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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]

Communication is fundamental in the animal kingdom, essential to interactions such as mating, defense, and parental care. Vibrational communication has often been overlooked in the past, but in recent decades, it has become clear that insects use substrate vibrations as a communication signal. In burying beetles of the genus Nicrophorus, which are known for their biparental brood care, both parents stridulate. Spending a considerable period of their lives underground, it is very likely the beetles utilize vibrations as part of their communication system. As playback experiments are challenging with this species, this study looked at the physical propagation of the signal of Nicrophorus vespilloides Herbst (Coleoptera: Siliphidae) through three soil types, as well as behavior, to see whether vibrational communication is possible. The aims were to determine: (1) whether the soils used in the laboratory compare to soil from the field, (2) whether the distance of propagation is enough for the range the beetles cover during brood care, (3) whether the two sexes show a difference in stridulation likelihood, (4) whether propagation of defensive signals differs from brood care signals, and (5) whether we can determine a behavior during stridulations that shows a clear and useable reaction to the signal. We manipulated beetles to induce stridulation and then used laser Doppler vibrometers to record the signals using three substrates and various distances, alongside behavioral observations. We showed that the three substrates tested, peat, coconut coir, and forest soil, displayed differences in terms of vibrational propagation, and that burying beetle stridulation signals can be transmitted up to about 25 cm in the soil. We also showed that the location where the animals stridulate exerts a significant influence on the total duration and number of stridulations. Overall, vibrational communication is in principle conceivable in this species, as the signals are transmitted far enough in the natural substrate to allow complex communication, opening possibilities for vibrational communication during this biparental brood care. [ABSTRACT FROM AUTHOR]

Investigating fundamental processes in biology requires the ability to ground broad questions in species‐specific natural history. This is particularly true in the study of behavior because an organism's experience of the environment will influence the expression of behavior and the opportunity for selection. Here, we provide a review of the natural history and behavior of burying beetles of the genus Nicrophorus to provide the groundwork for comparative work that showcases their remarkable behavioral and ecological diversity. Burying beetles have long fascinated scientists because of their well‐developed parenting behavior, exhibiting extended post‐hatching care of offspring that varies extensively within and across taxa. Despite the burgeoning success of burying beetles as a model system for the study of behavioral evolution, there has not been a review of their behavior, ecology, and evolution in over 25 years. To address this gap, we leverage a developing community of researchers who have contributed to a detailed knowledge of burying beetles to highlight the utility of Nicrophorus for investigating the causes and consequences of social and behavioral evolution. [ABSTRACT FROM AUTHOR]

Parental care strategies do not only vary greatly across species, but also within species there can be substantial between‐ and within‐individual variation in parental care behavior. To better understand the evolution of care strategies, it is crucial to determine how and when parents modify their behavior in response to internal as well as environmental factors. Here, we investigated the effect of brood size, resource size and an individual's quality on care strategies of uniparental males and examined the downstream consequences on offspring performance in the burying beetle Nicrophorus vespilloides. Burying beetles breed on small vertebrate cadavers and, on average, males invest much less in care than females. Nevertheless, we found that uniparentally caring males were responsive to their social and non‐social environment and adjusted the amount as well as the type of care to the size of the brood, the size of the cadaver and their own body size. Additionally, we show that the care strategies affected offspring performance. Specifically, males that cared longer had larger and more surviving larvae. Our results add to our understanding of plastic parenting strategies by showing that even the sex that provides less care can evolve a very flexible care behavior. [ABSTRACT FROM AUTHOR]

