Male Distribution and Mate Searching in the Yellow Dung Fly Scathophaga stercoraria: Comparison between Paired and Unpaired Males

Movements by individual males were examined in the yellow dung fly, Scathophaga stercoraria. Males were observed from their arrival until they found a female (paired males) or departed (unpaired males). The focal variables were the male mating status, body size and the number of males and females at the site. Paired males, independent of size, spent more time in the best mate-searching area (pat and the first up-wind zone) than unpaired males. Paired males in all size classes moved around and attacked other males more often than unpaired males. Among paired males, males that caught a single female and those that took over a female from another male were very similar in their mate-searching behaviour. The total time spent searching at the pat was positively related to its resource value as indicated by the number of pairs. Time spent in the best mate-searching area was negatively related with male numbers. The causes of differences in movements and aggression between paired and unpaired males are discussed. The male distribution around cow pats can be understood only if the differences in movement patterns by paired and unpaired males are taken into account.     

Altitudinal life history variation in the dung flies Scathophaga stercoraria and Sepsis cynipsea

 Field phenologies of high- (ca. 1500 m) and low- (ca. 500 m) altitude populations of the two most common European species of dung flies, Scathophaga stercoraria and Sepsis cynipsea, differ quite markedly due to differences in climate. To differentiate genetic adaptation due to natural selection and phenotypic plasticity, I compared standard life history characters of pairs of high- and low-altitude populations from three disjunctive sites in Switzerland in a laboratory experiment. The F1 rearing environment did not affect any of the variables of the F2 generation with which all experiments were conducted; hence, there were no carry-over or maternal effects. In Sc. stercoraria, high-altitude individuals were smaller but laid larger eggs; the latter may be advantageous in the more extreme (i.e. more variable and less predictable) high-altitude climate. Higher rearing temperature strongly decreased development time, body size and the size difference between males and females (males are larger), produced female-biased sex ratios and led to suboptimal adult emergence rates. Several of these variables also varied among the three sites, producing some interactions complicating the patterns. In Se. cynipsea, high-altitude females were marginally smaller, less long-lived and laid fewer clutches. Higher rearing temperature strongly decreased development time and body size but tended to increase the size difference between males and females (males are smaller); it also increased clutch size but decreased physiological longevity. Again, interpretation is complicated by variation across sites and some significant interactions. Overall, genetic adaptation to high-altitude conditions appears weak, probably prevented by substantial gene flow, and may be swamped by the effects of other geographic variables among populations. In contrast, phenotypic plasticity is extensive. This may be due to selection of flexible, multi-purpose genotypes. The results suggest that differences in season length between high- and low-altitude locations alone do not explain well the patterns of variation in phenology and body size. Received: 21 March 1996 / Accepted: 1 September 1996     

Sex and immunity in the yellow dung fly Scathophaga stercoraria

The immune system is of increasing interest to evolutionary biologists. Immunity may trade-off against other fitness components, with recent work suggesting reproduction in particular impinges on immune defence. There may also be sex differences in the immune system. Additionally, while life history traits typically have low heritability, little is known about additive genetic variance of immunity. An insect's major defence against multicellular pathogens is to encapsulate the invader. Phenoloxidase (PO) is a key enzyme in the cascade resulting in the melanized capsule, and is often used to estimate resistance to an immune insult. We examined the effects of copulation, egg laying, sex and age on PO in Scathophaga stercoraria. We also measured the heritability of PO activity. The sexes differed in haemolymph PO activity and PO was significantly affected by age, but not by copulation or egg laying. There was significant heritable variation for haemolymph PO.     

