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.     

Experimental warming disrupts reproduction and dung burial by a ball‐rolling dung beetle

1. Insects are sensitive to climate change. Consequently, insect‐mediated ecosystem functions and services may be altered by changing climates. 2. Dung beetles provide multiple services by burying manure. Using climate‐controlled chambers, the effects of warming on dung burial and reproduction by the dung beetle Sisyphus rubrus Paschalidis, 1974 were investigated. Sisyphus rubrus break up dung by forming and rolling away balls of manure for burial and egg deposition. 3. To simulate warming in the chambers, 0, 2 or 4 °C offsets were added to field‐recorded, diurnally fluctuating temperatures. We measured dung ball production and burial, egg laying, survival and residence times of beetles. 4. Temperature did not affect the size or number of dung balls produced; however warming reduced dung ball burial by S. rubrus. Because buried balls were more likely to contain eggs, warming could reduce egg laying via a reduction in ball burial. Warming reduced the humidity inside the chambers, and a positive relationship was found between the number of dung balls produced and humidity in two temperature treatments. Temperature did not affect survival, or whether or not a beetle left a chamber. Beetles that did leave the chambers took longer to do so in the warmest treatment. 5. This study demonstrates that climate warming could reduce reproduction and dung burial by S. rubrus, and is an important first step to understanding warming effects on burial services. Future studies should assess warming effects in field situations, both on individual dung beetle species and on aggregate dung burial services.     

Asymmetric competition in cattle dung between two species of Onthophagus dung beetle and the bush fly, Musca vetustissima

Abstract.

• 1 Competition in cattle dung pads between two dung beetles, Onthophagus ferox Harold and Onthophagus binodis Thunberg, and the bush fly, Musca vetustissima Walker, was investigated in laboratory experiments, to determine why spring fly abundance in the field did not fall following the introduction of O. binodis.
• 2 At low beetle densities, the number of eggs laid by each species was reduced by the second species. A similar amount of dung was buried by each species alone or by both together.
• 3 At high beetle densities O. binodis egg production was substantially affected by each additional O.ferox, but O.ferox egg production was not affected by each additional O.binodis. Asymmetric competition occurred because O.ferox buried more dung than O.binodis, and a greater proportion in day 1 (pre-emptive dung burial).
• 4 O.ferox caused greater M. vetustissima egg-puparia mortality than O. binodis. Mortality mostly occurred in young M. vetustissima larvae less than 1 day old. Total egg-puparia fly mortality was correlated better with the dung buried on day 1 than dung buried on day 8 (pre-emptive dung burial). O.binodis did not add to fly mortality by O.ferox at high densities because of asymmetric competition between the beetles.

     

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     

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.     

Functional implications of temperature‐correlated colour polymorphism in an iridescent,scarabaeine dung beetle

Abstract 1. In many Coleoptera, iridescent colouration is generated by exoskeleton ultra‐structure, within which multilayer interference reflects only certain wavelengths. Published work indicates that the colour polymorphism shown by some iridescent beetles is under genetic control. However, the present study suggests environmental involvement in the polymorphic southern African dung beetle, Gymnopleurus humanus Macleay. 2. At 24 study sites across a 1000‐km latitudinal temperature gradient, population samples of G. humanus were dominated by blue individuals in the cooler south, by cupreous individuals in the warmer north, and by locally co‐occurring blue, green and cupreous individuals in intermediate situations. 3. Using digital reflectance spectrophotometry, we measured wavelength intensity values across the visible spectrum (400–800 nm) and used the 70 measured specimens to estimate maximum reflectance from a further 3912 beetles. Differences in mean reflectance values between 24 populations were strongly correlated with average annual temperatures at study sites. 4. Much stronger correlations between mean reflectance values and average temperatures of the cool dry season months suggest that the cross‐climatic patterns may be related to interaction between breeding seasonality and development under different cooler temperatures. 5. Published evidence suggests that inherent physical properties of cholesteric liquid crystals and their responses to different thermal conditions could, potentially, generate the different exocuticle ultra‐structure responsible for different reflected colour wavelengths. Furthermore, colour polymorphism could be advantageous across a gradient from cooler to warmer climate as a result of the different thermal properties of different colours. 6. Given the correlation with temperature, it is predicted that the prevailing reflected colour balance in southern populations would shift in response to global climatic change.     

