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Recently, it has been demonstrated that heat-induced male sterility is likely to shape population persistence as climate change progresses. However, an under-explored possibility is that females may be able to successfully store and preserve sperm at temperatures that sterilise males, which could ameliorate the impact of male infertility on populations. Here, we test whether females from two fruit fly species can protect stored sperm from a high temperature stress. We find that sperm carried by female Drosophila virilis are almost completely sterilised by high temperatures, whereas sperm carried by female Zaprionus indianus show only slightly reduced fertility. Heat-shocked D. virilis females can recover fertility when allowed to remate, suggesting that the delivered heat-shock is damaging stored sperm and not directly damaging females in this species. The temperatures required to reduce fertility of mated females are substantially lower than the temperatures required to damage mature sperm in males, suggesting that females are worse than males at protecting mature sperm. This suggests that female sperm storage is unlikely to ameliorate the impacts of high temperature fertility losses in males, and instead exacerbates fertility costs of high temperatures, representing an important determinant of population persistence during climate change., (Copyright © 2022. Published by Elsevier Ltd.)

There is increasing emphasis on the use of new analytical approaches in subject analysis and classification, particularly in respect to minimal sample preparation. Here, we demonstrate that rapid evaporative ionization mass spectrometry (REIMS), a method that captures metabolite mass spectra after rapid combustive degradation of an intact biological specimen, generates informative mass spectra from several arthropods, and more specifically, is capable of discerning differences between species and sex of several adult Drosophila species. A model including five Drosophila species, built using pattern recognition, achieves high correct classification rates (over 90%) using test datasets and is able to resolve closely related species. The ease of discrimination of male and female specimens also demonstrates that sex-specific differences reside in the REIMS metabolite patterns, whether analysed across all five species or specifically for D. melanogaster. Further, the same approach can correctly discriminate and assign Drosophila species at the larval stage, where these are morphologically highly similar or identical. REIMS offers a novel approach to insect typing and analysis, requiring a few seconds of data acquisition per sample and has considerable potential as a new tool for the field biologist.

Environments vary stochastically, and animals need to behave in ways that best fit the conditions in which they find themselves. The social environment is particularly variable, and responding appropriately to it can be vital for an animal's success. However, cues of social environment are not always reliable, and animals may need to balance accuracy against the risk of failing to respond if local conditions or interfering signals prevent them detecting a cue. Recent work has shown that many male Drosophila fruit flies respond to the presence of rival males, and that these responses increase their success in acquiring mates and fathering offspring. In Drosophila melanogaster males detect rivals using auditory, tactile and olfactory cues. However, males fail to respond to rivals if any two of these senses are not functioning: a single cue is not enough to produce a response. Here we examined cue use in the detection of rival males in a distantly related Drosophila species, D. pseudoobscura, where auditory, olfactory, tactile and visual cues were manipulated to assess the importance of each sensory cue singly and in combination. In contrast to D. melanogaster, male D. pseudoobscura require intact olfactory and tactile cues to respond to rivals. Visual cues were not important for detecting rival D. pseudoobscura, while results on auditory cues appeared puzzling. This difference in cue use in two species in the same genus suggests that cue use is evolutionarily labile, and may evolve in response to ecological or life history differences between species.

Background: Populations of a species often differ in key traits. However, it is rarely known whether these differences are associated with genetic variation and evolved differences between populations, or are instead simply a plastic response to environmental differences experienced by the populations. Here we examine the interplay of plasticity and direct genetic control by investigating temperature-size relationships in populations of Drosophila pseudoobscura from North America. We used 27 isolines from three populations and exposed them to four temperature regimes (16°C, 20°C, 23°C, 26°C) to examine environmental, genetic and genotype-by-environment sources of variance in wing size., Results: By far the largest contribution to variation in wing size came from rearing temperature, with the largest flies emerging from the coolest temperatures. However, we also found a genetic signature that was counter to this pattern as flies originating from the northern, cooler population were consistently smaller than conspecifics from more southern, warmer populations when reared under the same laboratory conditions., Conclusions: We conclude that local selection on body size appears to be acting counter to the environmental effect of temperature. We find no evidence that local adaptation in phenotypic plasticity can explain this result, and suggest indirect selection on traits closely linked with body size, or patterns of chromosome inversion may instead be driving this relationship.

