Multiple male morphs in the leaf‐footed bug Mictis longicornis (Hemiptera: Coreidae)

Species within the coreid clade (Hemiptera: Coreidae) can often be observed competing in intrasexual competitions over access to mates and territories. Coreids that partake in these competitions typically possess sexually dimorphic hind legs that are used to strike and squeeze their rivals. In addition to their weaponized legs, some coreid species also possess sexually dimorphic abdominal tubercles, which are assumed to be sexually selected weapons. Still, much remains unknown about the morphology of these structures. Here, using the species Mictis longicornis Westwood, we investigate the frequency distribution and static allometry of abdominal thickness, a measure that includes tubercle length. Furthermore, we also investigate the morphological relationship between abdominal tubercles and weaponized hind legs. We find that male abdominal thickness is best explained by a bimodal distribution, thereby describing the first observed male polymorphism in the coreid clade; a phenomenon typically associated with alternative reproductive tactics. Additionally, we find that major males are characterized primarily by having large weaponized legs and abdominal tubercles, which further suggests that abdominal tubercles are used in male–male competition.     

Covariance among premating, post-copulatory and viability fitness components in Drosophila melanogaster and their influence on paternity measurement

In polyandrous mating systems, male fitness depends on success in premating, post-copulatory and offspring viability episodes of selection. We tracked male success across all of these episodes simultaneously, using transgenic Drosophila melanogaster with ubiquitously expressed green fluorescent protein (i.e. GFP) in a series of competitive and noncompetitive matings. This approach permitted us to track paternity-specific viability over all life stages and to distinguish true competitive fertilization success from differential early offspring viability. Relationships between episodes of selection were generally not present when paternity was measured in eggs; however, positive correlations between sperm competitive success and offspring viability became significant when paternity was measured in adult offspring. Additionally, we found a significant male × female interaction on hatching success and a lack of repeatability of offspring viability across a focal male's matings, which may underlay the limited number of correlations found between episodes of selection.     

Relationships among male sexually selected traits in the bean bug,Riptortus pedestris (Heteroptera: Alydidae)

Current concepts of sexual selection suggest that male reproductive success is determined by multiple sexual traits. As expression and production of multiple sexual traits are frequently associated with each other, positive or negative correlations among multiple sexual traits ensue. These relationships among traits associated with male reproductive success may be crucial in the evolution of male reproductive strategies. Here, we investigate phenotypic relationships among sexually selected traits in the armed bean bug Riptortus pedestris. In this insect, males with a larger body and weapon are more likely to win male–male competitions, and males with a larger weapon or higher courtship rate are more attractive to females. There was a significant positive correlation between body size and weapon size, whereas the courtship rate had significant negative correlations with body size and weapon size. Our results suggested that there was a phenotypic trade‐off between courtship rate and male morphology. In this insect, smaller males may make more effort in courtship behavior as an alternative mating tactic.     

Spatio-temporal variation of natural and sexual selection in breeding populations of the coreid bug,Colpula lativentris (Heteroptera: Coreidae)

Spatio-temporal variations of lifetime reproductive succes (LRS) of both male and female individuals of a coreid bugColpula lativentris were measured and analyzed using the multiple regression method of Arnold and Wade (1984a, b). The standardized variance of LRS was larger in males than that in females as males often to secure mates for a long period whereas females could easily find mates and oviposit simply dependent on ovarial maturation. LRS was partitioned into 4 consecutive fitness components: (1) reproductive lifespan, (2) copulating efficiency, (3) guarding efficiency (for males) or oviposition efficiency (for females), and (4) number of eggs per clutch. In males copulating efficiency was the largest determining factor of LRS, whereas in females reproductive lifespan was the most important factor. Such tendencies were stable on both a yearly and local basis. Patterns of relative contribution of natural selection (reproductive lifespan and number of eggs per clutch) and sexual selection (copulating efficiency and guarding or oviposition efficiency) to LRS were clearly different between males and females. This sexual difference is, at least to some extent, thought to be brought about by sexual selection among males for mating opportunity, though no physical fight was observed among males. Directional selection on body length was found only in relation to the clutch size of females because large females tended to lay larger clutches. No significant directional selection was found in other fitness components.     

