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.

     

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     

Patterns of daily flight activity in onitine dung beetles (Scarabaeinae: Onitini)

Different species of African dung beetles emerge from the soil at characteristic times of the day to fly and colonize the freshly-deposited dung of mammalian herbivores. Onitine dung beetles in their natural habitat displayed one of five distinctive daily flight behaviours: dusk crepuscular (Onitis alexis Klug, O. caffer Boheman, O. fulgidus Klug, O. tortuosus Houston, O. vanderkelleni Lansberge, O. westermanni Lansberge); dusk/dawn crepuscular (O. pecuarius Lansberge and O. viridulus Boheman); dusk/dawn crepuscular and nocturnal (O. aygulus (Fabricius), O. mendax Gillet, O. uncinatus Klug); late afternoon-dusk and dawn-early morning [Heteronitis castelnaui (Harold)]; or diurnal flight activity [O. belial (Fabricius), O. ion (Olivier)]. These diagnostic daily flight behaviours span a light intensity range of over 6 orders of magnitude and have been retained in selected species introduced into Australia. Ambient light intensity appears to be the primary determinant of the daily flight period in onitine dung beetles. Because the dung of mobile herbivores is rapidly exploited by onitine species for feeding and breeding purposes, different flight behaviours result in a spatial and temporal partitioning of species in the local dung beetle community. The timing of flight may contribute to, or lead to avoidance of, competition between species which may ultimately affect colonization success. Many onitines show a strong preference for dung of specific herbivores, which may further reduce interspecific competition. All crepuscular-nocturnal species examined raised their thoracic temperatures endothermically to between 35°C and 40°C before the onset of flight. In O. aygulus the thoracic temperature excess was as large as 19.3°C. The thermal threshold below which the frequency of flight onsets drops off rapidly is about 12°C for O. aygulus and 17°C for O. alexis and O. pecuarius. Radiant loss of body heat during cool nights and dawns may explain why smaller species (<0.4 g body weight), in particular, are adapted behaviourally so that they fly only during the day or early dusk.     

Diversity of dung and carrion beetles in a disturbed Mexican tropical montane cloud forest and on shade coffee plantations

This paper analyzes the diversity of dung and carrion beetles (Scarabaeinae and Silphidae) in four human-induced habitats of a disturbed tropical montane cloud forest: polyspecific shade coffee plantations, monospecific shade coffee plantations, tropical montane cloud forest fragments, and clear cuts. The four habitats had similar richness, species composition, and assemblage structure of dung and carrion beetles. Differences were found in abundance and biomass levels for the four dominant species in the landscape. Dung beetles were more abundant than carrion beetles, but the biomass was higher for the latter. Carrion beetles were seasonal, while dung beetles were clearly not. When forest fragments and shade coffee plantations were compared to other similar habitats in the region, the same general pattern was observed. However, forests with high disturbance and monospecific shade coffee plantations had lower species richness than forests with low and medium disturbance and polyspecific shade coffee plantations. Thus shade coffee plantations maintain connectivity between patches of cloud forest in a landscape that is strongly affected by human activities. Protecting landscape diversity appears to ensure high species richness.     

Using dung beetles to evaluate the effects of urbanization on Atlantic Forest biodiversity

We used dung beetles to evaluate the impact of urbanization on insect biodiversity in three Atlantic Forest fragments in Londrina, Paraná, Brazil. This study provides the first empirical evidence of the impact of urbanization on richness, abundance, composition and guild structure of dung beetle communities from the Brazilian Atlantic Forest. We evaluated the community aspects (abundance, richness, composition and food guilds) of dung beetles in fragments with different degrees of immersion in the urban matrix using pitfall traps with four alternative baits (rotten meat, rotten fish, pig dung and decaying banana). A total of 1 719 individuals were collected, belonging to 29 species from 11 genera and six Scarabaeinae tribes. The most urban‐immersed fragment showed a higher species dominance and the beetle community captured on dung presented the greatest evenness. The beetle communities were distinct with respect to the fragments and feeding habits. Except for the dung beetle assemblage in the most urbanized forest fragment, all others exhibited contrasting differences in species composition attracted to each bait type. Our results clearly show that the degree of urbanization affects Atlantic Forest dung beetle communities and that the preservation of forest fragments inside the cities, even small ones, can provide refuges for Scarabaeinae.     

