Oviposition of the dung beetle Aphodius ater in relation to the abundance of yellow dungfly larvae (Scatophaga stercoraria)

1. The dung beetle Aphodius ater and the yellow dungfly Scatophaga stercoraria are temporally co-occurring species in sheep dung, which they use for reproduction and nutrition ( A. ater ) or for reproduction only ( S. stercoraria ) during the spring in northern Germany. Scatophaga stercoraria uses fresh sheep dung pellets a few hours old for oviposition, whereas A. ater lays eggs into 2–10-day-old pellets. In the present study, the egg laying behaviour of A. ater in sheep dung in relation to the presence of larvae of S. stercoraria was investigated experimentally.
2. Choice experiments, based on examining the egg laying behaviour of beetles in 2- and 4-day-old pellets with and without high and low densities of fly larvae, showed the following. In 2-day-old pellets, the beetles did not distinguish between pellets without fly larvae or with fly larvae at low larval density but avoided laying eggs into pellets with a high larval density. In 4-day-old pellets, the beetles always preferred to lay their eggs into pellets without fly larvae, regardless of larval density.
3. The influence of different densities of larvae of S. stercoraria on dung depletion was examined by measuring the dry weight, organic matter content and organic nitrogen content of the remaining dung after larval development. The presence of the larvae led to a reduction in all three parameters.
4. The beetles' behaviour of laying eggs into older pellets, and their awareness of the presence of high densities of fly larvae, enables them to avoid egg laying into pellets that will have been depleted by fly larvae before the beetle larvae have finished their development.     

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     

Evidence that sex‐specific signals may support mate finding and limit aggregation in the dung beetle Aphodius fossor

1. In temperate climates, dung is often colonised by several species of endocoprid (dwelling) dung beetles which use pats for feeding, shelter, and reproduction. 2. Endocoprid beetles aggregate even when offered patches (dung pats) of consistent age, size, and origin, suggesting that beetles themselves might influence the attractiveness of patches to members of their own species. Both pheromones, and physical changes to the structure of dung pats caused by colonising beetles have been suggested as mechanisms facilitating intraspecific aggregation, but neither of these hypotheses have been empirically tested. 3. Using a common European dung beetle (Aphodius fossor L.), we conducted a simple choice experiment designed to test whether (i) earlier colonisation by conspecifics could alter dung attractiveness and (ii) whether attraction was influenced by sex‐specific signals. 4. We found that female beetles are repelled by dung colonised by conspecific females and are attracted to dung colonised by conspecific males. Male beetles show no evidence of attraction or repellence for dung colonised by either sex. Neither in females nor males was uncolonised dung found to be significantly more or less attractive than predicted by non‐preference. 5. Our results suggest that for A. fossor male‐produced signals may support mate finding in patchy environments, and that female‐produced signals may serve to discourage subsequent colonisation by additional females.     

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     

Larval population density affects female weight and fecundity in the dung beetle Aphodius ater

1. Competition for food at high densities during larval development leads to reduced adult weight in the northern temperate dung beetle Aphodius ater. 2. Analysis of female beetles caught in the field showed that numbers of eggs and total egg load per female were correlated positively with beetle size. 3. Female beetles reared at different population densities during larval development in the laboratory were analysed with regard to their lifetime fecundity and reproductive lifespan. 4. High population densities during development had a negative influence on the number of eggs per female and on reproductive lifespan. Lifetime fecundity was correlated positively with female weight. 5. It was concluded that competition during larval development in the first generation of offspring will result in a lower number of offspring in the second generation in Aphodius ater, and thereby reduce parental fitness.     

Abdominal ventilatory pattern in crickets depends on the stridulatory motor pattern

Abstract. Ventilatory motor patterns were recorded from abdominal muscles in crickets, Gryllus campestris L.and Teleogryllus commodus (Walker), at rest and during three types of stridulatory motor activity; calling, courtship and aggressive song.
Increases in ventilatory period were almost exclusively due to an increase of the pause between expiratory bursts, whereas abdominal ventilatory bursts remained constant at 200 ms.Ventilatory patterns depended on the stridulatory motor pattern and indicated that the same basic respiratory oscillator exists in both cricket species.
In G.campestris there was a strict 1:1 coupling between chirps and ventilatory bursts.In T.commodus such a relationship was also observed for the chirp part of the songs, but less strictly for the trill part of the calling song and not for the courtship song.In both species the onset of the ventilatory burst was within ± 100 ms of a stridulatory chirp.Ventilatory burst lasted longer the earlier they began before a stridulatory chirp.This suggests strongly that the stridulatory motor pattern terminates the expiratory burst, and thus influences the ventilatory motor pattern.     

