The variability of mating signals and preferences in the brown planthopper,Nilaparvata lugens (Homoptera: Delphacidae)

Male and female brown planthoppers, Nilaparvata lugens(Stål) (Homoptera: Delphacidae), exchange substrate-transmitted signals prior to mating. The pulse repetition frequency of the male song is known to be involved in mate recognition and also to vary among geographical populations. Here the variability of male signals, female signals, and female preferences has been examined within a population. Female preference variation has been partitioned into variation in mean preference and variation in the window of preference of individuals. The genetic component of variation has been examined using isofemale lines. Male signal variation was limited (CV=8%) and was mainly within individuals. Female signal variation was greater (CV=15%). Female mean preference varied little (CV=10%) and was closely matched to the male signal mean, but the preference window was wide (> 4 male signal standard deviations on average) and variable (CV=56%). There was evidence for genetic variation only for preference window. These results are discussed in relation to theories of signal system evolution.     

Genetic control and evolution of acoustic signals in planthoppers (Homoptera: Delphacidae)

Acoustic signals are part of the specific mate recognition system of planthoppers. The genetic control of acoustic signal characters was studied in the planthopperRibautodelphax imitans. Artificial selection for interpulse interval in the female call revealed a large additive genetic component for this polygenic character. Other female call characters showed a correlated response. Some male call characters also appeared to be genetically correlated with the female character selected for, despite the rather different structure of male and female calls. Parent-offspring regression provided significant heritability estimates for those male call characters that also responded to artificial selection in the female call, one of which appeared to be influenced by sex-linked genes. It is argued that the differentiation of this mate recognition system in planthopper populations and species could be the result of founder effects, enabled by the genetic plasticity of the call characters and the existence of a wing length dimorphism in these animals.     

The importance of male and female acoustic behaviour for reproductive isolation in Ribautodelphax planthoppers (Homoptera: Delphacidae)

The importance for reproductive isolation of species-specific acoustic differences between closely related Ribautodelphax planthopper species is tested by measuring responses to playbacks of both conspecific and heterospecific signals. Females respond to heterospecific male calls at about 80% of the conspecific response level, irrespective of the degree of cross-insemination of the combination involved. In a combination involving R. albostriatus , a taxonomically more distantly related species, female response levels are only 15–33% of the normal level. Study of the development of both female responsiveness and receptiveness shows that response levels correspond fairly well with insemination levels. Female heterospecific response is far too high to explain isolation between the species. Female answers to heterospecific males calls have normal response-delay times and durations. When offered a two-way choice between female playback calls, males significantly more often approach the conspecific call in almost all combinations tested. Offering only a heterospecific female signal induces males to call, but not to search. Males are capable of maintaining at least part of the sexual isolation by distinguishing between different female calls. This seems in conflict with the popular theory that the sex with the greater parental investment, here the female, should be exerting the choice.     

Host plant relations in the planthopper genus Ribautodelphax (Homoptera, Delphacidae)

ABSTRACT. 1. Host plant relations of closely related species and pseudogamous triploid forms of the planthopper genus Ribautodelphax were studied.
2. From field data and experimental results it is concluded that eleven European Ribautodelphax species and undescribed taxa are monophagous or oligophagous on different grass species. Only two Ribautodelphax species are found on the same grass species in the field.
3. Differential host plant relations indicate that four recently discovered taxa are probably true species.
4. Ribautodelphax species show a more restricted host plant range in choice experiments compared with breeding tests.
5. Choice experiments indicate no differences in host plant preference between males and females of two Ribautodelphax species.
6. No differences are found in host plant relations between bisexual species and associated pseudogamous triploid forms.     

