Experimental warming disrupts reproduction and dung burial by a ball‐rolling dung beetle

1. Insects are sensitive to climate change. Consequently, insect‐mediated ecosystem functions and services may be altered by changing climates. 2. Dung beetles provide multiple services by burying manure. Using climate‐controlled chambers, the effects of warming on dung burial and reproduction by the dung beetle Sisyphus rubrus Paschalidis, 1974 were investigated. Sisyphus rubrus break up dung by forming and rolling away balls of manure for burial and egg deposition. 3. To simulate warming in the chambers, 0, 2 or 4 °C offsets were added to field‐recorded, diurnally fluctuating temperatures. We measured dung ball production and burial, egg laying, survival and residence times of beetles. 4. Temperature did not affect the size or number of dung balls produced; however warming reduced dung ball burial by S. rubrus. Because buried balls were more likely to contain eggs, warming could reduce egg laying via a reduction in ball burial. Warming reduced the humidity inside the chambers, and a positive relationship was found between the number of dung balls produced and humidity in two temperature treatments. Temperature did not affect survival, or whether or not a beetle left a chamber. Beetles that did leave the chambers took longer to do so in the warmest treatment. 5. This study demonstrates that climate warming could reduce reproduction and dung burial by S. rubrus, and is an important first step to understanding warming effects on burial services. Future studies should assess warming effects in field situations, both on individual dung beetle species and on aggregate dung burial services.     

Dung burial strategies in some South African coprine and onitine dung beetles (Scarabaeidae: Scarabaeinae)

ABSTRACT.

• 1 The dung colonization and dung burial behaviour of twelve crepuscular/nocturnal tunnelling (paracoprid) species of beetles were examined in order to identify mechanisms which might facilitate resource (dung) partitioning. The species were selected from a diverse assemblage of dung beetles, the members of which coexist in the sandy-soil regions of Natal, South Africa.
• 2 The pattern of dung colonization in relation to dung age was examined in the field using baited pitfall traps. Some species, e.g. Onitis deceptor Peringuey, Catharsius tricornutus De Geer and Copris elphenor Klug, showed a marked preference for fresh dung (1–2 days old) whereas other species, e.g. O. viridulus Boheman and Copris fallaciosus Gillet, preferred older dung (3–7 days old).
• 3 Two distinct patterns of dung burial were recognized. In the Coprini, dung burial was complete within 24–48h of pad colonization, and the level of dung burial was similar in the laboratory and in the field. In the Onitini, dung burial occurred progressively over a 12-day period, although the timing of initiation of dung burial varied between species: in O. deceptor nearly all individuals had begun burial within 2 days of pad colonization, whereas only 20% of O. viridulus had commenced dung burial by that time. However, nearly all O. viridulus had buried substantial quantities of dung by day 12.
• 4 The mass of dung buried per pair by the larger coprine beetles (100–300 g) and onitine beetles (400–1000 g) suggests that there is potential for inter- and intraspecific competition, even in pads colonized by relatively few beetles. The colonization and use of dung of different ages are discussed as means of resource partitioning in relation to the relative abilities of species to compete for dung.

     

Effects of adding exotic dung beetles to native fauna on bush fly breeding in the field

Egg to pupal survival of bush fly,Musca vetustissima Walker, under field conditions was examined during 1987/88 in an area of south-eastern Australia that had not been colonised by exotic dung beetles. In pads of cattle dung containing only the native fauna, fly survival ranged from 0.3% to 12.5%. The addition of 2 species of exotic dung beetles,Euoniticellus fulvus (Goeze) andOnthophagus taurus (Schreber) to field pads, in numbers similar to those observed at the collection site, reduced fly survival to between 0.3% and 4.4%. Fly survival in the presence of the native and exotic dung fauna was sufficiently low to keep fly breeding below their mean replacement level of 3% for most of the season. Widespread dispersal and establishment of exotic dung beetles in south-eastern Australia, alongside the native fauna, should lead to long-term reduction of the bush fly problem.      

Sexual selection and condition dependence of courtship display in three species of horned dung beetles

Sexual selection has traditionally been divided into competitionover mates and mate choice. Currently, models of sexual selectionpredict that sexual traits are expressed in proportion to thecondition of their bearer. In horned beetles, male contestcompetition is well established, but studies on female preferencesare scarce. Here I present data on male mating success and condition dependence of courtship rate in three species of horn-dimorphicdung beetles, Onthophagus taurus, Onthophagus binodis, andOnthophagus australis. I found that in the absence of malecontest competition, mating success of O. taurus and O. australiswas unrelated to their horn length and body size, whereas inO. binodis horn size had a negative effect but body size hada positive effect on male mating success. Overall, in O. binodismajor morph males had greater mating success than minor morphmales. In all three species male mating success was affectedby courtship rate, and the courtship rate was condition dependent such that when males were manipulated to be in poor conditionthey had lower courtship rates than males that were manipulatedto be in good condition. My findings provide new insight intothe mating systems of horned dung beetles and support an importantassumption in indicator models of sexual selection.     

