Spatial distribution, resource utilisation and intraspecific competition in the dung beetle Aphodius ater

Competitive interactions in northern temperate dung beetles are poorly understood. This investigation therefore comprises a series of field and experimental work on a dung beetle species common in northern Europe, Aphodius ater, with special focus on intraspecific competitive interactions. The between-pat distribution of adult A. ater in relation to the age of sheep dung pats was studied in the field. The distribution of both sexes was contagious in the fresh pats but became more regular with increasing pat age. The successional occurrence of males and females did not differ, but immature females tended to occur in fresh pats while mature females were mainly found in older pats. With increasing age of pats, the egg load of females also increased. Egg-laying behaviour of the beetles was studied in laboratory experiments. The mean number of eggs laid per female per dung pat decreased with increasing beetle density. Thus, density-dependent processes seem to regulate resource utilisation with regard to breeding behaviour, resulting in equal exploitation of the available pats. Survival and weight of recently hatched beetles decreased with increasing initial density of eggs. Hence, in A. ater, competition between larvae for food within pats does occur. Received: 4 February 1998 / Accepted: 20 April 1998     

Use of food and space by tunneler dung beetles (Coleoptera; Scarabaeinae) during reproduction

At two temperate pasturelands in northern Mexico, we explored possible competition for food and space under pats during the simultaneous nesting periods of the univoltine species Dichotomius colonicus (Say), Phanaeus quadridens (Say), and Copris sierrensis Matthews. To simulate unlimited resources, 50 5-kg cow dung pats were placed at regular distance intervals in each pastureland. After building trenches around the pats, the number and depth of each nest, as well as larval development status, were documented once for a period of 1-8 mo. Analyses of variance and association tests were used to make a between-site comparison of dung pat occupation, nests occupied per species, nests per dung pat, and nest depth below each pat. The proportion of pats occupied by each species differed significantly between sites. C. sierrensis colonizing most pats at one site and D. colonicus at the other. There were no differences between sites in the frequency of pats occupied by more than one species. The association test and Ochiai index showed that each species colonized dung pats independently. The results suggest that pat occupation depended on their location by beetles and the relative abundance of each species. The species tended to dig nests at different depths, possibly reducing interspecific competition for space. It can therefore be concluded that, when food resources seem to be unlimited, they are shared following a "lottery dynamic" model if there is spatial differentiation among species.     

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.     

A review of competition in north temperate dung beetle communities

Abstract. 1. Studies of north temperate dung beetle communities frequently invoke competition as an influential ecological process. In this review, the evidence for competition in north temperate dung beetle communities is evaluated and the role of competition as a factor affecting community structure is assessed.
2. Resource limitation and the evidence for interspecific competition are assessed by collating the available experimental and observational evidence for both the adult and larval stages of the dung beetle life cycle. The role of competition as a structuring force in dung beetle communities is discussed under the following headings: niche dynamics, migration to and from individual pats, the aggregation model of co-existence, and metapopulations.
3. Some of the main conclusions are that competition for space is much more likely to occur than competition for food; the effects of competition on community structure are poorly understood; several of the influential studies of competition in north temperate dung beetle communities need to be evaluated carefully. The differences in ecology between tropical and temperate dung beetle communities are clarified.
4. As priorities for future research, resource utilisation and competition should be researched experimentally: density-dependent relationships should be investigated, particularly for the larval stages, as should competitive interactions with other dung fauna. If such experimental approaches establish convincingly the occurrence of competition, then the extent of competition in the field and under real world conditions needs to be established. A functional group classification of dung beetles and other dung fauna is described, which may improve the generality of interpretation from individual, site-specific results.     

