Structure and function of the mandibles of adult dung beetles (Coleoptera : Scarabaeidae)

The mouthparts of adult dung beetles (Coleoptera : Scarabaeidae) are adapted for manipulation of soft, pasty food—usually the excrement of vertebrates. Nutrients are derived primarily from micro-organisms contained in the food and these must be broken up before ingestion. The mandibles, particularly the molar lobes, are designed to finely grind these particulates; the molae function as a mortar-pestle system, which mills the organic “grist” contained in the food by a combination of squeezing and grinding actions. The ability of the molae to finely grind food particles resides primarily in the structure of the molar surface, which consists of a series of ridges bearing rows of submicroscopic scrapers (“tritors”). The incisor lobes of the mandibles scrape food from the surfaces of the galeae and lacinae, which bring food into the preoral cavity. The structure of the mouthparts of Canthon pilularius (L.), which is described in detail, is basically the same as that of 10 other species examined. Those of Aphodius and Geotrupes are similar in some respects to those of scarabaeines but are obviously less well specialized for microphagy.     

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.

     

Phylogeography of the Namib Desert dung beetles Scarabaeus (Pachysoma) MacLeay (Coleoptera: Scarabaeidae)

Aim Namib biogeography in many instances remains reliant on advanced and detailed systematic studies. This study attempts to combine molecular phylogenetic data, geology and palaeo‐climatic data to (i) resolve the relationships of the 13 morphological species of Scarabaeus (Pachysoma) and (ii) relate their evolution to past climatic and geological events. Location South Africa and Namibia. Methods Sequencing of a 1197 bp segment of the mitochondrial cytochrome oxidase I (COI) gene of the 13 species within Scarabaeus (Pachysoma) was undertaken. Analyses performed included Parsimony and Maximum Likelihood as well as imposing a molecular clock. Results The molecular phylogeny showed strong support for 11 of the 13 morphological species. The remaining two species, S. (P.) glentoni and S. (P.) hippocrates, formed a complex and could not be assigned specific status on the basis of the COI gene phylogeny. Strong support for the three species formerly classified within the genus Neopachysoma was consistently obtained. The subgenus appears to have arisen c. 2.9 Ma. Species within the subgenus arose at different times, with the common ancestor to Neopachysoma and the hippocrates complex having evolved 2.65 and 2.4 Ma, respectively. Scarabaeus (P.) denticollis, S. (P.) rotundigenus, S. (P.) rodriguesi and S. (P.) schinzi are some of the youngest species, having diverged between 2 million and 600,000 years ago. Main conclusions Scarabaeus (Pachysoma) is a derived monophyletic clade within the Scarabaeini. The subgenus appears to be young in comparison with the age of the Namib Desert, which dates back to the Miocene (c. 15 Ma). The psammophilous taxa are shown to disperse with their substratum and habitat, barchan dunes. Clear south/north evolutionary gradients can be seen within the species of this subgenus, which are consistent with the unidirectional wind regime. Species with a suite of mostly plesiomorphic characters have a southerly distribution while their derived psammophilous relatives have central to northern Namib distributions. Major rivers such as the Orange, Buffels and Holgat appear to be gene barriers to certain species as well as areas of origin of speciation events.     

Seasonal distribution and prevalence of dung beetles (Coleoptera: Scarabaeidae) in Kurukshetra,Northern India

A total of 8306 individuals as 29 species from 3 subfamilies (Hybosorinae, Aphodiinae, Scarabaeinae) were trapped by pitfall bait trapping. The maximum number of species of Scarabaeinae (68.97%) and number of individuals of Aphodiinae (56.84%) take part in the formation of a community. The number of species, number of individuals, diversity and dominance changes throughout the season. Seasonal prevalence of 82.76% of species and 50.81% of individuals was found in June. Aphodius campestris was found to be more attracted to the pitfall bait traps. The beetle community found in May shows the maximum Shannon's Diversity Index (2.46) but Margalef's Diversity Index was highest in August (3.06). Environmental factors play an important role in the occurrence and abundance of dung beetles. In the present study non-significant but positive correlations between temperature and species richness, abundance and diversity were found. Percent relative humidity showed a positive correlation with abundance only, and negative correlations with species richness and diversity.     