Harsh environmental conditions in form of low food availability for both offspring and parents alike can affect breeding behavior and success. There has been evidence that food scarce environments can induce competition between family members, and this might be intensified when parents are caring as a pair and not alone. On the other hand, it is possible that a harsh, food‐poor environment could also promote cooperative behaviors within a family, leading, for example, to a higher breeding success of pairs than of single parents. We studied the influence of a harsh nutritional environment on the fitness outcome of family living in the burying beetle Nicrophorus vespilloides. These beetles use vertebrate carcasses for reproduction. We manipulated food availability on two levels: before and during breeding. We then compared the effect of these manipulations in broods with either single females or biparentally breeding males and females. We show that pairs of beetles that experienced a food‐poor environment before breeding consumed a higher quantity of the carcass than well‐fed pairs or single females. Nevertheless, they were more successful in raising a brood with higher larval survival compared to pairs that did not experience a food shortage before breeding. We also show that food availability during breeding and social condition had independent effects on the mass of the broods raised, with lighter broods in biparental families than in uniparental ones and on smaller carcasses. Our study thus indicates that a harsh nutritional environment can increase both cooperative as well as competitive interactions between family members. Moreover, our results suggest that it can either hamper or drive the formation of a family because parents choose to restrain reproductive investment in a current brood or are encouraged to breed in a food‐poor environment, depending on former experiences and their own nutritional status. [ABSTRACT FROM AUTHOR]

Conflicts over parental investment are predicted to be common among family members, especially between parents and their offspring. Parent–offspring conflict has been studied in many brood-caring organisms, but whether its outcome is closer to the parental or offspring optimum is usually unknown, as is whether the presence of a second parent, a caring male partner, can affect the outcome. Here, we manipulated the initial brood size of single and paired female burying beetles to examine how many offspring are necessary to maintain parental care in the current brood. We found that mothers continued to invest in small broods even if their reproductive output would have been higher if they had discontinued their care and produced a second brood instead. Consequently, our data suggests that the offspring have the upper hand in the conflict. However, our results further show that paired females laid a second egg clutch more often and produced more offspring than single females, suggesting that the presence of a male partner shifts the conflict outcome towards the parental optimum. This latter result not only is a novel aspect of parent–offspring theory, but also represents an additional factor that might explain the evolution of biparental care. [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]

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]

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]

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 have fascinated scientists for centuries due to their elaborate form of biparental care that includes the burial and defense of a vertebrate carcass, as well as the subsequent feeding of the larvae. However, besides extensive research on burying beetles, one fundamental question has yet to be answered: what cues do males use to discriminate between the sexes? Here, we show in the burying beetle Nicrophorus vespilloides that cuticular lipids trigger male mating behavior. Previous chemical analyses have revealed sex differences in cuticular hydrocarbon (CHC) composition; however, in the current study, fractionated-guided bioassay showed that cuticular lipids, other than CHCs, elicit copulation. Chemical analyses of the behaviorally active fraction revealed 17 compounds, mainly aldehydes and fatty acid esters, with small quantitative but no qualitative differences between the sexes. Supplementation of males with hexadecanal, the compound contributing most to the statistical separation of the chemical profiles of males and females, did not trigger copulation attempts by males. Therefore, a possible explanation is that the whole profile of polar lipids mediates sex recognition in N. vespilloides. [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]

The necrophagous burying beetle Nicrophorus vespilloides reproduces on small carcasses that are buried underground to serve as food for their offspring. Cadavers that are too large to bury have previously been postulated to be important food sources for newly emerged beetles; however, the attractiveness of distinct successive stages of decomposition were not further specified. Therefore, we investigated the potential preference of newly emerged N. vespilloides females for odour bouquets of piglet cadavers at specific stages of decomposition. Analyses of walking tracks on a Kramer sphere revealed a significantly higher mean walking speed and, consequently, a higher mean total track length when beetles were confronted with odour plumes of the decomposition stages ‘post-bloating’, ‘advanced decay’ or ‘dry remains’ in comparison with the solvent control. Such a change of the walking speed of newly emerged N. vespilloides females indicates a higher motivation to locate such food sources. In contrast to less discriminating individuals this behaviour provides the advantage of not wasting time at unsuitable food sources. Furthermore, in the advanced decay stage, we registered a significantly higher preference of beetles for upwind directions to its specific odour plume when compared with the solvent control. Such a change to upwind walking behaviour increases the likelihood that a large cadaver will be quickly located. Our findings are of general importance for applied forensic entomology: newly emerged N. vespilloides females on large cadavers can and should be regarded as potential indicators of prolonged post mortem intervals as our results clearly show that they prefer emitted odour bouquets of later decomposition stages. [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]