Oviposition Site Choice and Life History Evolution

SYNOPSIS. Studies of life history evolution, as well as muchof life history theory, have typically focused on "hard" componentsof life histories; phenotypic characteristics that can be readilyobserved, quantified, and ultimately, connected rather directlyto fitness. Typical of these are propagule size, propagule number,and age and size at maturity. What is largely missing from thestudy of life history evolution is consideration of the roleof behavior, principally female oviposition site choice, inthe evolution of life histories. For oviparous organisms, naturalselection cannot produce locally optimized "hard" componentsof life history phenotypes without a consistent environmentalcontext (whether invariant orvariable); in a variable environment,that consistent environmental context can be most effectivelyprovided by interactive oviposition site choice. I present amodel of selection on oviposition site choice in the contextof the evolution of "hard" components of life history phenotypes,along with some experimental data illustrating oviposition sitechoice in response to predators. The model and data are thenrelated to the overall question of the role of oviposition sitechoice in life history evolution. The conclusion is that ovipositionsite choice must be under equally strong selection with eggsize, egg number and the other hard components of life historiesin order to generate and optimize locally adapted or ecologicallyspecialized life history phenotypes, and must therefore, playa significant role in the evolution of life histories.     

Post-copulatory sexual selection and female fitness in Scathophaga stercoraria

Whether sexual selection increases or decreases female fitness is determined by the occurrence and relative importance of sexual-conflict processes and the ability of females to choose high-quality males. Experimentally enforced polyandry and monogamy have previously been shown to cause rapid evolution in the yellow dung fly Scathophaga stercoraria. Flies from polyandrous lines invested more in reproductive tissue, and this investment influenced paternity in sperm competition, but came at a cost to immune function. While some fitness consequences of enforced polyandry or monogamy have been examined when flies mate multiply, the consequences for female fitness when singly copulated remain unexplored. Under a good-genes scenario females from polyandrous lines should be of higher general quality and should outperform females from monogamous lines even with a single copulation. Under sexual conflict, costly adaptations will afford no advantages when females are allowed to mate only once. We investigate the lifetime reproductive success and longevity of females evolving under enforced monogamy or polyandry when mating once with males from these selection regimes. Females from polyandrous lines were found to have lower fitness than their monogamous counterparts when mating once. They died earlier and produced significantly fewer eggs and offspring. These results suggest that sexual conflict probably drove evolution under enforced polyandry as female fitness did not increase overall as expected with purely good-genes effects.     

Size-Dependent Mating Success at Various Nutritional States in the Yellow Dung Fly

Mating success not only depends on genetic quality, but also equally on environmental factors, most prominently food availability. We investigated the interactive effects of nutritional state and body size on mating success and copula duration in yellow dung fly males ( Scathophaga stercoraria ; Diptera: Scathophagidae) of three body size selection lines in the laboratory in both non-competitive (single) and competitive (group) situations. Adults require protein and lipids from prey to reproduce, as well as sugars as an energy source for reproductive activity. We expected mating success to decrease with time because of sperm depletion (sugar treatment) and/or energy shortage (water treatment) relative to the control, prey plus sugar treatment. Based on physiological scaling, we also expected large-line males to become depleted either sooner because of their higher energy and sperm demands, or later because of their more efficient energy use. Average mating success indeed declined over a period of 5–7 d (or 5–15 potential copulations per male), but equally for all food treatments and body size classes. Surprisingly, water-fed and small-line males had the highest mating probability in the non-competitive setting, while in the competitive setting large-line males had the highest success. Energy-depleted males showed apparent terminal investment. Small males acquired females more readily but eventually lost them to larger males in the competitive situation. As shown before, copula duration was inversely related to body size and increased with copulation number, independent of food treatment. We conclude that sugar or prey shortage has little effect on mating success in the short term, and does not differentially affect males of different sizes.     

Phenoloxidase activity and pathogen resistance in yellow dung flies Scathophaga stercoraria

Phenoloxidase (PO) is an important component of the insect immune system and is frequently used to measure an individual's immune defence ability. However, evidence documenting positive correlations between the immune assay and resistance against pathogens is scarce and contradictory. We used replicate lines of yellow dung flies Scathophaga stercoraria (L.) with different PO levels to investigate whether PO levels affect resistance against parasitic mites and entomopathogenic fungi. Prevalence of flies exposed to pathogens was the same in all selection regimes, although pathogens clearly negatively affected fitness. PO measurements alone therefore do not necessarily predict overall resistance against pathogens. Furthermore, under starvation lines selected for high PO levels did not survive longer than those selected for low PO levels, irrespective of exposure to pathogens. This suggests that even if elevated immune levels increase an individual's ability to combat pathogens, the benefits may not outweigh the costs of increased investment in immunity.     