The genetic basis of female mate preferences

We review the evidence for genetic variation in female and male mate preferences. Genetic differences between species and partially isolated races show that preferences can evolve and were genetically variable in the past. Within populations there is good evidence of genetic variation, both of discrete genetic effects (8 cases) and quantitative genetic effects (17 cases), from a diverse range of taxa. We also review evidence for the presence of genetic covariance between mate preferences and sexual traits in the other sex. The 11 studies go a long way to validating the theoretical prediction of positive genetic covariance. The few negative results are best explained by a lack of appropriate experimental design. One unresolved question is whether genetic covariance is due to linkage disequilibrium between unlinked genes or physical linkage. Some evidence points to linkage disequilibrium but this is not yet conclusive.     

Female mate preferences in Drosophila simulans: evolution and costs

Female mate preference is central to sexual selection, and all indirect benefit models require that there is genetic variation in female preference. This has rarely been tested however, with relatively few studies documenting heritable variation in female preference and even fewer that have directly selected on mate preference to unequivocally show that it can evolve. Additionally, costs of mate preference are poorly understood even though these have implications for preference evolution. We selected on female preference for ebony‐males in replicate Drosophila simulans lines, and generated a rapid evolutionary response in both replicates, with the proportion of females mating with ebony‐males increasing from approximately 5% to 30% after five generations of selection. This increase was independent of changes in ebony‐males as only females were included in our selection regime. We could detect no cost to mate preference itself other than that associated with the fitness consequences of mating with ebony males.     

Karyotypic variation in bisexual species and pseudogamous forms of the planthopper genus Ribautodelphax (Homoptera,Delphacidae)

Chromosome numbers of Ribautodelphax collinus. R. angulosus, R. pungens, R. imitans, R. fanari, R. pallens, R. albostriatus, and of four undescribed species, provisionally indicated as Taxon 1 to 4, were determined. The chromosome number of all species was found to be 2n=30 (14II+XX-XO); however, in several species some females with 2n=31 or 32 also occurred. One female of R. collinus and one of Taxon 1 possessed eggs with 16 bivalents and one egg with 32 bivalents. One female of R. pungens contained only tetraploid eggs. Pseudogamous triploid females occurred in populations of R. pungens and Taxon 4. Their chromosome numbers ranged from 3n=40 to 46 in R. pungens and 3n=41 to 45 in Taxon 4. In several pseudogamous females from two R. pungens populations, eggs with 45 univalents and with various numbers of univalents, bivalents and trivalents were found. The mode of origin of pseudogamous triploid females is discussed and it is concluded that karyotypic data are consistent with an autopolyploid origin.     

Specificity of acoustic signals and mate choice in the brown planthopper Nilaparvata lugens

Mate choice experiments were made between populations of N. lugens from the Philippines, Solomon Is. and northern Australia. Significant barriers to mating were found between the Australian insects and the other two populations. The acoustic signals of successful males in hybridisation experiments were recorded and their pulse repetition frequencies (PRFs) were compared with the mean PRF for a random sample from their own population. In crosses between insects from Australia and the Philippines or Solomon Is. succesful males were characterised by PRFs significantly different from their own populations and closer to those of the female populations in each cross than to their own. It is concluded that PRF of male calls is an important species recognition signal in N. lugens.

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Résumé La communication sexuelle de N. lugens s'effectue par signaux acoustiques transmis par le substrat. Quelques populations géographiquement définissables diffèrent par la fréquence des répétitions des impulsions (PRF) produits par les appels des mâles. Des populations des Philippines, des Iles Salomon et de l'Australie du Nord ont été étudiées.Des expériences de sélection sexuelle ont montré que des obstacles au transfert de spermatozoïdes apparaissaient dans les croisements qui impliquent un parent originaire de la population australienne. Des expériences dhybridation ont été répétées entre individus males et females de chacune des 3 populations étudiées. Le taux de succès étrait faible quand l'un des parents était originaire d'Australie. Le PRF des males de chaque croisement réussi a été déterminé et comparé au PRF moyen d'un échantillon constitué au hasard de males provenant de la même population. Pour les croisements impliquant un parent australien, le PRF des males victorieux différait de la moyenne et se rapprochait de celui des males de la population dont dérivait la female.On en a conclu que le PRF des appels males est important dans la reconnaissance spécifique de N. lugens, mais que chaque population montre un polymorphisme important pour ce caractère.