Dung beetle community composition affects dung turnover in subtropical US grasslands

1. An important service in many ecosystems is the turnover and degradation of dung deposited by cattle. Dung beetles are the primary group of insects responsible for dung turnover, and factors affecting their abundance and distribution thus impact dung degradation. Lands lost to grazing due to dung buildup and pasture contamination total millions of acres per year in US pastures.
2. We evaluated the structural differences in dung beetle assemblages in natural grasslands versus a managed agroecosystem in subtropical southeastern Florida (USA). We measured the direct effect of dung longevity when dung beetle fauna normally inhabiting dung pats were excluded.
3. Our results indicate dung beetle abundance, functional diversity, and species richness have a substantial impact on the rate of dung turnover in subtropical pastoral lands with ~70% of dung removed from the soil surface after three months. Functional diversity and evenness did not have a significant positive effect on dung removal in managed, versus natural grasslands demonstrating a strong relationship between dung beetle assemblage composition and delivery of a key ecological process, dung degradation.
4. We suggest the importance of trees, which provide a thermal refuge for beetles, should be dispersed within matrixes of open pasture areas and within proximity to adjacent closed‐canopy hammocks to facilitate the exchange of dung beetles between habitats and therefore maintain the provisioning of dung degradation services by dung beetle assemblages.

     

Seasonal variations in numbers,biomass and breeding patterns of dung beettles [Coleoptera: Scarabaeidae] in Southern France

Seasonal variations in numbers and biomass and breeding of 28 species ofScarabaeidae-Scarabaeinae andGeotrupidae were studied at 2 temperate sites and 2 mediterranean sites in southern France in 1978 and 1979, using cattle-dung baited pitfall traps. Maximum dung burial occurred when females were parous and ovipositiong (i.e. during spring at the mediterranean sites and summer at the temperate sites). Using biomass as a measure of dung burial the most important spring active beetles at the mediterranean sites were:Budas bison, B. bubalus, Scarabaeus laticollis, Onthophagus lemur andCopris hispanus, and at the temperate sites:C. lunaris, O. vacca, andO. ovatus. These results confirmed the previous choice ofC. hispanus, C. lunaris, B. bison andO. vacca for introduction into Australia. In addition it is suggested thatB. bubalus, S. laticollis andO. lemur be considered for introduction into mediterranean areas of south western Australia to complement those species already introduced for bush fly control.      

Interactions between dung beetles (Coleoptera: Scarabaeidae) and the arbovirus vector Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae)

Abstract  The potential for dung beetles to reduce populations of the biting midge and arbovirus vector Culicoides brevitarsis in bovine dung was studied in the Hunter Valley, New South Wales (NSW) between 1999 and 2003 using natural populations of insects. Preliminary work to develop experimental procedures showed that: a few C. brevitarsis could survive in buried dung; dung beetles and C. brevitarsis coming to dung were unaffected by a background of shade-cloth used experimentally to prevent dung burial; the most abundant dung beetle, Onthophagus gazella L. and C. brevitarsis oviposition occurred concurrently in the first 2 d after dung deposition, and the potential for interaction between dung beetles and C. brevitarsis was greatest in open pasture adjacent to trees where cattle congregate at night. Laboratory experiments on dung burial showed that C. brevitarsis numbers decreased as numbers of dung beetles increased or as the dry weight of dung decreased due to burial. This was seldom reflected in the field where, although significant burial occurred experimentally in 9 of 20 trials over 3 years, a significant decline in C. brevitarsis numbers attributable to burial only occurred once. C. brevitarsis numbers in the field were lower in unburied dung in 70% of trials. Differences were significant twice and were considered the result of dung disturbance. In the laboratory, decreasing numbers of C. brevitarsis were related to three characteristics of disturbance: the flattening, spreading and reduction in wet weight of the dung. Evidence of C. brevitarsis activity throughout coastal NSW suggests that, while C. brevitarsis numbers may be modified by dung beetles, the interaction is insufficient to prevent their increase, spread and ability to transmit viruses to livestock.     

