Yoshitaka Kamimura

Possible Removal of Rival Sperm by the Elongated Genitalia of the Earwig, Euborellia plebeja

Abstract Sperm displacement is a sperm competition avoidance mechanism that reduces the paternity of males that have already mated with the female. Direct anatomical sperm removal or sperm flushing is known to occur in four insect orders: Odonata, Orthoptera, Coleoptera and Hymenoptera. In a fifth order, Dermaptera (earwigs), I found that the virga (the elongated rod of the male genitalia) of Euborellia plebeja seems to be used to remove rival sperm from the spermatheca (a fine-tubed female sperm storage organ). In this species, copulation lasted on average 4.6 minutes, during which time the male inserted the virga deep into the spermatheca, and then extracted it ejaculating semen from the opening of the virgal tip. The extraction of virgae (with its brim-like tip) appeared to cause removal of stored sperm in the spermatheca. The virga was as long as the body length of males, and the spermatheca was twice the female body length. The long length of the spermatheca and the possible removal function of the virga may select for virgal elongation.

Local mate competition with lethal male combat: effects of competitive asymmetry and information availability on a sex ratio game.

We constructed a sex allocation model for local mate competition considering the asymmetry of competitive abilities among sons. This model assumes two females of a parasitoid wasp oviposit on the same host in sequential order. The evolutionarily stable strategy will be in either Stackelberg or Nash equilibrium, depending on whether the females can recognize their opponents sex ratio or not, respectively. The Nash equilibrium predicts the second female produce more males than the first. If the second female is able to know and respond to the strategy of the first (a Stackelberg equilibrium), the first will decide an optimal sex ratio assuming that the second reply to it. Under such an assumption, our model predicts that not producing sons is adaptive for the second female when the sons she produces have low competitive ability. Males of parasitoid wasps Melittobia spp. are engaged in lethal male–male combat, indicating large asymmetry in mating success among sons. If females have the ability to recognize their opponents sex ratio, our model suggests that the severe lethal male–male combat may be one factor explaining their extremely female‐biased sex ratio that is unexplainable by pre‐existent models.

Effects of repeated mating and polyandry on the fecundity, fertility and maternal behaviour of female earwigs,Euborellia plebeja

Abstract I examined multiple mating and its function in female earwigs, Euborellia plebeja (Dermaptera: Anisolabididae). Like other earwigs, females of this species care for their eggs and intermittently lay eggs in clutches (iteroparity). Analysis of two polymorphic allozyme loci revealed that wild-caught adult females laid clutches with low within-brood genetic relatedness (0.210), indicating that females were promiscuous under natural conditions. Rearing experiments in the laboratory revealed that: (1) repeated mating with a single male increased female fecundity (number of clutches laid) and hence the number of hatchlings produced; (2) estimated sperm number was positively correlated with hatchability; (3) when frequency of mating was controlled, polyandry enhanced hatchability, although this effect was not statistically significant; (4) duration of maternal care varied for clutches with low hatchability, and sometimes exceeded the mean interclutch interval. Thus, although a possible benefit of polyandry is suggested, the greater beneficial effect of repeated mating on female fecundity can explain polyandrous mating in this species. Because female earwigs invest considerable effort in brooding their clutches, it may be adaptive to suppress oviposition unless stored sperm ensures high fertility. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.

A "spare" compensates for the risk of destruction of the elongated penis of earwigs (Insecta: Dermaptera)

Male animals in several groups have multiple intromittent organs that outnumber the corresponding female gonopore. In Dermaptera (earwigs), males of the family Anisolabididae have paired, elongated male intromittent organs (virgae), while females have a single sperm-storage organ (spermatheca). Several authors have assumed that one of the paired virgae is non-functional, because it points in the wrong direction. We investigated the mating success of handicapped males of Euborellia plebeja in which one of their paired virgae was removed experimentally. These handicapped males succeeded in inseminating a mate. Males with genital damage are found in the field, suggesting that the spare functions under natural conditions. Based on phylogenetic information on earwigs, we discuss possible evolutionary scenarios for this genital peculiarity.

Last-male paternity of Euborellia plebeja, an earwig with elongated genitalia and sperm-removal behavior

Both sexes of the earwig Euborellia plebeja (Dermaptera: Anisolabididae) mate frequently. The elongated intromittent organs of males are as long as their bodies. Previous studies have revealed that this organ is used to remove rival sperm from the female sperm-storage organ (spermatheca), the length of which is twice that of the female body. The fitness benefit of sperm removal was quantified using two mating experiments with paternity analysis. As expected, given that the sperm-removal organ is shorter than the sperm-storage organ, males gained only about 20% of paternity per single mating with sperm-saturated females. The significance of frequent repeated matings with the same female by males is discussed.

