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.     

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.     

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.     

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.     

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.     

Impacts of traditional land uses on biodiversity outside conservation areas: effects on dung beetle communities of Vaalbos National Park

Grazing is one of the key processes in terrestrial ecosystems and this can be provided by both indigenous and domestic ungulates. However, a question remains whether or not traditional forms of land use such as the grazing of domestic animals support the maintenance of biodiversity. If it does not, then the second question becomes to what extent does grazing of domestic animals alter the systems and processes that support biodiversity? This study demonstrates that in attempting to answer this question, small organisms like dung beetles are ideal indicators that can be used to express significant differences between conserved (indigenous) and non‐conserved (domestic) land. As a general trend, studies that investigated these differences displayed differences through analysis of the diversity indices. This method has in most cases demonstrated a lack of contrast between conserved and non‐conserved land. In the existence of such uncertainty this study has demonstrated that in such cases, where the analysis of biodiversity indices fail to demonstrate significant differences a closer examination of actual species such as guilds and functional groups could confirm significant differences between conserved and non‐conserved land. These apparently conflicting findings reflect the need to consider the actual elements of biodiversity (e.g. species) when assessing conservation issues rather than just the statistical measures of biodiversity.     

Climatic and biogeographical associations of southern African dung beetles (Coleoptera: Scarabaeidae s. str.)

Climatic and biogeographical associations of southern African dung beetles (Scarabaeinae, Coprinae) were analysed from a collection amassed between 1971 and 1986. Endemism to Africa south of 15°S was much greater in southwesterly climates (winter rainfall, bimodal spring/autumn rainfall, arid late summer rainfall) than to the north‐east in the moist, mid‐summer rainfall region. Major biogeographical groups centred to the south‐west comprised predominantly southern African endemics, Western/Eastern Cape coast, Karoo, Karoo/Namib, northern Namibia and the south‐western Kalahari. Biogeographical groups centred on the south‐eastern highlands and the subtropical east coast (mid‐summer rainfall) also comprised predominantly southern African endemics. All other major groups centred to the north‐east in the mid‐summer rainfall region comprised predominantly species with widespread tropical biogeographical affiliations, pan‐southern Africa but centred in the east, pan‐mid‐summer rainfall region, eastern mid‐summer rainfall region, tropical east Zimbabwe/central Moçambique, subtropical/tropical game reserves (non‐ruminant dung specialists). There were cross‐climatic differences in taxonomic composition of the fauna. Within the winter rainfall region, percentage species composition of Scarabaeinae was greater whereas that of the coprine tribe, Onthophagini, was lower than within the other three climatic regions. Percentage species richness of most other tribes of Coprinae differed little between most climates but that of Scarabaeinae declined from west to east (Canthonini, Scarabaeini), east to west (Sisyphini) or to either side of the late summer rainfall region (Gymnopleurini).     

Importance of point sources on regional nitrous oxide fluxes in semi-arid steppe of Inner Mongolia, China

The aim of the present work was to estimate the contribution of different point and diffuse sources to the regional N₂O emission strength of steppe in the Xilin river catchment, Inner Mongolia, People’s Republic of China. Transect studies showed that the topographic effect on N₂O emissions from upland soils was negligible and that upland steppe is only a very weak net source of N₂O during the growing season (0.8 ± 0.4 μg N₂O–N m⁻² h⁻¹). Slightly higher emissions were found for riparian areas (1.8 ± 0.3 μg N₂O–N m⁻² h⁻¹), which cover ∼4% of the landscape. Even faeces or urine additions stimulated N₂O emissions from steppe soils only weakly (<2.5 μg N₂O–N m⁻² h⁻¹ for a 5 days period). Due to low moisture contents, N₂O emissions from dung heaps were also rather low (6.2 ± 0.8 μg N₂O–N kg⁻¹ dry matter h⁻¹). In contrast, three orders of magnitude higher N₂O emissions were found at sheepfolds (2.45 mg N₂O–N m⁻² h⁻¹ on average). By calculating N₂O emissions on a landscape scale, we show that point sources, and especially sheepfolds, become the dominating regional N₂O source during the growing season if stocking rates are >1 sheep ha⁻¹. Our results indicate that the common grazing management in the Xilin river region leads to a translocation of nitrogen from large source areas towards defined spots. This finding is further supported by measurements of NH₃ concentrations at different sites. Since most of the nitrogen accumulated in these hot spots is finally lost through burning of the dried excrements by the farmers for heating and cooking purposes, the ecosystem faces a significant human perturbation of regional N cycling, which may contribute to an accelerated degradation of steppe in the Xilin river region. Responsible Editor: Per Ambus.     

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.

     

Discontinuous gas exchange in dung beetles: patterns and ecological implications

This study correlates a distinctive pattern of external gas exchange, referred to as the discontinuous gas exchange cycle (DGC), observed in the laboratory, with habitat associations of five species of telecoprid dung beetles. The beetles were chosen from a variety of habitats that would be expected to present different amounts of water stress. All five species exhibited DGC. Sisyphus fasciculatus has been recorded only in woodland areas, and does not have strict spiracular control during its DGC. Anachalcos convexus and Scarabaeus rusticus are associated with open mesic habitats. Both species exhibit a distinct DGC, previously found in some other insect species, but intermediate within this study group. Sc. flavicornis and Circellium bacchus are typically found in arid regions, and have the most unusual form of DGC, with spiracular fluttering during the burst phase. These results support the hypothesis that spiracular fluttering reduces respiratory water loss. From this study we conclude that the DGC is an ancestral adaptation, most probably as a result of anoxic environments in underground burrows, but that spiracular control is enhanced to reduce respiratory water loss in beetle species that live in arid habitats. Received 4 August 1999 / Accepted: 7 October 1999     

Relative abundance and species composition of some dung beetles (Coleoptera: Scarabaeinae) in Bangladesh

1. The occurrence and community structure of dung beetle species (Coleoptera: Scarabaeinae) in the faecal deposits of man and some domestic vertebrates varied according to dung type, seasonal conditions and soil type. 2. Of the twenty-four species belonging to six genera, Onthophagus was the prevalent genus and O. ramosellus the most abundant species. 3. Cattle dung was the dominant food type.