Effect of dung burial by the dung beetle Bubas bison on numbers and viability of Cryptosporidium oocysts in cattle dung

Cryptosporidium oocysts were inoculated into fresh dung (∼1.2 × 10⁴ oocysts per gram wet weight) and fed to dung beetles to assess the effect of dung burial by the dung beetle Bubas bison on the distribution of the oocysts in small cores of soil in the laboratory. The experiment consisted of five replicates of each of two treatments; controls (dung but no dung beetles) and the experimental treatment (inoculated dung and seven pairs of dung beetles). After 5 days, when approximately 90% of the dung was buried, the surface and buried dung was recovered and subsampled. The oocysts in the subsamples were recovered and enumerated using qPCR. Oocyst viability was evaluated using an assay based on the exclusion or inclusion of two fluorogenic vital dyes, 4′,6-diamidino-2-phenylindole (DAPI) and propidium iodide (PI). Results revealed that overall 13.7% of oocysts remained on the surface and 86.3% of oocysts were buried. The viability of oocysts in buried dung was only 10% compared to oocysts the surface dung (58%). Therefore, widespread dung burial by B. bison during the winter months could substantially reduce the numbers of Cryptosporidium oocysts available to be washed into waterways following winter rains.     

Influence ofOnthophagus gazella on hornfly,Haematobia irritans density in irrigated pastures

Haematobia irritans (L.) breeding in flood irrigated pastures of the lower Colorado Desert of southeastern California continues to remain unacceptably high during warm seasons (>1,000 adult flies per bovine head) despite the presence of moderately abundant populations ofOnthophagus gazella F. This study suggests that densities of > 40–70 adult beetles per dung pad and giving pronounced dung shredding activity, caused fly mortality of 38–56 %. The continued high abundance of adult horn flies on cattle suggests that at > 50% mortality, the pasture environment still produces sufficient flies to saturate cattle, although emigration might be reduced. Additional species of scarabs may be necessary to increase fly mortality. However, the dung drying activity of existingO. gazella significantly could interfere with resident staphylinid beetle breeding, which was significantly lower in pastures whereO. gazella reached densities of 40 per dung pad. Scarab beetle activity might also impede the introduction of superior predatory species for biological control.      

Duration and benefits of biparental brood care in the dung beetle Onthophagus vacca (Coleoptera: Scarabaeidae)

Abstract.

• 1 Single males, single females or pairs of dung beetles, Onthophagus vacca, were released on artificial small (100 g) or large (1000 g) dung pats in the laboratory. Emigrating beetles were trapped at 12 h intervals, and the number and size of the brood chambers were recorded after each replicate.
• 2 Emigration of males was delayed if females were present in the same dung pats, whereas emigration times of females were independent of the presence or absence of males.
• 3 A residency of 60 h proved to be a threshold value. Females emigrating before this time did not breed, whereas those emigrating later had built at least two brood chambers.
• 4 Females paired with males built more brood chambers than single females.
• 5 The reproductive success of pairs was not influenced by the size of the dung pats.

     

Longevity cost of reproduction for males but no longevity cost of mating or courtship for females in the male-dimorphic dung beetle Onthophagus binodis

Life history theory predicts a trade-off between current and future reproduction. Despite a wealth of research on the cost of reproduction for females, there have been very few studies that have looked at the cost of reproduction for males. Longevity is closely related to the opportunity for future reproduction, and thus decreased longevity in response to current reproductive effort has been used as a measure of the cost of reproduction. Here we examine the cost of reproduction for males and females in the dung beetle Onthophagus binodis. Like many onthophagines, O. binodis exhibit dimorphic male morphology; major males develop a large pronotal horn while minor males remain hornless. Alternative morphologies are associated with alternative reproductive tactics. Thus, we ask whether major and minor males pay different costs of reproduction. We found that in contrast to previous work on Diptera, mating is not costly in terms of reduced longevity for female dung beetles. Despite a longevity cost of reproduction for males, we found no evidence for differential longevity costs associated with alternative reproductive tactics.     

Patterns of alimentary resource use by dung beetles in introduced Brazilian pastures: Cattle versus sheep dung

Dung beetles (Coleoptera: Scarabaeidae) perform essential ecological functions in pastures, such as dung removal, nutrient recycling and parasite control. However, the patterns of alimentary use by dung beetles in introduced Brazilian pastures are poorly known. Here, we compared dung beetle species richness, abundance and species composition in cattle and sheep dung, and identified the dung beetle species preference by each dung type. In January 2019, dung beetles were sampled with pitfall traps baited with cattle and sheep dung in 12 introduced pastures (Urochloa spp.), in Aquidauana, Mato Grosso do Sul, Brazil. A total of 592 individuals belonging to 14 species of dung beetles were collected. Of the 14 species sampled, nine were recorded in both dung types, five were found exclusively in sheep dung and no species was exclusive to cattle dung. Species richness and abundance were higher in sheep dung. Species composition was different between the dung types. Dichotomius bos (Blanchard), Genieridium bidens (Balthasar), Onthophagus aeneus Blanchard and Trichillum externepunctatum Preudhomme de Borre were associated with sheep dung. Our results provide evidence that sheep dung is more attractive to dung beetles with a distinct community species between the two dung types, although the studied pastures have never been used before for sheep breeding. Thus, our data shows that the introduction of a new alimentary resource (e.g. sheep dung) can be an important strategy to help to obtain a more diverse dung beetle assemblage in introduced Brazilian pastures.     

