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

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

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

     

Survivorship and fertility schedules of two Sumatran tortoise beetles,Aspidomorpha miliaris andA. sanctaecrucis (Coleoptera: Chrysomelidae) under laboratory conditions

Two species of tortoise beetles, Aspidomorpha miliaris (AM) and A. sanctaecrucis (AS) feeding on a shrub-like morning glory, Ipomoea carnea, were reared under laboratory conditions to study their survivorship and fertility schedules. AM and AS required 34–39 days and 30–37, respectively, for the development of the immature stages. The mean longevity of the males was 88.4 days in AM and 63.8 in AS, and that of females was 87.9 days in AM and 83.3 in AS. The mean length of the pre-reproductive period (27.2 days in AM and 33.8 in AS) was much longer than that of the post-reproductive period (10.9 days in AM and 14.3 in AS). Females laid eggs at a nearly constant rate throughout their reproductive period. The reproductive value V_x/V₀ of the two species remained high for most of their adult life, as a result of prolonged survivorship and fertility periods. The total number of eggs produced per female was 442.9 (AM) and 80.1 (AS). The intrinsic rate of natural increase r was 0.070 (AM) and 0.044 (AS) per capita per day. The prolonged reproductive schedules, coupled with strong dispersal power, of these species no doubt have an adaptive value for living in highly disturbed tropical environments, where rainfall is ample but unpredictable and food resources are available throughout the year in a wide area, but distributed in widely flung patches.     

Sibling and non-sibling cannibalism by larvae of a lady beetleHarmonia axyridis Pallas (Coleoptera: Coccinellidae) in the field

Sibling (cannibalism among siblings) and non-sibling cannibalism (cannibalism among non-siblings) were studied in a natural population of a lady beetle, Harmonia axyridisPallas . Of all the eggs laid (n=2269), 24.76% (n=562) were killed by sibling cannibalism and 36.10% (n=819) were killed by non-sibling cannibalism. Sibling cannibalism occurred constantly and intensively in most egg batches throughout the entire oviposition period. On the other hand, non-sibling cannibalism was more intense in the middle and late oviposition periods, and when the egg batches were close to an aphid colony. This may be due to the high density of H. axyridis larvae relative to aphid density in the middle and late oviposition periods and also the larvae searching intensively near an aphid colony.     

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     

The effect of crowding on adult populations ofDelia (=Hylemya) antiqua (Meigen)

Using 5 density levels, the effect of increasing density was studied on several population statistics of adult Delia (=Hylemya) antiqua. Amongst the statistics studied were adult lifespan; female mating frequency, fecundity and egg hatchability. It was shown that females lived significantly longer than males and that increasing density significantly reduced lifespan. Density had no effect on mating frequency. Total fecundity/female was significantly reduced with increasing density but oviposition rate was density independent. Using these statistics, mean generation time T, net reproduction rate R₀ and capacity for increase r_c were calculated at the different density levels. Values of R₀ showed a one-tailed response but there was no clear effect of density on r_c. Multiple comparisons between variables revealed several important relationships.

1. Adult lifespan was the most important factor affecting egg production.
2. Number of mated females was more important in affecting total fecundity/cage than adult density.

It could also be calculated that to produce the effect of one single mated female on total fecundity/cage the overall density would have to be reduced by 0.0023 individuals/cm³. These results are discussed in the context of efficient laboratory rearing of the onion fly.     

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.     

Dispersal of Steinernema glaseri (Nematoda: Steinernematidae) in adult Japanese beetles, Popillia japonica (Coleoptera: Scarabaeidae)

Japanese beetle adults, Popillia japonica, can become infected with and disperse the entomopathogenic nematode, Steinernema glaseri, under laboratory and field conditions. After a 24-h exposure to 10 000 infective juveniles/20 adult beetles, 45% of the beetles died within 4 days post-treatment, but only 59% of these were infected with the nematode. Corresponding control mortality was 6.5%. An average of 238 infective juveniles were produced/beetle. Beetles exposed to 4000 and 10 000 infectives/10 adults carried with them an average of 17 and 59 infectives/adult on external body surfaces respectively. When beetles that had been exposed to 4000 infectives/20 adults were transferred to, and held in, cages containing soil for 2 weeks, up to 89% of the adults died, as did 74% of the P. japonica larvae that were subsequently placed in the cages. When adults that had been exposed to 50 000 infectives/250 beetles in moist sand for 16 h were released into screened cages in the field at soil temperatures of over 25 C, the soil beneath 83% of the cages tested positive for the nematode, using Galleria mellonella larvae as bait, 2 weeks after releasing the beetles. No nematodes were detected in control plots. The potential of infected adult P. japonica for dispersing S. glaseri by flight was investigated by exposing adults to 50 000 infectives/250 beetles, marking and releasing them in the field and recapturing them in lure-baited Japanese beetle traps. Less than 1% of the treated beetles were recaptured, but 33% of these had one or more nematodes in their hemocoels. Accordingly, this approach does not appear to be feasible for large-scale augmentation and dispersal of the nematode using currently developed methods of infection. If improvements in mass-inoculation methods can be made that enable a rapid high percentage of infection while still permitting flight, this concept could be employed to establish new foci of infection or for the introduction of other species of nematodes.     