Effects of ivermectin and doramectin faecal residues on the invertebrate colonization of cattle dung

Abstract: The effects of avermectin [ivermectin (IVM) and doramectin (DRM)] faecal residues on dung colonization and degradation by invertebrates were evaluated during late spring in the east of La Pampa province, Argentina. The study was conducted after collection of faecal material from animals (10 steers per group) allocated to the following groups: untreated control group (CG) and groups treated subcutaneously (200 μg/kg) with either DRM (DG) or a long‐acting formulation of IVM (IG). Fifty pats (550 g each) per group were collected, prepared and deposited on the field on days 3, 7, 16 and 29 post‐treatment (pt). Eight pats per group were recovered after 7, 14, 21, 42, 100 and 180 days post‐deposition (pd) on the field. The weight, percentage of dry matter, number of arthropods and nematodes from faeces were determined. The faecal concentrations of IVM and DRM were measured by high performance liquid chromatography (HPLC) throughout the trial period to correlate the pattern of drug degradation in dung with pd time. The total number of arthropods in dungs from CG was higher (P < 0.05) than those counted between days 3 and 29 pt in IG and DG. A decrease in the number of Coleoptera larvae (P < 0.05) between days 21 and 42 days pd was observed in both treated groups. Diptera larvae counts in CG pats were significantly higher (P < 0.05) than those obtained in treated groups in the 7‐ and 14‐day‐old pats. A lower number (P < 0.05) of Collembola, compared with pats from CG, was recovered from IG and DG pats deposited at days 3 and 7 pt and exposed from day 42. The counts of Acari in pats from treated animals were lower (P < 0.05) than those observed in CG pats at 3, 8 and 16 days pt. There were no differences neither in adult Scarabaeidae recovered nor in the proportions of dung buried and destroyed by great dung beetles. Dung specific nematodes were reduced (P < 0.05) in IG and DG pats from 3 and 7 days pt compared with those of CG pats. The comparative results shown here demonstrate that the negative effects of both IVM and DRM on dung colonization are similar. The pattern of drug degradation in the environment was very slow. High residual concentrations of both active parent compounds were recovered in dungs exposed in the field for up to 180 days pd. Concentrations as high as 13 ng/g (IVM) and 101 ng/g (DRM) were measured in faeces obtained from pats deposited on day 27 pt and exposed to the environment during 180 days. The results show a decrease in invertebrate colonization of dung recovered from IVM‐ and DRM‐treated cattle, which is in agreement with the large drug residual concentrations measured in faeces.     

Reproductive properties of two sympatric dung beetles,Aphodius haroldianus andA. Elegans (Coleoptera: Scarabaeidae)

1. Reproductive properties of two sympatric dung beetles, Aphodius haroldianus and A. elegans, were studied at a pasture in the central Japan.
2. Overwintered adults of A. haroldianus came flying to dung pats from May to early August and bred from June to July. A. elegans overwintered as larvae, new adults came to dung pats from mid May to mid June. After aestivation, they reproduced from October to November.
3. A. haroldianus was a species with low fecundity and large food reserve for larvae. Female of this species had short duration of residence and laid fewer number of eggs in one dung pat. On the other hand, A. elegans was a species with high fecundity and small food reserve for larvae. Female of this species had long duration of residence and laid larger number of eggs in one dung pat.
4. The daily egg production in A. haroldianus reached a peak (0.40 eggs/female/day) at middle of oviposition period, but, that in A. elegans reached a peak (6.49 eggs/female/ day) at the beginning of oviposition period. The mortality of A. haroldianus female occurred after the daily egg production reached a peak and then rapidly increased, whereas that of A. elegans began soon after the beginning of oviposition and then gradually increased.

     

Patch size and colonisation patterns: an experimental analysis using north temperate coprophagous dung beetles

The relationship between dung pad size and both adult colonisation and larval development was investigated in an assemblage of north temperate dung beetles ( Geotrupes. Aphodius and Sphaeridium ) using both dung pads and baited pitfall traps. Wet weight of 22-day-old natural dung pads was found to vary widely in the field (< 100 g – >1000 g). Across all sampling dates in field experiments, dung pad size had a significant influence on dung beetle biomass sampled from pads. Closer examination of experimental dung pads on the second day after deposition, when beetle biomass was at a maximum, revealed not only a general positive relationship between pad size and dung beetle biomass but, more importantly, a positive relationship between dung pad size and dung beetle density (dung beetle biomass per unit dung volume). There was a strong trend for Aphodius species richness to increase, and maintain higher values for longer periods of time, in larger pads. Although dung pad and pitfall trap samples could differ in the actual numbers of beetles captured, the relationship between different dung sizes and dung beetle biomass was similar, indicating that the phenomenon is largely related to immigration processes. Pat residence times oi A. rufipes in the laboratory were significantly positively correlated with dung pad size. In two field experiments, positive correlations were found between dung pad size and numbers of larvae in pads of different sizes and in one of these experiments, larval densities (numbers per unit dung volume) were significantly and positively correlated with dung pad size. In one experiment, Aphodius larvae in the early stages of development were found to preferentially occupy the basal area of dung pads. We discuss the implications of our findings in the context of resource utilisation by north temperate dung beetles.     