The extent of female multiple mating (polyandry) can strongly impact on the intensity of sexual selection, sexual conflict, and the evolution of cooperation and sociality. More subtly, polyandry may protect populations against intragenomic conflicts that result from the invasion of deleterious selfish genetic elements (SGEs). SGEs commonly impair sperm production, and so are likely to be unsuccessful in sperm competition, potentially reducing their transmission in polyandrous populations. Here, we test this prediction in nature. We demonstrate a heritable latitudinal cline in the degree of polyandry in the fruitfly Drosophila pseudoobscura across the USA, with northern population females remating more frequently in both the field and the laboratory. High remating was associated with low frequency of a sex-ratio-distorting meiotic driver in natural populations. In the laboratory, polyandry directly controls the frequency of the driver by undermining its transmission. Hence we suggest that the cline in polyandry represents an important contributor to the cline in sex ratio in nature. Furthermore, as the meiotic driver causes sex ratio bias, variation in polyandry may ultimately determine population sex ratio across the USA, a dramatic impact of female mating decisions. As SGEs are ubiquitous it is likely that the reduction of intragenomic conflict by polyandry is widespread.

Phenotypic plasticity is a key mechanism by which animals can cope with rapidly changeable environments, but the evolutionary lability of such plasticity remains unclear. The socio-sexual environment can fluctuate very rapidly, affecting both the frequency of mating opportunities and the level of competition males may face. Males of many species show plastic behavioural responses to changes in social environment, in particular the presence of rival males. For example, Drosophila pseudoobscura males respond to rivals by extending mating duration and increasing ejaculate size. Whilst such responses are predicted to be adaptive, the extent to which the magnitude of response is heritable, and hence selectable, is unknown. We investigated this using isofemale lines of the fruit fly D. pseudoobscura, estimating heritability of mating duration in males exposed or not to a rival, and any genetic basis to the change in this trait between these environments (i.e. degree of plasticity). The two populations differed in population sex ratio, and the presence of a sex ratio distorting selfish chromosome. We find that mating duration is heritable, but no evidence of population differences. We find no significant heritability of plasticity in mating duration in one population, but borderline significant heritability of plasticity in the second. This difference between populations might be related to the presence of the sex ratio distorting selfish gene in the latter population, but this will require investigation in additional populations to draw any conclusions. We suggest that there is scope for selection to produce an evolutionary response in the plasticity of mating duration in response to rivals in D. pseudoobscura, at least in some populations.

Background: The rate of female remating can have important impacts on a species, from affecting conflict and cooperation within families, to population viability and gene flow. However, determining the level of polyandry in a species can be difficult, with information on the mating system of many species being based on a single experiment, or completely absent. Here we investigate the mating system of the fruit fly Drosophila subobscura. Reports from England, Spain and Canada suggest D. subobscura is entirely monandrous, with no females remating. However, work in Greece suggests that 23% of females remate. We examine the willingness of female D. subobscura to remate in the laboratory in a range of conditions, using flies from both Greece and England. We make a distinction between pseudopolyandry, where a female remates after an ineffective first mating that is incapable of fertilising her eggs, and true polyandry, where a female remates even though she has received suitable sperm from a previous mating., Results: We find a low rate of true polyandry by females (4%), with no difference between populations. The rate of true polyandry is affected by temperature, but not starvation. Pseudopolyandry is three times as common as true polyandry, and most females showing pseudopolyandry mated at their first opportunity after their first failed mating. However, despite the lack of differences in polyandry between the populations, we do find differences in the way males respond to exposure to other males prior to mating. In line with previous work, English flies responded to one or more rivals by increasing their copulation duration, a response previously thought to be driven by sperm competition. Greek males only show increased copulation duration when exposed to four or more rival males. This suggests that the response to rivals in D. subobscura is not related to sperm competition, because sperm competition is rare, and there is no correlation of response to rivals and mating system across the populations., Conclusions: These results illustrate the difficulties in determining the mating system of a species, even one that is well known and an excellent laboratory species, with results being highly dependent on the conditions used to assay the behaviour, and the population used.