Maternal effects and their consequences for offspring fitness in the Yellow Dung Fly

1. Maternal adult diet and body size influence the fecundity of a female and possibly the quality and the performance of her offspring via egg size or egg quality. In laboratory experiments, negative effects in the offspring generation have often been obscured by optimal rearing conditions.
2. To estimate these effects in the Yellow Dung Fly, Scathophaga stercoraria , how maternal body size and adult nutritional status affected her fecundity, longevity and egg size were first investigated.
3. Second, it was investigated how female age and adult nutritional experience, mediated through the effects of egg size or egg quality, influenced the performance of offspring at different larval densities.
4. Maternal size was less important than maternal adult feeding in increasing reproductive output. Without food restriction, large females had larger clutch sizes and higher oviposition rates, whereas under food restriction this advantage was reversed in favour of small females.
5. Offspring from mothers reared under nutritional stress experienced reduced fitness in terms of egg mortality and survival to adult emergence. If the offspring from low-quality eggs survived, the transmitted maternal food deficiency only affected adult male body size under stressful larval environments.
6. Smaller egg sizes due to maternal age only slightly affected the performance of the offspring under all larval conditions.     

Size-dependent alternative male mating tactics in the yellow dung fly,Scathophaga stercoraria

Whenever males can monopolize females and/or resources used by females, the opportunity for sexual selection will be great. The greater the variation among males in reproductive success, the greater the intensity of selection on less competitive males to gain matings through alternative tactics. In the yellow dung fly, Scathophaga stercoraria, males aggressively compete for access to receptive, gravid females on fresh dung. Larger males are better able to acquire mates and to complete copulation successfully and guard the female throughout oviposition. Here we demonstrate that when an alternative resource is present where females aggregate (i.e. apple pomace, where both sexes come to feed), smaller males will redirect their searching for females from dung to the new substrate. In addition, we identify a class of particularly small males on the alternative substrate that appears never to be present searching for females on or around dung. Smaller males were found to have a mating ‘advantage’ on pomace, in striking contrast to the pattern observed on dung, providing further support for the existence of an alternative male reproductive tactic in this species.     

Sperm survival in the female reproductive tract in the fly Scathophaga stercoraria (L.)

While sperm competition risk favours males transferring many sperm to secure fertilizations, females of a variety of species actively reduce sperm numbers reaching their reproductive tract, e.g. by extrusion or killing. Potential benefits of spermicide to females include nutritional gains, influence over sperm storage and paternity, and the elimination of sperm bearing somatic mutations that would lower zygote fitness.We investigated changes in sperm viability after in vivo and in vitro exposure to the female tract in the polyandrous fly, Scathophaga stercoraria. Sperm viability was significantly lower in the females' spermathecae immediately after mating than in the experimental males' testes. Males also varied significantly in the proportion of live sperm found in storage in vivo. However, the exact mechanism of sperm degradation remains to be clarified. In vitro exposure to extracts of the female reproductive tract, including female accessory glands, failed to significantly lower sperm viability compared to controls. These results are consistent either with postcopulatory sperm mortality in vivo depending entirely on the male (with individual differences in sperm viability, motility or longevity) or with postcopulatory sperm mortality being subtly affected by female effects which were not detected by the in vitro experimental conditions. Importantly, we found no evidence in support of the hypothesis that female accessory glands contribute to sexual conflict via spermicide. Therefore, female muscular control remains to date the only ascertained mechanism of female influence on sperm storage in this species.     