Effects of broken male intromittent organs on the sperm storage capacity of female earwigs, Euborellia plebeja

Abstract. In earwigs of the family Anisolabididae, male intromittent organs (virgae) sometimes break off inside female sperm-storage organs (spermathecae) during mating. I examined the effects of this genital breakage on the sperm storage capacity of females using Euborellia plebeja as a representative species. When genital breakage was artificially induced in virgin females, subsequent males successfully inseminated these females. However the sperm-storage capacity of these females was limited by the presence of broken virgae in their spermathecae. In another experiment, genital breakage was experimentally induced in the spermathecae of inseminated females, and their reproductive performance was then monitored for 60xa0days. In all of four cases where the entire piece of the broken virga remained inside the spermatheca, females deposited fertile eggs (more than 60% hatchability). The average number of clutches, that of eggs laid, and that of hatchlings were similar to those of controls. On the other hand, females laid no eggs in the other two cases where the broken virgae protruded from the spermathecal opening. I discuss the relevance of the results to the mating system and possible removal of rival sperm, which has been reported for E. plebeja.

The potential for incorporation of male derived proteins into developing eggs in the leafhopper Bothrogonia ferruginea

The leafhoppers, Bothrogonia ferruginea (Homoptera: Cicadellidae), eclose to adults in summer with immature reproductive organs. The adults live for 10 months including a hibernation of 4 months. Overwintered females mate multiply in spring. Eggs develop rapidly and are laid continuously in this mating period. Males produce sperm-bundles in which sperm are attached in a row to a rope-like hyaline material, and transfer them to the female via a large spermatophore that is placed in her bursa copulatrix. After mating, sperm are separated and removed to a spermatheca for storage prior to fertilization, but the sperm-binding material (trypsin degradable proteins) and the spermatophore disappear in the bursa and an enlarged portion of the genital duct. An injection of rhodamine B-dyed proteins into the female bursa with a microsyringe results in the production of intensely fluorescent eggs developing in the ovaries. This suggests that females could incorporate proteinaceous material derived from male spermatophores and/or sperm-binding material into their oocytes.

Mating behaviour and insemination in Diplatys flavicollis , an earwig with double-barrelled penises and a variable number of female sperm-storage organs

Earwigs (Insecta, Dermaptera) show astonishing diversity in penis morphology. In several families, males have a single penis (termed virga), whereas males of other families possess two functional virgae. Taxonomists have assumed that the two-virgae state is ancestral; however, ecological reasons why the ancestor acquired two virgae have not been explored. This study investigated in detail male and female genital structures, mating behaviour and insemination processes in the earwig Diplatys flavicollis (Diplatyidae). Diplatyidae are considered to be the most primitive family of earwigs. SEM and light-microscopy revealed that males of this species have two gonopores on each of two virgae, similar to those reported in other diplatyids (i.e. two double-barrelled penises), while females have four to six independent sperm-storage organs (spermathecae). Rapid fixation of mating pairs and insemination success of males from which one virga had been removed clearly revealed that only one virga was used for mating and was usually sufficient for inseminating multiple spermathecae. This finding rejects the one-to-one correspondence between male gonopores and female spermathecae. Based on allometric analysis of spermathecal variation, the possible significance of multiple spermathecae in relation to sperm-storage strategies of females is discussed. Compared to studies of male genital morphology, few studies have described spermathecal morphology. Based on compiled data of spermathecal and virgal morphology among earwigs, a parallel evolutionary trend between spermathecae and virgae from complicated (multiple) to simple (single) ones is suggested, and further investigation of the phylogeny, female genital morphology and insemination processes among earwigs is encouraged.

Promiscuity and elongated sperm storage organs work cooperatively as a cryptic female choice mechanism in an earwig

Female genitalia often show complex morphologies that cannot be explained by sperm reception and storage functions. However, our understanding of the forces underlying genital exaggeration in females is limited. Female earwigs, Euborellia plebeja , are promiscuous and their highly elongated sperm storage organs allow only partial removal and displacement of stored sperm by shorter male genitalia, resulting in only a 20% gain in paternity per copulation with a sperm-saturated female. This study examined the significance and optimality of restricted sperm displacement for females. A staged mating experiment with a paternity success analysis revealed that large males dominated male–male competition for burrows housing females, resulting in repeated copulations with the same female. Despite the low paternity gain per copulation, such repeated copulations resulted in a significant increase in paternity for larger males with higher resource-holding potential. A numerical simulation based on the relationship between male body size and copulation frequency showed that restricted sperm displacement (about 20% per copulation) is optimal for promiscuous females to accumulate sperm effectively from larger males. Because male body size is heritable in this species, females were estimated to benefit from a 1.4% increase in their sons mating success. This genetic benefit disappeared when only a single copulation per male–female encounter was assumed. Since no measurable costs of mating for females have been detected in this species, the combination of promiscuity and restricted sperm displacement is best explained by the cryptic female choice hypothesis.

Origin of the Transition from Aquatic to Terrestrial Habits in Nothopsyche Caddisflies (Trichoptera: Limnephilidae) Based on Molecular Phylogeny

Abstract The larvae and pupae of most caddisflies (Trichoptera) are aquatic, whereas the adults are terrestrial. However, the trichopteran genus Nothopsyche includes species with terrestrial pre-pupal and pupal stages and with an entirely terrestrial life cycle. The phylogenetic relationships within this genus, inferred from sequences of mitochondrial 16S and nuclear 18S ribosomal RNA genes, suggest that Nothopsyche species were originally aquatic and that a single lineage acquired tolerance to terrestrial habitats at the pre-pupal and pupal stages. In this lineage, N. montivaga became completely terrestrial. In addition, the larval case materials changed from plant matter to sand at one point in the phylogeny of this genus.