Dung Beetle Onthophagus Taurus Research Articles

Experimental evidence that dung beetles benefit from reduced ivermectin in targeted treatment of livestock parasites

Anthelmintic residues in livestock dung can adversely affect beneficial organisms. Targeted selective treatment (TST) of a reduced proportion of livestock with anthelmintics can slow resistance development in gastrointestinal nematodes by providing residue-free dung which could also benefit non-target organisms. We tested effects of TST on survival and reproduction of the dung beetle Onthophagus taurus (Scarabaeidae) in a factorial glasshouse experiment (Experimental treatments: five TST levels, 0.00, 0.25, 0.50, 0.75, 1.00 x four ivermectin concentrations, 125, 250, 375, 500 ppb). Each mesocosm comprised a 60 L bin containing sand, four dung pats and six pairs of adult beetles (F0 generation). No effects of TST level and ivermectin concentration on mortality of F0 adults after one week were observed. F0 adult brood ball production was affected by TST level, particularly at high ivermectin concentrations. Brood ball production increased as more untreated pats became available, with greater increases at higher ivermectin concentrations. We tested for evidence of a reported attraction of dung beetles to ivermectin-treated dung using a novel glitter-marker to trace the origin of dung used in brood balls. Where mesocosms contained both dung types, the proportion of brood balls created from untreated dung showed no statistical difference from the null expectation based on untreated dung availability in the mesocosm. Emergence of F1 adults was affected by the increase in TST, with this effect dependent on concentration. Treatments with concentrations of 250–500 ppb had the lowest emergence rates (ca. 5–20 % in mesocosms where all dung pats were treated) but emergence rates increased with TST level, reaching 68–88 % emergence where no dung pats were treated with ivermectin. Ivermectin-induced mortality occurred predominantly at egg and first instar stages. TST can provide refuges for dung beetles offering a strategy for livestock producers to maintain livestock welfare whilst benefiting from ecosystem services provided by important insects.

Diurnal temperature variation impacts energetics but not reproductive effort across seasons in a temperate dung beetle.

Temperature varies on multiple timescales and ectotherms must adjust to these changes to survive. These adjustments may lead to energetic trade-offs between self-maintenance and reproductive investment. However, we know little about how diurnal and seasonal temperature changes impact energy allocation. Here we used a combination of empirical data and modeling of both thermoregulatory behaviors and body temperature to examine potential energetic trade-offs in the dung beetle Onthophagus taurus. Beginning in March 2020, universities and laboratories were officially closed due to the COVID-19 pandemic. We thus performed experiments at a private residence near Knoxville, Tennessee, USA, leveraging the heating, ventilation and air conditioning of the home to manipulate temperature and compare beetle responses to stable indoor temperatures versus variable outdoor temperatures. We collected O. taurus beetles in the early-, mid-, and late-breeding seasons to examine energetics and reproductive output in relation to diurnal and seasonal temperature fluctuations. We recorded the mass of field fresh beetles before and after a 24-h fast and used the resulting change in mass as a proxy for energetic costs of self-maintenance across seasons. To understand the impacts of diurnal fluctuations on energy allocation, we held beetles either indoors or outdoors for 14-day acclimation trials, fed them cow dung, and recorded mass change and reproductive output. Utilizing biophysical models, we integrated individual-level biophysical characteristics, microhabitat-specific performance, respirometry data, and thermoregulatory behaviors to predict temperature-induced changes to the allocation of energy toward survival and reproduction. During 24 h of outdoor fasting, we found that beetles experiencing reduced temperature variation lost more mass than those experiencing greater temperature variation, and this was not affected by season. By contrast, during the 14-day acclimation trials, we found that beetles experiencing reduced temperature variation (i.e., indoors) gained more mass than those experiencing greater temperature variation (i.e., outdoors). This effect may have been driven by shifts in the metabolism of the beetles during acclimation to increased temperature variation. Despite the negative relationship between temperature variation and energetic reserves, the only significant predictor of reproductive output was mean temperature. Taken together, we find that diurnal temperature fluctuations are important for driving energetics, but not reproductive output.