Local extinctions may be evidenced by the holes of the morphometric hypervolume in dung beetle communities

The body shape of a species is associated with its evolutionary history and can reflect behavioural peculiarities related to the ecological niche of each species. Morphology can characterise the morphometric niche of species and can be represented as body shape points within a morphometric universe. This information can be to calculate the morphometric diversity of communities through hypervolume metrics, and the hole sizes that remain in the morphometric hypervolume, which are empty areas with no species. Such holes may be ‘natural’ or caused by a local extinction. In this study, we evaluate the ecological community of dung beetles through the lens of morphometric diversity. We evaluated 38 dung beetle species from 30 subtropical communities in southern Brazil sampled in the summer of 2015, including 15 forest remnant communities from the Atlantic Forest and 15 communities from adjacent maize cultivations. The shape of 495 dung beetle specimens was measured using geometric morphometric and hypervolume techniques to calculate the morphometric diversity and the hole size of each of the 30 communities. We found that the taxonomic diversity positively correlated with the morphometric diversity and negatively correlated with the size of the holes. We also found that forest communities had higher values of morphometric diversity and smaller holes in the hypervolume than the maize cultivation communities, suggesting that local extinction may reduce community body shape spaces.     

Maternal and paternal effects on offspring phenotype in the dung beetle Onthophagus taurus

Abstract.— Parents often have important influences on the development of traits in their offspring. One mechanism by which parents are able to influence offspring phenotype is through the level of care they provide. In onthophagine dung beetles, parents typically provision their offspring by packing dung fragments into a brood mass. Onthophagus taurus males can be separated into two discrete morphs: Large, "major" males have head horns, whereas "minor" males are hornless. Here we show that a switch in parental provisioning strategies adopted by males coincides with the switch in male morphology. Male provisioning results in the production of heavier brood masses than females will produce alone. However, unlike females in which the level of provisioning increases with body size in a continuous manner, the level of provisioning provided by males represents an "all-or-none" tactic with all major males providing a fixed level of provisioning irrespective of their body size. Offspring size is determined largely by the quantity of dung provided to the developing larvae so that paternal and maternal provisioning affects the body size and horn size of offspring produced. The levels of provisioning by individual parents are significantly repeatable, suggesting paternal and maternal effects as candidate indirect genetic effects in the evolution of horn size in the genus Onthophagus .     

A stable,genetically determined colour dimorphism in the dung beetle Aphodius depressus: patterns and mechanisms

1. Melanism – the occurrence of dark morphs – in insects has been attributed to differences in, among other things, thermoregulation and immune defence. Dark individuals are hypothesised to perform better in colder areas, and to exhibit stronger melanin‐based immune defence. 2. In the present study, the geographical distribution of two colour morphs in Aphodius depressus (Kugelann), its climatic correlates, and temporal stability was described. Underlying mechanisms were then targeted through experiments: the inheritance of colour through controlled crosses, heating rates by thermal imaging, physiological tolerance by critical thermal limits, and immune efficiency by melanisation of implants. 3. In A. depressus, colour appears inherited by simple Mendelian principles, with red dominating over black. The frequency of two colour morphs forms a large‐scale cline. In the South West of Finland, all individuals are black, whereas, in the North East, most are red. This pattern has remained constant over 13 years (1996–2008). 4. The geographical pattern was not attributable to thermoregulation: black morphs were more abundant in warmer rather than colder parts of the country. In experiments, we found no differences in the heating rate of the two morphs, or in their upper temperature maxima. Neither did the morphs differ in their response to artificial objects inserted in their haemolymph. 5. Overall, colour variation in A. depressus occurs as a stable, genetically determined dimorphism, governed by Mendelian inheritance. Yet, no support for prevailing theory of factors sustaining melanism was found. The reasons for colour polymorphism in insects may thus be complex, and should be sought on a case‐by‐case basis.     

Genetic diversity of the dung beetle,Copris tripartitus (Coleoptera: Scarabaeidae), that is endangered in Korea

The dung beetle, Copris tripartitus (Coleoptera: Scarabaeidae), is an endangered insect in Korea. In order to establish a conservation strategy, a preliminary investigation on the genetic diversity of Korean populations was performed using mitochondrial COI (658 bp), CytB (433 bp), and nuclear ITS2 (411 nucleotide positions). Sequencing of 69 individuals collected from five localities showed substantially higher variability (5.02% for COI, 4.62% for CytB, and 8.03% for ITS2). The resulting networks for mitochondrial DNA (mtDNA) haplotypes exhibited two star‐like phylogenies, which might indicate that Korean populations have recently expended from two small populations. The ITS2 network, which was presented in the form of a star‐like phylogeny, confirmed that a recent population expansion occurred. Considering the high genetic diversity and gene flow in C. tripartitus populations, one issue regarding conservation seems to be the recovery of previous habitats.     

