Bolitotherus Cornutus Research Articles

The influence of resource patch condition and size on social interactions, reproductive behaviors, and oviposition in a fungivorous beetle

Variation among resource patches can structure spatial patterns of social behavior. Individual preferences and requirements for heterogeneous resources can lead to differences in where behaviors take place and which kinds of individuals interact. In this study we examined whether properties of habitat patches predict presence of and interactions among wild forked fungus beetles (Bolitotherus cornutus). B. cornutus use shelf-like fruiting bodies of wood decaying fungi (brackets) as social arenas, courtship and mating sites, oviposition locations, and food sources. Brackets on a single log vary in size and condition and represent a dynamic resource as individual brackets decay over the years that they persist on a subpopulation. We found that size and condition significantly predicted how a bracket was used by B. cornutus. Both males and females were found more often on larger, living brackets. Larger brackets hosted proportionally more social interactions and mating behaviors relative to the overall visitation by individuals. The relationship between these resource properties and frequencies of behaviors was stronger for reproductive, between-sex social interactions. Live brackets also attracted more oviposition events by females, but bracket size had no effect on this behavior. Our results indicate that the dynamics of growth and aging of a critical resource can impact how social and reproductive behaviors are distributed in time and space, which in turn could affect the social structure of subpopulations.Significance statementSpecies that depend on critical host resources are often faced with a patchy landscape of resource quality. Those patches represent the infrastructure within which social behaviors take place, and can have formative effects on where, when, and how social interactions occur. We demonstrated that the size and condition of a fungal resource predicted male and female forked fungus beetle presence and behavior. Larger, living brackets hosted more beetles overall, while larger brackets in any condition hosted more social interactions and mating behaviors. Female beetles were more likely to lay eggs on living brackets, regardless of size. This study demonstrates how ecological dynamics of aging resource patches can indirectly structure social landscapes by influencing where and how individuals interact.

Multilevel selection on social network traits differs between sexes in experimental populations of forked fungus beetles

Both individual and group behavior can influence individual fitness, but multilevel selection is rarely quantified on social behaviors. Social networks provide a unique opportunity to study multilevel selection on social behaviors, as they describe complex social traits and patterns of interaction at both the individual and group levels. In this study, we used contextual analysis to measure the consequences of both individual network position and group network structure on individual fitness in experimental populations of forked fungus beetles (Bolitotherus cornutus) with two different resource distributions. We found that males with high individual connectivity (strength) and centrality (betweenness) had higher mating success. However, group network structure did not influence their mating success. Conversely, we found that individual network position had no effect on female reproductive success but that females in populations with many social interactions experienced lower reproductive success. The strength of individual-level selection in males and group-level selection in females intensified when resources were clumped together, showing that habitat structure influences multilevel selection. Individual and emergent group social behavior both influence variation in components of individual fitness, but impact the male mating success and female reproductive success differently, setting up intersexual conflicts over patterns of social interactions at multiple levels.

Mycophagous beetle females do not behave competitively during intrasexual interactions in presence of a fungal resource.

Intrasexual interactions can determine which individuals within a population have access to limited resources. Despite their potential importance on fitness generally and mating success especially, female–female interactions are not often measured in the same species where male–male interactions are well‐defined. In this study, we characterized female–female interactions in Bolitotherus cornutus, a mycophagous beetle species native to Northeastern North America. We used dyadic, behavioral assays to determine whether females perform directly aggressive or indirectly exclusionary competitive behaviors. Polypore shelf fungus, an important food and egg‐laying resource for B. cornutus females, is patchily distributed and of variable quality, so we tested for competition over fungus as a resource. Behavior of females was assessed in three sets of dyadic trials with randomly paired female partners. Overall, females did not behave aggressively toward their female partner or perform exclusionary behaviors over the fungal resource. None of the behaviors performed by females were individually repeatable. Two scenarios may explain our lack of observed competition: our trial context may not induce competition, or female B. cornutus simply may not behave competitively in the wild. We compare our results to a similar study on male–male interactions in the same species and propose future studies on female–female interactions under different competitive contexts to expand the understanding of female competition.

Group and individual social network metrics are robust to changes in resource distribution in experimental populations of forked fungus beetles.