Onthophagus taurus Increases Soil Microbes Associated with Nutrient Cycling in California Pastureland Soils

When grazed intensively, grasslands can result in biodiversity loss and increased greenhouse gas emissions. However, when grasslands are maintained using agroecological practices, they can create diverse habitats and act as carbon sinks. Carbon sequestration in soil is mediated by the soil microbial community, which is largely affected by influxes of nutrients and the soil arthropod community. One such arthropod, the tunneling dung beetle Onthophagus taurus (Schreber 1759) incorporates animal dung into the soil, influencing both the soil microbial community and nutrient cycling. While dung beetles do affect the soil microbial community, there is a dearth of studies that examine the effect of tunneling dung beetles on soil microbial communities that may be associated with nutrient cycling. This study looks at the effect of the tunneling dung beetle O. taurus on the fungal, bacterial, and archaeal community of grassland soil in an in situ experiment in California's Central Coast region, using eDNA metabarcoding of the 16S locus for bacteria and archaea and the ITS1 locus for fungi. We find that O. taurus has a clear effect on the soil microbial community and its presence increases the abundance of soil microbes affiliated with degrading plant materials and carbohydrate metabolism.

Evaluating old truths: Final adult size in holometabolous insects is set by the end of larval development.

For centuries, it has been understood that the final size of adult holometabolous insects is determined by the end of the larval stage, and that once they transform to adults, holometabolous insects do not grow. Despite this, no previous study has directly tested these "old truths" across holometabolous insects. Here, we demonstrate that final adult size is set at the end of the last larval stage in species representing each of the four orders of holometabolous insects: the fruit fly Drosophila melanogaster (Diptera), the tobacco hornworm Manduca sexta (Lepidoptera), the dung beetle Onthophagus taurus (Coleoptera), and the Florida carpenter ant Camponotus floridanus (Hymenoptera). Furthermore, in both D. melanogaster and C. floridanus, we show that the size of adult individuals fluctuates but does not significantly change. Therefore, our study finally confirms these two basic assumptions in the biology of insects, which have for centuries served as the foundation for studies of insect growth, size, and allometry.

Comparative metabolomic analysis of polyphenic horn development in the dung beetle Onthophagus taurus.

Organisms alter their phenotypes in response to changing environmental conditions. The developmental basis of this phenomenon, phenotypic plasticity, is a topic of broad interest in many fields of biology. While insects provide a suitable model for studying the genetic basis of phenotypic plasticity, the physiological aspects of plasticity are not fully understood. Here, we report the physiological basis of polyphenism, an extreme form of phenotypic plasticity by utilizing a dung beetle species, Onthophagus taurus. We highlighted the metabolome between sexes as well as two distinct male morphs—large and small horns. Unlike results from previous transcriptomic studies, the comparative metabolomic study revealed that differences in metabolite level were more prominent between animals with different body sizes than different sexes. Our results also indicate that specific metabolites and biochemical pathways may be active during horn size determination.

Doublesex mediates species-, sex-, environment- and trait-specific exaggeration of size and shape.

Context-dependent trait exaggeration is a major contributor to phenotypic diversity. However, the genetic modifiers instructing development across multiple contexts remain largely unknown. We use the arthropod tibia, a hotspot for segmental differentiation, as a paradigm to assess the developmental mechanisms underlying the context-dependent structural exaggeration of size and shape through nutritional plasticity, sexual dimorphism and segmental differentiation. Using an RNAseq approach in the sexually dimorphic and male-polyphenic dung beetle Digitonthophagus gazella, we find that only a small portion (3.7%) of all transcripts covary positively in expression level with trait size across contexts. However, RNAi-mediated knockdown of the conserved sex-determination gene doublesex suggests that it functions as a context-dependent master mediator of trait exaggeration in D. gazella as well as the closely related dung beetle Onthophagus taurus. Taken together, our findings suggest (i) that the gene networks associated with trait exaggeration are highly dependent on the precise developmental context, (ii) that doublesex differentially shapes morphological exaggeration depending on developmental contexts and (iii) that this context-specificity of dsx-mediated trait exaggeration may diversify rapidly. This mechanism may contribute to the resolution of conflict arising from environment-dependent antagonistic selection among sexes and divergent developmental contexts in a wide range of animals.