     

Oxalic acid as a sucking inhibitor of the brown planthopper in rice (Delphacidae, Homoptera)

Oxalic acid, isolated from leaf sheath extract of rice, has been identified as a potent sucking inhibitor against the brown planthopper. Potassium and sodium oxalate at concentrations of 0.1%–0.2% completely inhibited sucking on a 15% sucrose solution. Of twelve aliphatic acids commonly present in plants, only oxalic, maleic, andtrans-aconitic acids at 0.1%, significantly reduced sucking activity while Krebs cycle acids, such as citric, malic, and succinic, did not have any inhibitory effect.

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Résumé L'acide oxalique, isolé des gaines foliaires du riz, a été identifie comme un puissant inhibiteur de la succion deNilaparvata lugens. Les oxalates de potassium et de sodium aux concentrations de 0,1% et 0,2% inhibent complètement la succion d'une solution de sucrose à 15%. Parmi les douze acides aliphatiques généralement présents dans la plante, seuls les acides oxalique, maleique et trans-aconique réduisent significativement à 0,1% l'activité de succion, tandis que les acides cycliques comme les acides citrique, malique et succinique n'ont pas d'effet inhibiteur.

     

Factors affecting the divergence of mate recognition systems in the Limnocytherinae (Crustacea,Ostracoda)

Specific Mate Recognition Systems (SMRS) consist of a set of morphological, behavioural and physiological traits which allow mate recognition. The Limnocytherinae, a lineage of non-marine podocopid Ostracoda, have a relatively wide diversity of copulatory modules, a concept largely congruent with the morphological part of the SMRS. The present paper describes the various copulatory modules in some detail and discusses potential mechanisms responsible for the divergence of these modules. Although none of the processes was thus far demonstrated directly, resulting patterns provide indirect evidence that four different mechanisms contribute. Stochastic processes (chance) as well as developmental and other phylogenetic constraints are involved in the initial selection (choice) of modified structures. Subsequent (positive) directional sexual selection on traits of the recognition systems causes radiative speciation within lineages. At all times, natural selection acts on the development of these structures, either stabilising or negative directional. A number of potential tests for these hypotheses are suggested.     

Environmental variability and acoustic signals: a multi-level approach in songbirds

Among songbirds, growing evidence suggests that acoustic adaptation of song traits occurs in response to habitat features. Despite extensive study, most research supporting acoustic adaptation has only considered acoustic traits averaged for species or populations, overlooking intraindividual variation of song traits, which may facilitate effective communication in heterogeneous and variable environments. Fewer studies have explicitly incorporated sexual selection, which, if strong, may favour variation across environments. Here, we evaluate the prevalence of acoustic adaptation among 44 species of songbirds by determining how environmental variability and sexual selection intensity are associated with song variability (intraindividual and intraspecific) and short-term song complexity. We show that variability in precipitation can explain short-term song complexity among taxonomically diverse songbirds, and that precipitation seasonality and the intensity of sexual selection are related to intraindividual song variation. Our results link song complexity to environmental variability, something previously found for mockingbirds (Family Mimidae). Perhaps more importantly, our results illustrate that individual variation in song traits may be shaped by both environmental variability and strength of sexual selection.     

Divergent selection on,but no genetic conflict over,female and male timing and rate of reproduction in a human population

The sexes often have different phenotypic optima for important life-history traits, and because of a largely shared genome this can lead to a conflict over trait expression. In mammals, the obligate costs of reproduction are higher for females, making reproductive timing and rate especially liable to conflict between the sexes. While studies from wild vertebrates support such sexual conflict, it remains unexplored in humans. We used a pedigreed human population from preindustrial Finland to estimate sexual conflict over age at first and last reproduction, reproductive lifespan and reproductive rate. We found that the phenotypic selection gradients differed between the sexes. We next established significant heritabilities in both sexes for all traits. All traits, except reproductive rate, showed strongly positive intersexual genetic correlations and were strongly genetically correlated with fitness in both sexes. Moreover, the genetic correlations with fitness were almost identical in men and women. For reproductive rate, the intersexual correlation and the correlation with fitness were weaker but again similar between the sexes. Thus, in this population, an apparent sexual conflict at the phenotypic level did not reflect an underlying genetic conflict over the studied reproductive traits. These findings emphasize the need for incorporating genetic perspectives into studies of human life-history evolution.     