Reduced splash dispersal of bovine parasitic nematodes from cow pats by the dung beetle Diastellopalpus quinquedens.

From a thoroughly mixed portion of cattle feces with Cooperia sp. eggs, 1-kg artificial pats were placed in 6 buckets containing 6 kg of soil each. Ten dung beetles, Diastellopalpus quinquedens, were added to each of 3 buckets. The remaining 3 buckets served as controls without beetles. When infective parasite larvae (L3) had developed in the cow pats indoors, the following procedure was followed. During occasions of rainfall each bucket was placed outdoors in the center of a wider and higher container. When the rain stopped all buckets were brought indoors, and infective larvae spread by splash droplets during the rain were collected in the containers and counted. After 33 days, the remaining dung on the soil surface in buckets with dung beetles constituted only 38% of that in the controls. Moreover, the number of L3 in feces left on the soil surface in the buckets with beetles was reduced by 88%, presumably due to beetle activity. This may explain the 70-90% reduction in splash dispersal of L3 of Cooperia sp. from cow pats attacked by beetles. The dung-burying activity of the beetles did not result in increased numbers of L3 in the soil under the cow pats, suggesting that many of the parasites in buried feces were destroyed.     

Dung feeding in adult scarabaeines (tunnellers and endocoprids): even large dung beetles eat small particles

Abstract 1. The maximum size of ingested particles was determined in 15 species of adult dung beetle (Scarabaeidae: Scarabaeinae) by mixing small latex or glass balls of known diameter into the dung used as food. Twelve species (tribes Coprini, Onitini, Oniticellini, and Onthophagini) were tunnellers (making dung stores for feeding and breeding in the soil below the pat) and three species (tribe Oniticellini) were endocoprids (feeding and breeding in the dung pat itself).
2. The test species, covering a wide range of body size (fresh weights 0.05–7.4 g), ingested minute particles only (maximum diameter 8–50 µm), and there was a statistically significant but numerically small increase in particle size with body weight.
3. When the effect of body size was taken into account, taxon (tribe), ecological group (tunneller/endocoprid), and dung preference (coarse/fine) had no significant effect on the size of ingested particles.
4. Tests using two tunnelling species did not indicate that beetles use their mandibles to grind dung particles prior to ingestion.
5. The results suggest essentially the same feeding mechanism in all adult tunnelling or endocoprid scarabaeines that eat fresh dung. Larger, indigestible plant fragments are avoided by filtration, and ingestion is confined to very small particles of higher nutritional value.     

Heterotrophic succession within dung-inhabiting beetle communities in northern Spain

Successional patterns of beetles inhabiting dung pats were examined during May and July 1993 in a mountain area in northern Spain (Picos de Europa). Beetles belonging to six families were caught during the course of succession (30 d). Coprophagous beetles were more abundant in dung pats than predatory beetles (89 and 11 %, respectively). A trophic sequence was observed in relation to age of the dung, coprophagous beetles occurring earlier in the dung than predatory beetles. The pattern was observed on two occasions during the season, though succession proceeded somewhat faster in July than in May. These results suggest that food availability and microclimatic conditions in dung pats appear to determine the successional occurrence of beetle taxa. On the other hand, coprophagous species (Aphodius) were poorly segregated along the successional axis. Null models failed to support the hypothesis that successional overlap and differences in successional mean occurrence between species could be the result of competition. Successional patterns at the specific level probably reflect differences in behaviour, such as pat location, feeding, mating, egg-laying and larva requirements, rather than competitive replacement.     