Laboratory investigations of Aphodius beetles (Scarabaeidae) as biological control agents of the Cattle Lungworm,Dictyocaulus viviparus (Nematoda: Dictyocaulidae)

The effect of Aphodius fossor, A. luridus and A. foetidus adults on Dictyocaulus viviparus larvae in cattle faeces was investigated. In chambers containing A. fossor and A. luridus, a significant decrease (at 5% level) in the median number of D. viviparus larvae occurred after 90 h at a density of one beetle per g of faeces. The possibility of using dung beetles as biocontrol agents for D. viviparus is discussed.     

Getting the most out of waste: how dung beetles boost the nitrogen content in their food

The life cycle of almost all dung beetles revolve around mammalian dung, the feed on dung, look for mating partners on dung and lay eggs in the dung. We know they feed on dung, but we still do not understand how exactly they filter‐feed on the dung and which particles size range they are ingesting. The aim on this study was to investigate the filter feeding by particle selection by adult dung beetles using Scarabaeus goryi and how that improves the nutrient quality of the ingested particles. We compared the particle sizes and nutrient content of the dung with the ingested material in the foregut, hindgut and the faeces of the dung beetle. Adult dung beetles do select smaller dung particles when feeding, we found the maximum particle size for the ingested particle to be around 1400 μm. The average particle size increased through the gut length. Dung beetles also selected particles with higher nitrogen content when feeding, the nitrogen content increased from about 1.5% in the dung to just over 5% in the foregut which then decreased to the level of the unprocessed dung in the dung beetle faeces. Carbon content did not increase from the unprocessed dung to the foregut but decreased through the gut length. Feeding by particle size selection by dung beetles helps in selecting particles with higher nitrogen content to compensate for the low levels found in dung.     

Global dung webs: high trophic generalism of dung beetles along the latitudinal diversity gradient

At the global scale, species diversity is known to strongly increase towards the equator for most taxa. According to theory, a higher resource specificity of consumers facilitates the coexistence of a larger number of species and has been suggested as an explanation for the latitudinal diversity gradient. However, only few studies support the predicted increase in specialisation or even showed opposite results. Surprisingly, analyses for detritivores are still missing. Therefore, we performed an analysis on the degree of trophic specialisation of dung beetles. We summarised 45 studies, covering the resource preferences of a total of 994503 individuals, to calculate the dung specificity in each study region. Our results highlighted a significant (4.3‐fold) increase in the diversity of beetles attracted to vertebrate dung towards the equator. However, their resource specificity was low, unrelated to diversity and revealed a highly generalistic use of dung resources that remained similar along the latitudinal gradient.     

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.     

Dung beetles in a Central Amazonian rainforest and their ecological role as secondary seed dispersers

Abstract 1. The role of several factors that affect the composition of the dung beetle assemblages in an Amazonian rainforest was quantified, together with the effect of these factors on the role that dung beetles play as secondary seed dispersers. 2. A total of 61 dung beetle species was captured during 3360 h of trapping. During nocturnal trapping periods, more dung beetles, of larger mean size, and more species were captured per trap than during diurnal trapping periods. 3. During the rainy season, more dung beetle species were captured per trap than during the dry season, but the number of individuals and their mean size did not vary between seasons. 4. Bait size had a significant effect on the mean number of beetles and mean number of species but not on mean beetle size. As bait size increased from 5, 10, 25, to 50 g, more beetles and more species were captured per trap. 5. Between 6 and 73% of plastic beads, used as seed mimics, were buried by dung beetles at depths that ranged from 0.5 to 7 cm. Both the proportion of beads buried and burial depth decreased with increasing bead size, and increased with increasing amounts of dung surrounding each bead (5, 10, and 25 g). 6. The proportion of buried seeds for three species varying in size between 5 and 27 mm, increased with increasing dung beetle size, using beetles of seven sizes, varying between 10 and 25 mm. 7. Seeds surrounded by dung were buried more often and more deeply when placed on the forest floor during the late afternoon than when placed during the early morning. Seeds were also buried more often when placed on the forest floor during the rainy season than when placed during the dry season, but season had no effect on burial depth. 8. Forests in Central Amazonia hold a rich dung beetle community that plays an active role in secondary seed dispersal, and consequently in plant regeneration. The interaction between seeds and beetles is complex because it is affected by many factors.     