Social interactions drive many important ecological and evolutionary processes. It is therefore essential to understand the intrinsic and extrinsic factors that underlie social patterns. A central tenet of the field of behavioural ecology is the expectation that the distribution of resources shapes patterns of social interactions.We combined experimental manipulations with social network analyses to ask how patterns of resource distribution influence complex social interactions.We experimentally manipulated the distribution of an essential food and reproductive resource in semi‐natural populations of forked fungus beetles Bolitotherus cornutus. We aggregated resources into discrete clumps in half of the populations and evenly dispersed resources in the other half. We then observed social interactions between individually marked beetles. Half‐way through the experiment, we reversed the resource distribution in each population, allowing us to control any demographic or behavioural differences between our experimental populations. At the end of the experiment, we compared individual and group social network characteristics between the two resource distribution treatments.We found a statistically significant but quantitatively small effect of resource distribution on individual social network position and detected no effect on group social network structure. Individual connectivity (individual strength) and individual cliquishness (local clustering coefficient) increased in environments with clumped resources, but this difference explained very little of the variance in individual social network position. Individual centrality (individual betweenness) and measures of overall social structure (network density, average shortest path length and global clustering coefficient) did not differ between environments with dramatically different distributions of resources.Our results illustrate that the resource environment, despite being fundamental to our understanding of social systems, does not always play a central role in shaping social interactions. Instead, our results suggest that sex differences and temporally fluctuating environmental conditions may be more important in determining patterns of social interactions.

Group composition of individual personalities alters social network structure in experimental populations of forked fungus beetles.

Social network structure is a critical group character that mediates the flow of information, pathogens and resources among individuals in a population, yet little is known about what shapes social structures. In this study, we experimentally tested whether social network structure depends on the personalities of individual group members. Replicate groups of forked fungus beetles (Bolitotherus cornutus) were engineered to include only members previously assessed as either more social or less social. We found that individuals expressed consistent personalities across social contexts, exhibiting repeatable numbers of interactions and numbers of partners. Groups composed of more social individuals formed networks with higher interaction rates, higher tie density, higher global clustering and shorter average shortest paths than those composed of less social individuals. We highlight group composition of personalities as a source of variance in group traits and a potential mechanism by which networks could evolve.

Phenotypic Assortment Changes the Landscape of Selection.

Social interactions with conspecifics can dramatically affect an individual's fitness. The positive or negative consequences of interacting with social partners typically depend on the value of traits that they express. These pathways of social selection connect the traits and genes expressed in some individuals to the fitness realized by others, thereby altering the total phenotypic selection on and evolutionary response of traits across the multivariate phenotype. The downstream effects of social selection are mediated by the patterns of phenotypic assortment between focal individuals and their social partners (the interactant covariance, Cij', or the multivariate form, CI). Depending on the sign and magnitude of the interactant covariance, the direction of social selection can be reinforced, reversed, or erased. We report estimates of Cij' from a variety of studies of forked fungus beetles to address the largely unexplored questions of consistency and plasticity of phenotypic assortment in natural populations. We found that phenotypic assortment of male beetles based on body size or horn length was highly variable among subpopulations, but that those differences also were broadly consistent from year to year. At the same time, the strength and direction of Cij' changed quickly in response to experimental changes in resource distribution and social properties of populations. Generally, interactant covariances were more negative in contexts in which the number of social interactions was greater in both field and experimental situations. These results suggest that patterns of phenotypic assortment could be important contributors to variability in multilevel selection through their mediation of social selection gradients.

Male competition reverses female preference for male chemical cues.

Females must choose among potential mates with different phenotypes in a variety of social contexts. Many male traits are inherent and unchanging, but others are labile to social context. Competition, for example, can cause physiological changes that reflect recent wins and losses that fluctuate throughout time. We may expect females to respond differently to males depending on the outcome of their most recent fight. In Bolitotherus cornutus (forked fungus beetles), males compete for access to females, but copulation requires female cooperation. In this study, we use behavioral trials to determine whether females use chemical cues to differentiate between males and whether the outcome of recent male competition alters female preference. We measured female association time with chemical cues of two size‐matched males both before and after male–male competition. Females in our study preferred to associate with future losers before males interacted, but changed their preference for realized winners following male competitive interactions. Our study provides the first evidence of change in female preference based solely on the outcome of male–male competition.

