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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]

Family life in animals is often considered as beneficial for offspring but costly for parents. However, parents might also profit from remaining aggregated within a family unit, especially if a nutrient-rich resource is used for reproduction. We aimed to reveal the potential personal benefits of breeding within a family environment for male Nicrophorus vespilloides, a species of burying beetles that use small vertebrate cadavers to raise their larvae. We previously hypothesized that males obtain an advantage from remaining with their family, because they themselves can feed from the cadaver. This, in turn, enables them to produce more sex pheromone, thereby making them more attractive to females after leaving their brood. However, whether such personal benefits arise under natural conditions is currently unclear because we have no knowledge of the nutritional condition of wild beetles. If carrion is abundant anyways, feeding from a vertebrate cadaver during breeding might not have a noticeable positive effect on the males' body condition. In the current study, we caught wild males with a natural feeding history and compared their body mass and attractiveness before and after participating in family life. We show that wild males gain weight during breeding and attract more and larger females afterwards. Our study suggests that access to a highly nutrient-rich meal can be a driver of the evolution of family life and eventually biparental care. Males benefit indirectly from defending the resource and offspring against competitors and benefit personally by a higher chance of mating again after breeding. In wild male burying beetles, family living on a vertebrate cadaver leads to a gain in body mass and higher attractiveness to females. Our results highlight that scarce resources such as carcasses or dung can be considered as drivers of the evolution of family life not only because the parents remain with their brood to protect the resource, but also because they can profit personally from an improved somatic condition. [ABSTRACT FROM AUTHOR]

There is growing evidence that a wide range of insect sex pheromones are condition dependent and play a fundamental role in mate choice. However, the effectiveness of pheromonal communication might not only depend on internal factors of the sender, but also on attributes of the microhabitat, in which the signaler chooses to emit its chemical signal. For example, the degree of anthropogenic land use might affect how successful the signal is transmitted, as land use has been shown to affect animal communities and the complexity of biotic interactions. To test the hypothesis that parameters of the microenvironment determine males' ability to attract females via their sex pheromone, we used the burying beetle Nicrophorus vespilloides Herbst (Coleoptera: Silphidae) as our model system. We exposed 144 males across differently managed forest stands and analyzed the impact of 29 environmental parameters. Our data revealed that human land use intensity had no effect on a male's attractiveness. However, the harvested tree biomass positively affected the proportion of competitors attracted. Furthermore, we found that soil characteristics were important factors determining the amount and body size of females a male was able to attract. Consequently, we present evidence that the environmental context of a signaling male influences the effectiveness of chemical signaling either because it affects the transmission process or the prevailing abundance of potential signal receivers. Thus, our results demonstrate that males need to make careful decisions about the location where they emit their pheromone, as this choice of microhabitat has an impact on their fitness. [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]

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]

Intrasexual competition for mates or resources important for reproduction often leads to the establishment of dominance relationships that influence an individual’s reproductive success. Although dominance can be an honest indicator of health or immunocompetence, the attainment and maintenance of dominance status can also influence an individual’s ability to invest in immunity, making it difficult to disentangle cause and effect. Here we examine the relationship between intrasexual competition and the nutritional condition and immunity of burying beetles, Nicrophorus orbicollis, insects that reproduce on small vertebrate carcasses that serve as a larval food source. We staged intrasexual contests on carcasses in both sexes and compared the nutritional state and immunity of dominant and subordinate individuals with those of beetles reproducing on a carcass without competitors. The nutritional state and immunity of dominant beetles were not significantly different from beetles without competitors, but subordinates were characterized by a lower weight gain and reduced encapsulation response. In addition, we found a clear sex effect, with females gaining more weight than males and having superior immunity. We conclude that the subordinate’s exclusion from the carcass plays an important role in mediating the difference in encapsulation. Our data suggest that this is not entirely a nutritional effect because better-fed subordinates did not exhibit higher immune responses. Rather, subordinates may have no need to invest in improved immunity because they do not participate in carcass maintenance and defense. [ABSTRACT FROM PUBLISHER]

Intense intrasexual competition between males often results in the establishment of a dominance hierarchy, promoting differential access to females. Dominant males are expected to evolve more prudent sperm allocation strategies, whereas subordinate males are expected to counteract their mating disadvantage by increasing their sperm allocation. Burying beetles, insects that breed on small vertebrate carcasses, offer an ideal model for testing these predictions because aggressive interactions between males typically result in one dominant male that monopolizes access to the female, relegating subordinates to sneak matings. Males that have not found a carcass use pheromones to attract females and only obtain a small number of matings per day. We measured the mating frequency of males competing on a carcass and confirmed that dominant males mate significantly more often than subordinates. We then examined the paternity of competing males, with a 5:1 mating advantage for the dominant male, to determine whether relative mating frequency is sufficient to explain the severe reproductive skew in polyandrous broods in the field. When competing on a carcass, dominant males did not sire significantly more offspring than subordinates. However, when males mated without a carcass and did not interact, significantly more offspring were sired by the dominant male, but even here, the subordinate male’s paternity was greater than his share of matings. Our results show that when males compete on a carcass, dominant males have a clear mating advantage, whereas subordinates appear to transfer greater numbers of sperm per mating to compensate for their limited mating opportunities. [ABSTRACT FROM PUBLISHER]