Environmental gradients and community attributes underlying biodiversity patterns of semi-arid Mediterranean grasslands

Protection and/or establishment of forest plantation have been used as a management strategy to conserve and stop the deterioration of semi-arid Mediterranean grasslands ecosystems, producing a mosaic of vegetation types. This study was intended to investigate the changes in grassland community in response to protection and forest tree plantation practice as well as to explore the underlying environmental gradients responsible for the observed differences or similarities among these vegetation types. Two multivariate analysis methods including discriminate analysis and non-metric multi-dimensional scaling were used to quantify changes in community composition and attributes following different management practices (free grazing, protection with open grassland, sparse and dense forest tree plantations). This was investigated using species frequency, species abundance, or habitat characteristics. The study results showed that habitat types differed significantly between each other and were significantly separated using multivariate approaches. Discrimination based on habitat characteristics and species composition indicated that protection (or grazing) and light (or shade) explained more than 90% of the observed variability in community changes in response to the protection and forest tree plantation. Also, results indicated that shade effect can be attributed to tree canopy cover and/or litter accumulation on the ground. It could be hypothesized that protection from grazing and afforestation resulted in complex environmental gradients of which shade, litter accumulation as well as protection from grazing disturbance are major constituents. A careful manipulation of protection and afforestation can be used to create a spatially different environmental gradients leading to greater habitat diversity as well as a greater species diversity, and better conservation means of grassland in semi-arid areas.     

The effects of rainforest canopy loss on arboreal dung beetles in Borneo: implications for the measurement of biodiversity in derived tropical ecosystems

We examine the effect of selective timber extraction, and corresponding forest canopy loss, on arboreal dung beetles in the tropical rainforests of Sabah, Malaysian Borneo. Changes in vertical distribution of beetles are measured through differences in the abundance of beetles in ground-based pitfall traps in primary, logged and plantation forest. Previous research has demonstrated that arboreal dung beetles are not generally collected in pitfall traps in primary forest, but are present in large numbers above the ground in canopy vegetation: the presence of arboreal beetles in pitfalls in areas of reduced or modified canopy cover may therefore reflect a response to the absence or modification of their usual habitat, and the proliferation of these beetles nearer to the ground. In this paper, statistically significant differences are found in the abundance of beetles in ground pitfall traps from logged forest compared to primary forest. Results show that virtually no arboreal dung beetles are recorded in primary forest traps, with an increased abundance of arboreal dung beetles in traps from logged and plantation forest, with 1.72% of the total number of arboreal beetles recorded from primary forest, 22.32% from logged forest, and 75.96% from plantation forest. The presence of arboreal dung beetles in plantations demonstrates that arboreal dung beetles can survive outside their normal habitat, and we relate these observations to adaptations to upper rainforest canopy conditions, and proliferation of these microclimatic conditions in man-made habitats. Results are also discussed in terms of their relevance to the measurement of species richness and diversity in logged and other derived ecosystems, where mixing of the ground-based and arboreal faunas occurs.     

Are ball-rolling (Scarabaeini,Gymnopleurini, Sisyphini) and tunnelling scarabaeine dung beetles equally choosy about the size of ingested dung particles?