On the copulation duration of the yellow dung fly (Scathophaga stercoraria)

We model the optimal copulation duration in the yellow dungflyScathophaga stercoraria, assuming that males optimize their reproductive success per day. The independent state-variables of a male are the actual sperm reserves, the female encounterrate and the time of the day. We used stochastic dynamic programming to predict the optimal copulation duration. The model predicts that copulation duration should increase (i) for larger males, (ii) for males with a better previous diet (iii) for males accepting more females (iv) for males staying away from the dung during a whole day. It also predicts that (v) males on a restricted diet should accept fewer matings, (vi) testes size should shrivel during the day, (vii) the correlation coefficient between copulation duration and sequence number should decrease with increasing initial sperm reserve and with increasing number of copulations performed on a given day.There are two essential differences between our model and a model of Parker (1992). (i) Our model assumes that males optimize per day. Parker's model assumes that males optimize per mating bout, one bout comprising many days. (ii) Since all the important state variables depend only on the moment of the copulation, our model does not assume a long-time memory in dungflies, and it is more flexible than Parker's model. Overall, both models explain the data available equally well.     

Aggressive Behavior in Olive Fruit Fly Females: Oviposition Site Guarding against Parasitic Wasps

Among Tephritidae flies, the females display agonistic behavior to maintain single oviposition sites and reduce larval competition for food. In the olive fruit fly, Bactrocera oleae, female-female aggressive interactions are characterized by reciprocal wing waving, chasing, head butting and boxing with forelegs. Little is known on tephritid aggressive behaviors directed towards natural enemies, with special reference to parasitoids attacking their young instars. In this study, we quantified the aggressive behavior of B. oleae females guarding their oviposition site against the braconid parasitoid Psyttalia concolor. The fly aggressive behavior displayed against the parasitoids was compared that directed towards paper dummies mimicking P. concolor adults. When a P. concolor female came close (<20 mm) to a B. oleae female guarding the oviposition site, the 91.67% of the flies displayed wing-waving, the 63.34% chased the parasitoid, the 45% showed head-butting, while boxing was observed only in the 26.67% of the aggressions. When paper dummies were tested, only the 66.67% of the flies displayed wing-waving, the 8.33% performed chasing, followed by head butting (5.00%) and boxing (3.33%). B. oleae displayed longer aggression bouts towards live wasps over dummies. Overall, this is the first evidence pointing out that tephritid aggressive acts, besides their role in intraspecific interactions, were also highly effective to displace parasitic wasps from the fly oviposition site. Further research on potential consequences on fitness traits arising from the above-discussed behaviors, as well as on parasitoid learning-mediated responses to tephritid aggressions, is urgently required.     

The heritability of sexually dimorphic traits in the yellow dung fly Scathophaga stercoraria (L.)

A precise method was used for estimating the proportion of heritable variation in two life history parameters of the yellow dung fly, whereby environmental components of variance were minimized. Significant heritable variation for body size was revealed for father to son and mother to daughter relationships. Variation in development time was not significantly heritable. There is a marked sexual dimorphism in body size in this species which is discussed in the light of the observed sex-genotype interaction in heritabilities and low genetic correlation for size between the sexes. It is suggested that opposing pressures of sexual and natural selection and/or genetic pleotropy may be responsible for the maintenance of heritable variation, and the evolution of sexual dimorphism in these two traits.     

Conspecifics and Their Posture Influence Site Choice and Oviposition in the Damselfly Argia moesta

Finding a suitable oviposition site can be costly because of energy and time requirements, and ovipositioning can be dangerous because of the risk of predation and harassment by males. The damselfly Argia moesta oviposits, contact-guarded by her mate, on vegetation in streams. Oviposition aggregations are commonly observed in this species, despite their territorial nature during other behaviors. We conducted experiments in the field to test the hypothesis that aggregations are the result of conspecific attraction. In the first experiment, two oviposition sites (sycamore leaves) were provided, one with models of ovipositing pairs, and one without. In the second experiment, one leaf again had ovipositing models, while the other had models of uncoupled males and females in a resting posture. In both experiments, damselfly pairs preferred the site with ovipositing models. In general, they visited the ovipositing models first more often than expected by chance, stayed longer there, were more likely to oviposit there, and laid a greater total number of eggs there. These results support the hypothesis that conspecific attraction is responsible for ovipositing aggregations in A. moesta and that posture is an important cue for attraction. Using conspecific cues could be a beneficial strategy to save in search costs while taking advantage of the presence of ovipositing conspecifics to dilute the effects of harassment and predation.     