Courtship in long-legged flies (Diptera: Dolichopodidae): function and evolution of signals

Analyzing the courtship behavior of long-legged flies (Diptera:Dolichopodidae), we focus on the evolutionary development ofcourtship signals. Long-legged flies may serve as a model forthis evolutionary process, because males of some species presentsexually dimorphic badges during courtship, whereas others donot exhibit such conspicuous signals but present lavish courtshipbehavior, including dynamic flight maneuvers. A comparison ofthese two groups within a single taxonomic family provides insightinto the evolution of courtship signals and the correspondingbehavior. Males of the closely related Empididae do not possesssuch badges. Within the super-family Empidoidea, we proposean evolutionary shift from dynamically courting and mating onthe wing (in Empididae) to courting and mating on ground (inDolichopodidae), accompanied by signaling through badge-waving.By comparing previously published data and observations on courtshipbehavior in Dolichopodidae, we present the hypothesis that thelatter replaced the former energetically expensive behavioras a case of automimicry and sensory trap.     

Indirect mate choice, direct mate choice and species recognition in a bower-building cichlid fish lek

Sexual selection arising through female mate choice typically favours males with larger, brighter and louder signals. A critical challenge in sexual selection research is to determine the degree to which this pattern results from direct mate choice, where females select individual males based on variation in signalling traits, or indirect mate choice, where male competition governs access to reproductively active females. We investigated female mate choice in a lekking Lake Malawi cichlid fish, Hemitilapia oxyrhynchus, in which males build and aggressively defend sand 'bowers'. Similar to previous studies, we found that male reproductive success was positively associated with bower height and centrality on the lek. However, this pattern resulted from males holding these territories encountering more females, and thus their greater success was due to indirect mate choice. Following initial male courtship, an increase in the relative mating success of some males was observed, but this relative increase was unrelated to bower size or position. Crucially, experimentally manipulating bowers to resemble those of a co-occurring species had no appreciable effect on direct choice by females or male spawning success. Together, these results suggest indirect mate choice is the dominant force determining male-mating success in this species, and that bowers are not signals used in direct mate choice by females. We propose that, in this species, bowers have a primary function in intraspecific male competition, with the most competitive males maintaining larger and more central bowers that are favoured by sexual selection due to higher female encounter rates.     

The evolution, maintenance and adaptive function of genetic colour polymorphism in birds

The hypothesis that ornaments can honestly signal quality only if their expression is condition-dependent has dominated the study of the evolution and function of colour traits. Much less interest has been devoted to the adaptive function of colour traits for which the expression is not, or is to a low extent, sensitive to body condition and the environment in which individuals live. The aim of the present paper is to review the current theoretical and empirical knowledge of the evolution, maintenance and adaptive function of colour plumage traits for which the expression is mainly under genetic control. The finding that in many bird species the inheritance of colour morphs follows the laws of Mendel indicates that genetic colour polymorphism is frequent. Polymorphism may have evolved or be maintained because each colour morph facilitates the exploitation of alternative ecological niches as suggested by the observation that individuals are not randomly distributed among habitats with respect to coloration. Consistent with the hypothesis that different colour morphs are linked to alternative strategies is the finding that in a majority of species polymorphism is associated with reproductive parameters, and behavioural, life-history and physiological traits. Experimental studies showed that such covariations can have a genetic basis. These observations suggest that colour polymorphism has an adaptive function. Aviary and field experiments demonstrated that colour polymorphism is used as a criterion in mate-choice decisions and dominance interactions confirming the claim that conspecifics assess each other's colour morphs. The factors favouring the evolution and maintenance of genetic variation in coloration are reviewed, but empirical data are virtually lacking to assess their importance. Although current theory predicts that only condition-dependent traits can signal quality, the present review shows that genetically inherited morphs can reveal the same qualities. The study of genetic colour polymorphism will provide important and original insights on the adaptive function of conspicuous traits.     