The production, maintenance and transmission of chemical signals often entail costs. Costs can arise, for instance, if signal production depends on the availability of limited nutritional resources or if signal transmission leads to attraction of predators. Many species effectively reduce these costs by signaling at specific times or in certain contexts only. We previously reported that breeding burying beetle females facultatively adjust their pheromone emission in response to their social environment, emitting high amounts of their chemical signal in the presence of a male partner, but not when providing uniparental care. Here we present data showing that chemical signaling is costly, and that higher investments in signaling result in reduced clutch sizes, but not a shorter life span, in the burying beetle Nicrophorus vespilloides. [ABSTRACT FROM AUTHOR]

Summary 1. Immunity may trade-off against other important life history traits, with recent work suggesting that reproduction and parental care in particular impinge on immune defence. However, whereas the effect of parental care on immunocompetence has been intensively studied in birds and mammals, virtually nothing is known about how it affects insect immunity. 2. Burying beetles provide extensive biparental care that includes the burial, preparation and defence of a carcass, as well as the subsequent feeding of the larvae. In addition, they cover the carcass with anal exudates that have been shown to serve an antimicrobial function (social immunity sensu Behavioral Ecology, 21, 663-668). We examined the effect of sex, mating and parental care on measurements of individual and social immunity in the burying beetle Nicrophorus orbicollis. 3. Both males and females showed a rapid upregulation of the encapsulation response upon discovery of a carcass. The high encapsulation rate was maintained during the entire period of parental care. Lytic activity in anal exudates, a measure of social immunity, likewise increased. Mating had no effect on individual or social immunity, but females generally exhibited higher individual immunity than male N. orbicollis. 4. Our results suggest that the unusual breeding environment of burying beetles - a microbe-rich carcass - has selected for an atypical pattern of immune defence, with a significant upregulation of individual and social immunity during the physically demanding period of reproduction and parental care. The simultaneous investment in two life history traits that normally compete for resources may be an adaptive response in species that breed in environments with high densities of micro-organisms. [ABSTRACT FROM AUTHOR]

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]

Burying beetles exhibit elaborate biparental care, in which a male and a female bury a carcass upon which they rear their brood and defend it against conspecific competitors. If an intruder succeeds in taking over the carcass, it kills the entire brood and rears its own young on the remaining resource. However, the resident male and female are able to discriminate between an intruder and their breeding partner, and assist their mate in driving off intruders of either sex. Efficient brood guarding requires effective recognition and discrimination mechanisms. Theory predicts that the setting of the optimal acceptance threshold should be affected by the rates of interaction with desirable and undesirable individuals, and shift towards a less permissive threshold when the probability of encountering intruders increases. Here we provide a critical test of acceptance-threshold theory by investigating discrimination rules during reproduction in the burying beetle Nicrophorus vespilloides. The beetles typically discriminate between their breeding partners and conspecific intruders based on a class-specific discrimination rule: they generally accept beetles that are also caring for larvae (whether their own partner or not), but are aggressive towards all nonbreeding beetles. However, we demonstrate that this recognition system is flexible at the level of individual experience and changes with context. After the introduction of an intruder, rejection rate increased, which was manifested as a shift from class-specific to individual-specific discrimination: males suddenly began discriminating between their own breeding partner and novel breeding females, and were more aggressive towards the latter. We show that the plasticity in the recognition system is not caused by template updating, but instead by a flexible acceptance threshold. Our experiments suggest that breeding males learn the individual features of their own breeding partner, but set their acceptance threshold depending on the risk of encountering an intruder. [Copyright &y& Elsevier]

