Constanze Reim

Correlated responses to artificial body size selection in growth, development, phenotypic plasticity and juvenile viability in yellow dung flies.

Most life history traits are positively influenced by body size, whereas disadvantages of large body size are poorly documented. To investigate presumed intrinsic costs of large size in the yellow dung fly (Scathophaga stercoraria; Diptera: Scathophagidae), we established two replicates each of three body size laboratory selection lines (small, control and large; selection on males only), and subjected flies of the resulting extended body size range to various abiotic stresses. Response to selection was symmetrical in the small and large lines (realized h2 = 0.16–0.18). After 24 generations of selection body size had changed by roughly 10%. Female size showed a correlated response to selection on male size, whereas sexual size dimorphism did not change. Development time also showed a correlated response as, similar to food limited flies, small line flies emerged earlier at smaller body size. At the lowest larval food limit possible, flies of all lines emerged at the same small body size after roughly the same development time; so overall phenotypic plasticity in body size and development time strongly increased following selection. Juvenile mortality increased markedly when food was extremely limited, large line flies showing highest mortality. Winter frost disproportionately killed large (line) flies because of their longer development times. Mortality at high temperatures was high but size‐selective effects were inconsistent. In all environments the larger males suffered more. Initial growth rate was higher for males and at unlimited food. Small line individuals of both sexes grew slowest at unlimited larval food but fastest at limited larval food, suggesting a physiological cost of fast growth. Overall, extension of the natural body size range by artificial selection revealed some otherwise cryptic intrinsic juvenile viability costs of large size, mediated by longer development or faster growth, but only in stressful environments.

Sexual selection on morphological and physiological traits and fluctuating asymmetry in the yellow dung fly.

Previous univariate studies of the yellow dung fly (Scathophaga stercoraria) have demonstrated strong sexual selection, in terms of mating success, on male size (estimated as hind tibia length). To identify specific target(s) of selection on body size and possible conflicting selection pressures on particular body parts, two multivariate field studies of sexual selection were conducted. In one study using point samples from three populations, we assessed several morphological traits, including genital traits and measures of fluctuating asymmetry (FA) of all paired traits. There was sexual selection for large male size in general, confirming previous, univariate studies. With the possible exception of thorax width, which was selected in the opposite direction, no main target of selection was identified, as most morphological traits were highly correlated. There was no detectable sexual selection on the male external genital structures assessed. In a second study using multiple samples from one population, we included physiological measures of energy reserves (lipids, glucose and glycogen) known to affect mating success, in addition to trait size and FA of wings and legs. Inclusion of physiological traits is rare in phenomenological studies of selection. This study again confirmed the mating advantage of large males, and additionally showed independent positive influences of lipid and glucose but not glycogen levels. FA in paired traits generally did not affect male mating success, but was negatively correlated with energy reserves. Our study suggests that inclusion of physiological measures and genital traits in phenomenological studies of selection would be fruitful in other species.

Sexual selection on morphological and physiological traits and fluctuating asymmetry in the black scavenger fly Sepsis cynipsea

Previous univariate studies of the fly Sepsis cynipsea (Diptera: Sepsidae) have demonstrated spatiotemporally variable and consequently overall weak sexual selection favouring large male size, which is nevertheless stronger on average than fecundity selection favouring larger females. To identify specific target(s) of selection on body size and additional traits possibly affecting mating success, two multivariate field studies of sexual selection were conducted. In one study using seasonal replicates from three populations, we assessed 15 morphological traits. No clear targets of sexual selection on male size could be detected, perhaps because spatiotemporal variation in selection was again strong. In particular, there was no (current) selection on male abdomen length or fore coxa length, the only traits for which S. cynipsea males are not smaller than females. Interestingly, copulating males had a consistently shorter fore femur base, a secondary sexual trait, and a wider clasper (hypopygium) gap, an external genital trait. In a second study using daily and seasonal replicates from one population, we included physiological measures of energy reserves (lipids, glucose, glycogen), in addition to hind tibia length and fluctuating asymmetry (FA) of all pairs of legs. This study again confirmed the mating advantage of large males, and additionally suggests independent positive influences of lipids (the long‐term energy stores), with effects of glucose and glycogen (the short‐term energy stores) tending to be negative. FA of paired traits was not associated with male mating success. Our study suggests that inclusion of physiological measures and genital traits in phenomenological studies of selection, which is rare, would be fruitful in other species.

Size-dependent energy reserves, energy utilization and longevity in the yellow dung fly

Abstract Teneral reserve components (soluble sugars, glycogen, lipids), as well as proteins, of laboratory‐reared yellow dung flies (Scathophaga stercoraria) are shown to increase hypo‐allometrically (slope < 1) with body size. This is also true for sugars and glycogen of field‐caught reproductive flies, the lipid content of which however, increases hyper‐allometrically (slope > 1), probably related to foraging or reproductive activity. The longevity, or starvation resistance, of adults subsisting exclusively on their teneral reserves equally increases with body size, and this positive relationship remains when flies have access to sugar, pollen or prey for 24 h after emergence. Energy use, by contrast, is generally not body size dependent, except for lipid (and possibly glycogen) utilization when flies were completely starved. All the results obtained are independent of sex after controlling for sexual size dimorphism. These net results clearly support the greater energy efficiency of larger flies (Kleiber’s law), whereas only weak evidence is obtained in support of the contrasting hypothesis of greater absolute energy demands of larger individuals, possibly because such effects are likely masked by stronger size‐dependent anabolic effects.

A cost of being large: genetically large yellow dung flies lose out in intra-specific food competition

Most life history traits are positively influenced by body size, while disadvantages of large size are poorly documented. To investigate presumed intrinsic costs of large body size in yellow dung flies (Scathophaga stercoraria; Diptera: Scathophagidae), we allowed larvae from replicate lines artificially selected for small and large body size for 21 generations to compete directly with each other at 20°C (benign) and 25°C (stressful) and low and high food (dung) availability. Greater mortality of large line flies was evident at low food independent of temperature, suggesting a cost of fast growth and/or long development for genetically large flies during larval scramble competition under food limitation. Our results are congruent with a previous study assessing mortality when competing within body size lines, so no additional mechanisms affecting scramble or contest behavior of larvae need be invoked to explain the results obtained beyond the costs of longer development and faster growth. Thus, artificial selection producing larger yellow dung flies than occur in nature revealed some, albeit weak mortality costs of large body size that otherwise might have remained cryptic. We conclude, however, that these costs are insufficient to explain the evolutionary limits of large body size in this species given persistently strong fecundity and sexual selection favoring large size in both sexes.

A cost of cryptic female choice in the yellow dung fly

Female dung flies Scathophagastercoraria (L.) store sperm from several males in three or four spermathecae. Selection on the number of spermathecae was successful and the morphological intermediate stages in the evolution from three to four spermathecae are illustrated. The genetic quality of a male from a female’s perspective depends on an interaction between their genotypes and the microhabitat in which the offspring will grow. Females influence the paternity pattern of their offspring, and do this differently in different microhabitats. Females with four spermathecae are better able to influence paternity than are those with three spermathecae. However, there must be a cost to building and maintaining an extra spermatheca. We estimate, using the animal model on pedigree data, that this cost is approximately five eggs per clutch, i.e. around 8% of the mean clutch size. This is a substantial cost and such costs should not be ignored in discussions of the benefits to females of assessing the genetic qualities of their mating partners. We suggest that the number of spermathecae in the study population is stable because the relative benefits in quality of offspring through cryptic female choice is balanced by the costs in total numbers of offspring.