Developmental temperature affects the expression of ejaculatory traits and the outcome of sperm competition in Callosobruchus maculatus

The outcome of post‐copulatory sexual selection is determined by a complex set of interactions between the primary reproductive traits of two or more males and their interactions with the reproductive traits of the female. Recently, a number of studies have shown the primary reproductive traits of both males and females express phenotypic plasticity in response to the thermal environment experienced during ontogeny. However, how plasticity in these traits affects the dynamics of sperm competition remains largely unknown. Here, we demonstrate plasticity in testes size, sperm size and sperm number in response to developmental temperature in the bruchid beetle Callosobruchus maculatus. Males reared at the highest temperature eclosed at the smallest body size and had the smallest absolute and relative testes size. Males reared at both the high‐ and low‐temperature extremes produced both fewer and smaller sperm than males reared at intermediate temperatures. In the absence of sperm competition, developmental temperature had no effect on male fertility. However, under conditions of sperm competition, males reared at either temperature extreme were less competitive in terms of sperm offence (P₂), whereas those reared at the lowest temperature were less competitive in terms of sperm defence (P₁). This suggests the developmental pathways that regulate the phenotypic expression of these ejaculatory traits are subject to both natural and sexual selection: natural selection in the pre‐ejaculatory environment and sexual selection in the post‐ejaculatory environment. In nature, thermal heterogeneity during development is commonplace. Therefore, we suggest the interplay between ecology and development represents an important, yet hitherto underestimated component of male fitness via post‐copulatory sexual selection.     

Intra‐specific variance in sperm morphometry: a comparison between wild and domesticated Zebra Finches Taeniopygia guttata

The Zebra Finch Taeniopygia guttata is a model bird species for the experimental study of behavioural and evolutionary concepts in captivity and especially sexual selection. The validity of sexual selection studies of domesticated birds is of long‐standing concern as little is known about the influence of domestication on sexually selected traits. Most domesticated Zebra Finch populations are maintained under a strict breeding regime to avoid potential inbreeding. However, these breeding regimes may interfere with the processes of sexual selection and influence the evolution of sexually selected traits because they may limit or prohibit active mate choice. Here, we investigated the potential impact of a monogamous breeding scheme in a domesticated population in which active mate choice is largely inhibited, on the evolution of sperm morphometry as a sexually selected trait. We compared sperm morphometric traits (total sperm length and length of sperm head, midpiece and flagellum), and the variance thereof, between a domesticated and two wild Zebra Finch populations. Although we found significant differences between the three populations for certain sperm traits (head length, midpiece length), which may be of importance in postcopulatory sexual selection, overall, variance in sperm morphometry did not differ between the domesticated and the wild Zebra Finch populations. Our results validate the use of domesticated Zebra Finches for further studies of postcopulatory sexual selection and sperm competition.     

Heritable variation in the timing of spermatophore removal, a mechanism of post-copulatory female choice in crickets

Female crickets can exert post-copulatory mating preferences by prematurely removing a male's spermatophore after copulation, which terminates sperm transfer. Although most models of sexual selection assume that female mating preferences are heritable, there has been little work addressing genetic variation underlying post-copulatory mate choice. We used a paternal half-sib design, in which different males were randomly assigned as mates to several females to create half-sib families, to determine the heritability of spermatophore retention time in female house crickets, Acheta domesticus. There was significant additive genetic variance in the timing of spermatophore removal by females [h(2) = 0.50 +/- 0.19 (+/- SE)], suggesting that the timing of spermatophore removal is determined, in part, by the female's own genotype independent of the quality of her mate. The relatively high heritability of spermatophore retention time may be reflective of the absence of strong selection on this trait, consistent with previous work showing no difference in the fitness of females permitted to freely remove the spermatophore of their mates and those forced to accept complete ejaculates.     

Sexual size dimorphism decreases with temperature in a blowfly,Chrysomya megacephala (Fabricius) (Diptera: Calliphoridae)

1. There is wide intra‐specific variation in sexual size dimorphism (SSD). Much of this variation is probably as a result of sexual differences in the selective pressure on body size. However, environmental variables could affect males and females differently, causing variation in SSD. 2. We examined the effects of two temperatures (20 and 30 °C) on SSD in six populations of the blowfly, Chrysomya megacephala. 3. We found that body size increased with temperature in all the populations studied, and the sexes differed in phenotypic plasticity of body size in response to rearing temperature. This created substantial temperature‐induced variation in SSD (i.e. sex × temperature interaction). Males were often smaller than females, but the degree of dimorphism was smaller at the higher temperature (30 °C) and larger at the lower temperature (20 °C). This change in SSD was not because of a gender difference in the effect of temperature on development time. Further studies should address whether this variation can be produced by adaptive canalisation of one sex against variation in temperature, or whether it may be a consequence of non‐adaptive developmental differences between the sexes. 4. Although most studies assume that the magnitude of SSD is fixed within a species, the present study demonstrates that rearing temperature can generate considerable intra‐specific variation in the degree of SSD.     

