Dung beetles increase plant growth: a meta-analysis.

The ecosystem services provided by dung beetles are well known and valued. Dung beetles bury dung for feeding and breeding, and it is generally thought that the process of burying dung increases nutrient uptake by plant roots, which promotes plant growth. Many studies have tested the effects of dung beetles on plant growth, but there has been no quantitative synthesis of these studies. Here we use a multi-level meta-analysis to estimate the average effect of dung beetles on plant growth and investigate factors that moderate this effect. We identified 28 publications that investigated dung beetle effects on plant growth. Of these, 24 contained the minimum quantitative data necessary to include in a meta-analysis. Overall, we found that dung beetles increased plant growth by 17%; the 95% CI for possible values for the true increase in plant growth that were most compatible with our data, given our statistical model, ranged from 1% to 35%. We found evidence that the dung beetle-plant growth relationship is influenced by the plant measurement type and the number of beetles accessing the dung. However, beetles did not increase plant growth in all quantitative trials, as individual effect sizes ranged from -72% to 806%, suggesting important context-dependence in the provision of ecosystem services.

Indirect parental effects on offspring viability by egg-derived fluids in an external fertilizer.

The capacity for parents to influence offspring phenotypes via nongenetic inheritance is currently a major area of focus in evolutionary biology. Intriguing recent evidence suggests that sexual interactions among males and females, both before and during mating, are important mediators of such effects. Sexual interactions typically extend beyond gamete release, involving both sperm and eggs, and their associated fluids. However, the potential for gamete-level interactions to induce nongenetic parental effects remains under-investigated. Here, we test for such effects using an emerging model system for studying gamete interactions, the external fertilizer Mytilus galloprovincialis. We employed a split-ejaculate design to test whether exposing sperm to egg-derived chemicals (ECs) from a female would affect fertilization rate and offspring viability when those sperm were used to fertilize a different female's eggs. We found separate, significant effects of ECs from non-fertilizing females on both fertilization rate and offspring viability. The offspring viability effect indicates that EC-driven interactions can have nongenetic implications for offspring fitness independent of the genotypes inherited by those offspring. These findings provide a rare test of indirect parental effects driven exclusively by gamete-level interactions, and to our knowledge the first evidence that such effects occur via the gametic fluids of females.

Natural and sexual selection on cuticular hydrocarbons: a quantitative genetic analysis.

While the reproductive benefits of sexual displays have been widely studied, we have relatively limited evidence of the fitness costs associated with most display traits. Insect cuticular hydrocarbon (CHC) profiles are sexually selected traits that also protect against desiccation. These two functions are thought to oppose each other, with investment in particular compounds believed to increase attractiveness at the expense of compounds that protect against water loss. We investigated this potential trade-off in a quantitative genetic framework using the Australian field cricket, Teleogryllus oceanicus. Several compounds were significantly genetically correlated with either attractiveness or desiccation resistance. Of these compounds, one was negatively genetically correlated with attractiveness but positively genetically correlated with desiccation resistance. Furthermore, scoring each individual's overall CHC profile for its level of attractiveness and desiccation resistance indicated a negative genetic correlation between these multivariate phenotypes. Together, our results provide evidence for a genetic trade-off between sexually and naturally selected functions of the CHC profile. We suggest that the production of an attractive CHC profile may be costly for males, but highlight the need for further work to support this finding experimentally. Genetic covariation between the CHC profile and attractiveness suggests that females can gain attractive sons through female choice.

Experimental evidence for the role of sexual selection in the evolution of cuticular hydrocarbons in the dung beetle, Onthophagus taurus.

A role for sexual selection in the evolution of insect cuticular hydrocarbons (CHCs) is suggested by observations of selection acting on male CHCs during female mate choice. However, evidence that CHCs evolve in response to sexual selection is generally lacking, and there is a need to extend our understanding beyond well-studied taxa. Experimental evolution offers a powerful approach to investigate the effect of sexual selection on the evolution of insect CHCs. We conducted such an experiment using the dung beetle, Onthophagus taurus. After six, 12 and 21 generations of experimental evolution, we measured the CHCs of beetles from three populations subject to sexual selection and three populations within which sexual selection had been removed via enforced monogamy. We found that the male CHC profile responded to the experimental removal of sexual selection. Conversely, the CHC profile of females responded to the presence of sexual selection but not to its removal. These results show that sexual selection can be an important mechanism affecting the evolution of insect CHCs and that male and female CHCs can evolve independently.