Associations between environmental stress, selection history, and quantitative genetic variation in Drosophila melanogaster

Stressful environments may increase quantitative genetic variation in populations by promoting the expression of genetic variation that has not previously been eliminated or canalized by natural selection. This “selection history” hypothesis predicts that novel stressors will increase quantitative genetic variation, and that the magnitude of this effect will decrease following continued stress exposure. We tested these predictions using Drosophila melanogaster and sternopleural bristle number as a model system. In particular, we examined the effect of high temperature stress (31°C) on quantitative genetic variation before and after our study population had been reared at 31°C for 15 generations. High temperature stress was found to increase both additive genetic variance and heritability, but contrary to the selection history hypothesis prediction, the magnitude of this effect significantly increased after the study population had been reared for 15 generations under high temperature stress. These results demonstrate that high temperature stress increases quantitative genetic variation for bristle number, but do not support the selection history hypothesis as an explanation for this effect.     

Mate choice and the genetic basis for colour variation in a polymorphic dart frog: inferences from a wild pedigree

Understanding how reproductive barriers evolve during speciation remains an important question in evolution. Divergence in mating preferences may be a common first step in this process. The striking colour pattern diversity of strawberry dart frog (Dendrobates pumilio) populations has likely been shaped by sexual selection. Previous laboratory studies have shown that females attend to male coloration and prefer to court with males of their own colour, suggesting that divergent morphs may be reproductively isolated. To test this hypothesis, we used molecular data to estimate pedigree relationships from a polymorphic population. Whereas in the laboratory both red and yellow females preferred to court with males of their own phenotype, our pedigree shows a pattern of assortative mating only for red females. In the wild, yellow females appear to be less choosy about their mates, perhaps because they incur higher costs associated with searching than females of the more common red phenotype. We also used our pedigree to investigate the genetic basis for colour-pattern variation. The phenotype frequencies we observed were consistent with those expected if dorsal background coloration is controlled by a single locus, with complete dominance of red over yellow. Our results not only help clarify the role of sexual selection in reducing gene flow, but also shed light on the mechanisms underlying colour-pattern variation among sympatric colour morphs. The difference we observed between mating preferences measured under laboratory conditions and the pattern of mate choice observed in the wild highlight the importance of field studies for understanding behavioural reproductive isolation.     

Determination and distribution of a female-specific protein in the brown planthopper, Nilaparvata lugens Stal (Homoptera: Delphacidae)

Electrophoretic analyses of hemolymph and body or ovary homogenates from reproducing females, males, and 5th instar nymphs of the brown planthopper, Nilaparvata lugens Stal, revealed a protein band of 175 kDa in females. An immunoblot test using antibody against this protein showed a positive reaction with a 175 kDa protein from female body or ovary homogenates. It is likely that this protein in hemolymph is vitellogenin (Vg). Distribution of Vg was determined by immunofluorescence and immunogold labeling techniques. The results showed that the positive immunofluorescence reactions were present in yolk particles, the intercellular space of follicle cells, hemolymph, and the epithelial plug of ovarioles. In addition, the yeast-like symbiotes (YLS) in mycetocytes of adults and various nymphal instars as well as those free in hemolymph or entering oocytes also exhibited a positive reaction. Electron micrographs showed that immunogold particles were found most in yolk mass and YLS over other tissues. Especially the YLS in various developmental stages all contained immunogold particles, implying that the symbiote is somewhat related with production of the female-specific protein.