Abstract.  1. The maximum size of ingested particles was determined in 11 species of ball-rolling, adult dung beetle (Scarabaeidae: Scarabaeinae) by mixing small latex or glass balls of known diameter into their food. The tribes Scarabaeini, Gymnopleurini, and Sisyphini (four, four, and three species respectively) were represented, with mean body sizes ranging from 0.33 to 4.0 g fresh weight.
2. Only particles with maximum diameters of 4–85 µm were ingested. Hence rollers, like other known beetles feeding on fresh dung, filter out larger, indigestible plant fragments and confine ingestion to small particles of higher nutritional value.
3. The maximum diameter of ingested particles increased significantly with body weight, whereas taxon (tribe) had no additional effect. Because big rollers accept larger particles than do tunnellers (which make dung stores for feeding and breeding in the soil immediately below the pat) of similar weight, the slope of the diameter-against-weight regression for rollers was significantly higher than that found earlier for tunnellers.
4. An explanation could be that a typical food ball made by a roller is considerably smaller than the amount of dung available to a feeding tunneller of the same size. If the roller were as choosy about particle size as the tunneller, it might not get enough food. This applies to large rollers in particular because their food balls contain a higher proportion of coarse fibres than those made by small species.     

Age and size of European saltmarshes and the population genetic consequences for ground beetles

Enzyme and dispersal polymorphisms of the saltmarsh carabid beetles Pogonus chalceus and Dicheirotrichus gustavii were studied in European populations varying in size and in isolation in space and time. D. gustavii, a constantly fullwinged species, has a larger genetic diversity and a smaller genetic differentiation between populations than the wing-polymorphic P. chalceus. Clear relationships between population or site characteristics and genetic structure were not observed, except for the special position taken by some small populations in both species. The dispersal power of P. chalceus in small populations is larger than in large populations, suggesting that these populations are unstable and/or young. Small populations, however, do not always show a lower genetic diversity than large populations, as would be expected from genetic drift. Dispersal power in P. chalceus declines with increasing age of the saltmarsh, probably due to continuous emigration of winged individuals. Age and size of saltmarshes, although difficult to study independently, both appear to be important in determining the genetic structure of saltmarsh beetles. Maximum diversity in both parameters is therefore recommended as an optimal nature conservation strategy. Received: 22 September 1997 / Accepted: 29 December 1997     

Sub-lethal effects of moxidectin on the Neotropical dung beetle Onthophagus landolti Harold (Coleoptera: Scarabaeinae)

Macrocyclic lactones can have adverse effects in dung beetles exposed to manure containing them. An assessment of the survival and fertility of adult Onthophagus landolti Harold fed with manure from cows treated with moxidectin was performed, as well as the emergence rate of the imagoes from the masses. Three cows (Bos indicus x B. taurus) were subcutaneously injected with 1% moxidectin (0.2 mg kg^?1 b.w.) and another three were injected with 10% moxidectin (1.0 mg/kg^?1 b.w.). Manure was collected from these animals one day prior to moxidectin administration, five days post-treatment in the 1% and 10% treatments, and fourteen days in the 10% treatment. Four bioassays were done: a control using manure without moxidectin; 1% moxidectin at five days post-treatment; 10% moxidectin at five days post-treatment; and 10% moxidectin at fourteen days post-treatment. In each replicate, for each pair of adult O. landolti was daily fed with 30 g manure according to the treatments. No lethal effects were observed in any of the four treatments. Sub-lethal effects (P < .05) were present in the 10% moxidectin treatments at five and fourteen days post-treatment. Fecundity was reduced by 78.2% at five days and 54.9% at fourteen days, and imago emergence was negatively affected at both times. Current moxidectin application methods may have negative effects on the environmental services provided by dung beetles, and therefore need to be modified to minimize any impacts they might have on these vital members of tropical livestock systems.     

Effects of herd management and the use of ivermectin on dung arthropod communities in grasslands

Agroecosystems represent a large geographical footprint in most terrestrial landscapes, and management decisions within these systems affect their function in species conservation. We evaluated the effects that rangeland management systems (based on stocking density, rotation frequency, and the number of avermectin applications) have on conserving the dung arthropod community in the Northern Great Plains of North America. Comprehensive bioinventories of arthropods were collected from 16 rangelands using core samples of dung pats. Ivermectin was quantified in pats from each ranch using enzyme-linked immunosorbant assay (ELISA). Arthropods in dung were abundant (116,244 specimens) and diverse in eastern South Dakota (172 morphospecies). Rangelands managed with more regenerative practices (frequent rotation at high stocking densities and lack of ivermectin applications) had greater species richness, diversity, predator species abundance, and dung beetle abundance than more conventionally managed rangelands. Ivermectin quantity in cattle pats was negatively correlated with dung beetle abundance and diversity. This work shows that herd management (specifically high-intensity, frequent rotational grazing and eliminating prophylactic ivermectin use) that aims to mimic intensive grazing of large migrating herds of ruminants can foster dung arthropod community structure, a key trait correlated with nutrient cycling, pest suppression, and productivity of cattle-grazed rangelands.     

