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 h(2) = 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.     

Effects of temperature on growth, development and diapause in the yellow dung fly – against all the rules?

The effects of rearing temperature (and photoperiod) on growth, development, body size, and diapause induction and termination in the yellow dung fly, Scathophaga stercoraria, were investigated by allowing replicate families of larvae to develop in the field along a time sequence approaching the onset of winter. This was supplemented with extensive laboratory rearing. At constant laboratory temperatures, growth rates were maximal between 15°C and 20°C and decreased at higher (25°C) and lower (10°C) temperatures, while the development rate was maximal at 25°C. Perhaps related to this, yellow dung flies reached a given size faster at naturally variable, as opposed to constant, temperatures. In the field, lower temperatures towards the end of the season resulted in larger individuals that grew faster. Adult body size increased as development time, expressed in calendar days, increased, a positive relationship commonly taken for granted in life history theory, but decreased as development time expressed in degree-days increased. The effect of temperature on growth, development and body size can thus change or even reverse if individuals can alter their growth rate independently of development time, and if the physiological effects of temperature are factored out by converting development time into degree-days above a lower development threshold. Therefore, supposedly well-established trends possibly need to be re-examined along these lines. Pupal winter diapause towards the end of the season was highly reversible by temperature. Pre- and post-winter emergence patterns together suggest that the minimum time for yellow dung flies to successfully complete development, at any time of the year, is about 230–250 degree-days. Received: 2 September 1996 / Accepted: 26 February 1997     

ADAPTIVE PHENOTYPIC PLASTICITY IN GROWTH,DEVELOPMENT, AND BODY SIZE IN THE YELLOW DUNG FLY

Life-history theory predicts that age and size at maturity of organisms should be influenced by time and food constraints on development. This study investigated phenotypic plasticity in growth, development, body size, and diapause in the yellow dung fly, Scathophaga stercoraria. Full-sib families were allowed to develop under predator-free field conditions. The time before the onset of winter was varied and each brood was split into three environments differing in the amount of dung a set number of larvae had as a resource. When resources were abundant and competition was minimal, individuals of both sexes grew to larger body sizes, took longer time to mature, and were able to increase their growth rates to attain large body sizes despite shorter effective development periods later in the season. In contrast, limited larval resources and strong competition constrained individuals to mature earlier at a smaller adult size, and growth rates could not be increased but were at least maintained. This outcome is predicted by only two life-history optimality models, which treat mortality due to long development periods and mortality due to fast growth as independent. Elevated preadult mortality indicated physiological costs of fast growth independent of predation. When larval resources were limited, mortality increased with heritable variation in development time for males, and toward the end of the season mortality increased as larval resources became more abundant because this induced longer development periods. Sexual and fecundity selection favoring large body size in this species is thus opposed by larval viability selection favoring slower growth in general and shorter development periods when time and resources are limited; this overall combination of selective pressures is presumably shaping the reaction norms obtained here. Flexible growth rates are facilitated by low genetic correlations between development time and body size, a possible consequence of selection for plasticity. Heritable variation was evident in all traits investigated, as well as in phenotypic plasticity of these traits (genotype X interactions). This is possibly maintained by unpredictable spatiotemporal variation in dung abundance, competition, and hence selection.     

Temperature and clinal variation in larval growth efficiency in Drosophila melanogaster

Geographic clines in ectotherm species including Drosophila melanogaster have been found throughout the world, with genetically larger body size and shorter development time occurring at high latitudes. Temperature is thought to play a major role in the evolution of this clinal variation. Laboratory thermal selection has effects similar to those seen in geographical clines. Evolution at low temperatures results in more rapid development to larger adult flies. This study investigated the effects of geographical origin and experimental temperature on larval growth efficiency in D. melanogaster. Larvae from populations that had evolved at high latitudes were found to use limited food more efficiently, so that the overall adult body size achieved was larger. Larvae reared at a lower experimental temperature (18 °C) used food more efficiently than those reared at a higher temperature (25 °C). The increases in growth efficiency found in populations from high latitudes could explain their increased body size and more rapid development.     