INSECTARY STUDIES ON THE CONTROL OF DUNG BREEDING FLIES BY THE ACTIVITY OF THE DUNG BEETLE, ONTHOPHAGUS GAZELLA F. (COLEOPTERA: SCARABAEINAE)

i
The Afro-Asian dung beetle, Onthophagus gazella F., buries bovine dung as food for its larvae so rapidly that when beetle populations are of the order of 4 insects per 100 c.c. dung, entire cow pads are completely broken up and buried within 30 to 40 hours.
Insectary studies show that this rate of dung disposal caused 80 to 100 per cent. reduction in the numbers of the bushfly Musca vetustissima Walker emerging from the pads. Surviving maggots gave rise to small, stunted flies of low or nil reproductive capacity. Viable fly eggs or maggots were never found in brood balls, and it is presumed that they were destroyed in the course of the elaborate process by which the beetles convert lumps of dung into brood balls.
Speed of dung burial is the critical factor in fly control. If half of a cow pad is buried within the first 24 hours, few or no adult flies emerge. More rapid burial within this period resulting in the complete removal of the cow pad, produces complete fly control.     

Oviposition of the dung beetle Aphodius ater in relation to the abundance of yellow dungfly larvae (Scatophaga stercoraria)

1. The dung beetle Aphodius ater and the yellow dungfly Scatophaga stercoraria are temporally co-occurring species in sheep dung, which they use for reproduction and nutrition ( A. ater ) or for reproduction only ( S. stercoraria ) during the spring in northern Germany. Scatophaga stercoraria uses fresh sheep dung pellets a few hours old for oviposition, whereas A. ater lays eggs into 2–10-day-old pellets. In the present study, the egg laying behaviour of A. ater in sheep dung in relation to the presence of larvae of S. stercoraria was investigated experimentally.
2. Choice experiments, based on examining the egg laying behaviour of beetles in 2- and 4-day-old pellets with and without high and low densities of fly larvae, showed the following. In 2-day-old pellets, the beetles did not distinguish between pellets without fly larvae or with fly larvae at low larval density but avoided laying eggs into pellets with a high larval density. In 4-day-old pellets, the beetles always preferred to lay their eggs into pellets without fly larvae, regardless of larval density.
3. The influence of different densities of larvae of S. stercoraria on dung depletion was examined by measuring the dry weight, organic matter content and organic nitrogen content of the remaining dung after larval development. The presence of the larvae led to a reduction in all three parameters.
4. The beetles' behaviour of laying eggs into older pellets, and their awareness of the presence of high densities of fly larvae, enables them to avoid egg laying into pellets that will have been depleted by fly larvae before the beetle larvae have finished their development.     

Differences in temporal utilization patterns of dung pats among three scarabaeid dung beetles

1. Differences in the temporal utilization patterns of dung pats were investigated among three dung beetle species, Aphodius haroldianus, Onthophagus lenzii and Liatongus phanaeoides, and density effects of three species on emigration from a dung pat were compared.
2. A. haroldianus preferred fresh dung pats, immigrated to fresh pats and then emigrated from those pats quickly (mean duration of residence 48.4 h), whereas L. phanaeoides immigrated to older dung pats (over 7 days after deposition) as well as fresh ones and remained in the pats for the longest time (165.6 h).O. lenzii showed intermediate behavior in this respect (63.3 h).
3. In A. haroldianus, the presence of conspecifics induced faster emigration than presence of the other two species. L. phanaeoides showed the opposite relationship. In O. lenzii, the response to conspecifics was intermediate to those of the other two species.

     

Diversity and seasonal fluctuations in dung beetle (Coleoptera) community in Kurukshetra,India

Five pitfall traps baited with 150–200 g of fresh cattle dung were installed for 24 h at weekly intervals. A total of 991 dung beetles from 11 genera, 31 species and three subfamilies was obtained. The community was dominated by Oniticellus spinipes individuals by 32.3%. Ten species appeared only once during the collection period and species composition and dominance changed throughout the period. The overall pattern we detected in the organization of the dung beetle community is that the species richness, abundance and diversity rise in September and the 2^nd week of October. The dung beetle community was found to be affected by season.     