Discontinuous gas-exchange cycles in Scarabaeus dung beetles (Coleoptera: Scarabaeidae): mass-scaling and temperature dependence.

Although discontinuous gas exchange cycles (DGC) are known from many insects, the effects of body size and temperature on DGC have not been widely examined. Here, these effects are investigated in five Scarabaeus dung beetle species from mesic and xeric habitats. The investigation tests two hypotheses: that previous estimates of the scaling exponents for the DGC and its characteristics are more broadly applicable to insects, and that, in response to temperature, both DGC frequency and the quantity of CO2 emitted during the open (O) phase (O-phase emission volume) are modulated. Like previous workers, we find that V&d2;co2 scaled as mass0.968 and that O-phase emission volume scaled as mass0.833. However, temperature-associated increases in .Vco2 (Q10's of 2.19-2.65) were modulated mostly by increases in DGC frequency since O-phase volumes remained constant across temperature. Flutter (F)-phase and O-phase durations were closely coupled to DGC duration, although the relationship between closed (C)-phase duration and DGC duration was less pronounced. We show that ventilation phase coefficients, previously considered a measure of the proportional duration of each phase of the DGC, calculated from the slopes of these relationships are a measure of change in phase duration with change in DGC duration and not a measure of the way in which total DGC duration is apportioned among phases. We suggest that proportions be used to estimate the contribution of each of the phases to the total duration of the DGC.     

Inferring species decline from collection records: roller dung beetles in Italy (Coleoptera, Scarabaeidae)

The decline of roller dung beetles (Coleoptera, Scarabaeidae) in Italy, at national and regional level, was described using a database of both literature and unpublished data, since the late of 19th century. The probability of finding roller species was assessed for each decade of the 20th century in order to detect changes in their distribution and status. An analysis of decline was performed using a complementary approach of different extinction indexes, according to IUCN criteria. During the 20th century, both the absolute number of records and the UTM cells where roller beetles were found increased clearly as a consequence of the escalation of entomological surveys. Nevertheless, comparing R species (rollers) with all the scarab dung beetles from the first to the last quarter of the 20th century, the relative frequency of roller records decreased by 31%, while the relative number of occupied UTM cells dropped by 23%. Six roller species showed a significant decline in Italy from the first to the second part of the 20th century, and nine in the last 30years. Other two species disappeared from the majority of the northern regions. Finally, the values of all the extinction indexes were strongly correlated and detected a high risk of extinction in Italy for six species. A gradual change in land use during the last three decades, consisting of a reduced extension of grasslands in favour of either intensive agriculture or reforestation after abandonment of livestock raising, was likely the main factor inducing decline and local extinction of roller species.     

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.     

Edge and land use effects on dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) in Brazilian cerrado vegetation

The Edge Influence is one of the most pervasive effects of habitat fragmentation, as many forest remnants in anthropogenic landscapes are within 100 m of edges. Forest remnants may also affect the surrounding anthropogenic matrix, possibly resulting in a matrix–edge–remnant diversity gradient for some species groups. We sampled dung beetles in 15 agricultural landscapes using pitfall traps placed along transects in matrix–edge–remnant gradients. The remnants were a native savanna-like vegetation, the cerrado, and the matrix was composed of three human-dominated environments (sugarcane, eucalyptus, pasture). More species were observed in cerrado remnants than in adjacent land uses. Dung beetles were also more abundant in the cerrado than in the landscape matrix of sugarcane and eucalypt, but not of pasture. Dung beetles were severely affected by anthropogenic land uses, and notwithstanding their high abundance in some land uses such as pasture, the species richness in these areas tended to be smaller than in the cerrado remnants. We also found that the influence of the edge was evident only for abundance, particularly in landscapes with a pasture matrix. However, this land use disrupts the species composition of communities, indicating that communities located in cerrado and pasture have a distinct species composition, and that both communities are affected by edge distance. Thus, anthropogenic land uses may severely affect dung beetles, and this impact can extend to communities located in cerrado remnants as well as to those in matrices, with possible consequences for ecological processes such as decomposition and nutrient cycling.     

