUTHOR]

Microbial volatiles provide essential information for animals, which compete to detect, respond to, and perhaps control this information. Burying beetle parents have the opportunity to influence microbially derived semiochemicals, because they monopolize a small carcass for their family, repairing feeding holes and applying exudates that alter the microbial community. To study adaptive manipulation of microbial cues, we integrated mechanistic and functional approaches. We contrasted gas chromatography–mass spectrometry (GC-MS) volatile profiles from carcasses that were or were not prepared by a resident pair of Nicrophorus orbicollis. Methyl thiocyanate (MeSCN), the primary attractant for burying beetles seeking a fresh carcass, was reduced 20-fold by carcass preparation, while dimethyl trisulfide (DMTS), which deters breeding beetles, was increased 20-fold. These results suggest that parental care serves to make previously public information more private (crypsis, MeSCN) and to disinform rivals with a deterrent (DMTS). Functional tests in the field demonstrated that carcass preparation reduced discovery and use by congeners (threefold) as well as by dipteran rivals. Because microbes and their chemicals influence nearly every aspect of animal ecology, animal manipulation of microbial cues may be as widespread as manipulation of their own signals. [ABSTRACT FROM AUTHOR]

Abstract: Parental care is thought to be costly, as it consumes time and energy. Such costs might be reduced in animal parents that raise their young on valuable food sources such as dung or carcasses, as parents are able to invest in self‐maintenance by feeding from the same resource. However, this might lower the nutritional value for other family members and, as a consequence, food competition might arise. To promote our understanding of the outcome of such competition, we manipulated the necessity of parents to feed from the resource. Using a full factorial design, we paired food‐deprived or well‐fed males with food‐deprived or well‐fed females of burying beetles, which are known to raise their young on vertebrate cadavers. We found that food‐deprived parents consumed more of the carrion than those that were well‐fed and this had a negative impact on other family members. However, the outcome of the competition depended on the sex of the parents, with females suffering when males fed more and offspring suffering when females fed more. Thus, family life involves selfish elements, as both parents remove resources for the purpose of self‐maintenance. However, females show altruistic aspects, as they appear to restrict their food consumption for the benefit of their offspring when paired with a food‐deprived male. Interestingly, males extend their stay with the brood when having faced food scarcity prior to reproduction, presumably to replenish their energy reserves. Our study therefore reveals that breeding on shared resources can promote family living, but also results in competition. [ABSTRACT FROM AUTHOR]

Abstract: Recent studies demonstrate that pheromones can be costly to produce and emit and, therefore, the types and quantities that they express are likely to covary with individual condition. Previous experiments reveal that, when given the opportunity to breed and care for young, male burying beetles Nicrophorus vespilloides go on to produce a higher amount of their sex pheromone and attract more females than control males that do not exhibit parental care. This finding is surprising because parental care is usually assumed to be energetically costly, reducing the future capacity to invest in sexual signalling. However, burying beetles reproduce on dead vertebrates and the carrion meal might enable males to acquire resources that can subsequently be allocated to pheromone signalling. Alternatively, males might accumulate pheromone precursors because they do not emit their sex pheromone during brood care. To shed light on the mechanisms of enhanced pheromone emission after brood care, in the present study, we test the effect of diet quality, social condition during care (biparental versus uniparental care) and an experimentally enforced calling pause on subsequent male pheromone emission, body weight and energy storage components. The experimentally enforced calling pause and social condition during brood care demonstrate no impact on pheromone quantity. However, the results of the present study suggest that the vertebrate carrion meal during brood care partly explains enhanced pheromone release after care. Unravelling the biosynthetic pathways of the pheromone components and analyzing the impact of potential microbial symbionts on pheromone production represents a fruitful avenue for future research. [ABSTRACT FROM AUTHOR]

In animal species in which parents provide food to their dependent young, offspring often display conspicuous begging signals. These solicitation behaviors are important components of parent-offspring communication, but it is currently unclear how they and the parental response covary with offspring dependency on parental food provisioning across species. Burying beetles (Nicrophorus) are well known for providing elaborate biparental care, including provisioning of begging larvae. By using a multispecies approach, we show that larval begging intensity, as well as the time parents spend provisioning, differ greatly between individuals of the 3 species: N. orbicollis, N. pustulatus, and N. vespilloides. Our results demonstrate that the most dependent offspring of N. orbicollis invest the most time in begging, whereas the most independent offspring of N. pustulatus invest the least amount of time in begging. Thus, we suggest that begging intensity differs due to intrinsic differences in nutritional need between the species rather than because of an arbitrary divergence in begging behavior. We further show that in all 3 species, females spend significantly more time provisioning than males, although there is considerable divergence between species in the extent to which females and males contribute to the provisioning of larvae. We discuss the potential selective factors leading to this diversification of offspring begging and parental provisioning in relation to the distinct variation in offspring dependence between the 3 species. [ABSTRACT FROM AUTHOR]

Studying the relationship between parental and mating effort helps us to understand the evolution of parental care and, consequently, has been the subject of many theoretical and empirical investigations. Using burying beetles as a model, we found no correlation between the intensity of a sexual signal (sex pheromone quantity) and the amount of care provided by males. However, males that were given the opportunity to breed and care for young went on to produce a higher amount of their sexual signal and attracted three times more females in the field than control males that were not given the opportunity to breed. The likely explanation for our finding is that specific aspects of care in burying beetles, that is the defense and preservation of a nutrient rich breeding resource, a small vertebrate cadaver, is not only beneficial for the offspring but also for the adults themselves. Obtaining a good carrion meal possibly enables males to store resources that they can subsequently allocate toward sexual signaling. Collectively, our results highlight that conditions can exist where male participation in brood care has a positive effect on its sexual attractiveness. This in turn might have facilitated the evolution of male assistance in parental care. [ABSTRACT FROM AUTHOR]