Size‐dependent insect flight energetics at different sugar supplies

Under most circumstances, large body size confers a higher fitness and is positively selected, whereas selection against large size is empirically poorly documented. Physiologically, according to the ¾ power law, larger animals have lower relative but higher absolute energy demands, such that large body size may become disadvantageous, particularly under fast locomotion in food‐limited environments. After a period of initial feeding on different sugar concentrations, we investigated size‐dependent energy content (reserves) at baseline and of females unflown (i.e. resting) or flown for 18 h in two (replicate) insect species: the yellow dung fly Scathophaga stercoraria and the yellow fever mosquito Aedes aegypti. Tethered adults of various sizes were tested in a flight mill. In both species, teneral glycogen, sugar, and lipid content increased with sugar availability, and isometrically or even hyper‐allometrically (slope > 1) with body size. Activity treatment also revealed the expected consumption effects. Both species increased their flight distance with sugar supply, although only larger mosquitoes flew longer. Crucially, larger females of both species disproportionately exhausted more glycogen and sugars (but not lipid) during flight. The mosquitoes appeared to adjust their flight more finely to their size‐dependent energy reserves at all sugar availabilities, whereas, in the dung flies, size‐dependent energy demands were detectable only with a low but not with an overly high sugar supply. Although we found a greater absolute and relative locomotory energy demand for the larger flies, which is in agreement with interspecific patterns in insects, this was (more than) compensated by their greater baseline energy reserves, resulting in the greater net flying capacity of larger individuals. Consequently, we found no evidence for energetic mechanisms limiting the performance of large flying insects under food limitation. The differences between the two species presumably relate to mosquitoes inherently being long distance flyers and dung flies being short distance flyers. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Different growth responses to temperature and resource limitation in three fly species with similar life histories

Growth responses to temperature and resource limitation in three dipteran species with similar life histories were compared. With respect to current life history theory, two points are raised. First, growth rate in real time increased steeply with temperature in all species, following the standard pattern. However, when expressed in physiological time growth rate increased as temperature decreased in the yellow dung fly Scathophaga stercoraria, remained approximately constant in Sepsis cynipsea, and increased in Drosophila melanogaster. These responses can be understood as adaptations to climate and seasonality. It is concluded that some patterns of adaptation may be more easily interpreted if, and some may even go undetected unless, they are analysed in physiological time. Second, a decrease in body size, development rate and growth rate when resources are limited is believed to be nearly universal and generally predicted by life history models. Despite their similar life histories, the three species investigated showed qualitatively different growth responses to larval food shortage. At unlimited resources, yellow dung flies showed the fastest initial larval body mass gain per unit time, while those of S. cynipsea and D. melanogaster were lower and about equal. The period of no body mass gain at the end of larval development was longest in S. stercoraria and shortest in S. cynipsea. When facing resource limitation, S. stercoraria emerged smaller but earlier (thus nearly maintaining their growth rate), S. cynipsea smaller after the same development period, and D. melanogaster smaller and later (showing reduced and much reduced growth, respectively). It is concluded that whether growth really slows when resources are limited depends on the precise ecological circumstances of the species in question. More refined models, particularly those where mortality costs are independent of time, and more experiments are necessary to account for the variation in growth and size and age at maturity present in nature. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effects of temperature on growth and efficiency of food utilization in fifth-instar caterpillars of the tobacco hornworm, Manduca sexta

Fifth-instar larvae of Manduca sexta were reared from hatching on artificial diet at 15, 20, 25, 30 and 35°C. Total development time decreased with increasing temperature. Very few larvae (12%) survived at 15°C, so this temperature was not considered further. There was some mortality at 30°C (11%), and at 35°C (50%).The absolute rate of growth in the fifth instar was faster at 25 than at 20°C, but was similar at 25, 30 and 35°C. This was true both for caterpillars that were chronically exposed to experimental temperatures (i.e. since hatching) and for those acutely exposed (i.e. reared up to fifth instar at 25°C).There was a progressive decrease with higher rearing temperatures in both the initial and final sizes of chronically exposed fifth-instar larvae. Acutely exposed caterpillars matched for initial size showed smaller temperature related differences in final size. Because of these size differences there were differences in relative growth rate which did not reflect true differences in absolute growth rate.Total food consumed by chronically exposed caterpillars was greatest at the lowest temperature (20°C), and decreased progressively with increasing temperature. The absolute rate of food consumption increased from 20 to 25°C, but did not vary significantly between 25 and 35°C. Differences in the sizes of the insects at the different temperatures meant that there were differences among relative measures of consumption that did not reflect absolute food consumption.For chronically exposed caterpillars, none of the three usual indices of food conversion efficiency (AD, ECI and ECD) varied significantly with temperature between 20 and 35°C. This implies that the effects of temperature on metabolic costs are closely matched to food consumption.Oxygen consumption increased with temperature between 20 and 25°C but was temperature compensated between 25 and 35°C.These findings are discussed in terms of their implications for the optimal temperature for growth in Manduca.     