Thermoregulation in endothermic dung beetles (Coleoptera: Scarabaeidae): effect of body size and ecophysiological constraints in flight

We explore the physiological constraints of body temperature as related to body mass and ambient temperature during flight in endothermic dung beetles showing a mass-related breakpoint where species show strong vs. weak endothermy. We found two different strategies in the dung beetles prior to flight; larger beetles (>1.9 g) elevate and maintain their body temperature (T(b)) at levels well above ambient temperature (T(a)) whereas smaller beetles' (<1.9 g) T(b) tends to conform with T(a). Physiological constraints analysis revealed a constant maximum tolerated temperature (in flight) of 42 degrees C and a minimum temperature for flight of around 25 degrees C. These, with body mass, may play a role in thermal niche partitioning and geographical distribution patterns.     

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.

     

Roles of endothermy in niche differentiation for ball-rolling dung beetles (Coleoptera: Scarabaeidae) along an altitudinal gradient

Abstract.  1. An analysis of whether niche differentiation in ball-rolling dung beetles can be explained by the way in which they regulate their body temperature was conducted.
2.  A priori assumptions were: (i) if thermoregulation affects niche partitioning, sympatric species must have different endothermic strategies that minimise encounters; or, alternatively (ii) if two co-occurring species show the same thermoregulation pattern and their flight periods overlap, they might be avoiding competition by exhibiting different resource preferences or different food relocation behaviour.
3. The ball-rolling dung beetles studied showed a hierarchical structure based on the species' endothermic capacity, measured as temperature excess [ T _ex= difference between body temperature ( T _b) and ambient temperature ( T _a)]. Those with a high T _ex (10–15 °C) were located exclusively at altitudes >1000 m a.s.l. On the coastal plains, species with a high T _ex were restricted to flying at night when the T _a was lower. Species with a lower T _ex (less than 10 °C higher than T _a) were found in the coastal plains zone.
4. Where there was sympatry with similar trophic habits, the species involved showed very different thermal niches, and where there was significant overlap of thermal niches between sympatric species, trophic habits of species were very different.
5. The results suggest that it is possible to use the concept of the thermal niche as a tool to explain interspecific interactions and the spatial distribution of species.     

Life history traits and resource utilisation in an assemblage of north temperate Aphodius dung beetles (Coleoptera: Scarabaeidae)

To help understand and interpret the structure and function of Aphodius dung beetle assemblages, life history traits and resource utilisation were studied for the ten species comprising the local assemblage of intensively grazed pastures in southern Ireland Most species were univoltine but one species ( A fimetarius ) was at least partly bivoltine However, temporal overlap m adult flight periods does not necessarily imply overlap in resource use Three different strategies of ovarian development were distinguished and were related to the preferred oviposition site and successional occurrence of the various species Evidence suggested that absence of mature eggs in a female's ovaries did not necessarily imply that a female was in a non-reproductive state Two species ( A prodromus and A sphacelatus ) did not breed in dung, in the laboratory larvae were reared in decaying vegetation One species ( A errattcus ) developed m brood masses beneath the dung pat Larvae of all the other species developed within the dung pat There were consistent Interspecific differences in the larval development rates, with two species ( A rufipes and A rufus ) overwintering mainly as prepupae and the other species mainly as adults Previous studies have considered Aphodius assemblages as single guilds but the detailed natural histories of these species may affect guild designation     

Attractiveness of dung beetles (Coleoptera: Scarabaeinae) to faeces from native mammals in different trophic guilds