Can Sexual Selection Drive the Evolution of Sperm Cell Structure?

Sperm cells have undergone an extraordinarily divergent evolution among metazoan animals. Parker recognized that because female animals frequently mate with more than one male, sexual selection would continue after mating and impose strong selection on sperm cells to maximize fertilization success. Comparative analyses among species have revealed a general relationship between the strength of selection from sperm competition and the length of sperm cells and their constituent parts. However, comparative analyses cannot address causation. Here, we use experimental evolution to ask whether sexual selection can drive the divergence of sperm cell phenotype, using the dung beetle Onthophagus taurus as a model. We either relaxed sexual selection by enforcing monogamy or allowed sexual selection to continue for 20 generations before sampling males and measuring the total length of sperm cells and their constituent parts, the acrosome, nucleus, and flagella. We found differences in the length of the sperm cell nucleus but no differences in the length of the acrosome, flagella, or total sperm length. Our data suggest that different sperm cell components may respond independently to sexual selection and contribute to the divergent evolution of these extraordinary cells.

Dung beetles show metabolic plasticity as pupae and smaller adult body size in response to increased temperature mean and variance

Though organisms may use thermal plasticity to cope with novel temperature regimes, our understanding of plastic responses is limited. Research on thermal plasticity has traditionally focused on the response of organisms to shifts in mean temperatures. However, increased temperature variation can have a greater impact on organismal performance than mean temperature alone. In addition, thermal plasticity studies are often designed to investigate plasticity in response to more extreme temperatures despite the fact that organisms make physiological adjustments to diurnal temperature fluctuations that they experience. Using pupae of the dung beetle Onthophagus taurus, we investigated the potential for plasticity in response to increasing temperature mean and variance using thermal regimes that were well within the species critical thermal limits. We reared 40 beetles from egg to pupae (n = 20) or adults (n = 20) at one of nine incubation treatments, including all combinations of three mean temperatures (22, 24, 26 °C) and three amplitudes of fluctuation (±2, ±4, ±8 °C). To measure thermal plasticity of pupae, we quantified CO2 production across a range of temperatures (i.e., 15, 20, 25, and 30 °C) for 20 beetles per treatment. The relationship between CO2 production and temperature provides an estimate of energetic costs at a given temperature (i.e., using the intercept) and thermal sensitivity (i.e., using the slope). We reared the remaining O. taurus in each treatment (n = 20) to adulthood and then recorded mass (g) to determine body size, a proxy for fitness. Pupae exhibited thermal plasticity in response to the additive and interactive effects of temperature mean and variance. Pupae reared in the warmest and most variable treatment (26 ± 8 °C) showed the greatest decrease in overall metabolism compared to all other treatments, and adult beetles from this treatment (26 ± 8 °C) were also significantly smaller than adult beetles from any other treatment. We found that both temperature mean and variance contributed to thermal plasticity of pupae and had consequences for adult body size, a trait related to dung beetle fitness. Importantly, the temperatures we used in our treatments are not extreme and are likely well below the critical thermal maxima of the species, demonstrating that organisms can make adjustments to temperatures they experience across diurnal or seasonal timescales.

Evolution and plasticity of morph-specific integration in the bull-headed dung beetle Onthophagus taurus.