Responses to selection on phenoloxidase activity in yellow dung flies

Maintaining an immune system is costly. Resource allocation to immunity should therefore trade off against other fitness components. Numerous studies have found phenotypic trade-offs after immune challenge, but few have investigated genetic correlations between immune components and other traits. Furthermore, empirical evidence for the costs of maintaining an innate immune system in the absence of challenges is rare. We examined responses to artificial selection on phenoloxidase (PO) activity, an important part of the insect innate defense against multicellular pathogens, in yellow dung flies, Scathophaga stercoraria (L.). After 15 generations of successful selection on PO activity, we measured reproductive characters: clutch size, egg hatching rates, adult emergence rates, and adult longevity. We found no evidence for negative genetic correlations between PO activity and reproduction. In fact, flies of lines selected for increased PO activity had larger first clutches, and flies of lines selected for decreased PO activity had smaller ones. However, flies from high-PO lines died earlier than did low-PO flies when no food was available; that is, there is a survival cost of running at high PO levels in the absence of challenge. Variation in resource acquisition or use may lead to positive genetic correlations between PO and fertility and fecundity. The negative correlation between PO and longevity under starvation may indicate that variation for resource acquisition is maintained by a cost of acquisition, based on a genotype-environment interaction.     

The function of female accessory reproductive gland secretion and a cost to polyandry in the yellow dung fly

Abstract Female yellow dung flies have large paired accessory reproductive glands, the function of which remains unclear. However, gland contents are secreted during copula and egg laying. Other female flies produce a range of anti-bacterial substances in their accessory reproductive glands that protect them and their eggs from pathogens, and it is possible that gland secretion acts similarly in yellow dung flies. A series of experiments was conducted to test this idea. Because the volume of secretion remaining in the glands is negatively related to copula duration, egg hatchability and longevity of females was compared in groups that copulated experimentally once or three times. A zone inhibition assay was used to see if gland extract inhibited bacterial growth. Egg number was positively associated with female body size, but the proportion of eggs hatching was not. Neither copula number nor duration influenced egg number or hatch number or proportion. In accordance with this, gland extract did not inhibit bacterial growth. However, female longevity was reduced in females that copulated with more males. This suggests that gland secretion does not serve to protect eggs, but as with a number of taxa, copula is costly to female yellow dung flies.     

Seasonal changes in characteristics of cattle dung as a resource for an insect in southwestern Australia

The value of cattle dung as a food resource for the bush fly Musca vetustissima (Walker) in the winter rainfall agricultural region of southwestern Australia was assessed by bioassay in the laboratory. The size (headwidth) of adult females was measured from flies reared on different samples of dung. Variation in size correlated with seasonal patterns of pasture growth, larger flies being produced during the growing season from autumn to spring. Size declined with senescence of annual pastures in late spring and early summer, occurring later in southern areas where the growing season was longer. After pasture senescence, dung from shorter growing season areas usually produced larger flies, apparently a result of the inverse relationship between digestibility of feed and length of growing season. Dung from irrigated perennial pastures never produced flies as large as that from annual pastures but generally high values were sustained during summer. Grazing of cereal stubble and feeding of hay in annual pasture areas during summer usually caused some increase in fly size. A spontaneous resurgence in the size of flies often occurred several weeks after pasture senescence and was attributed to more thorough digestion as a result of reduced intake of less palatable dry pasture.     