Male body size and condition affects sperm number and production rates in mosquitofish,Gambusia holbrooki

Sperm number is an important predictor of paternity when there is sperm competition. Sperm number is often measured as maximum sperm reserves, but in species where mating is frequent, males will often be replenishing their reserves. Thus, variation in how quickly males can produce sperm is likely to be important in determining male success in sperm competition. Despite this, little is known about how male size, body condition or diet affects sperm production rates. We counted sperm number in large and small Gambusia holbrooki (eastern mosquitofish) after 3 weeks on either a high or low food diet. Sperm number was significantly higher in both larger males and in well‐fed males. We then stripped ejaculates again either 1, 2, 3, 4 or 5 days later to investigate subsequent sperm production. The rate of sperm replenishment was influenced by an interaction between size and diet. Large, well‐fed males had consistently high levels of sperm available over the 5 days (i.e. rapid replenishment), whereas small poorly fed males showed consistently low levels of sperm availability over the 5 days (i.e. slow replenishment). In contrast, large, poorly fed and small, well‐fed males increased their sperm numbers over the first 3 days (i.e. intermediate replenishment). Our study highlights that when mating is frequent and sperm competition is high, size and condition dependence of maximal sperm number and of sperm production rate might both contribute to variation in male reproductive success.     

Genetic and nutritional effects on male traits and reproductive performance in Tribolium flour beetles

In Tribolium flour beetles and other organisms, individuals migrate between heterogeneous environments where they often encounter markedly different nutritional conditions. Under these circumstances, theory suggests that genotype-by-environment interactions (GEI) may be important in facilitating adaptation to new environments and maintaining genetic variation for male traits subject to directional selection. Here, we used a nested half-sib breeding design with Tribolium castaneum to partition the separate and joint effects of male genotype and nutritional environment on phenotypic variation in a comprehensive suite of life-history traits, reproductive performance measures across three sequential sexual selection episodes, and fitness. When male genotypes were tested across three nutritional environments, considerable phenotypic plasticity was found for male mating and insemination success, longevity and traits related to larval development. Our results also revealed significant additive genetic variation for male mating rate, sperm offence ability (P(2)), longevity and total fitness and for several traits reflecting both larval and adult resource use. In addition, we found evidence supporting GEI for sperm defence ability (P(1)), adult longevity and larval development; thus, no single male genotype outperforms others in every nutritional environment. These results provide insight into the potential roles of phenotypic plasticity and GEI in facilitating Tribolium adaptation to new environments in ecological and evolutionary time.     

Effect of male age on sperm traits and sperm competition success in the guppy (Poecilia reticulata)

Deleterious mutations can accumulate in the germline with age, decreasing the genetic quality of sperm and imposing a cost on female fitness. If these mutations also affect sperm competition ability or sperm production, then females will benefit from polyandry as it incites sperm competition and, consequently, minimizes the mutational load in the offspring. We tested this hypothesis in the guppy (Poecilia reticulata), a species characterized by polyandry and intense sperm competition, by investigating whether age affects post‐copulatory male traits and sperm competition success. Females did not discriminate between old and young males in a mate choice experiment. While old males produced longer and slower sperm with larger reserves of strippable sperm, compared to young males, artificial insemination did not reveal any effect of age on sperm competition success. Altogether, these results do not support the hypothesis that polyandry evolved in response to costs associated with mating with old males in the guppy.     

Seasonal phenology and natural enemies of the squash bug (Hemiptera: Coreidae) in Kentucky

The squash bug, Anasa tristis (De Geer), is a major indigenous pest of Cucurbita species across the United States and a vector of cucurbit yellow vine disease. The seasonal phenology of the squash bug in central Kentucky and its natural enemies were studied using summer squash planted sequentially throughout the 2005 and 2006 growing seasons. The squash bug was first detected on 5 June 2005 and 3 June 2006. In both years, peak numbers of all squash bug stages occurred in July and August. Our field data, substantiated by published degree-day models for squash bug development, suggest one complete and a partial second generation of squash bugs in 2005 and one complete generation of squash bugs in 2006. The most abundant ground-active predators in squash fields included Araneae, Carabidae, Staphylinidae, and Geocoridae. Coleomegilla maculata (De Geer) and Geocoris punctipes (Say) were the most abundant foliage-inhabiting predators. Direct field observations of predators feeding on squash bugs or their eggs included G. punctipes, Pagasa fusca (Stein), and Nabis sp. The parasitoids Trichopoda pennipes (Fabricius) and Gyron pennsylvanicum (Ashmead) were found also. Squash bug egg masses were monitored to determine predation and parasitism rates in the field. In four studies during 2005 and 2006, predation rates were low (7% or less), and parasitism ranged from 0 to 31%. Overall, squash bug egg mortality increased as the season progressed.     