The brown-world role of insectivores: Frogs reduce plant growth by suppressing detritivores in an alpine meadow

Predators of plant-suppressing herbivores have long been known to indirectly enhance plant biomass, while more recent work has revealed that predators of plant-facilitating detritivores can have the opposite effect on plant biomass. Generalist predators, such as frogs that typically facilitate plant growth in green food webs, may potentially negatively affect plant growth by consuming prey from brown food webs. In a marshy Tibetan alpine meadow, we tested the hypothesis that the locally abundant frog Rana kukunoris could negatively affect plant growth through suppression of dung-decomposing detritivores that promote plant growth by enhancing nutrient recycling. We conducted a factorial experiment (presence/absence of predators × presence/absence of dung) using replicate field enclosures over a growing season. Where dung was present, frogs significantly reduced the number of dung-feeding beetles and dung-feeding flies (including fly eggs and maggots) per dung pat, thereby decreasing dung mass loss and, indirectly, aboveground plant biomass (by 22%) surrounding dung pats. Where dung was absent, frogs did not affect plant biomass. Moreover, the number of dung beetles was positively associated with dung mass loss and soil soluble N concentration (but not total N concentrations), which in turn positively correlated with aboveground plant biomass. These results indicate that a generalist predator species standing in green food webs may play a contrasting role in brown food webs and indicate that a more nuanced appreciation of the functional role of predators in tri-trophic systems is required to accurately predict their cascading effects. Future studies must assess the relative strength of cascading effects mediated through brown and green channels in order to assess the net cascading effects of generalist predators in natural food webs.     

Aggregated distribution of resources creates competition refuges for rainforest dung beetles

This study examines the effects of resource distribution on colonisation, intra- and interspecific aggregation, and the occurrence of low-density, competition refuges for tropical dung beetles. In field experiments from central Peru, using dung pats (resource patches) of different volumes, the numbers of interacting species and total beetle biomass at individual pats increased with increasing pat volume. In two of three separate experiments (including an experiment that also varied patch density), this represented a decrease in the biomass of beetles per unit volume (biomass-density) at larger patches. The numbers of interacting tunneller species and tunneller biomass-density were also related to the distance between pats (patch density) in one of two experiments with constant numbers of pats. Closely positioned pats had generally fewer interacting species and a lower biomass-density of beetles. For the most abundant Dichotomius species, interspecific associations increased as distances increased between dung pats. The numbers of interacting species and biomass-density declined linearly under the combined effects of increasing patch density and local patch abundance in 25 m² plots. In experimentally placed grids with large numbers of pats, colonisation of pats at the edge of the grids was generally higher than at the centre of the grids for tunnellers and Eurysternus spp. but not for ball rollers; however, at least tunnellers did not readjust to avoid patches with high densities of competitors. These results indicate that an aggregated distribution of dung and natural variability in patch size contribute to species coexistence by creating low-density refuges for weaker competitors.     

Effect of population density on reproduction of two sympatric dung beetle species,Aphodius haroldianus andA. Elegans (Coleoptera: Scarabaeidae)

The effect of population density on reproduction as well as the oviposition modes of two sympatric dung beetle species, Aphodius haroldianus (a low fecundity/high parental effort species) and A. elegans (a high fecundity/low parental effort species) was studied at a pasture in central Japan from 1982 to 1986.

1. The adult population density of A. haroldianus was high (>40 per dung pat). The density of A. elegans was low (<5 per dung pat). Oviposition of A. haroldianus was suppressed above the density of about 10 adults of conspecifics but not affected by the density of other species. The intra- and interspecific density effects on oviposition were not detected in A. elegans.
2. The number of A. haroldianus adults per pat decreased with dung age, while that of A. elegans increased until sixth day after deposition and then decreased. The number of eggs laid per pat was not different between 1-day-and 3-day-old pats for A. haroldianus. However, more eggs of A. elegans were found in 3-day-old pats than in 1-day-old ones.
3. In the both species, the amplitude of population fluctuations was not remarkable; the maximum/minimum ratio for five years being 2.3 for A. haroldianus and 2.9 for A. elegans. Different density dependent processes were suggested to function for the small fluctuation of population density between the two species, i.e. intraspecific density effect on oviposition for A. haroldianus and contest type competition for food among larvae of A. elegans.