Developmental and evolutionary processes underlying phenotypic variation frequently target several traits simultaneously, thereby causing covariation, or integration, among phenotypes. While phenotypic integration can be neutral, correlational selection can drive adaptive covariation. Especially, the evolution and development of exaggerated secondary sexual traits may require the adjustment of other traits that support, compensate for, or otherwise function in a concerted manner. Although phenotypic integration is ubiquitous, the interplay between genetic, developmental, and ecological conditions in shaping integration and its evolution remains poorly understood. Here, we study the evolution and plasticity of trait integration in the bull‐headed dung beetle Onthophagus taurus which is characterized by the polyphenic expression of horned (‘major’) and hornless (‘minor’) male morphs. By comparing populations subject to divergent intensities of mate competition, we tested whether mating system shifts affect integration of traits predicted to function in a morph‐specific manner. We focussed on fore and hind tibia morphology as these appendages are used to stabilize major males during fights, and on wings, as they are thought to contribute to morph‐based differences in dispersal behavior. We found phenotypic integration between fore and hind tibia length and horn length that was stronger in major males, suggesting phenotypic plasticity in integration and potentially secondary sexual trait compensation. Similarly, we observed that fore tibia shape was also integrated with relative horn length. However, although we found population differentiation in wing and tibia shape and allometry, populations did not differ in integration. Lastly, we detected little evidence for morph differences in integration in either tibia or wing shape, although wing allometries differed between morphs. This contrasts with previous studies documenting intraspecific differentiation in morphology, behavior, and allometry as a response to varying levels of mate competition across O. taurus populations. We discuss how sexual selection may shape morph‐specific integration, compensation, and allometry across populations.

Phenotypic plasticity but no adaptive divergence in cuticular hydrocarbons and desiccation resistance among translocated populations of dung beetles

Translocated populations must adapt to their new environment to survive. A key aspect of survival for insects is the maintenance of water balance. It is thought that insects can adapt to dry environments by adjusting their cuticular hydrocarbon (CHC) profile to reduce water loss, though there is limited empirical support for this, and studies generally focus on other roles of CHCs, such as chemical communication. We tested for phenotypic adaptation in introduced populations of the Mediterranean dung beetle Onthophagus taurus, which have become established along a climatic gradient from dry northern to wet southern locations in southwestern Australia. We compared CHC profiles and desiccation resistance between northern and southern populations of the species. To quantify desiccation resistance, we measured both the rate of weight loss and time until death in beetles incubated at 35 °C. We tested for associations between these measures of desiccation resistance and CHC profiles, which were obtained through gas chromatography mass spectrometry. The abundance of CHCs was positively associated with desiccation resistance, and individuals that underwent the desiccation treatment produced a greater quantity of several CHC compounds indicating their ability to plastically adjust their CHC profile in response to desiccation stress. However, northern populations did not produce more CHCs than southern populations, and southern populations were better able to tolerate desiccation. Our results suggest that CHCs are an important component of desiccation resistance in O. taurus. However, the lack of evidence for local adaptation to the drier northern climate suggests there may be constraints to increasing desiccation resistance, and demonstrates the importance of considering local environmental conditions before translocating populations to new locations.

Notch signaling patterns head horn shape in the bull-headed dung beetle Onthophagus taurus.

Size and shape constitute fundamental aspects in the description of morphology. Yet while the developmental-genetic underpinnings of trait size, in particular with regard to scaling relationships, are increasingly well understood, those of shape remain largely elusive. Here we investigate the potential function of the Notch signaling pathway in instructing the shape of beetle horns, a highly diversified and evolutionarily novel morphological structure. We focused on the bull-headed dung beetle Onthophagus taurus due to the wide range of horn sizes and shapes present among males in this species, in order to assess the potential function of Notch signaling in the specification of horn shape alongside the regulation of shape changes with allometry. Using RNA interference-mediated transcript depletion of Notch and its ligands, we document a highly conserved role of Notch signaling in general appendage formation. By integrating our functional genetic approach with a geometric morphometric analysis, we find that Notch signaling moderately but consistently affects horn shape, and does so differently for the horns of minor, intermediate-sized, and major males. Our results suggest that the function of Notch signaling during head horn formation may vary in a complex manner across male morphs, and highlights the power of integrating functional genetic and geometric morphometric approaches in analyzing subtle but nevertheless biologically important phenotypes in the face of significant allometric variation.