The total opportunity for sexual selection and the integration of pre‐ and post‐mating episodes of sexual selection in a complex world

It is well known that sexual selection can target reproductive traits during successive pre‐ and post‐mating episodes of selection. A key focus of recent studies has been to understand and quantify how these episodes of sexual selection interact to determine overall variance in reproductive success. In this article, we review empirical developments in this field but also highlight the considerable variability in patterns of pre‐ and post‐mating sexual selection, attributable to variation in patterns of resource acquisition and allocation, ecological and social factors, genotype‐by‐environment interaction and possible methodological factors that might obscure such patterns. Our aim is to highlight how (co)variances in pre‐ and post‐mating sexually selected traits can be sensitive to changes in a range of ecological and environmental variables. We argue that failure to capture this variation when quantifying the opportunity for sexual selection may lead to erroneous conclusions about the strength, direction or form of sexual selection operating on pre‐ and post‐mating traits. Overall, we advocate for approaches that combine measures of pre‐ and post‐mating selection across contrasting environmental or ecological gradients to better understand the dynamics of sexual selection in polyandrous species. We also discuss some directions for future research in this area.     

Functional morphology of the hind legs as weapons for male contests inLeptoglossus australis (Heteroptera: Coreidae)

Functional morphology of the hind legs as weapons in male contests was analyzed in the leaf-footed plant bug, Leptoglossus australis Fabricius. Measurement of some metrical tratis of the hind femur and tibia indicated that the weapon was the hind femur. Sexual dimorphism in the femoral length and width, and in the number of spines on the lower side of the femur, indicated that these parts play a significant role in male contests. It was also suggested that the length from the base to the widest part of the femur had a functional significance for male-male combat behaviors ofL. australis.     

Reaction norm variants for male calling song in populations of Achroia grisella (Lepidoptera: Pyralidae): toward a resolution of the lek paradox

Significant additive genetic variance often occurs for male advertisement traits in spite of the directional selection imposed by female choice, a problem generally known in evolutionary biology as the lek paradox. One hypothesis, which has limited support from recent studies, for the resolution of this paradox is the role of genotype x environment interaction in which no one genotype exhibits the superior performance in all environments--a crossover of reaction norms. However, these studies have not characterized the actual variation of reaction norms present in natural populations, and the extent to which crossover maintains genetic variance remains unknown. Here, we present a study of genotype x environment interaction for the male calling song in populations of Achroia grisella (Lepidoptera: Pyralidae; lesser waxmoth). We report significant variance among reaction norms for male calling song in two North American populations of A. grisella as measured along temperature, food availability, and density gradients, and there is a relatively high incidence of crossover of the temperature reaction norms. This range of reaction norm variants and their crossover may reflect the co-occurrence of plastic and canalized genotypes, and we argue that the different responses of these variants along environmental gradients may contribute toward the maintenance of genetic variance for male song.     

Sensitive periods during the development and expression of vertebrate sexual signals: A systematic review

Many sexually selected traits exhibit phenotypic plasticity. Despite a growing appreciation for the ecological context in which sexual selection occurs, and for the role of plasticity in shaping traits associated with local adaptation and divergence, there is an important gap in knowledge about the onset and duration of plasticity in sexual trait expression. Integrating this temporal dimension of plasticity into models of sexual selection informs our understanding of the information conveyed by sexual traits and our predictions related to trait evolution, and is critical in this time of unprecedented and rapid environmental change. We conducted a systematic review of 869 studies to ask how trait modalities (e.g., visual and chemical) relate to the onset and duration of plasticity in vertebrate sexual signals. We show that this literature is dominated by studies of coloration in birds and fish, and most studies take place during the breeding season. Where possible, we integrate results across studies to link physiology of specific trait modalities with the life stage (e.g., juvenile, breeding, or nonbreeding) during which plasticity occurs in well‐studied traits. Limitations of our review included a lack of replication in our dataset, which precluded formal analysis. We argue that the timing of trait plasticity, in addition to environmental context, is critical for determining whether and how various communication signals are associated with ecological context, because plasticity may be ongoing or occur at only one point in an individual''s lifetime, and determining a fixed trajectory of trait expression. We advocate for careful consideration of the onset and duration of plasticity when analyzing how environmental variation affects sexual trait expression and associated evolutionary outcomes.     