Male responses to risk of sperm competition play an important role in sexual selection, sexual conflict, and the evolution of mating systems. Such responses can combine behavioural and physiological processes, and can be mediated through different components of the ejaculate such as sperm numbers and seminal proteins. An additional level of ejaculate complexity is sperm heteromorphism, with the inclusion of non-fertilising parasperm in the ejaculate. We now test the response to rivals in a sperm heteromorphic species, Drosophila pseudoobscura, measuring the behavioural response and sperm transfer and, crucially, relating these to short-term fitness. Males respond to exposure to conspecific rivals by increasing mating duration, but do not respond to heterospecific rivals. In addition, after exposure to a conspecific rival, males increased the transfer of fertilising eusperm, but not non-fertilising parasperm. Males exposed to a conspecific rival also achieve higher offspring production. This suggests that the evolution of parasperm in flies was not driven by sperm competition and adds to the increasing evidence that males can make extremely sophisticated responses to mating competition., (Copyright © 2012. Published by Elsevier Ltd.)

Selfish genetic elements occur in all living organisms and often cause reduced fertility and sperm competitive ability in males. In the fruit fly Drosophila pseudoobscura, the presence of a sex-ratio distorting X-chromosome meiotic driver Sex Ratio (SR) has been shown to promote the evolution of increased female remating rates in laboratory populations. This is favored because it promotes sperm competition, which decreases the risk to females of producing highly female-biased broods and to their offspring of inheriting the selfish gene. Here, we show that non-SR males in these SR populations evolved an increased ability to suppress female remating in response to the higher female remating rates, indicating male-female coevolution. This occurred even though SR was rare in the populations. This was further supported by a correlation between females' remating propensity and males' ability to suppress female remating across populations. Thus SR can generate sexual conflict over female remating rate between females and the noncarrier males that make up the majority of the males, promoting evolution of increased ability of males to suppress female remating.

Females of most animal species are polyandrous, with individual females usually mating with more than one male. However, the ubiquity of polyandry remains enigmatic because of the potentially high costs to females of multiple mating. Current theory to account for the high prevalence of polyandry largely focuses on its benefits to individual females. There are also higher-level explanations for the high incidence of polyandry-polyandrous clades may speciate more rapidly. Here we test the hypothesis that polyandry may also reduce population extinction risk. We demonstrate that mating with multiple males protects populations of the fruit fly Drosophila pseudoobscura against extinction caused by a "selfish" sex-ratio-distorting element. Thus, the frequency of female multiple mating in nature may be associated not only with individual benefits to females of this behavior but also with increased persistence over time of polyandrous species and populations. Furthermore, we show that female remating behavior can determine the frequency of sex-ratio distorters in populations. This may also be true for many other selfish genetic elements in natural populations.

Selfish genetic elements (SGEs) are ubiquitous in animals and often associated with low male fertility due to reduced sperm number in male carriers. In the fruit fly Drosophila pseudoobscura, the meiotic driving X chromosome "sex ratio" kills Y-bearing sperm in carrier males (SR males), resulting in female only broods. We competed SR males against the ejaculates of noncarrying standard males (ST males), and quantified the number of sperm transferred by SR and ST males to females. We show that SR males are very poor sperm competitors, which is partly related to transfer of fewer sperm during mating. However, sperm numbers alone cannot explain the observed paternity reduction, indicating SR males' sperm may be of reduced quality, possibly due to damage during the killing of the noncarrying Y-sperm. The reduction in sperm competitive ability due to SR is large enough to potentially stabilize the spread of sex ratio drive through populations. The poor sperm competitive ability of SR males coupled with their low fitness as mates could favor increased remating by females to reduce paternity by SR males. Given the generally poor performance of SGE-carrying males in sperm competition, this may generate strong selective pressure favoring polyandry in many species.