     

Modelled impact of insecticide-contaminated dung on the abundance and distribution of dung fauna

Deterministic models assessed the effects that contaminated dung from insecticide-treated cattle had on populations of three hypothetical species of dung fauna that dispersed randomly and could double their numbers every 1-28 weeks at low density. Insecticide was allowed to kill 2-98 % of adults and prevent 16-100% of breeding in pats produced immediately after cattle treatment, with toxicity declining to < 1% in pats produced 2-23 days later. Treatment intervals were 10-40 days. The modelled impact of insecticide was affected little by approximately four-fold variations in: length and density dependence of the attractive life span of pats, frequency of pat occupation by immature adults, distribution of pat toxicity during treatment interval, and changes in dispersal rates due to age and population density. Of greater importance were variations in: pat toxicity, treatment interval, frequency of pat occupation by breeding adults, density dependence of recruitment and death, natural adversity and mortality in dormancy, general rate of dispersal, and the size and shape of the area with treated cattle. Overall, it seemed that wide variations in the impact of contamination will occur in the field, but in many situations the risk to dung fauna can be substantial, especially for slow breeding beetles, and muscoids contacting insecticide on cattle. Risk extends outside the treated areas, for a distance equal to several daily displacements of the insects. Untreated refuges for species survival should be compact blocks at least 25 daily displacements wide.     

Respiratory loss and bulk export of organic matter from cattle dung pats: a field study

Respiratory loss and bulk transport of organic matter from dung pats in a Danish pasture were measured by a chromic oxide marker technique. Measurements covered the period until a dung age of 7-8 weeks. On average, respiration and transport contributed 46% and 54% of organic matter disappearance in three spring experiments: in three autumn experiments the corresponding values were 25% and 75%. Thus, both processes are important. The rate of transport was highest at the bottom of the pat, whereas the highest respiratory rate occurred in the top and periphery. During the first 10–14 days transport predominated, probably owing to the activity of dung insects, whereas respiration was remarkably low. During the remaining period, transport was probably affected mainly by earthworms. Respiration seemed to be mostly microbial and to proceed during dung pat decay in essentially the same way as did cellulolytic activity. Earthworms did not significantly affect the respiration of pats.     

Dividing the pie: differential dung pat size utilization by sympatric Haematobia irritans and Musca autumnalis

Horn flies [Haematobia irritans (Diptera: Muscidae) (L.)] and face flies [Musca autumnalis (Diptera: Muscidae) De Geer] use the same larval resource, but their interactions are poorly studied. Dung pats (n = 350) were core sampled in the summers of 2012 and 2013 from irrigated pastures in Pomona, California, U.S.A. (34°03′N, 117°48′W) and held for face fly and horn fly emergence. Surface areas and estimated weights were recorded for each whole pat. Almost half (42.0%) of the pat cores yielded neither fly, 29.7% yielded horn flies only, 12.9% yielded face flies only and 15.4% yielded both flies. Of the fly‐positive pats, surface area and mass were larger for face fly‐occupied pats, whereas horn fly‐occupied pats were smaller. Pats shared by the two species were intermediate. Horn flies per positive core were unaffected by the absence/presence of face flies, but half as many face flies emerged when pats were co‐inhabited by horn flies. Face flies inhabited larger pats, which might better resist heating and drying, to which they are susceptible; horn flies inhabited a broad pat size range. Horn fly tolerance of lower dung moisture probably allows horn flies to colonize and survive in a wide range of pats in dry areas like southern California.     

The capability of the predacious fungus Arthrobotrys oligospora (Hyphomycetales) to reduce numbers of infective larvae of Ostertagia ostertagi (Trichostrongylidae) in cow pats and herbage during the grazing season in Denmark

Artificially prepared cow pats containing Ostertagia ostertagi eggs were deposited on two pasture plots in May, June and July 1986. Half of the cow pats, placed on one plot, were inoculated with 2000 conidia per gram faeces of the predacious fungus Arthrobotrys oligospora. On the other plot fungus-free control cow pats were placed at the same time. In the faeces generally fewer infective O. ostertagi larvae developed in the inoculated than in the control cow pats. On the herbage around the control cow pats deposited in May, June and July a maximum concentration of infective larvae was found at the same time on the 6th of August 1986. At that time the herbage larval infectivity around inoculated cow pats deposited in May, June and July was subject to a reduction of 48%, 89% and 46%, respectively, compared with fungus-free control cow pats. This experiment indicates that a concentration of 2000 A. oligospora conidia per gram faeces results in a significant lowering of the herbage larval infectivity during the grazing season in Denmark.     