Social environment: Trait, context and agent for selection in a meta‐population

In Focus: Formica, V., Donald, H., Marti, H., Irgebay, Z., Brodie III, E. Social network position experiences more variable selection than weaponry in wild subpopulations of forked fungus beetles. Journal of Animal Ecology, 90, 168-182, https://doi.org/10.1111/1365-2656.13322. That social network traits can exhibit consistent-individual differences among individuals and confer a fitness benefit or cost is increasingly well-established. However, how selection-natural or sexual-affects those social traits and at what scale remains an open question. In this Special Feature, Formica and colleagues employ a meta-population of forked fungus beetles to test and contrast whether sexual selection on social network traits contrasted to morphological traits occurs at the local (soft) or global (hard) scales. The authors demonstrate that morphological traits are largely under hard directional positive selection, whereas social traits are under soft and variable selection. The findings are compelling and raise interesting discussion of multi-level selection and the evolution of social traits in a meta-population.

Social network position experiences more variable selection than weaponry in wild subpopulations of forked fungus beetles.

The phenotypic expression and fitness consequences of behaviours that are exhibited during social interactions are especially sensitive to their local social context. This context-dependence is expected to generate more variation in the sign and magnitude of selection on social behaviour than that experienced by static characters like morphology. Relatively few studies, however, have examined selection on behavioural traits in multiple populations. We estimated sexual selection in the wild to determine if the strength and form of selection on social phenotypes is more variable than that on morphology. We compared selection gradients on social network position, body size, and weaponry of male forked fungus beetles Bolitotherus cornutus as they influenced mating success across nine natural subpopulations. Male horn length consistently experienced positive sexual selection. However, the sign and magnitude of selection on individual measures of network centrality (strength and betweenness) differed significantly among subpopulations. Moreover, selection on social behaviours occurred at a local scale ('soft selection'), whereas selection on horn length occurred at the metapopulation scale ('hard selection'). These results indicate that an individual with a given social phenotype could experience different fitness consequences depending on the network it occupies. While individuals seem to be unable to escape the fitness effects of their morphology, they may have the potential to mediate the pressures of selection on behavioural phenotypes by moving among subpopulations or altering social connections within a network.

Contest interactions and outcomes: relative body size and aggression independently predict contest status

In species with pronounced male armaments, body and weapon size often determine success in agonistic interactions. However, the behavioural components of interactions also play a significant role in determining outcomes and individuals that win agonistic interactions may not be the ones that start contests. In this study, we used dyadic, intrasexual assays to characterize agonistic behaviours and determine whether body size and/or the propensity to start interactions influences contest outcome. We characterized agonistic interactions from start to end in the sexually dimorphic forked fungus beetle, Bolitotherus cornutus. Males of the species are known to use their thoracic horns in intrasexual combat, but other aspects of male–male behaviour are undescribed. We created an ethogram that described transitions between behaviours and categorized actions as aggressive, nonaggressive and mounting. Individual aggressive and nonaggressive behaviours were highly repeatable between trials while mounting behaviours were not repeatable. The initiation of nonaggressive and mounting behaviours was not predictive of contest outcome. Relative body size and absolute aggression independently predicted contest outcome. Our results indicate that traits important for establishing contest outcome are not always correlated. Considering either aggression or body size alone may be misleading when determining competitor abilities.

Constraints Imposed by a Natural Landscape Override Offspring Fitness Effects to Shape Oviposition Decisions in Wild Forked Fungus Beetles.

Oviposition site decisions often maximize offspring fitness, but costs constraining choice can cause females to oviposit in poor developmental environments. It is unclear whether these constraints cumulatively outweigh offspring fitness to determine oviposition decisions in wild populations. Understanding how constraints shape oviposition in natural landscapes is a critical step toward revealing how maternal behavior influences fundamental phenomena like the evolution of specialization and the use of sink environments. Here, we used a genetic capture-recapture technique to reconstruct the oviposition decisions of individual females in a natural metapopulation of a beetle (Bolitotherus cornutus) that oviposits on three fungus species. We measured larval fitness-related traits (mass, development time, survival) on each fungus and compared the oviposition preferences of females in laboratory versus field tests. Larval fitness differed substantially among fungi, and females preferred a high-quality (high larval fitness) fungus in laboratory trials. However, females frequently laid eggs on the lowest-quality fungus in the wild. They preferred high-quality fungi when moving between oviposition sites, but this preference disappeared as the distance between sites increased and was inconsistent between study plots. Our results suggest that constraints on oviposition preferences in natural landscapes are sufficiently large to drive oviposition in poor developmental environments even when offspring fitness consequences are severe.

