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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]

To link to full-text access for this article, visit this link: http://dx.doi.org/10.1016/j.anbehav.2010.06.018 Byline: Sandra Steiger (a)(b), Josef K. Muller (b) Abstract: 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. Author Affiliation: (a) Behavior, Ecology, Evolution & Systematics Section, School of Biological Sciences, Illinois State University, U.S.A. (b) Faculty of Biology, Department of Evolutionary Biology and Ecology, University of Freiburg, Germany Article History: Received 25 March 2010; Revised 20 April 2010; Accepted 19 May 2010 Article Note: (miscellaneous) MS. number: A10-00208

To link to full-text access for this article, visit this link: http://dx.doi.org/10.1016/j.anbehav.2009.09.019 Byline: Wolf Haberer (a), Sandra Steiger (a)(b), Josef K. Muller (a) Abstract: 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. Author Affiliation: (a) Faculty of Biology, Department of Evolutionary Biology and Ecology, University of Freiburg, Germany (b) Behavior, Ecology, Evolution and Systematics Section, School of Biological Sciences, Illinois State University, USA Article History: Received 25 June 2009; Revised 4 August 2009; Accepted 9 September 2009 Article Note: (miscellaneous) MS. number: 09-00428

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]

When burying beetles first emerge as adults, they search for well-rotted carcasses with fly maggots on which to feed. After attaining reproductive competence, they switch their search and respond to a small, fresh carcass to prepare for their brood. Because the cues used to locate a feeding versus a breeding resource both originate from carrion, the beetles must respond to subtle changes in volatiles during decomposition. We investigated cues used to locate a fresh carcass in the field by (1) a general subtractive method, applying an antibacterial or antifungal compound to reduce microbially derived volatiles, and (2) a specific additive method, placing chemical supplements near a fresh carcass. Five sulfur-containing compounds, known to result from bacterial metabolism of sulfur-containing amino acids, were studied: dimethyl sulfide (DMS), dimethyl disulfide (DMDS), dimethyl trisulfide (DMTS), methyl thiolacetate (MeSAc, also known as S-methyl thioacetate), and methyl thiocyanate (MeSCN). When a carcass aged for 48 h was treated with an antibacterial compound to reduce volatiles, there was a 59% decrease in beetles discovering the resource. The addition of the chemical supplement MeSAc had no effect on discovery of a fresh carcass, while DMS and DMDS had a limited ability to attract breeding beetles. The chemical that was least well known, MeSCN, increased beetle numbers by 200–800% on a fresh carcass and almost guaranteed discovery. DMTS, which is known to attract a variety of carrion insects, was the only compound to significantly reduce beetle presence at a fresh carcass. A laboratory experiment demonstrated that DMTS does not directly inhibit breeding, suggesting that DMTS deters breeding beetles while they fly. [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]

The decomposition of dead mammalian tissue involves a complex temporal succession of epinecrotic bacteria. Microbial activity may release different cadaveric volatile organic compounds which in turn attract other key players of carcass decomposition such as scavenger insects. To elucidate the dynamics and potential functions of epinecrotic bacteria on carcasses, we monitored bacterial communities developing on still-born piglets incubated in different forest ecosystems by combining high-throughput Illumina 16S rRNA sequencing with gas chromatography-mass spectrometry of volatiles. Our results show that the community structure of epinecrotic bacteria and the types of cadaveric volatile compounds released over the time course of decomposition are driven by deterministic rather than stochastic processes. Individual cadaveric volatile organic compounds were correlated with specific taxa during the first stages of decomposition which are dominated by bacteria. Through best-fitting multiple linear regression models, the synthesis of acetic acid, indole and phenol could be linked to the activity of Enterobacteriaceae, Tissierellaceae and Xanthomonadaceae, respectively. These conclusions are also commensurate with the metabolism described for the dominant taxa identified for these families. The predictable nature of in situ synthesis of cadaveric volatile organic compounds by epinecrotic bacteria provides a new basis for future chemical ecology and forensic studies. [ABSTRACT FROM AUTHOR]

Chemical communication is the most ancient and widespread form of communication. Yet we are only beginning to grasp the complexity of chemical signals and the role they play in sexual selection. Focusing on insects, we review here the recent progress in the field of olfactory-based sexual selection. We will show that there is mounting empirical evidence that sexual selection affects the evolution of chemical traits, but form and strength of selection differ between species. Studies indicate that some chemical signals are expressed in relation to an individual's condition and depend, for example, on age, immunocompetence, fertility, body size or degree of inbreeding. Males or females might benefit by choosing based on those traits, gaining resources or "good genes". Other chemical traits appear to reliably reflect an individual's underlying genotype and are suitable to choose a mating partner that matches best the own genotype. [ABSTRACT FROM AUTHOR]