Making the cryptic visible – resolving the species complex of Aphodius fimetarius (Linnaeus) and Aphodius pedellus (de Geer) (Coleoptera: Scarabaeidae) by three complementary methods

Species in cryptic complexes are, per definition, difficult to identify using morphological characters. One such complex was recently detected in the dung beetle Aphodius fimetarius (Linnaeus) sensu lato, an abundant dung beetle with a wide distribution. While the two component taxa, Aphodius fimetarius sensu stricto and Aphodius pedellus (De Geer) exhibit distinctly different karyotypes, the validity of subtle morphological characters proposed to distinguish between them has been debated. Given the variability and minor interspecific differences in external characters, the large‐scale distribution of respective taxa has remained unknown, as have potential differences in ecology and habits. In this study, we ask how A. fimetarius and A. pedellus can best be distinguished, whether the use of different types of characters (karyotypes, DNA sequences and morphological traits) results in consistent species identification, where these species occur and whether they exhibit ecological differences. In total, we inspected a material of 4401 individuals from across the globe, of which 183 were examined for both mtDNA sequences and morphology, 154 for both morphology and karyotype, and 9 (including the recently proposed neotype of Aphodius fimetarius) for all three types of characters. As a marker gene, we sequenced a 590 bp region of the cytochrome c oxidase I gene for 183 individuals. Overall, DNA sequences offered a clear‐cut distinction between taxa: sequences of A. fimetarius and A. pedellus differed by an average pairwise distance of 8.2%, whereas variation within species was only 0.9% for A. fimetarius and 0.5% for A. pedellus. Morphological and chromosomal characters offered species identifications consistent with that of molecular characters: karyotypes identified as A. pedellus consistently fell within one of the molecular clades, whereas karyotypes identified as A. fimetarius fell within the other clade. Likewise, the majority of individuals identified by morphological characters were assigned to the same species by sequence‐based characters. Both taxa thus defined were found to be Holarctic in distribution, with major sympatry within Central and Southern Europe and mixed patterns of sympatry within the US. Northern areas of Europe, Asia and North America are dominated by A. pedellus alone. Within A. pedellus, patterns of sequence diversity were indicative of a recent population expansion. In the western US, the phenology of a population of A. fimetarius was observed to significantly differ from that of a sympatric population of A. pedellus, thereby revealing an ecological difference between the two cryptic taxa. Overall, we conclude that all types of characters offered a consistent classification of the two species. Thus, the laborious karyotyping techniques used to originally establish the presence of two cryptic taxa can now be substituted by characters more easily applied to large ecological samples. Using this approach of integrative taxonomy, we were able to establish the global distribution and species‐specific ecology of these ecologically important cryptic taxa. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:4033473E-8BF7-40F4-852D-916E4F858593 .     

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.     

Neuropharmacological evidence for inhibitory cephalic control mechanisms of stridulatory behaviour in grasshoppers

In gomphocerine grasshoppers the neuromuscular patterns of stridulatory hindleg movements are produced by metathoracic rhythm generators under the control of cephalic command neurons. Injections of cholinergic agonists into the protocerebrum activate this command system which induces the performance of stridulatory sequences, resembling natural species specific movements. Injections of GABA, glycine and picrotoxin into the central protocerebrum of the species Omocestus viridulus, Chorthippus mollis and Ch. biguttulus revealed a contribution of inhibitory mechanisms to the control of the stridulatory behaviour. The experiments suggest that inhibition interferes with the cephalic command systems at three levels: (1) sustained inhibition through picrotoxin sensitive receptors acting on all command units while grasshoppers are at rest, and during stridulation on all command units except the one activating the pattern generators of the currently performed movements; (2) premature termination of song sequences, experimentally induced by injections of GABA and glycine; and (3) coupling of a timing mechanism that terminates a song sequence or its subunits with a particular movement pattern after specific durations. These results together with those from previous studies on the pharmacological activation of stridulatory behaviour suggest that a balance of inhibitory and excitatory inputs to the command system selects the appropriate song type and controls its performance. Accepted: 11 June 1998     

Separate localization of sound recognizing and sound producing neural mechanisms in a grasshopper

Summary In the two acridid speciesChorthippus parallelus andCh. montanus, the sound template by which females recognize male song varies with temperature, as does the song itself. At relatively high temperatures the females respond best to simulated songs with high syllable frequencies, and at lower temperatures songs with lower syllable frequencies are preferred.The temperature around the supraesophageal and metathoracic ganglia of female grasshoppers was monitored by implanted thermocouples, and either the head or the thorax was warmed selectively while the animal was free to move (within the imits of the wires). Then simulations of the conspecific song varying in syllable frequency corresponding to different song temperatures were presented, and the stridulatory responses of the animals were observed.The results were as follows. 1. Song recognition (in particular, the position of the peak of the response curve) depended on the temperature of the head. 2. The rate of stridulatory hindleg movement was determined by the temperature of the thoracic ganglia.This result provides strong evidence against the genetic coupling hypothesis.     