Burying beetles reproduce on small vertebrate carcasses by exhibiting elaborate biparental brood care. Partner recognition in breeding Nicrophorus species ( Coleoptera: Silphidae) relies substantially on information encoded in cuticular hydrocarbon profiles. Until recently, it was unknown whether breeding burying beetles also produce volatile low molecular weight substances and, if so, which functions can be attributed to such volatiles. The present study reports a survey of the volatiles released by males and females of Nicrophorus vespilloides Herbst in nonbreeding status and at different stages of breeding. Headspace analyses are performed by using solid phase micro-extraction fibres and gas chromatography-mass spectrometry. The volatiles released by nonbreeding males and females include phenolic compounds, alcohols, aldehydes and ketones and are quite similar in both sexes. With the onset of breeding, the volatile profiles of males and females become distinct, with a number of female-specific compounds occurring. An analysis of the anal secretions reveals the presence of some of the compounds previously detected in the headspace analysis. The specific chemical properties suggest that some of the volatiles may function against competitors and parasites, such as bacteria, fungi, nematodes and arthropods at the carcass breeding resource. By contrast, the emission of 4-methyl branched esters by the females closely parallels the emission of the terpenoid methyl geranate and they may function together as a complex signal by the females. Signalling traits associated with biparental care and specific constraints associated with the ephemeral nature of the breeding resource may explain the occurrence of both groups of compounds in the volatile profiles. [ABSTRACT FROM AUTHOR]

Few studies have attempted to distinguish between cues and signals in the context of chemical communication. A number of chemical substances have been shown to vary with physiological state, such as stage of oestrus cycle, fertility, dominance status or nutritional condition, but little is known about whether this variation is incidental or adaptive. Here, we provide evidence of a substance whose emission varies with breeding state, but is not merely an incidental by-product of physiological state, but rather, an evolved signal. Breeding females of the facultative biparental burying beetle, Nicrophorus vespilloides, releasemethyl geranate, a substance that helps males to identify breeding status and to distinguish between their female partners and non-breeding intruders. We demonstrate that females respond flexibly to their social environment and emit high amounts of methyl geranate only in the presence of a male partner, i.e. a receiver. In contrast, cuticular hydrocarbons, which also have been shown to change with breeding status, are not modulated and do not differ between single and paired breeding females. Receiver-dependent chemical signalling is expected to evolve when costs are involved in the production or transmission of the signal; such signal modulation might be more common than previously thought. [ABSTRACT FROM AUTHOR]

Despite a vast literature on the factors controlling adult size, few studies have investigated how maternal size affects offspring size independent of direct genetic effects, thereby separating prenatal from postnatal influences. I used a novel experimental design that combined a cross-fostering approach with phenotypic manipulation of maternal body size that allowed me to disentangle prenatal and postnatal maternal effects. Using the burying beetle Nicrophorus vespilloides as model organism, I found that a mother's body size affected egg size as well as the quality of postnatal maternal care, with larger mothers producing larger eggs and raising larger offspring than smaller females. However, with respect to the relative importance of prenatal and postnatal maternal effects on offspring growth, only the postnatal effects were important in determining offspring body size. Thus, prenatal effects can be offset by the quality of postnatal maternal care. This finding has implications for the coevolution of prenatal and postnatal maternal effects as they arise as a consequence of maternal body size. In general, my study provides evidence that there can be transgenerational phenotypic plasticity, with maternal size determining offspring size leading to a resemblance between mothers and their offspring above and beyond any direct genetic effects. [ABSTRACT FROM AUTHOR]

The theory of prudent sperm allocation predicts that males should reduce their investment in sperm transfer when their mate's propensity to produce offspring is low. Furthermore, if mating is costly for females, they would benefit from signalling this propensity to males to keep them from attempting to mate. Here we document a remarkable flexibility in insect mating tactic during the period of parental care. Extensive video observations in the biparental burying beetle Nicrophorus vespilloides revealed that pairs mate frequently before and during oviposition, temporarily discontinue their sexual activity while young larvae are present, and resume mating when males are close to leaving the brood. An experiment, in which we switched males at different time intervals and determined their P2 value (i.e. the proportion of progeny sired by the second male), confirmed the lack of mating activity during the first few days of larval care and its resumption later in parental care. Females cease laying eggs when caring for larvae, but females will resume egg laying if none of the larvae from their first clutch reach the carcass. We manipulated females into producing such replacement clutches by withholding their larvae, and we found that in this situation males did not cease to mate, but continued to engage in frequent copulations. Hence, males have the ability to assess a female's propensity for oviposition during the period of parental care. Further experiments demonstrated that males do not use the presence or absence of larvae as a cue to adjust copulation rate, but instead use female-produced cues of reproductive state. Therefore, our study reveals that female beetles express cues that display their reproductive condition and males have evolved the ability to detect these cues and respond with prudent sperm allocation. Our findings further contribute to the developing picture of the sophisticated mating tactics that insects employ. [Copyright &y& Elsevier]