Sampling the rarest: threatened beetles in boreal forest biodiversity inventories

Presence or absence of threatened species in samples is information that is widely used in designing and implementing conservation actions. We explored the effectiveness of beetle (Coleoptera) inventories and contribution of different sampling methods in revealing occurrences of threatened and near threatened species in boreal forests. The number of species caught using traps in a particular area proved to be a useful indicator of the representativeness of data, the relationship between total number of species and the number of threatened and near threatened species being almost exponential. Samples containing less than 200 trapped species (or 2000 individuals) are almost useless in surveying threatened and near threatened species. The probability of finding such species increases considerably when the number of trapped species exceeds 400. Window traps attached directly on the trunks of dead trees proved to be the most efficient sampling method in trapping threatened beetles, whereas many other standard methods gave relatively poor results. We suggest that the best alternative in surveying threatened species in boreal forests is a combination of intensive direct searching and trunk window traps. Finding threatened beetles with rigorous probability requires very large sample sizes, even if the most effective sampling methods are used. For example, ranking 10 boreal forest areas to be protected according to the occurrence of threatened species with some reliability may require trapping of over 100000 beetle individuals. Collecting and identifying these large samples routinely in conservation actions is not feasible, which means that shortcuts (indicators etc.) are necessary. However, a lot of good-quality inventories with appropriate sampling efforts are needed before these shortcuts can be identified and elaborated. Such inventories are also crucial for the improvement of the classification of threatened species and full assessment on how past forest management has eventually affected the biota.     

Field effects of faecal residues from ivermectin slow‐release boluses on the attractiveness of cattle dung to dung beetles

A 2‐year study was performed in two sites in southern France to assess the effect of ivermectin residues on the attractiveness of cattle dung to colonizing insects. Insect captures were compared between pitfall traps baited with dung from untreated cattle and dung from cattle that had been treated with a slow‐release (SR) bolus of ivermectin. Cattle dung was collected at different times after treatment (4, 14, 42, 70 and 98 days). Excretion showed a plateau, with levels ranging between 0.688 µg and 1.123 µg ivermectin per gram of wet dung. Faecal residues affected insect captures at both sites. Effects were independent of the time dung was collected after treatment, except for one result subsequent to a severe drought during the baiting period. Ivermectin‐contaminated dung showed a significant attractive effect, with increased captures regardless of the guild to which beetles belonged. This study demonstrates the attractiveness of ivermectin residues over a long period after the treatment of animals. It draws attention to the danger of widespread use of this endectocide‐based SR bolus, which is attributable to the preferential attraction of insects to treated dung, which potentially puts at risk the survival of their offspring.     

Evidence of developmental niche construction in dung beetles: effects on growth,scaling and reproductive success

Niche construction occurs when organisms modify their environments and alter selective conditions through their physiology and behaviours. Such modifications can bias phenotypic variation and enhance organism–environment fit. Yet few studies exist that experimentally assess the degree to which environmental modifications shape developmental and fitness outcomes, how their influences may differ among species and identify the underlying proximate mechanisms. Here, we experimentally eliminate environmental modifications from the developmental environment of Onthophagus dung beetles. We show that these modifications (1) differentially influence growth among species, (2) consistently shape scaling relationships in fitness‐related traits, (3) are necessary for the maintenance of sexual dimorphism, (4) influence reproductive success among females of at least one species and (5) implicate larval cultivation of an external rumen as a possible mechanism for environmental modification. Our results present evidence that Onthophagus larvae engage in niche construction, and that this is a fundamental component of beetle development and fitness.