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.     

Elongated hindguts in desert‐living dung beetles (Scarabaeidae: Scarabaeinae) feeding on dry dung pellets or plant litter

Most adult dung beetles (Scarabaeidae: Scarabaeinae) feed on fresh, wet dung of larger herbivorous or omnivorous mammals. As refractory plant fragments are selected out before ingestion, the food is presumed easily digestible. However, members of the desert‐living scarabaeine genus Pachysoma (probably evolved from an ancestor closely related to the wet‐dung feeding genus Scarabaeus) select dry dung pellets and/or plant litter. Thus, they ingest a much higher proportion of structural plant material, which nevertheless appears to be digested rather efficiently. This study investigates morphological modifications of the gut for this digestion in adults of eight Pachysoma species, both pellet and litter feeders. To ascertain hypothesized ancestral conditions, four fresh‐dung feeding Scarabaeus species were also examined. The latter have the usual dung beetle gut consisting of a long, simple midgut, followed by an equally simple, much shorter hindgut of the same width. Lengths of midguts (M) and hindguts (H) divided by body length (B) for comparison between species of different size are: 4.9–6.3 (M/B) and 0.7–0.8 (H/B), which is normal for dung feeders. In Pachysoma, lengths are 6.3–6.5 (M/B) and 1.0–1.4 (H/B) in pellet feeders, and 4.4–5.0 (M/B) and 2.0–2.5 (H/B) for litter feeders. Hindguts are still morphologically undifferentiated and of midgut width, but clearly longer, particularly in litter feeders. Presumably, plant fragments in the food are digested, at least partly, in the hindgut. If so, the morphological adaptation is unusual: simple elongation rather than the expansion of part of the hindgut, as found in several other plant‐ or detritus‐feeding scarabaeids. J. Morphol. 2013. © 2013 Wiley Periodicals, Inc.     

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.     

Evolutionary patterns of hind wing morphology in dung beetles (Coleoptera: Scarabaeinae)

The shape of wings can be a good predictor of adaptations to different selective pressures and the value of wing features in taxonomy and phylogeny has long been recognized. In our investigation of the hind wing evolution of dung beetles (Scarabaeinae) we use geometric morphometrics combined with a cladistic approach. The variations of entire hind wings and of three specific regions of 80 dung beetle species were investigated using 19 landmarks and outline data. Extensive evidence indicates that the wing as a whole and the three separate regions were under different selective pressures. The detailed evolutionary patterns of the three regions and the reconstruction of the ancestral forms were computed by mapping the geometric morphometrics data onto a tree based on a cladistic character analysis.     

Evolutionary constraints in hind wing shape in Chinese dung beetles (Coleoptera: Scarabaeinae)

This study examines the evolution hindwing shape in Chinese dung beetle species using morphometric and phylogenetic analyses. Previous studies have analyzed the evolution of wing shape within a single or very few species, or by comparing only a few wing traits. No study has analyzed wing shape evolution of a large number of species, or quantitatively compared morphological variation of wings with proposed phylogenetic relationships. This study examines the morphological variation of hindwings based on 19 landmarks, 119 morphological characters, and 81 beetle species. Only one most parsimonious tree (MPT) was found based on 119 wing and body characters. To better understand the possible role of the hindwing in the evolution of Scarabaeinae, additional phylogenetic analyses were proposed based on the only body features (106 characters, wing characters excluded). Two MPT were found based on 106 body characters, and five nodes were collapsed in a strict consensus. There was a strong correlation between the morphometric tree and all phylogenetic trees (r>0.5). Reconstructions of the ancestral wing forms suggest that Scarabaeinae hindwing morphology has not changed substantially over time, but the morphological changes that do occur are focused at the base of the wing. These results suggest that flight has been important since the origin of Scarabaeinae, and that variation in hindwing morphology has been limited by functional constraints. Comparison of metric disparity values and relative evolutionary sequences among Scarabaeinae tribes suggest that the primitive dung beetles had relatively diverse hindwing morphologies, while advanced dung beetles have relatively similar wing morphologies. The strong correlation between the morphometric tree and phylogenetic trees suggest that hindwing features reflect the evolution of whole body morphology and that wing characters are suitable for the phylogenetic analyses. By integrating morphometric and cladistic approaches, this paper sheds new light on the evolution of dung beetle hind wings.     

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.

     

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.     

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.     

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.     

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.     

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.     

Spatial and temporal segregation in dung beetles (Coleoptera: Scarabaeinae) in the Cerrado of Central Brazil

Biodiversity and Conservation - In dung beetles, spatial and temporal segregation is determined by the niche and evolutionary history of the species and is mediated by competition. Different...     

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.     

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.     

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.     

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.     

Influence of land use on the taxonomic and functional diversity of dung beetles (Coleoptera: Scarabaeinae) in the southern Atlantic forest of Argentina

The degradation and replacement of native ecosystems affects both their taxonomic and functional biodiversity. However, native species may find a gradient of habitat suitability in different land uses within a region. The aim of this study was to evaluate the effect of land use on the taxonomic and functional diversity of dung beetle assemblages in the southern Atlantic forest of Argentina. Dung beetles were sampled in both the native forest (control) and different land uses (Pine and Yerba mate plantations and cattle pastures) during the 2014 summer, using pitfall traps baited with human feces and rotten meat. Samplings were taken from 20 different sites, with five replicates of each land use and the native forest (100 pitfall traps in total). A total of 1699 beetles of 27 species were captured. Canthon quinquemaculatus, Canthon conformis and Dichotomius sericeus were the most abundant species. Cattle pastures were the land use most negatively affected in their taxonomic and functional diversity, particularly large paracoprid dung beetles. Pine plantations maintained their taxonomic and functional diversity in relation to the native forest and Yerba mate plantations showed, in general, an intermediate situation. Microclimatic conditions (average temperature and humidity and maximum temperature) were correlated with functional diversity (the proportion of large paracoprid dung beetles decreased with increasing temperature) and are probably good predictors to explain the observed patterns of functional diversity of dung beetles. The development of sustainable production systems that preserve the native biodiversity requires the conservation of key components from the ecological niche of native species, especially microclimatic conditions.