A link between heritable parasite resistance and mate choice in dung beetles

AbstractParasites play a central role in the adaptiveness of sexual reproduction. Sexual selection theory suggests a role for parasite resistance in the context of mate choice, but the evidence is mixed. The parasite-mediated sexual selection (PMSS) hypothesis derives a number of predictions, among which that resistance to parasites is heritable, and that female choice favors parasite resistance genes in males. Here, we tested the PMSS hypothesis using the dung beetle Onthophagus taurus, a species that can be heavily parasitized by Macrocheles merdarius mites, which are known to affect adult survival. We investigated the heritability of resistance to M. merdarius, as well as whether female O. taurus impose a mating bias against males susceptible to mite infestation. Female choice for parasite resistance is difficult to disentangle from the possibility that females are simply choosing less parasitized males due to naturally selected benefits of avoiding contracting those parasites. This is especially likely for ectoparasites, such as mites. We tackled this problem by performing a mate choice trial first, and then measuring a male’s resistance to mite infestation. Resistance to mite infestation exhibited significant levels of additive genetic variance. Although we found no relationship between mating success and parasite resistance, males with greater resistance to infestation mated for longer. If females control copula duration, given that short copulations often result in mating failure, female choice could act on parasite resistance.

Diplogastrellus nematodes are sexually transmitted mutualists that alter the bacterial and fungal communities of their beetle host

A recent accumulation of studies has demonstrated that nongenetic, maternally transmitted factors are often critical to the health and development of offspring and can therefore play a role in ecological and evolutionary processes. In particular, microorganisms such as bacteria have been championed as heritable, symbiotic partners capable of conferring fitness benefits to their hosts. At the same time, parents may also pass various nonmicrobial organisms to their offspring, yet the roles of such organisms in shaping the developmental environment of their hosts remain largely unexplored. Here, we show that the nematode Diplogastrellus monhysteroides is transgenerationally inherited and sexually transmitted by the dung beetle Onthophagus taurus By manipulating artificial chambers in which beetle offspring develop, we demonstrate that the presence of D. monhysteroides nematodes enhances the growth of beetle offspring, empirically challenging the paradigm that nematodes are merely commensal or even detrimental to their insect hosts. Finally, our research presents a compelling mechanism whereby the nematodes influence the health of beetle larvae: D. monhysteroides nematodes engineer the bacterial and fungal communities that also inhabit the beetle developmental chambers, including specific taxa known to be involved in biomass degradation, possibly allowing larval beetles better access to their otherwise recalcitrant, plant-based diet. Thus, our findings illustrate that nongenetic inheritance can include intermediately sized organisms that live and proliferate in close association with, and in certain cases enhance, the development of their hosts' offspring.

Adaptive maternal behavioral plasticity and developmental programming mitigate the transgenerational effects of temperature in dung beetles

Phenotypic plasticity allows organisms to cope with rapid environmental change. Yet exactly when during ontogeny plastic responses are elicited, whether plastic responses produced in one generation influence phenotypic variation and fitness in subsequent generations, and the role of plasticity in shaping population divergences, remains overall poorly understood. Here, we use the dung beetleOnthophagus taurusto assess plastic responses to temperature at several life stages bridging three generations and compare these responses across three recently diverged populations. We find that beetles reared at hotter temperatures grow less than those reared at mild temperatures, and that this attenuated growth has transgenerational consequences by reducing offspring size and survival in subsequent generations. However, we also find evidence that plasticity may mitigate these consequences in two ways: 1) mothers modify the temperature of their offspring's developmental environment via behavioral plasticity and 2) in one population, offspring exhibit accelerated growth when exposed to hot temperatures during very early development (‘developmental programming’). Lastly, our study reveals that offspring responses to temperature diverged among populations in fewer than 100 generations, possibly in response to range‐specific changes in climatic or social conditions.