Dung beetles are predominantly coprophagous and use mammalian faeces as their main food resource and for offspring rearing. However, these resources are generally ephemeral and scarce. The objective of this study was to verify the attractiveness to dung beetles of the faeces of nine native mammal species of the Cerrado biome in different trophic guilds. This study was conducted during the rainy season (October 2016 to March 2017) in two sensu stricto areas of the Cerrado in the Brasília National Park (PNB), Federal District, Brazil. Pitfalls containing faeces of nine mammals native to the region were used as bait (from three herbivores, three carnivores and three omnivores) for the collection of dung beetles. We collected 203 individuals and 19 species, and the most abundant species were Uroxys aff. thoracalis, Oxysternon palemo, Coprophanaeus spitzi and Diabroctis mirabilis. The most attractive faecal bait was from the omnivorous Chrysocyon brachyurus (32.67% of the individuals and 13 species). This study reveals variation in the attraction of dung beetles to native mammal dung of the Cerrado, with more abundance in the faeces of omnivores and carnivores and less in the faeces of herbivores. The species composition in the carnivore baits did not differ from that of the omnivore baits. These data indicate that changes in Scarabaeinae assemblages result from changes in mammal species such as those that have occurred in the Cerrado biome with the conversion to cattle production, which in recent years has caused intense fragmentation of habitats, expansion of agricultural and livestock activities and the introduction of exotic species. The mammal C. brachyurus, vulnerable to extinction in Brazil, should be a focus for future research, since its faeces support a great diversity of dung beetles in the Cerrado.     

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).     

Four new species from the Agrilus wittemani species-group (Coleoptera: Buprestidae: Agrilini) from Silent Valley National Park,Southern Western Ghats,Kerala, India

The Agrilus wittemani species-group comprising six species from South and Southeast Asia; two of these are endemic to south India. Four new species of A. wittemani species-group Jendek & Nakládal, 2017 are described in this paper: Agrilus keralensis Seena, Anand and Shibu-Vardhanan sp. nov.; A. palakkadensis Seena, Anand and Shibu-Vardhanan sp. nov.; Agrilus sahyadriensis Seena, Anand and Shibu-Vardhanan sp. nov.; A. silentvalleyensis Seena, Anand and Shibu-Vardhanan sp. nov. A key to all species is provided and complemented with illustrations of habitus and genitalia.     

Not all dung beetles feed on dung: Scarabaeinae (Coleoptera: Scarabaeidae) attracted to different carrion types in contrasting habitats at Brazilian Amazon

By understanding how assemblages segregate according to food types, it is possible to depict and understand species distribution and exploitation of similar food resources. Although it is well known that dung beetles may feed on carrion, but the attractiveness of different carrion types for these beetles is still poorly understood. In this study, we compared the dung beetle (Coleoptera: Scarabaeidae: Scarabaeinae) diversity attracted to two carrion types in different habitats in an Amazonian landscape. Dung beetles were captured by using pitfall traps baited with decaying cattle beef and cattle liver in native forest, peach palm plantation, teak plantation and exotic pasture. Overall, we captured 582 dung beetles of 43 species, where Canthidium aff. lentum was classified as a specialist of cattle beef, Deltochilum sp. One was considered specialist of liver carrion, 11 were considered generalists and for the 30 remaining species, it was not possible to determine their carrion preference due to the low number. Abundance, but not species richness, was affected by carrion type, and a higher number of beetles was recorded in beef-baited traps compared to liver-baited traps. According to functional groups, endocoprid beetles were more abundant in traps baited with decaying beef, while paracoprid and telecoprid beetles did not show preference for any carrion type. This study suggests a novel scenario of intra-resource segregation, where species vary their preferences depending to the part of the dead corpse being exploited. The amount of food resource, the maintenance of stable populations and trophic preference mediated by chemical cues are some mechanisms that may explain the segregation of dung beetles among carrion types.