First discovery of fossil dung beetle brood balls and nests in the Chadian Pliocene Australopithecine levels

The Pliocene continental formations of the paleo-lake Chad system are known because of the recent discovery of the first australopithecine known west of the Rift Valley. The structures under study are found in sandstone levels associated with a rich fauna, including mammals, birds, reptiles and fishes. Analysis of the depositional environment and fauna indicates a mosaic landscape of gallery forest, savannah, grassland and ephemeral rivers interrupted by lacustrine episodes. This sandstone facies contains bioturbation in the form of sandstone balls 4-12 cm in diameter, slightly flattened at the poles. These structures are characterized by an external husk or crust and by a decimetric cavity in the upper part of the ball. Between the husk and the cavity are a number of concave laminae similar to those of a bulb, whose concavity is directed toward the upper cavity. The comparison between these structures and the brood balls of modern Scarabaeidae shows great similarity, especially in the external husk, the concave internal laminae and the chamber of the grub in the upper part of the structures. This bioturbation is interpreted as fossil brood balls of dung beetles (Coleoptera, Scarabaeidae). Fossil dung beetle brood balls are generally rare, but have been known since the 1940s. They can be very abundant in any series, as described by several authors in South America. The first fossil dung balls were described in this area in 1938 by Frenguelli and by Roselli. These authors describe elementary spherical forms of 35 mm in diameter on average, flattened at the poles and with an upper cavity. The fossil dung beetle brood balls discovered in Chad are the first in which all the internal characteristic structures are preserved. Many of them are connected by a remarkably large net of tunnels which has no parallel in the past or the present.     

Evidence for sex‐specific selection in brain: a case study of the nine‐spined stickleback

Theory predicts that the sex making greater investments into reproductive behaviours demands higher cognitive ability, and as a consequence, larger brains or brain parts. Further, the resulting sexual dimorphism can differ between populations adapted to different environments, or among individuals developing under different environmental conditions. In the nine‐spine stickleback (Pungitius pungitius), males perform nest building, courtship, territory defence and parental care, whereas females perform mate choice and produce eggs. Also, predation‐adapted marine and competition‐adapted pond populations have diverged in a series of ecologically relevant traits, including the level of phenotypic plasticity. Here, we studied sexual dimorphism in brain size and architecture in nine‐spined stickleback from marine and pond populations reared in a factorial experiment with predation and food treatments in a common garden experiment. Males had relatively larger brains, larger telencephala, cerebella and hypothalami (6–16% divergence) than females, irrespective of habitat. Females tended to have larger bulbi olfactorii than males (13%) in the high food treatment, whereas no such difference was found in the low food treatment. The strong sexual dimorphism in brain architecture implies that the different reproductive allocation strategies (behaviour vs. egg production) select for different investments into the costly brains between males and females. The lack of habitat dependence in brain sexual dimorphism suggests that the sex‐specific selection forces on brains differ only negligibly between habitats. Although significance of the observed sex‐specific brain plasticity in the size of bulbus olfactorius remains unclear, it demonstrates the potential for sex‐specific neural plasticity.     

Genetic correlations and sex‐specific adaptation in changing environments

Females and males have conflicting evolutionary interests. Selection favors the evolution of different phenotypes within each sex, yet divergence between the sexes is constrained by the shared genetic basis of female and male traits. Current theory predicts that such “sexual antagonism” should be common: manifesting rapidly during the process of adaptation, and slow in its resolution. However, these predictions apply in temporally stable environments. Environmental change has been shown empirically to realign the direction of selection acting on shared traits and thereby alleviate signals of sexually antagonistic selection. Yet there remains no theory for how common sexual antagonism should be in changing environments. Here, we analyze models of sex‐specific evolutionary divergence under directional and cyclic environmental change, and consider the impact of genetic correlations on long‐run patterns of sex‐specific adaptation. We find that environmental change often aligns directional selection between the sexes, even when they have divergent phenotypic optima. Nevertheless, some forms of environmental change generate persistent sexually antagonistic selection that is difficult to resolve. Our results reinforce recent empirical observations that changing environmental conditions alleviate conflict between males and females. They also generate new predictions regarding the scope for sexually antagonistic selection and its resolution in changing environments.     

