Published papers

Background: Many male animals donate nutritive materials during courtship or mating to their female mates. Donation of large-sized gifts, though costly to prepare, can result in increased sperm transfer during mating and delayed remating of the females, resulting in higher paternity. Nuptial gifting sometimes causes severe female-female competition for obtaining gifts (i.e., sex-role reversal in mate competition) and selection on females to increase their mating rate, changing the intensity of sperm competition and the resultant paternity gains. We built a theoretical model to simulate such coevolutionary feedbacks between nuptial gift size (male trait) and propensity for multiple mating (female trait). Donation of nuptial gifts sometimes causes development of female persistence trait for gift acquisition. We also analyzed the causes and consequences of this type of traits, taking double receptacles for nutritious seminal gifts, which are known to occur in an insect group with a “female penis” (Neotrogla spp.), as an illustrative example. Results: Our individual-based simulations demonstrated that female-female competition for male-derived nutrients always occur when the environment is oligotrophic and mating costs are low for females. However, a positive correlation between donated gift size and the resultant paternity gain was a requisite for the co-occurrence of large gifts and females’ competitive multiple mating for the gifts. When gift donation satisfied female demands and thus resulted in monandry, exaggeration of nuptial gift size also occurred under the assumption that the last male monopolizes paternity. The evolution of double slots for gift acquisition and digestion (female persistence trait) always occurred when males could not satisfy the demands of females for gifts. However, through coevolutionary reduction in male gift size, fixation of this trait in a population drastically reduced the average female fitness. Conclusion: Sperm usage patterns, which have rarely been examined for animals with nuptial gifts, can be a critical factor for determining the extent of exaggeration in nuptial gifting. Sex-role reversals in mate competition, as a result of donation of nuptial gifts from males to females, can involve the evolution of male-like, persistent traits in females that reduce population productivity, as is the case with persistence traits in males.

During mating, male bed bugs (Cimicidae) pierce the female abdomen to inject sperm using their needle-like genitalia. Females evolved specialized paragenital organs (the spermalege and associated structures) to receive traumatically injected ejaculates. In Leptocimex duplicatus, the spermalege is duplicated, but the evolutionary significance of this is unclear. In Cimex hemipterus and C. lectularius, in which females normally develop a single spermalege on the right side of the abdomen, similar duplication sometimes occurs. Using these aberrant morphs (D-females) of C. hemipterus, we tested the hypothesis that both of the duplicated spermaleges are functionally competent. Scars on female abdominal exoskeletons indicated frequent misdirected piercing by male genitalia. However, the piercing sites showed a highly biased distribution towards the right side of the female body. A mating experiment showed that when the normal insemination site (the right-side spermalege) was artificially covered, females remained unfertilized. This was true even when females also had a spermalege on the left side (D-females). This result was attributed to handedness in male mating behavior. Irrespective of the observed disuse of the left-side spermalege by males for insemination, histological examination failed to detect any differences between the right-side and left-side spermaleges. Moreover, an artificial insemination experiment confirmed that spermatozoa injected into the left-side spermalege show apparently normal migration behavior to the female reproductive organs, indicating an evolutionary potential for functionally-competent duplicated spermaleges. We discuss possible mechanisms for the evolutionary maintenance of D-females and propose a plausible route to the functionally-competent duplicated spermaleges observed in L. duplicatus.

Although several evolutionary forces have been proposed to contribute to genital morphological diversification, it is unclear which might act early during the evolution of novel structural traits. We test the hypothesis that mismatch between interacting male and female secondary sexual structures gives rise to increased harm to females, consistent with the outcome predicted from a history of sexual conflict. We mate Drosophila sechellia females to males from a collection of D. mauritiana-D. sechellia interspecific genetic introgression lines that possess quantitative morphological variation in the posterior lobe of the genital arch, an external genital structure that can cause wounds to the female abdomen during mating. We find that males with smaller posterior lobes, and those that possess lobes with similarities in shape to D. mauritiana, cause more severe wounding compared to either D. sechellia males with strain-specific morphologies or introgression males that possess larger lobes or lobes with more pronounced D. sechellia features. These results suggest a possible history of sexual conflict during the evolution of the posterior lobe in D. sechellia, but also suggest a potential contribution of divergence in sensory recognition mechanisms to posterior lobe evolution.