Population density mediates the interaction between pre- and postmating sexual selection.

When females mate with more than one male, sexual selection acts both before and after mating. The interaction between pre- and postmating episodes of selection is expected to be context dependent, but few studies have investigated how total sexual selection changes under different ecological conditions. We examined how population density mediates the interaction between pre- and postmating sexual selection by establishing replicate populations of the horned dung beetle Onthophagus taurus at low, medium, and high densities, and then using microsatellite-based parentage analyses to measure male fitness. We found that mating success and fertilization success were positively correlated at all three densities, but the strength of the correlation decreased with increasing density. We also found a shift from negative to positive linear selection on testes mass as density increased, and opposing selection on weapons and testes at high densities. These patterns suggest that the importance of postmating processes increases with increasing population density, which reduces the selective advantage of weapons for premating contest competition, and increases the selective advantage of large ejaculates for postmating sperm competition. We expect that density-dependent selection on testes mass has contributed to the phenotypic variation observed between natural populations of O. taurus that differ in density.

The role of ancestral phenotypic plasticity in evolutionary diversification: population density effects in horned beetles

Plasticity's role in shaping phenotypic diversification continues to receive considerable attention. One especially debated issue concerns the significance of genetic accommodation in diversification, and the proposed role of ancestrally plastic responses in facilitating or biasing subsequent genetically canalized differentiation among taxa. Here, we investigated whether pre-existing plasticity in response to variation in population density present in the ancestral Mediterranean range of the bull-headed dung beetle Onthophagus taurus may have mediated previously documented rapid canalized divergences among descendent exotic populations that have been subject to dramatically different levels of competition for mates and resources in the field. We focused on two maternal behavioural traits, two life history traits and two morphological traits. We find that (1) Mediterranean O. taurus exhibited plasticity in response to adult densities for four of our six focal traits; (2) in two of those, plastic responses matched the direction of canalized divergences among natural populations; and (3) the presence and direction of plasticity appeared unrelated to trait type. More generally, our results provide partial support for the hypothesis that evolution by genetic accommodation could have contributed to the very early stages of population differentiation in a subset of traits in O. taurus.

Doublesex alters aggressiveness as a function of social context and sex in the polyphenic beetle Onthophagus taurus

Despite sharing nearly the same genome, individuals within the same species can vary drastically in both morphology and behaviour as a function of developmental stage, sex or developmental plasticity. Thus, regulatory processes must exist that enable the stage-, sex- or environment-specific expression of traits and their integration during ontogeny, yet exactly how trait complexes are co-regulated and integrated is poorly understood. In this study, we explore the developmental genetic basis of the regulation and integration of environment-dependent sexual dimorphism in behaviour and morphology in the horn-polyphenic dung beetle Onthophagus taurus through the experimental manipulation of the transcription factor doublesex (dsx). The gene dsx plays a profound role in the developmental regulation of morphological differences between sexes as well as alternative male morphs by inhibiting horn formation in females but enabling nutrition-responsive horn growth in males. Specifically, we investigated whether experimental downregulation of dsx expression affects male and female aggressive and courtship behaviours in two social contexts: interactions between individuals of the same sex and interactions between males and females. We find that dsx downregulation significantly alters aggressiveness in both males and females, yet does so differently for both sexes as a function of social context: dsxRNAi males exhibited elevated aggression towards males but showed reduced aggression towards females, whereas dsxRNAi females became more aggressive towards males, while their aggressiveness towards other females was unaffected. Moreover, we document unexpectedly high levels of female aggression independent of dsx treatment in both wild-type and control-injected individuals. Lastly, we found no effects of dsxRNAi on courtship and mating behaviours. We discuss the role of dsx in the regulation of sex-specific and plastic behaviours, the unexpectedly high levels of aggression of hornless dsxRNAi males in relation to the well-established description of the hornless sneaker phenotype and the potential ecological function of high female aggression.