Acoustic defence in an insect: characteristics of defensive stridulation and differences between the sexes in the tettigoniid Poecilimon ornatus (Schmidt 1850)

Many insects exhibit secondary defence mechanisms upon contact with a predator, such as defensive sound production or regurgitation of gut contents. In the tettigoniid Poecilimon ornatus, both males and females are capable of sound production and of regurgitation. However, wing stridulatory structures for intraspecific acoustic communication evolved independently in males and females, and may result in different defence sounds. Here we investigate in P. ornatus whether secondary defence behaviours, in particular defence sounds, show sex-specific differences. The male defence sound differs significantly from the male calling song in that it has a longer syllable duration and a higher number of impulses per syllable. In females, the defence sound syllables are also significantly longer than the syllables of their response song to the male calling song. In addition, the acoustic disturbance stridulation differs notably between females and males as both sexes exhibit different temporal patterns of the defence sound. Furthermore, males use defence sounds more often than females. The higher proportion of male disturbance stridulation is consistent with a male-biased predation risk during calling and phonotactic behaviour. The temporal structures of the female and male defence sounds support a deimatic function of the startling sound in both females and males, rather than an adaptation for a particular temporal pattern. Independently of the clear differences in sound defence, no difference in regurgitation of gut content occurs between the sexes.     

Habitat and weather affect reproduction and size of the dung beetle Aphodius fossor

1. The influence of sun exposure and moisture on reproduction and size of a north temperate dung beetle Aphodius fossor was investigated. Adult beetles were added to cow pats that were either shaded or exposed and treated with different watering regimes. 2. After 26 days, more eggs and larvae were recovered from shaded and wetter pats than from exposed and drier pats. 3. At the end of the experiment, more beetles had developed in shaded and wetter pats, and these beetles were larger than those from exposed and drier pats. 4. There was no between‐treatment difference in the ratio between adults in final sampling and larvae in first sampling. 5. It was concluded that exposure and moisture can influence both reproduction and size of A. fossor. It is suggested that weather may be a factor regulating reproduction in Aphodius not only through heat‐ or drought‐induced mortality, as suggested earlier, but also by limiting the time available for egg laying. Further studies are needed to clarify this suggestion.     

Patch size and colonisation patterns: an experimental analysis using north temperate coprophagous dung beetles

The relationship between dung pad size and both adult colonisation and larval development was investigated in an assemblage of north temperate dung beetles ( Geotrupes. Aphodius and Sphaeridium ) using both dung pads and baited pitfall traps. Wet weight of 22-day-old natural dung pads was found to vary widely in the field (< 100 g – >1000 g). Across all sampling dates in field experiments, dung pad size had a significant influence on dung beetle biomass sampled from pads. Closer examination of experimental dung pads on the second day after deposition, when beetle biomass was at a maximum, revealed not only a general positive relationship between pad size and dung beetle biomass but, more importantly, a positive relationship between dung pad size and dung beetle density (dung beetle biomass per unit dung volume). There was a strong trend for Aphodius species richness to increase, and maintain higher values for longer periods of time, in larger pads. Although dung pad and pitfall trap samples could differ in the actual numbers of beetles captured, the relationship between different dung sizes and dung beetle biomass was similar, indicating that the phenomenon is largely related to immigration processes. Pat residence times oi A. rufipes in the laboratory were significantly positively correlated with dung pad size. In two field experiments, positive correlations were found between dung pad size and numbers of larvae in pads of different sizes and in one of these experiments, larval densities (numbers per unit dung volume) were significantly and positively correlated with dung pad size. In one experiment, Aphodius larvae in the early stages of development were found to preferentially occupy the basal area of dung pads. We discuss the implications of our findings in the context of resource utilisation by north temperate dung beetles.     

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.

     

Dung beetle movements at two spatial scales

To understand the dynamics of spatially structured populations, we need to know the level of movements at different spatial scales. This paper reports on Aphodius dung beetle movements at two scales: movements between dung pats within pastures, and movements between pastures. First, I test an assumption common to many recent models of spatially structured populations – that the probability of an individual moving between habitat patches decreases exponentially with distance. For dung beetles, I find sufficient evidence to reject this assumption. The distribution of dispersal distances was clearly leptokurtic, with more individuals moving short and long distances than expected on the basis of an exponential function. In contrast, the data were well described by a power function. I conclude that dung beetle movements include an element of non-randomness not captured by the simplistic exponential model. The power function offers a promising alternative, but the actual mechanisms behind the pattern need to be clarified. Second, I compare several species of Aphodius to each other. Although these species occur in the same network of habitat patches, their movement patterns are different enough to result in a mixture of different spatial population structures. Movements between pastures were more frequent the larger the species, the more specific its occurrence in relation to pat age, and the more specialized it is on cow dung and open pasture habitats. Within pastures, all species form "patchy" populations, with much movement among individual pats.     