Seasonal abundance and reproductive output of the dung flies Neomyia cornicina and N. viridescens (Diptera: Muscidae)

The seasonal abundance and reproductive output of two common, but little studied, dung-breeding flies, Neomyia cornicina and N. viridescens, were examined in artificial cow pats in pastures in southwest England in 2001 and 2004. In 2001, the numbers of both Neomyia species increased slowly over summer to show a sharp seasonal peak in late August and early September. There was no significant effect of mean temperature, mean relative humidity or dung water content on abundance or seasonally de-trended abundance. High levels of aggregation were seen between pats and, when present, greater numbers of N. cornicina emerged than N. viridescens. Neomyia cornicina was present in 13% of 240 artificial standardized pats put out in 2001, at a median of 19 adults per colonized pat; N. viridescens was present in 8% of artificial pats at a median of three adults per colonized pat. In 2004, N. cornicina emerged from 46% of the 94 artificial pats put out at a median of three adults per colonized pat, while N. viridescens emerged from only 12% of pats at a median of one adult per colonized pat. Flies were also collected in 2004, using sticky-traps and hand nets. Again, free-flying N. cornicina appeared to be more abundant in the field than N. viridescens; 162 N. cornicina were caught compared to 44 N. viridescens over the same sampling period. The size of each adult female was recorded and ovarian dissection was used to determine the numbers of eggs matured. Female N. viridescens were significantly larger than the N. cornicina and matured significantly higher numbers of eggs. Gravid N. viridescens matured a mean of 37.1 (+/-16.9) eggs, whereas gravid N. cornicina matured a mean of 28.8 (+/-13.2) eggs. The reasons why the larger, more fecund, N. viridescens adults are less abundant in the field or emerging from pats than N. cornicina are unknown. Further work is required to identify the nature and cause of the mortality experienced by the larvae of these species and the ecological differences and functional specialisation which allows co-existence to be maintained.     

Oxygen uptake in coprophilous beetles (Aphodius, Geotrupes, Sphaeridium) at low oxygen and high carbon dioxide concentrations

Abstract. The rate of O₂ consumption was measured in five coprophilous beetle species (common in Denmark) at O₂ concentrations from 1–21%. With the exception of the mainly soil-living Geotrupes spiniger (Marsham) (Geotrupidae), these beetles are probably exposed to severe hypoxia in fresh cattle pats. Aphodius fossor (Linnaeus), A. contaminatus (Herbst) (Aphodiidae) and Sphaeridium lunatum Fabricius (Hydrophilidae) maintained normal movements and a normal rate of 0₂ uptake (for at least 30 min) at only 1% O₂. There is no evidence, therefore, that the beetles switch to anaerobic metabolism under these conditions. This ability to regulate respiration, and hence to extract 0₂ at very low concentrations, is exceptional even among terrestrial arthropods living in soil or other potentially hypoxic substrates. In A. rufipes (Linnaeus), respiration declined at ambient concentrations below 2% O₂, and in G. spiniger the ability to regulate respiration seemed to fail at even higher concentrations. In four of the species (G. spiniger was not tested), about 11% CO₂ (the level in a dung pat at 2% O₂) did not affect the O₂ uptake at 2% O₂.     

Nest construction and larval behaviour of Onitis belial and Onitis ion (Coleoptera, Scarabaeidae)

Abstract. 1. Female beetles working alone or in cooperation with a male buried dung to make brood masses. O.belial brood masses were packed close together in clusters; each mass was constructed as a horizontal thick-walled tube of dung which was filled with a dung sausage containing two to six eggs. O.ion made vertical sausage-shaped brood masses with one to four eggs.
2. The larvae of both species were able to survive in artificial brood balls as well as in multi-egg brood masses because of their ability to repair larval chambers with their own excrement.
3. The multi-egg brood mass of Onitis has probably evolved from a simple one-egg brood mass. It does not resemble the underground dung mass from which brood balls are made by certain Coprini.