In recent years, studies have shown that animals can communicate their physiological state or condition by means of chemical signals. However, as the chemicals involved in the condition-dependent signals have rarely been identified, evolutionary mechanisms that ensure their reliability are not well understood. We identified a volatile chemical signal that may serve as a reliable indicator of hormonal state in burying beetles, and is involved in their partner recognition system. Burying beetles reproducing on carcasses are able to distinguish between their breeding partners and infanticidal conspecifics. This discrimination depends on breeding status, which is positively linked to juvenile hormone III (JH III) titre. Breeding Nicrophorus vespilloides beetles, in contrast to nonbreeding ones, emitted considerable amounts of (E)-methylgeranate. The amount of emitted (E)-methylgeranate was positively correlated with juvenile hormone titres known from other burying beetle studies. Moreover, our behavioural experiments showed that dummies treated with methylgeranate induced tolerant behaviour, whereas control dummies were treated aggressively. The fact that (E)-methylgeranate and JH III share a conspicuous structural similarity and the same biosynthetic pathway may explain how the reliability of the signal for JH titre is ensured. We discuss the implications of our results in the light of theoretical work on the evolution of chemical communication, particularly on the origin of chemical signals. [Copyright &y& Elsevier]

The multicomponent nature of chemical cues and signals are not very well understood. One reason for the often found complexity of chemical blends might be that they provide multiple messages. Burying beetles which use vertebrate carcasses as food for their larvae and defend these carcasses against inter- and intraspecific competitors are able to recognise the sex and breeding status of conspecifics. Studies have shown that the chemical composition of cuticular lipids is correlated with sex and breeding status, but there is no definitive evidence that these chemicals function in recognition. In the present study, we performed behavioural bioassays to directly asses the role of chemical cues in the recognition system of the burying beetle Nicrophorus vespilloides. After finding a carcass, females were more tolerant of dead males than of females. The behaviour was reversed when a solvent extract from the opposite sex was applied. An earlier experiment had shown that females breeding on a carcass treat non-breeding males more aggressively than breeding ones. In the present study, we could trigger the same dichotomous behaviours by presenting a single elytron from a breeding and a non-breeding beetle. In an additional experiment, females tolerated the elytra of non-breeding beetles if we had first applied an extract from a breeding beetle to these elytra. Our study is the first behavioural proof that female burying beetles obtain multiple information from chemical cues. [ABSTRACT FROM AUTHOR]

Asynchronous hatching is an important component of animal reproductive strategies, yet it has been studied almost exclusively in altricial birds. In this study, we provide evidence on the adaptive consequences and the heritable basis of asynchronous hatching in an insect, the burying beetle Nicrophorus vespilloides. Parents of this species breed on carcasses of small vertebrates and provide food in the form of predigested carrion for their offspring. We found that the size of the carcass used for breeding had a significant effect on hatching skew towards the earlier part of the hatching period, suggesting that female parents adjust hatching skew facultatively to the amount of resources available for breeding. Using a full sibling breeding design, we also found that parent family had a significant effect on both hatching skew and hatching spread, suggesting that there is a heritable basis to asynchronous hatching. Finally, we found that hatching spread affected offspring survivorship, providing evidence that asynchronous hatching patterns have adaptive consequences in N. vespilloides. Our study provides valuable new insights into the evolution and ecological significance of asynchronous hatching by providing evidence on the adaptive consequences and the heritable basis of asynchronous hatching in a non-avian species. [ABSTRACT FROM AUTHOR]