Sound production in Parapellopedon instabilis (Rehn, 1906) (Orthoptera: Gomphocerinae)

The sounds produced by Parapellopedon instabilis (Rehn, 1906), are described for the first time on the basis of recordings made, in captivity, with an analogical tape recorder. The signals were digitized in the laboratory and analyzed using a software. Three types of song are described: the male calling song, typical of the gomphocerinae species, the female’s agreement song, less organized temporally and unusually loud for a gomphocerinae species, and disturbance songs among males and among females, which follow the typical structure of these signals in the subfamily. Oscillograms and frequency spectra of all songs are given. The stridulatory file of both sexes, male and female, are described.     

Recognition of sex in the acoustic communication of the grasshopper Chorthippus biguttulus (Orthoptera, Acrididae)

Many gomphocerine grasshoppers communicate acoustically: a male's calling song is answered by a female which is approached phonotactically by the male. Signals and recognition mechanisms were investigated in Chorthippus biguttulus with regard to the cues which allow sex discrimination. (1) The stridulatory files on the hindfemur of both sexes are homologous in that they are derived from the same row of bristles, but convergent with respect to the “pegs”. In males the pegs are derived from the bristles, and in females from the wall of the bristle's cup. (2) Male and female songs are generated by similar, probably homologous motor programs, but differ in the duration, intensity, “gappyness” of syllables, risetime of pulses, and the frequency spectra. The hindleg co-ordination during stridulation and the resulting temporal song patterns are less variable in males than in females. (3) For both sexes, recognition of a mate's signal depends on species-specific syllable structure. For males it is essential that the female syllables consist of distinct short pulses, whereas females reject “gappy” syllables. Males strongly prefer “ramped” pulses, females respond to syllables irrespective of steeply or slowly rising ramps. Males react only to the low-frequency component, whereas females prefer spectra containing both, low and high frequency components. Accepted: 20 November 1996     

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.     

A role of her own: female cowbirds, Molothrus ater, influence the development and outcome of song learning

Previous work has shown that captive female cowbirds, Molothrus ater, can influence the outcome of male song development by affecting retention or deletion of song elements and by stimulating improvization. Here we looked for evidence of female influence during the process of learning, as males progress from subsong to plastic song to stereotyped song. In a longitudinal study, we measured the rate and timing of vocal development in captive, juvenile male brown-headed cowbirds, M. a. artemisiae. Half the young males were housed with female cowbirds from their own population (South Dakota: SD) and half with female cowbirds from a M. a. ater population (Indiana: IN). Both populations of females prefer local songs and differ in the time of breeding, with SD females breeding 2 weeks later than IN females. The results showed significant effects of female presence on the age at which males advanced through stages of vocal development: the SD males with SD females, as opposed to SD males with IN females, developed stereotyped song earlier, reduced motor practise earlier, and produced more effective playback songs. Longitudinal observations of social interactions showed that the two groups of females reliably differed in social responses to males. Degree of social proximity of females to males in the winter predicted song maturity, rate of rehearsal and song potency. Thus, females can stimulate the progression of song learning, as well as prune song content. Copyright 2000 The Association for the Study of Animal Behaviour.     

Response of the cercus-to-giant interneuron system in crickets to species-specific song

Summary In addition to their high frequency stridulatory sound, crickets (Gryllus campestris) also produce low frequency airborne vibrations resulting from the strokes of the crickets' wings closing and opening during stridulation. Giant interneurons of the ventral cord, which receive inputs from cereal hairs, respond to these low frequencly components of cricket song up to a distance of some ten cm. The discharges are correlated to the time course of the acoustic stimulation, therefore allowing the transmission of the time patterns (syllables, chirps) of calling song, rivalry song and even courtship song. With simultaneous recording from both sides of the abdominal nerve cord, synchronous or alternating discharges can be detected, correlated either with syllables or intervals. The response mode depends on the position of the two individuals in relation to each other. Recording directly from singing males demonstrates interneuron response to selfgenerated signals, too. A possible communicative function of the system is discussed.Supported by the Deutsche Forschungsgemeinschaft as part of the program Neurale Mechanismen des Verhaltens (Da 61/8, 61/9)We thank Dr. F. Huber and Dr. H. Scharstein for helpful comments.