Temperature effects on egg size and their fitness consequences in the yellow dung fly Scathophaga stercoraria

Organisms and parts of an organism like eggs or individual cells developing in colder environments tend to grow bigger. A unifying explanation for this Bergmann's rule extended to ectotherms has not been found, and whether this is an adaptive response or a physiological constraint is debated. The dependence of egg and clutch size on the mother's temperature environment were investigated in the yellow dung fly Scathophaga stercoraria. Smaller eggs were laid at warmer temperatures in the field and the laboratory, where possible confounding variables were controlled for. As clutch size at the same time was unaffected by temperature, this effect was not due to a trade-off between egg size and number. Temperature-dependent egg sizes even persisted within individuals: when females were transferred to a cooler (warmer) environment, they laid third-clutch eggs that were larger (smaller) than their first-clutch eggs. The fitness consequences of these temperature-mediated egg sizes were further investigated in two laboratory experiments. Neither egg and pre-adult survivorship nor larval growth rate were maximized, nor was development time minimized, at the ambient temperature corresponding to the mother's temperature environment. This does not support the beneficial acclimation hypothesis. Instead, this study yielded some, but by no means conclusive indications of best performance by offspring from eggs laid at intermediate temperatures, weakly supporting the optimal temperature hypothesis. In one experiment the smaller eggs laid at 24 °C had reduced survivorship at all ambient temperatures tested. Smaller eggs thus generally performed poorly. The most parsimonious interpretation of these results is that temperature-mediated variation in egg size is a maternal physiological response (perhaps even a constraint) of unclear adaptive value. This revised version was published online in November 2006 with corrections to the Cover Date.     

Sperm length is not influenced by haploid gene expression in the flies Drosophila melanogaster and Scathophaga stercoraria

Recent theoretical models have postulated a role for haploid–diploid conflict and for kin selection favouring sperm cooperation and altruism in the diversification and specialization of sperm form. A critical assumption of these models—that haploid gene expression contributes to variation in sperm form—has never been demonstrated and remains contentious. By quantifying within-male variation in sperm length using crosses between males and females from populations that had been subjected to divergent experimental selection, we demonstrate that haploid gene expression does not contribute to variation in sperm length in both Drosophila melanogaster and Scathophaga stercoraria. This finding casts doubt on the importance of haploid–diploid conflict and kin selection as evolutionary influences of sperm phenotypes.     

Temporal and microspatial variation in the intensities of natural and sexual selection in the yellow dung fly Scathophaga stercoraria

Studies of phenotypic selection in natural populations often concentrate only on short time periods and do not quantify selection intensities. We quantified temporal and microspatial variation in the intensities of natural and sexual selection for body size in the yellow dung fly over 2 years. Female fecundity selection intensity remained approximately constant over the season with an overall mean ± SE of 0.187 ± 0.014. Selection intensity for male reproductive success, defined as eggs obtained by mating males, did not differ from zero, indicating there was no assortative mating by size. Sexual selection intensity for male mating success favouring large males was variable but overall strong in the two years (0.499 ± 0.053 and 0.510 ± 0.051). As theoretically expected for male–male competition, sexual selection intensity increased with competitor density and reached an asymptote at about 250 males per pat; it also decreased with time in spring and increased again in autumn as a function of density. Small males had the best chance of obtaining a female at very low male densities. Greater selection intensity for large size in males than females is consistent with, and might be responsible for, the observed sexual size dimorphism in this species, as males are larger. The seasonal pattern of mean male body size (smallest at the beginning and end of the season) most likely reflects mere environmental (primarily temperature) influences on phenotypic size.