Using age grading by wing injuries to estimate size-dependent adult survivorship in the field: a case study of the yellow dung fly Scathophaga stercoraria

Abstract 1. Studies of natural selection depend on estimates of longevity and mortality in the wild. In small and mobile species such as insects, direct, mark–recapture (resight), studies are difficult to perform because individuals cannot be tracked easily.
2. It was investigated whether age grading based on wing injuries alone can be used to estimate size-specific survivorship in the field in the yellow dung fly Scathophaga stercoraria (L.) (Diptera: Scathophagidae).
3. The accumulation of different types of wing injury throughout the spring and autumn flight seasons for both sexes was recorded: tears, notches (both reflecting regular wear), and large missing areas (probably due to intra- and inter-specific interactions).
4. Female longevity increased with body size in both spring and autumn, whereas male longevity increased slightly with size in spring but decreased in autumn.
5. The two sexes and males of different size classes accumulated the various types of wing injury differentially, presumably due to differential patterns of intraspecific interactions. Additionally, body size exhibited a seasonal pattern, complicating interpretation of the relationship between body size and wing injuries.
6. It is therefore concluded that estimating adult viability selection on body size using wing injuries is problematic in dung flies, and potentially also in other species. It is suggested that before this method is applied in any particular species, pilot studies should be conducted to verify whether wing injuries accumulate equally in all classes of individuals of interest. In addition, it is necessary to investigate the causes of different types of wing injury.     

The effects of temperature on development of the large narcissus fly (Merodon equestris)

Eggs, larvae, pupae and adults of the large narcissus fly (Merodon equestris) were reared at a series of constant temperatures between 9–24°C. Egg development required from 37 days at 9°C to 7 days at 21.5°C. The low-temperature threshold for development was 6.7°C. Larvae reared at 1424°C were fully-grown after 18 weeks, but it took much longer for such insects to pupate, and adult flies emerged only after about 45 weeks of development. Large narcissus flies enter diapause during the larval stage and overwinter as fully-fed larvae, forming pupae in the following spring. Post-winter pupation and pupal development took from 169 days at 10°C to 36 days at 21.5°C. Of this, pupal development required from 91 days at 10°C to 19 days at 21.5°C. The low-temperature threshold for post-winter pupation and pupal development was 7.1°C, and for pupal development alone, 7.2°C. Females maintained at or below 19°C laid few eggs, whereas some females kept at or above 21.5°C laid more than 100 eggs (mean 69 ± 36). Approximately 50% of females maintained at or above 21.5°C laid less than 10 eggs during their lifetime. The mean egg-laying time was 6 to 9 days. Although temperatures at or below 19°C inhibited mating, once a female had mated, such temperatures did not prevent oviposition.     

Temperature-induced changes in the life cycle of Leuctra nigra(Plecoptera: Leuctridae) from a Lake District stream

1. The life cycle of Leuctra nigra (Olivier) took 2 years in a small stream in the English Lake District and the exponential growth of the larvae was scarcely affected by variations in water temperature (range 4.2-14.0°C). Mean growth rates for three year-classes were 0.43±0.01, 0.42±0.01, 0.39±0.05% body length day^?1. There were thirteen or fourteen larval instars for males and fourteen or fifteen for females. The ratio between successive instars was a constant 1.20 (conformed with Dyar's rule). 2. Larval growth and mortality were exponential at six constant temperatures (5.9, 8.2, 12.1, 15.8, 18.2, 19.8°C) in the laboratory. Mean growth rates (% body length day^?1) increased directly with temperature from 0.37 (5.9°C) to 0.55 (19.8°C). Mean mortality rates (% day^?1) increased directly with temperature from 0.20 (5.9°C) to 0.26 (12.1°C) and then markedly increased to 0.54-0.58 at the three higher temperatures. Only 7-10% of animals completed their life cycle at the three higher temperatures compared with 23–27% at the three lower temperatures. Egg production also decreased considerably at the higher temperatures. 3. As growth rates in the stream and laboratory were similar at similar temperatures (<14°C), the optimum conditions for growth in the laboratory were probably similar to those in the stream; therefore resources such as food and space were not restricting growth in the stream. 4. The implications of the temperature-induced changes in growth and mortality are discussed and it is concluded that although the life cycle can change from semivoltine to univoltine with increasing temperature, the costs of a univoltine life cycle are high in terms of survival and egg production, both of which decreased markedly between 12.1 and 15.8°C. Therefore the optimum habitat for this species appears to be a summer cool stream (maximum temperature <14°C) and the optimal life cycle appears to be about 2 years from egg to adult.     