The puzzling sex ratio behavior of Melittobia wasps has long posed one of the greatest questions in the field of sex allocation. Laboratory experiments have found that, in contrast to the predictions of theory and the behavior of numerous other organisms, Melittobia females do not produce fewer female-biased offspring sex ratios when more females lay eggs on a patch. We solve this puzzle by showing that, in nature, females of Melittobia australica have a sophisticated sex ratio behavior, in which their strategy also depends on whether they have dispersed from the patch where they emerged. When females have not dispersed, they lay eggs with close relatives, which keeps local mate competition high even with multiple females, and therefore, they are selected to produce consistently female-biased sex ratios. Laboratory experimentsmimic these conditions. In contrast, when females disperse, they interact with nonrelatives, and thus adjust their sex ratio depending on the number of females laying eggs. Consequently, females appear to use dispersal status as an indirect cue of relatedness and whether they should adjust their sex ratio in response to the number of females laying eggs on the patch.

Animal terminalia represent some of the most diverse and rapidly evolving structures in the animal kingdom, and for this reason have been a mainstay in the taxonomic description of species. The terminalia of Drosophila melanogaster, with its wide range of experimental tools, have recently become the focus of increased interest in the fields of development, evolution, and behavior. However, studies from different disciplines have often used discrepant terminologies for the same anatomical structures. Consequently, the terminology of genital parts has become a barrier to integrating results from different fields, rendering it difficult to determine what parts are being referenced. We formed a consortium of researchers studying the genitalia of D. melanogaster to help establish a set of naming conventions. Here, we present a detailed visual anatomy of male genital parts, including a list of synonymous terms, and suggest practices to avoid confusion when referring to anatomical parts in future studies. The goal of this effort is to facilitate interdisciplinary communication and help newcomers orient themselves within the exciting field of Drosophila genitalia.

To explore diversity of earwigs (Dermaptera) in different agricultural ecosystems of South India, an extensive taxonomic survey was conducted in 2020 during which an undescribed species of Diplatys was collected. Twenty-one species of the genus Diplatys (Diplatyidae, Diplatyinae) have been reported to date from India, of which six species are known from Karnataka, South India. Based on a male specimen collected from a sugarcane field in Karnataka, a new species, Diplatys sahyadriensis sp. nov., is described as the twenty-second species of this genus from India. A revised key to the males of Diplatys species from India and Sri Lanka is provided. This new record adds to the known species diversity in the Sahyadri Ranges of the Western Ghats in Shivamogga District, Karnataka, part of the Southern Plateau and Hills agro-climatic region of India.

We surveyed natural population of the Drosophila ananassae species complex on Penang Island, Malaysia. Analyses of phenotypic traits, chromosome arrangements, molecular markers, and reproductive isolation suggest the existence of two species: D. ananassae and D. cf. parapallidosa. Molecular marker analysis indicates that D. cf. parapallidosa carries chromosome Y and 4 introgressions from D. ananassae. Thus, D. cf. parapallidosa seems to be a hybrid descendant that recently originated from a natural D. parapallidosa female x D. ananassae male cross. Furthermore, D. cf. parapallidosa behaves differently from authentic D. parapallidosa with respect to its reproductive isolation from D. ananassae. Premating isolation is usually seen in only the D. ananassae female x D. parapallidosa male cross, but we observed it in crosses of both directions between D. ananassae and D. cf. parapallidosa. In addition, hybrid males from the D. ananassae female x D. parapallidosa. cross are usually sterile, but they were fertile when D. ananassae female were mated with D. cf. parapallidosa male. We attempted an artificial reconstruction of the hybrid species to simulate the evolutionary process(es) that produced D. cf. parapallidosa. This is a rare case of natural hybrid population in Drosophila and may be a useful system for elucidating speciation with gene flow.

The pygidicranid earwigs (Dermaptera) of Penang Island, Peninsular Malaysia were re-examined based on material collected in extensive field surveys in 2012-2013 and 2015. Echinosoma roseiventre Kamimura & Nishikawa, sp. n. is described and illustrated, and Cranopygia pallidipennis (de Haan, 1842) is reported from the island for the first time. The taxonomic and nomenclatural problems of the genus Cranopygia sensu Hincks (1959) [A Systematic Monograph of the Dermaptera of the World. Part II. Pygidicranidae excluding Diplatyinae. British Museum (Natural History)] are also discussed. For the members of the subfamily Pygidicraninae from Indo-Austral and Oriental regions, the system, definitions of genera, and key of Hincks (1959) are followed. The genus Mucrocranopygia Steinmann, 1986 is synonymized with Cranopygia Burr, 1908. A key to the males of small Echinosoma from the Oriental region is provided.