Contextualized niche shifts upon independent invasions by the dung beetle Onthophagus taurus

The historical contingencies of biological invasions may have important consequences for final invasion outcomes. Here, we characterize the variations in the realized niche during the invasions of the bull-headed dung beetle Onthophagus taurus (Coleoptera: Scarabaeidae) from its native Mediterranean range following accidental (Eastern North America) as well as deliberate (Western North America, Western Australia, and Eastern Australia) releases into novel, exotic ranges approximately 50 years ago. Specifically, we examined whether the climatic responses of exotic O. taurus have diverged from those characterizing their native range, and if so, to what degree and in what dimensions. We found that when compared to the native range, all exotic populations exhibited similar overlap proportions regardless of invasion history. However, more detailed analysis of climatic niche features showed that all three deliberately established populations were characterized by overall similar climatic niche features, whereas the accidentally-established Eastern North American populations have undergone significant changes in their climatic niche. Specifically, when analog climates were considered on the background of each pairwise range comparison, accidentally-established Eastern North American populations showed a different climatic niche expansion than their deliberately introduced Australian or Western North American counterparts, in particular towards colder and more humid climates. We discuss our results in the context of the widely divergent introduction histories of O. taurus in Australia and North America, and highlight the possible roles of contrasting propagule sizes, disparate genetic profiles and variances, adaptive processes and invadable landscapes in shaping invasion outcomes in the different exotic ranges.

The transcriptomic basis of tissue- and nutrition-dependent sexual dimorphism in the beetle Onthophagus taurus.

Sexual dimorphism accounts for a large fraction of intraspecific diversity. However, not all traits are equally sexually dimorphic; instead, individuals are mosaics of tissues that vary in their ability to exhibit dimorphism. Furthermore, the degree of a trait's sexual dimorphism is frequently environment‐dependent, with elaborate sexual dimorphism commonly being restricted to high nutritional conditions. Understanding the developmental basis and evolution of condition‐dependent sexual dimorphism can be critically informed by determining – across tissues and nutritional conditions – what sex‐biased genes are deployed and how they interact and translate into functional processes. Indeed, key theories concerning the evolution of condition‐dependent sexually dimorphic traits rest on assumptions regarding their developmental genetic underpinnings, yet, have largely gone unexamined by empirical studies. Here, we provide such evidence by investigating the transcriptomic basis of tissue‐ and nutrition‐dependent sexual dimorphism in the bull‐headed dung beetle Onthophagus taurus. Our findings suggest (1) that generating morphological sexual dimorphism requires sex‐biased gene expression in and developmental remodeling of both sexes, regardless of which sex exhibits externally visible trait exaggeration, (2) that although sexually dimorphic phenotypes are comprised of traits underlain by independent repertoires of sex‐biased gene expression, they act similarly at a functional level, and (3) that sexual dimorphism and condition‐dependence share common genetic underpinnings specifically in sexually‐selected traits.

Selection on male physical performance during male–male competition and female choice

Male contest competition and female choice are key features of the mating systems of many species, and whole-organism performance may be targeted by both mechanisms given the dynamic body movements required during fighting and courtship displays. Using the dimorphic horned dung beetle Onthophagus taurus as a model system, we tested whether physical performance was important in determining a male’s victory in fights, and whether successful fighters were preferred by females as mates. We found that physical strength, horn length, and body mass were significant predictors of male fighting success, but males that won fights were not more attractive to females. Rather, females preferred males that delivered a high courtship rate, which was not correlated with strength, horn length, or body mass, but previously has been shown to be genetically correlated with body condition. The fact that there was no relationship between fighting success and mating success suggests that selection on traits favored by male–male competition and female choice can act relatively independently in this species, although both mechanisms appear to favor traits (strength and courtship, respectively) that are linked to a male’s ability to acquire and allocate resources for mass gain. Future work is needed to determine the relative contributions of these processes to the total strength of sexual selection acting on male phenotypes.