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.

     

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.     

Adjustment of parental expenditure in sympatric <Emphasis Type="Italic">Onthophagus</Emphasis> beetles (Coleoptera: Scarabaeidae)

Many organisms adjust their parental expenditure to offspring in response to resource quality. However, the mechanisms underlying the adjustment in parental expenditure are not well understood. We examined the adjustments in parental expenditure and subsequent offspring performance in two sympatric, closely related dung beetles, Onthophagus ater and O. fodiens, that were provided either monkey, deer, horse, or cow dung. The egg contained within each dung brood mass provisioned by the parent beetles develops to adulthood underground. Thus, the size of the brood mass roughly represents the amount of parental expenditure. The brood mass size differed between the two species and among the four dung types. Results of offspring performance suggested that O. ater parents optimally adjusted the brood mass size in response to dung quality, whereas O. fodiens parents did not. We hypothesized that brood mass size in O. ater may increase with prolonged egg maturation caused by the lower nutrition level of cow dung. In addition, our complex results may be explained in part by the specific threshold concept of dung quality (i.e., water content and nutritional level).     

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.     

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.     

Population genetic characterization of the endangered dung beetle Copris tripartitus (Coleoptera: Scarabaeidae) using novel microsatellite markers

The dung beetle Copris tripartitus (Coleoptera: Scarabaeidae) has long been considered an endangered insect in South Korea; the detection of recent population increases leaves its endangered status uncertain. Population genetic analysis subsequent to development of molecular markers is essential for establishing proper conservation strategies. In this study, we developed ten microsatellite markers specifically for C. tripartitus. Sixty-eight individuals of C. tripartitus collected from six South Korean localities were genotyped to validate these markers and preliminarily assess population genetic characteristics. Per-locus observed number of alleles, observed heterozygosity (H_O), and expected heterozygosity (H_E) ranged from 5–12, 0.499–0.958, and 0.54–0.743, respectively. All populations showed higher H_O than H_E, negative values of inbreeding coefficient, and, overall, no sign of recent population bottlenecks (excluding one population, Seosan). This suggests that C. tripartitus did not suffer from genetic drift and inbreeding, which are typically severe in small, isolated populations. Nevertheless, detection of only one of the two gene pools in some populations and resultant genetic subdivision into two population groups may suggest that the population size is not enough to cover both gene pools. Thus, a more extended period of protection may be required to ensure higher genetic diversity of widespread populations and achieve the long-term conservation goal.     

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.     

A revision of Dichotomius (Homocanthonides) Luederwaldt, 1929 (Coleoptera: Scarabaeidae: Scarabaeinae)

The monospecific subgenus Dichotomius (Homocanthonides) is revised and its single species, Dichotomius (H.) smaragdinus (Perty, 1830) is redescribed and distinguished from other Dichotomius species. Dichotomius (H.) smaragdinus is a polymorphic species and its distribution comprises the Brazilian Cerrado. For the first time, morphological variation and male genital organ are described and illustrated. We believe that this species might be endangered due to extensive deforestation of Brazilian Cerrado by agricultural and pasture expansion. Thus, we highlight the importance of preserving this very unusual Dichotomius species.     

Endocoprid activity of Aphodius fossor (Coleoptera: Scarabaeidae) related to bovine dung decomposition in a mixed grass prairie

Aphodius fossor (L.) (Coleoptera: Scarabaeidae), a common endocoprid dung beetle in southeastern Wyoming, may have a survival strategy to maintain dung pad integrity and moisture crucial to larval survival in an arid climate (annual precipitation <30 cm). Typically, A. fossor seems to contribute little to dung pad decomposition, because inhabited dung pads seem to be intact and weigh approximately the same as uninhabited pads, even after 1 yr on pasture. To assess the role of A. fossor in dung pad decomposition and nutrient recycling, artificially formed bovine dung pads were inoculated with five pairs of adult A. fossor. After 40 d, A. fossor activity had no measurable effect on external surface area or moisture retention within the dung pad cores. Pads inhabited by A. fossor weighed significantly more than did control pads on most weigh dates of the experiment, possibly because of incorporation of soil particles at the dung/soil interface. Externally, A. fossor-inhabited dung pads seemed intact; however extensive tunneling was evident throughout the core of the pad leaving an intact, protective crust. A. fossor activity increased microbial biomass carbon in the soil beneath the dung pad. Levels of total nitrogen (N) and carbon (C) decreased in the pads but increased in soil beneath the pads. Dung in the core and in the crust of pads with A. fossor had significantly less total N than pads with no beetles and total C was significantly lower in the crust.