Topical treatment of calves with synthetic pyrethroids: effects on the non-target dung fly Neomyia cornicina (Diptera: Muscidae)

Dung from calves treated with synthetic pyrethroids negatively influenced, in varying degrees, survival, reproduction and size of the common dung fly Neomyia cornicina (Fabricius). This was documented in assays where the coprophagous larvae and adults of N. cornicina were exposed to dung collected from calves dosed with topical preparations of deltamethrin, flumethrin, cyfluthrin, and alpha-cypermethrin. Larval mortality was significantly increased in dung collected up to at least seven days after treatment with deltamethrin, alpha-cypermethrin and cyfluthrin. Alpha-cypermethrin caused significant mortality of adults allowed to feed on moist dung. Nulliparous flies fed for six days on dung collected three days after treatment of calves with alpha-cypermethrin or deltamethrin showed little or no ovarian development. A tendency for a comparable effect with flumethrin was also observed. A connection between ovarian development and inhibition of feeding was indicated by the observation of significantly lowered excretion rates in flies exposed to residues of deltamethrin, alpha-cypermethrin and flumethrin. Larvae that survived exposure to dung from calves dosed with deltamethrin, alpha-cypermethrin, or cyfluthrin gave rise to smaller flies. The effect on adult fly size decreased when larvae were exposed to dung collected at longer times after treatment of the calves. Adult fly size was significantly reduced in dung collected up to 14 days (alpha-cypermethrin) or up to 28 days after treatment (deltamethrin and cyfluthrin). Fluctuating asymmetry of a wing vein character did not reflect the anticipated levels of exposure. The study strongly indicated that the use of synthetic pyrethroids affected the insect dung fauna and that such use may reduce dung decomposition.     

Intraspecific competition of Glyptotendipes paripes (Diptera: Chironomidae) larvae under laboratory conditions

Glyptotendipes paripes larvae were reared in wells of tissue culture plates, in groups of 2, 4, 8, 16, and 32 (representing densities of about 1,300, 2,600, 5,200, 10,400, and 20,800 larvae per m², respectively). Larval groups were supplied with one of two concentrations (low or high) of food and larvae were individually observed to evaluate the effects of density on mortality, growth, development, behavior, and adult body size. Increased larval densities resulted in higher mortality, as well as slower larval growth and development. The distribution of developmental time became flatter at higher density, with a wider range of values, or even became bimodal. This was a consequence of the most rapidly developing individuals at higher densities emerging as adults sooner than the fastest developing individuals at lower densities, although overall mean developmental time was longer at higher densities. At higher densities, growth and development of smaller larvae were slowed, based on the relative difference in body length between competitors. When larger competitors emerged as adults or died, the growth of smaller larvae may have accelerated, resulting in increased variability of developmental times. The effect of larval density on adult body size was complex, with the largest body size found at the lowest density and a second peak of adult size at high-middle densities, with smaller adult body sizes found at low-middle, and high densities. Similarly, as with developmental time, the range of body size increased with increasing density. Examined food concentrations had no effect on larval mortality, but significantly affected developmental time, growth rate, and adult body size. At higher densities, larvae spent more time gathering food and were engaged in aggressive or antagonistic behaviors.     

CHROMOSOMAL ANALYSIS OF HEAT-SHOCK TOLERANCE IN DROSOPHILA MELANOGASTER EVOLVING AT DIFFERENT TEMPERATURES IN THE LABORATORY

We investigated the heat tolerance of adults of three replicated lines of Drosophila melanogaster that have been evolving independently by laboratory natural selection for 15 yr at “nonextreme” temperatures (18°C, 25°C, or 28°C). These lines are known to have diverged in body size and in the thermal dependence of several life-history traits. Here we show that they differ also in tolerance of extreme high temperature as well as in induced thermotolerance (“heat hardening”). For example, the 28°C flies had the highest probability of surviving a heat shock, whereas the 18°C flies generally had the lowest probability. A short heat pretreatment increased the heat tolerance of the 18°C and 25°C lines, and the threshold temperature necessary to induce thermotolerance was lower for the 18°C line than for the 25°C line. However, neither heat pretreatment nor acclimation to different temperatures influenced heat tolerance of the 28°C line, suggesting the loss of capacity for induced thermotolerance and for acclimation. Thus, patterns of tolerance of extreme heat, of acclimation, and of induced thermotolerance have evolved as correlated responses to natural selection at nonextreme temperatures. A genetic analysis of heat tolerance of a representative replicate population each from the 18°C and 28°C lines indicates that chromosomes 1, 2, and 3 have significant effects on heat tolerance. However, the cytoplasm has little influence, contrary to findings in an earlier study of other stocks that had been evolving for 7 yr at 14°C versus 25°C. Because genes for heat stress proteins (hsps) are concentrated on chromosome 3, the potential role of hsps in the heat tolerance and of induced thermotolerance in these naturally selected lines is currently unclear. In any case, species of Drosophila possess considerable genetic variation in thermal sensitivity and thus have the potential to evolve rapidly in response to climate change; but predicting that response may be difficult.     