The impact of rising global temperatures on survival and reproduction is putting many species at risk of extinction. In particular, it has recently been shown that thermal effects on reproduction, especially limits to male fertility, can underpin species distributions in insects. However, the physiological factors influencing fertility at high temperatures are poorly understood. Key factors that affect somatic thermal tolerance such as hardening, the ability to phenotypically increase thermal tolerance after a mild heat shock, and the differential impact of temperature on different life stages are largely unexplored for thermal fertility tolerance. Here, we examine the impact of high temperatures on male fertility in the cosmopolitan fruit fly Drosophila virilis. We first determined whether temperature stress at either the pupal or adult life history stage impacts fertility. We then tested the capacity for heat‐hardening to mitigate heat‐induced sterility. We found that thermal stress reduces fertility in different ways in pupae and adults. Pupal heat stress delays sexual maturity, whereas males heated as adults can reproduce initially following heat stress, but become sterile within seven days. We also found evidence that while heat‐hardening in D. virilis can improve high temperature survival, there is no significant protective impact of this same hardening treatment on fertility. These results suggest that males may be unable to prevent the costs of high temperature stress on fertility through heat‐hardening, which limits a species' ability to quickly and effectively reduce fertility loss in the face of short‐term high temperature events. [ABSTRACT FROM AUTHOR]

Cannibalism is taxonomically widespread and can have large impacts on individual fitness and population-level processes. As such, identifying how cannibalism rates vary in response to ecological cues is important for predicting species evolution and population dynamics. 2. In this study, we aimed to identify several eco-evolutionary factors that affect cannibalism rate and measure how they interacted with one another. 3. To do this, we conducted two experiments using complimentary methods to measure how cannibalism rates varied among larval Drosophila melanogaster and Drosophila simulans in response to changes in conspecific relatedness, social familiarity and density. 4. We found that larvae were more likely to cannibalise non-related larval victims in both species, and that this effect increased at high densities in D. simulans. We found no evidence that Drosophila larvae use social familiarity to assess relatedness. Finally, in D. melanogaster, cannibalistic larvae prefer to cannibalise larvae that are being attacked by a greater number of conspecifics, implying that cues linked to conspecific abundance encourage cooperative cannibalism. 5. By showing that cannibalism frequency in Drosophila spp. is sensitive to relatedness and several other factors, we reveal the complex relationship between cannibalism frequency and species ecology. Also, by showing that the effect of relatedness on cannibalism frequency is density dependent, we advance the current understanding of how ecological variables interact to affect kin selection. [ABSTRACT FROM AUTHOR]

Age is well known to be a basis for female preference of males. However, the mechanisms underlying age-based choices are not well understood, with several competing theories and little consensus. The idea that the microbiota can affect host mate choice is gaining traction, and in this study we examine whether the male microbiota influences female preference for older individuals in the fruit fly Drosophila pseudoobscura. We find that an intact microbiota is a key component of attractiveness in older males. However, we found no evidence that this decrease in older male attractiveness was simply due to impaired microbiota generally reducing male quality. Instead, we suggest that the microbiota underlies an honest signal used by females to assess male age, and that impaired microbiota disrupt this signal. This suggests that age-based preferences may break down in environments where the microbiota is impaired, for example when individuals are exposed to naturally occurring antibiotics, extreme temperatures, or in animals reared in laboratories on antibiotic supplemented diet. [ABSTRACT FROM AUTHOR]

Transposable elements (TEs) are genomic parasites that proliferate within host genomes, and which can also invade new species. The P-element, a DNA-based TE, recently invaded two Drosophila species: Drosophila melanogaster in the 20th century, and D. simulans in the 21st. In both species, lines collected before the invasion are susceptible to "hybrid dysgenesis", a syndrome of abnormal phenotypes apparently due to P-element-inflicted DNA damage. In D. melanogaster, lines collected after the invasion have evolved a maternally acting mechanism that suppresses hybrid dysgenesis, with extensive work showing that PIWI-interacting small RNAs (piRNAs) are a key factor in this suppression. Most of these studies use lines collected many generations after the initial P-element invasion. Here, we study D. simulans collected early, as well as late in the P-element invasion of this species. Like D. melanogaster, D. simulans from late in the invasion show strong resistance to hybrid dysgenesis and abundant P-element-derived piRNAs. Lines collected early in the invasion, however, show substantial variation in how much they suffer from hybrid dysgenesis, with some lines highly resistant. Surprisingly, although, these resistant lines do not show high levels of cognate maternal P-element piRNAs; in these lines, it may be that other mechanisms suppress hybrid dysgenesis. [ABSTRACT FROM AUTHOR]