Forked Fungus Beetle (suggested common name) Bolitotherus cornutus (Panzer) 1794 (Insecta: Coleoptera: Tenebrionidae)

The forked fungus beetle, Bolitotherus cornutus (Panzer), is a cryptic horned beetle in the family Tenebrionidae that is found throughout much of eastern North America. The most distinctive feature of the forked fungus beetle is a pair of forward-facing horns that emerge from the thorax of the adult male (Figure 1). Males use these horns in competitions with rivals for access to reproductive opportunities with females. These beetles spend most of their lives on or within shelf fungi that grow on decaying logs. The fascinating life history of the forked fungus beetle makes this insect an excellent model for studying behavior, population dynamics, and sexual selection.

Case 3723 — Bolitophagus cornutus Fabricius, 1801 (currently Bolitotherus cornutus) (Insecta: Coleoptera: Tenebrionidae): proposed conservation of the specific name

Abstract. The purpose of this application, under Article 23.9.3 of the Code, is to conserve the specific name Bolitophagus cornutus Fabricius, 1801 (currently Bolitotherus cornutus) for a common and well-known North American darkling beetle by giving it precedence over its senior synonym Opatrum bifurcum Fabricius, 1798.

Consistent patterns of male mate preference in the laboratory and field

Mate preference by males is a widely observed phenomenon; however, it is not well understood how these preferences are expressed in wild populations. Most studies that describe the expression of male mate preference in the wild tend to observe assortative mating, where higher quality males express a strong preference for high quality females while low quality males express either no preference or a preference for low quality females. We examined male mate preference in Bolitotherus cornutus, the forked fungus beetle, a system that exhibits male–male combat and protracted male courtship. We conducted no-choice trials in the laboratory and monitored male courtship in a wild population. In the laboratory, we found a strong preference for larger, more fecund females across all sizes of males. Results from the field were similar to those in the lab; larger females were courted more often, and there was no relationship between the size of a male and the mean size of females he courted (i.e., no assortative mating). This study documents an unusual pattern of expression of male mate preference in the wild and suggests that mate preference by males may be underappreciated in species with intense male–male competition and non-assortative mating. While males of many species express a preference for high-quality females, their preferences can change in complex social situations such as those in wild populations, often resulting in pairing for quality. However, recent mathematical theory suggests that male preference could evolve in a variety of social contexts. We studied the mate preferences of male forked fungus beetles in the laboratory and a wild population. In the lab, males of all sizes preferred larger females. In the complex social environment of a wild population, males still preferred larger females even though there was a strong male-male competition. There was no pairing for quality as often observed in other species. These results suggest that male mate preferences may evolve in a variety of systems not just those with flexible preferences and pairing for quality.

No Effect of Host Species on Phenoloxidase Activity in a Mycophagous Beetle.

Ecological immunology is an interdisciplinary field that helps elucidate interactions between the environment and immune response. The host species individuals experience have profound effects on immune response in many species of insects. However, this conclusion comes from studies of herbivorous insects even though species of mycophagous insects also inhabit many different host species. The goal of this study was to determine if fungal host species as well as individual, sex, body size, and host patch predict one aspect of immune function, phenoloxidase activity (PO). We sampled a metapopulation of Bolitotherus cornutus, a mycophagous beetle in southwestern Virginia. B. cornutus live on three species of fungus that differ in nutritional quality, social environment, and density. A filter paper phenoloxidase assay was used to quantify phenoloxidase activity. Overall, PO activity was significantly repeatable among individuals (0.57) in adult B. cornutus. While there was significant variance among individuals in PO activity, there were surprisingly no significant differences in PO activity among subpopulations, beetles living on different host species, or between the sexes; there was also no effect of body size. Our results suggest that other factors such as age, genotype, disease prevalence, or natal environment may be generating variance among individuals in PO activity.

The effect of ecological context and relatedness on larval cannibalism in a fungus-associated beetle

The fitness consequences of extreme competitive interactions such as cannibalism are often governed by the environment because the costs and benefits of such behaviors depend on the ecological and social context of the participants. However, most studies of context-dependent cannibalism are conducted under extreme circumstances or examine only a single environmental context, conditions that are unlikely to exist in natural populations. In this study, we tested the effect of multiple environmental contexts on the frequency of cannibalism in forked fungus beetle larvae (Bolitotherus cornutus), which develop in 1 of 3 different fungus species. We paired larvae in laboratory trials and measured the effect of 1) ecological context (the 3 fungi) and 2) the relatedness of the paired larvae on the frequency of cannibalism. We found a strong effect of ecological context on cannibalism: larvae in 1 fungus cannibalized nearly twice as often as larvae in the other two. We did not detect an effect of relatedness on cannibalism in the 1 species of fungus in which trials were conducted. Cannibalism conferred benefits in the form of accelerated growth rates in all measured traits relative to noncannibals. However, contrary to most studies, cannibalism was most common in the highest quality fungus, contradicting the hypothesis that cannibalism occurs in poor environments to compensate for resource deficiencies. We discuss alternative mechanisms that may drive the ecological context dependence of cannibalism in B. cornutus and emphasize the importance of studying context-dependent behavior in naturally occurring environments.