SELECTION FOR HEAT-SHOCK RESISTANCE IN LARVAL AND IN ADULT DROSOPHILA BUZZATII: COMPARING DIRECT AND INDIRECT RESPONSES

Direct and correlated responses in selection for heat-shock resistance in adult and in larval Drosophila buzzatii were studied. Two lines were artificially selected for higher survival to heat stress as adults, and two other lines were reared under a fluctuating thermal environment as larvae, 35°C for 6 h and 25°C for 18 h, to “naturally” select for higher resistance as larvae. The latter two lines were duplicated after nine generations to yield additional lines to be “naturally” selected as larvae at a higher temperature, 38.2°C for 6 h. Control lines were maintained separately for the adult and larval selection lines. A significant direct response to selection was found for the adult selection lines. However, larvae of these adult selection lines were no more heat resistant than were larvae of the control lines. One of the two larval selection lines increased significantly in heat resistance as larvae. However, adult heat resistance was similar for lines selected as larvae and the corresponding control lines maintained at 25°C. Changes in developmental time accompanied changes in survival after stress in both sets of lines selected for increased heat resistance.     

Artificial selection on larval growth curves in Tribolium: correlated responses and constraints

Body size is often constrained from evolving. Although artificial selection on body size in insects frequently results in a sizable response, these responses usually bear fitness costs. Further, these experiments tend to select only on size at one landmark age, rather than selecting for patterns of growth over the whole larval life stage. To address whether constraints may be caused by larval growth patterns rather than final size, we implemented a function‐valued (FV) trait method of selection, in which entire larval growth curves from Tribolium were artificially selected. The selection gradient function used was previously predicted to give the maximal response and was implemented using a novel selection index in the FV framework. Results indicated a significant response after one generation of selection, but no response in subsequent generations. Correlated responses included increased mortality, increased critical weight, and decreased development time (DT). The lack of response in size and development time after the first generation was likely caused by increased mortality suffered in selected lines; we demonstrated that the selection criterion caused both increased body size and increased mortality. We conclude that artificial selection on continuous traits using FV methods is very efficient and that the constraint of body size evolution is likely caused by a suite of trade‐offs with other traits.     

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.     

What do dung beetles eat?

Abstract.  1. Most adult coprophagous beetles feed on fresh dung of mammalian herbivores, confining ingestion to small particles with measured maximum diameters from 2–5 to 130 μm, according to body size and kind of beetle. This study explores benefits and costs of selective feeding in a 'typical' dung beetle with a maximum diameter of ingested particles (MDIP) of 20 μm.
2. Examined dung types (from Danish domestic sheep, cattle and horse, and African wild buffalo, white rhino and elephant) contained 76–89% water. Costs of a 20 μm MDIP were often low, since 69–87% of the total nitrogen in bulk dung other than that of elephant and rhino (40–58%) was available to selective feeders.
3. Nitrogen concentrations were high – and C/N ratios low – in most types of bulk dung compared with the average food of terrestrial detritivores or herbivores. Exceptions were elephant and rhino dung with low nitrogen concentrations and high C/N ratios.
4. Estimated C/N ratios of 13–39 in bulk dung (sheep–elephant) were decreased by selective feeding to 7.3–12.6 in the ingested material. In assimilated food, ratios are probably only 5–7, as most assimilable nitrogen and carbon may be of microbial origin. If so, the assimilable food contains a surplus of nitrogen relative to carbon.
5. The primary advantage of selective feeding, particularly in dung with a high C/N ratio, may be to concentrate assimilable carbon in the ingested food. Effects of changing the MDIP within 20–106 μm are modest, especially in dung with a low C/N ratio.