Ecosystem services provided by dung beetles in Australia

Although there are nearly 500 species of native dung beetles in Australia, most are adapted to small, hard, dry, pelletised marsupial droppings and not to dealing with the large, moist deposits of cattle. In 1788, Governor Arthur Phillip arrived at Botany Bay with five cows, two bulls, 44 sheep and seven horses: this signalled major changes in Australia. Now there are about 27 million cattle, whose annual dung production has a dry matter content of about 42 million tonnes. Until CSIRO introduced exotic dung beetles in the 1960s, the dung of these herbivores sat on the soil surface, sometimes for years, locking up organic matter, smothering pasture and polluting waterways. CSIRO introduced 53 exotic dung beetle species, of which 43 were released to the Australian mainland between 1965 and 1985. Twenty-three of these have become established, many of which have reached the natural limits of their distribution. I consider the reason for the failure of the other 30 species to establish and briefly review previous contributions to examining the role of dung beetles in delivering ecosystem services, noting that much of the published literature concerns laboratory studies. New field data are then examined on the way in which introduced species are transforming dung communities and the ecosystem services they provide. The capacity of deep-tunnelling dung beetles to transform the soil profile is examined along with their effects on pasture production and the flow of nutrients from dung on pasture. The biocontrol capacity of dung beetle activity is considered in relation to the native bush fly, Musca vetustissima, the introduced buffalo fly, Haematobia irritans exigua, and dung-borne intestinal parasites (helminths and Cryptosporidium). The rationale for introducing additional species to Australia is considered.     

Brood ball size but not egg size correlates with maternal size in a dung beetle,Onthophagus atripennis

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A comparison of dung beetle (Coleoptera: Scarabaeidae) attraction to native and exotic mammal dung

Although the preference of dung beetles (Coleoptera: Scarabaeidae) for specific types and conditions of dung has been given substantial attention, little has been done to investigate the potential effects of exotic mammal introduction for game farms or rewilding projects. We used pitfall traps baited with various native and exotic herbivore, carnivore, and omnivore dung to evaluate dung beetle preference in the Great Plains of North America. Additionally, we analyzed of the nutrient quality of each dung type. In total, 9,089 dung beetles from 15 species were captured in 2 yr of sampling. We found significant differences (P < 0.05) in mean dung beetle capture among omnivore, herbivore, and carnivore dung, as well as differences in individual species preference for dung type. Omnivore dung was the most attractive with chimpanzee and human dung having the highest mean capture (291.1 ± 27.6 and 287.5 ± 28.5 respectively). Carrion also was highly attractive with a mean of 231.9 ± 20.6 beetles per trap (N = 8). Our results suggest definitive local preference of carrion in Phanaeus vindex Macleay and Onthophagus hecate (Panzer), while the congener, O. pennsylvanicus (Harold), was rarely captured in carrion and highly preferred omnivore dung. Preference for a specific bait type does not appear to be correlated with dung quality, mammalian diet, or origin of mammal. Results suggest niche segregation by dung type among dung beetle species.     

Ivermectin residues in cattle dung impair insect-mediated dung removal but not organic matter decomposition

1. Veterinary parasiticide residues in livestock dung have been repeatedly shown to negatively affect the abundance and diversity of dung-associated insects. While these losses are concerning from a conservation perspective, they can also translate to impairment of ecosystem functions in agricultural landscapes (e.g. nutrient cycling, primary productivity and greenhouse gas mitigation). 2. Most research focusing on decomposition-related ecosystem functioning has focused on dung beetles (Coleoptera: Scarabaeoidea) while other insects, particularly flies, have received comparatively less attention. 3. Using mesocosms, this study manipulated the insect groups colonising cow dung (beetles only, flies only, beetles and flies together, and an insect-free control). Half of the insects were exposed to 1 mg kg⁻¹ ivermectin in dung, while the other half were exposed to ivermectin-free dung. Dung decomposition (mass of organic matter lost) and dung removal (change in the dry mass of the dung pat attributed to both dung decomposition and burial) were measured. 4. Comparison of beetles and flies in ivermectin-free dung showed that beetles removed nearly twice as much dung as did flies. Comparison of dung removal across all treatments showed that ivermectin residues significantly reduced dung removal provided by beetles by 47% and dung removal provided by beetles and flies together by 32%. 5. Organic matter decomposition was not significantly affected by insect colonists or by the presence of ivermectin, indicating that organic matter decomposition can occur independently of insect activity and chemical perturbations.