Surprisingly little population genetic structure in a fungus‐associated beetle despite its exploitation of multiple hosts

In heterogeneous environments, landscape features directly affect the structure of genetic variation among populations by functioning as barriers to gene flow. Resource-associated population genetic structure, in which populations that use different resources (e.g., host plants) are genetically distinct, is a well-studied example of how environmental heterogeneity structures populations. However, the pattern that emerges in a given landscape should depend on its particular combination of resources. If resources constitute barriers to gene flow, population differentiation should be lowest in homogeneous landscapes, and highest where resources exist in equal proportions. In this study, we tested whether host community diversity affects population genetic structure in a beetle (Bolitotherus cornutus) that exploits three sympatric host fungi. We collected B. cornutus from plots containing the three host fungi in different proportions and quantified population genetic structure in each plot using a panel of microsatellite loci. We found no relationship between host community diversity and population differentiation in this species; however, we also found no evidence of resource-associated differentiation, suggesting that host fungi are not substantial barriers to gene flow. Moreover, we detected no genetic differentiation among B. cornutus populations separated by several kilometers, even though a previous study demonstrated moderate genetic structure on the scale of a few hundred meters. Although we found no effect of community diversity on population genetic structure in this study, the role of host communities in the structuring of genetic variation in heterogeneous landscapes should be further explored in a species that exhibits resource-associated population genetic structure.

Morphological Correlates of a Combat Performance Trait in the Forked Fungus Beetle, Bolitotherus cornutus

Combat traits are thought to have arisen due to intense male-male competition for access to females. While large and elaborate weapons used in attacking other males have often been the focus of sexual selection studies, defensive traits (both morphological and performance) have received less attention. However, if defensive traits help males restrict access to females, their role in the process of sexual selection could also be important. Here we examine the morphological correlates of grip strength, a defensive combat trait involved in mate guarding, in the tenebrionid beetle Bolitotherus cornutus. We found that grip strength was repeatable and differed between the sexes. However, these differences in performance were largely explained by body size and a non-additive interaction between size and leg length that differed between males and females. Our results suggest that leg size and body size interact as part of an integrated suite of defensive combat traits.

Nondestructive sampling of insect DNA from defensive secretion

Nondestructive techniques to obtain DNA from organisms can further genetic analyses such as estimating genetic diversity, dispersal and lifetime fitness, without permanently removing individuals from the population or removing body parts. Possible DNA sources for insects include frass, exuviae, and wing and leg clippings. However, these are not feasible approaches for organisms that cannot be removed from their natural environment for long periods or when adverse effects of tissue removal must be avoided. This study evaluated the impacts and efficacy of extracting haemolymph from a defensive secretion to obtain DNA for amplification of microsatellites using a nondestructive technique. A secretion containing haemolymph was obtained from Bolitotherus cornutus (the forked fungus beetle) by perturbation of the defensive gland with a capillary tube. A laboratory experiment demonstrated that the sampling methodology had no impact on mortality, reproductive success or gland expression. To evaluate the quality of DNA obtained in natural samples, haemolymph was collected from 187 individuals in the field and successfully genotyped at nine microsatellite loci for 95.7% of samples. These results indicate that haemolymph-rich defensive secretions contain DNA and can be sampled without negative impacts on the health or fitness of individual insects.

Fitness consequences of social network position in a wild population of forked fungus beetles (Bolitotherus cornutus)

Social networks describe the pattern of intraspecific interactions within a population. An individual's position in a social network often is expected to influence its fitness, but only a few studies have examined this relationship in natural populations. We investigated the fitness consequences of network position in a wild beetle population. Copulation success of male beetles positively covaried with strength (a measure of network centrality) and negatively covaried with clustering coefficient (CC) (a measure of cliquishness). Further analysis using mediation path models suggested that the activity level of individuals drove the relationships between strength and fitness almost entirely. In contrast, selection on CC was not explained by individual behaviours. Although our data suggest that social network position can experience strong sexual selection, it is also clear that the relationships between fitness and some network metrics merely reflect variation in individual-level behaviours.