Abstract: In most animal species, brood size and body size exhibit some variation within and between populations. This is also true for burying beetles (genus Nicrophorus), a group in which the body size of offspring depends critically on the number of offspring competing for food due to the discrete nature of resource used for larval nutrition (vertebrate carcasses). In one species, brood size and body size are correlated with population density, and appear to be phenotypically plastic. We investigated potential proximate causes of between-population variation in brood size and body size in two species, Nicrophorus vespilloides and Nicrophorus defodiens. Our first experiment supported the notion that brood size is phenotypically plastic, because it was affected by environmental variation in adult nutritional condition. We found that the pre-breeding nutritional status of female N. vespilloides affected the number of eggs they laid, the number of surviving larvae in their broods, and the body size of their offspring. We do not know whether this plasticity is adaptive because greater offspring body size confers an advantage in contests over breeding resources, or whether starved females are constrained to produce smaller clutches because they cannot fully compensate for their poor pre-breeding nutritional status by feeding from the carcass. Our second experiment documents that brood size, specifically the infanticidal brood-size adjustment behavior, has undergone genetic differentiation between two populations of N. defodiens. Even under identical breeding conditions with identical numbers of first-instar larvae, females descended from the two populations produced broods of different size with corresponding differences in offspring body size. [Copyright &y& Elsevier]

Communication plays a fundamental role in the evolution of any form of cooperative behaviour, such as parental care. However, it can be challenging to understand the specific role of certain signals and how they might have evolved into complex communication systems. To investigate what effect a lack of acoustic communication can have on brood care and offspring performance, we silenced parents of biparentally caring burying beetles with a noninvasive method and studied the effect on clutch and offspring performance. Moreover, by analysing three species with varying degrees of offspring dependency on parental care, we aimed to investigate how differing acoustic communication is related to the level of their dependency and if those two align in some way. We used Nicrophorus pustulatus , a nondependent species, Nicrophorus orbicollis , a highly dependent species and Nicrophorus vespilloides , an intermediately dependent species. We found strong effects of silencing parents on offspring performance in all three species. The lack of stridulations impacted offspring weight across all three species. However, our results point towards a difference between species in which development stage communication had the most substantial impact. Looking at larval weight at dispersal, the effects seem to be in line with the larval dependency in the way we would have expected, with N. orbicollis being the most strongly affected, N. vespilloides being also affected and N. pustulatus not being affected. However, looking more closely, we found various differences at other time points and also that larval survival was strongly affected in N. pustulatus. Few studies have looked at the exact function of acoustic signals during brood care with most of them focusing on what type of different signals are emitted rather than what effect they have. Our study is one of the first to start disentangling the interplay of communication and offspring performance. • Silencing parents has a strong effect on offspring performance in burying beetles. • Different species of Nicrophorus differ in communication during brood care. • The larvae of silenced parents are at a disadvantage. • More dependent species do not, per se, rely more on communication. [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]

Long-range sex pheromones have been subjected to substantial research with a particular focus on their biosynthesis, peripheral perception, central processing and the resulting orientation behaviour of perceivers. Fundamental to the research on sex attractants was the assumption that they primarily coordinate species recognition. However, especially when they are produced by the less limiting sex (usually males), the evolution of heightened condition dependence might be expected and long-range sex pheromones might, therefore, also inform about a signaller's quality. Here we provide, to our knowledge, the first comprehensive study of the role of a male's long-range pheromone in mate choice that combines chemical analyses, video observations and field experiments with a multifactorial manipulation of males' condition. We show that the emission of the long-distance sex pheromone of the burying beetle, Nicrophorus vespilloides is highly condition-dependent and reliably reflects nutritional state, age, body size and parasite load -- key components of an individual's somatic state. Both, the quantity and ratio of the pheromone components were affected but the time invested in pheromone emission was largely unaffected by a male's condition. Moreover, the variation in pheromone emission caused by the variation in condition had a strong effect on the attractiveness of males in the field, with males in better nutritional condition, of older age, larger body size and bearing less parasites being more attractive. That a single pheromone is influenced by so many aspects of the somatic state and causes such variation in a male's attractiveness under field conditions was hitherto unknown and highlights the need to integrate indicator models of sexual selection into pheromone research. [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]