New occurrence of the invasive alien leaf‐footed bug Leptoglossus gonagra (Hemiptera: Coreidae) in South Korea

The invasive alien leaf‐footed bug Leptoglossus gonagra (Fabricius 1775), native to the Americas, is herein recorded from South Korea for the first time. Forty nymphs and adults were collected on 22 September 2016 from arable lands of two separate sites. These individuals were identified based on the analysis of morphological characters and cytochrome oxidase c subunit 1 sequences. A neighbor‐joining tree indicated the existence of two clades: Group A from Brazil, and Group B from Asia and Oceania. Mean genetic distance between clades was 8.11%, range 7.60–8.57%. Within clades, mean genetic distance was 0.42% (0.17–0.83%) in Group A and, 0.45% (0.00–0.83%) in Group B. Colonies of 2nd‐final instar nymphs and adults were found, on top of leaves, flowers, fruits, and between the stems of pumpkins (Cucurbita moschata). Nymphs and adults damaged host plants by sucking their leaves, stems, fruits, and buds. After the first discovery in 2016, we were unable to find the bugs again (in 2017–2019). Therefore, the invasion of this species seems to have occurred only temporarily, during the warm season. We suggest that the species may have failed to overwinter as it is normally distributed in regions warmer than Korea. Nevertheless, possible appearances of this species must be continuously monitored as it is a potential pest of various plants cultivated in South Korea.     

Sexual coloration and sperm performance in the Australian painted dragon lizard,Ctenophorus pictus

Theory predicts trade‐offs between pre‐ and post‐copulatory sexually selected traits. This relationship may be mediated by the degree to which males are able to monopolize access to females, as this will place an upper limit on the strength of post‐copulatory selection. Furthermore, traits that aid in mate monopolization may be costly to maintain and may limit investment in post‐copulatory traits, such as sperm performance. Australian painted dragons are polymorphic for the presence or absence of a yellow gular patch (‘bibs’), which may aid them to monopolize access to females. Previous work has shown that there are physiological costs of carrying this bib (greater loss of body condition in the wild). Here, we show that male painted dragons use this bright yellow bib as both an inter‐ and intrasexual signal, and we assess whether this signal is traded off against sperm performance within the same individuals. We found no relationship between aspects of bib colour and sperm swimming velocity or percentage of motile sperm and suggest that the bib polymorphism may be maintained by complex interactions between physiological or life‐history traits including other sperm or ejaculate traits and environmental influences.     

Evaluating potential trap crops for managing leaffooted (Hemiptera: Coreidae) and phytophagous stink bug (Hemiptera: Pentatomidae) species in peaches

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The evolution of phenotypic plasticity in spatially structured environments: implications of intraspecific competition, plasticity costs and environmental characteristics

We model the evolution of reaction norms focusing on three aspects: frequency-dependent selection arising from resource competition, maintenance and production costs of phenotypic plasticity, and three characteristics of environmental heterogeneity (frequency of environments, their intrinsic carrying capacity and the sensitivity to phenotypic maladaptation in these environments). We show that (i) reaction norms evolve so as to trade adaptation for acquiring resources against cost avoidance; (ii) maintenance costs cause reaction norms to better adapt to frequent rather than to infrequent environments, whereas production costs do not; and (iii) evolved reaction norms confer better adaptation to environments with low rather than with high intrinsic carrying capacity. The two previous findings contradict earlier theoretical results and originate from two previously unexplored features that are included in our model. First, production costs of phenotypic plasticity are only incurred when a given phenotype is actually produced. Therefore, they are proportional to the frequency of environments, and these frequencies thus affect the selection pressure to avoid costs just as much as the selection pressure to improve adaptation. This prevents the frequency of environments from affecting the evolving reaction norm. Secondly, our model describes the evolution of plasticity for a phenotype determining an individual's capability to acquire resources, and thus its realized carrying capacity. When individuals are distributed randomly across environments, they cannot avoid experiencing environments with intrinsically low carrying capacity. As selection pressures arising from the need to improve adaptation are stronger under such extreme conditions than under mild ones, better adaptation to environments with low rather than with high intrinsic carrying capacity results.