Genomic evidence that resource‐based trade‐offs limit host‐range expansion in a seed beetle

Trade‐offs have often been invoked to explain the evolution of ecological specialization. Phytophagous insects have been especially well studied, but there has been little evidence that resource‐based trade‐offs contribute to the evolution of host specialization in this group. Here, we combine experimental evolution and partial genome resequencing of replicate seed beetle selection lines to test the trade‐off hypothesis and measure the repeatability of evolution. Bayesian estimates of selection coefficients suggest that rapid adaptation to a poor host (lentil) was mediated by standing genetic variation at multiple genetic loci and involved many of the same variants in replicate lines. Sublines that were then switched back to the ancestral host (mung bean) showed a more gradual and variable (less repeatable) loss of adaptation to lentil. We were able to obtain estimates of variance effective population sizes from genome‐wide differences in allele frequencies within and between lines. These estimates were relatively large, which suggests that the contribution of genetic drift to the loss of adaptation following reversion was small. Instead, we find that some alleles that were favored on lentil were selected against during reversion on mung bean, consistent with the genetic trade‐off hypothesis.     

Duration and benefits of biparental brood care in the dung beetle Onthophagus vacca (Coleoptera: Scarabaeidae)

Abstract.

• 1 Single males, single females or pairs of dung beetles, Onthophagus vacca, were released on artificial small (100 g) or large (1000 g) dung pats in the laboratory. Emigrating beetles were trapped at 12 h intervals, and the number and size of the brood chambers were recorded after each replicate.
• 2 Emigration of males was delayed if females were present in the same dung pats, whereas emigration times of females were independent of the presence or absence of males.
• 3 A residency of 60 h proved to be a threshold value. Females emigrating before this time did not breed, whereas those emigrating later had built at least two brood chambers.
• 4 Females paired with males built more brood chambers than single females.
• 5 The reproductive success of pairs was not influenced by the size of the dung pats.

     

Experimental evidence for the effects of dung beetle functional group richness and composition on ecosystem function in a tropical forest

Much of the literature on the relationship between species richness or functional group richness and measures of ecosystem function focuses on a restricted set of ecosystem function measures and taxonomic groups. Few such studies have been carried out under realistic levels of diversity in the field, particularly in high diversity ecosystems such as tropical forests. We used exclusion experiments to study the effects of dung beetle functional group richness and composition on two interlinked and functionally important ecological processes, dung removal and secondary seed dispersal, in evergreen tropical forest in Sabah, Malaysian Borneo. Overall, both dung and seed removal increased with dung beetle functional group richness. However, levels of ecosystem functioning were idiosyncratic depending on the identity of the functional groups present, indicating an important role for functional group composition. There was no evidence for interference or competition among functional groups. We found strong evidence for overyielding and transgressive overyielding, suggesting complementarity or facilitation among functional groups. Not all mixtures showed transgressive overyielding, so that complementarity was restricted to particular functional group combinations. Beetles in a single functional group (large nocturnal tunnellers) had a disproportionate influence on measures of ecosystem function: in their absence dung removal is reduced by approximately 75%. However, a full complement of functional groups is required to maximize ecosystem functioning. This study highlights the importance of both functional group identity and species composition in determining the ecosystem consequences of extinctions or altered patterns in the relative abundance of species.     

No evidence for size‐assortative mating in the wild despite mutual mate choice in sex‐role‐reversed pipefishes

Size‐assortative mating is a nonrandom association of body size between members of mating pairs and is expected to be common in species with mutual preferences for body size. In this study, we investigated whether there is direct evidence for size‐assortative mating in two species of pipefishes, Syngnathus floridae and S. typhle, that share the characteristics of male pregnancy, sex‐role reversal, and a polygynandrous mating system. We take advantage of microsatellite‐based “genetic‐capture” techniques to match wild‐caught females with female genotypes reconstructed from broods of pregnant males and use these data to explore patterns of size‐assortative mating in these species. We also develop a simulation model to explore how positive, negative, and antagonistic preferences of each sex for body size affect size‐assortative mating. Contrary to expectations, we were unable to find any evidence of size‐assortative mating in either species at different geographic locations or at different sampling times. Furthermore, two traits that potentially confer a fitness advantage in terms of reproductive success, female mating order and number of eggs transferred per female, do not affect pairing patterns in the wild. Results from model simulations demonstrate that strong mating preferences are unlikely to explain the observed patterns of mating in the studied populations. Our study shows that individual mating preferences, as ascertained by laboratory‐based mating trials, can be decoupled from realized patterns of mating in the wild, and therefore, field studies are also necessary to determine actual patterns of mate choice in nature. We conclude that this disconnect between preferences and assortative mating is likely due to ecological constraints and multiple mating that may limit mate choice in natural populations.