A costly chemical trait: phenotypic condition dependence of cuticular hydrocarbons in a dung beetle.

Chemical traits are increasingly recognized as important cues used in mate choice. For example, the cuticular hydrocarbons (CHCs) of insects have been shown to influence mating success in a range of taxa. Less is known, however, about how CHCs are expressed in proportion to an individual's condition, and consequently whether CHCs can function as condition-dependent signals of quality. We investigated this question using the dung beetle, Onthophagus taurus. CHCs are subject to sexual selection in this species through mate choice. A dietary manipulation revealed condition dependence of CHC expression for both sexes: dietary restriction decreased overall CHC production and altered the composition of CHCs. Furthermore, CHC production was associated with a measure of condition in beetles fed a limited diet but not those fed ad libitum. These results implicate a resource cost to CHC production that is likely to result in trade-offs with other fitness components in this species, as these respond similarly to a dietary restriction. The CHC profiles showed sexual dimorphism: males produced more CHCs and the sexes differed in the blend of compounds they produced. There was evidence for a male dimorphism in the CHC profile, in line with the presence of the alternative reproductive tactics (minor sneaks and major fighters) in this species. However, rather than mimicking a female CHC profile, minor males differed more from females than major males. Our results suggest that CHCs are a costly trait in O. taurus that has the potential to act as a condition-dependent signal of quality.

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.

Deliberately introduced dung beetles in Australia: 12 years of occurrence and abundance records from 2001 to 2022.

Since 1968, the Australian Dung Beetle Project has carried out field releases of 43 deliberately introduced dung beetle species for the biological control of livestock dung and dung-breeding pests. Of these, 23 species are known to have become established. For most of these species, sufficient time has elapsed for population expansion to fill the extent of their potential geographic range through both natural and human-assisted dispersal. Consequently, over the last 20 years, extensive efforts have been made to quantify the current distribution of these introduced dung beetles, as well as the seasonal and spatial variation in their activity levels. Much of these data and their associated metadata have remained unpublished, and they have not previously been synthesized into a cohesive dataset. Here, we collate and report data from the three largest dung beetle monitoring projects from 2001 to 2022. Together, these projects encompass data collected from across Australia, and include records for all 23 species of established dung beetles introduced for biocontrol purposes. In total, these data include 22,718 presence records and 213,538 absence records collected during 10,272 sampling events at 546 locations. Most presence records (97%) include abundance data. In total, 1,752,807 dung beetles were identified as part of these data. The distributional occurrence and abundance data can be used to explore questions such as factors influencing dung beetle species distributions, dung beetle biocontrol, and insect-mediated ecosystem services. These data are provided under a CC-BY-NC 4.0 license and users are encouraged to cite this data paper when using the data.

Female cuticular hydrocarbons can signal indirect fecundity benefits in an insect.

Male choosiness of prospective female mating partners provides an increasingly recognized pathway through which males can increase their fitness. For example, males may increase their number of offspring by targeting more fecund females as mating partners. If fecundity is heritable, males that mate with more fecund females can also receive the indirect benefit of more fecund daughters. In species where female fecundity is not directly assessable, female ornaments may act as signals of fecundity. However, whether female ornaments reliably signal the indirect benefit of more fecund daughters is not well understood. We investigated this question using the field cricket, Teleogryllus oceanicus. Previous work had identified the cuticular hydrocarbon (CHC) profile as a female sexual display trait in T. oceanicus. To examine whether CHCs can provide a reliable signal of fecundity, we tested whether individual CHC compounds and the first principal axis of CHC variation (PC1) are genetically correlated with ovary mass, a reliable proxy for fecundity in this species. We found significant genetic correlations between ovary mass and three individual CHC compounds, as well as PC1. This result indicates that by targeting females as mating partners based on their CHC profile, males can sire more fecund daughters.