The effect of size,age, and recent egg laying on copulatory choice of the hermaphroditic mollusc Aplysia juliana

Summary

Aplysia juliana is a cross-fertilizing, simultaneous hermaphrodite. During copulation, an individual may act as either a sperm donor or a sperm recipient, or both, when a pair copulates reciprocally. Experiments were conducted with A. juliana to determine if an animal's size, age, or recent egg-laying activity influenced its choice of copulatory role. Animals were isolated except when paired during daily, half-hour trials. In the first experiment, mature animals of different size (weight) but similar age were randomly paired. Animal size had no effect on the initial copulatory role chosen. In the second experiment, young, maturing A. juliana were paired with older animals. Young animals showed no preference in initial copulatory role either as a group or individually. Older A. juliana showed no copulatory preference as a group, but over half of the individuals demonstrated a consistent choice of one role. Some individuals acted almost exclusively as sperm donors, others as sperm recipients, suggesting that as an A. juliana ages, it is likely to develop a preference for a single copulatory role. A record of daily egg mass production was kept for all animals in the second experiment. Production of an egg mass since the last copulation did not affect the copulatory role chosen in the subsequent copulation.     

Copulation in the hermaphroditic snail Lymnaea stagnalis: a review

Summary

Male copulatory behavior of the hermaphroditic snail Lymnaea stagnalis is a complex one: the appetitive behavior consists of a number of elements which do not always appear in the same sequence and have variable durations. Backfills of the penis nerve revealed the neurons that send projections to the male copulatory apparatus. Immunocytochemical experiments have demonstrated that these neurons contain at least ten different messenger molecules. Based on in situ hybridization and chemical purification data, it is suspected that this number will probably be doubled. How the different neurons and the molecules they contain might be involved in generation of the different elements of male copulatory behavior is discussed.     

Female response to predation risk alters conspecific male behaviour during pre‐copulatory mate guarding

Mating behaviour often increases predation risk, but the vulnerability within mating pairs differs between the sexes. Such a sex difference is expected to lead to differences in responses to predation risk between the sexes. In the two‐spotted spider mite Tetranychus urticae, males engage in pre‐copulatory mate guarding because only the first mating results in fertilisation. We investigated (i) whether pre‐copulatory pairs are more conspicuous to the predatory mite Phytoseiulus persimilis than solitary females, (ii) whether the vulnerability to the predator differs between sexes within the pre‐copulatory pair, (iii) whether each sex of T. urticae responds to predation risk during pre‐copulatory mate guarding and (iv) whether T. urticae's response to predation risk affects predator behaviour. Because T. urticae females are immobile during pre‐copulatory mate guarding, we observed male behaviour to evaluate effects of predation risk. We found that the predators detect more pre‐copulatory pairs than solitary females and that more females than males of the pre‐copulatory pairs are preyed upon by the predators. The preference of spider mite males for pre‐copulatory pairs versus solitary females was affected by whether or not the female had been exposed to predators during development. Male T. urticae exposed to predation risk did not alter their behaviour. These results suggest that only the most vulnerable sex, that is the female, responds to predation risk, which modifies male behaviour. Regardless of T. urticae females’ experience, however, P. persimilis detected more T. urticae pre‐copulatory pairs than solitary females, suggesting that pre‐copulatory mate guarding itself is dangerous for T. urticae females when these predators are present. We discuss our results in the context of sex‐dependent differences in predation risk.     

Mating Behavior, Sexual Selection, and Copulatory Courtship in a Promiscuous Beetle

The function of male movements during copulation is unclear. These movements may be a result of the necessary mechanics of insemination, or they may also have further function, for instance, stimulating or courting a female during mating, perhaps influencing female mate choice. We present data from three experiments exploring the mating behavior and copulatory movements of the highly promiscuous beetle Psilothrix viridicoeruleus. Male mating success in the struggle over mating was not related to male or female size (measured by weight) but successful males were more vigorous in terms of copulatory movements. These males took longer to mount females but copulated longer and remained mounted longer. We discuss these results in terms of the mating system of Psilothrix and also in terms of observations of the timing of insemination during copulation. We suggest that copulatory movements in this species are best understood as copulatory courtship.     

Copulatory Plug Displacement Evidences Sperm Competition in Lemur catta

Male displacement of copulatory (sperm) plugs from female vaginas provides further evidence for sperm competition in ring-tailed lemurs (Lemur catta), a gregarious prosimian species with a multimale, multifemale mating system. During two mating seasons, I studied two groups of free-ranging ring-tailed lemurs on St. Catherines Island, GA, USA. I observed 22 mating pairs in which males achieved penile intromission. Copulatory plug displacement by males occurred in 9 cases. Plugs were displaced during copulation by male penes upon withdrawl following deep vaginal thrusting. In every case of copulatory plug displacement, the male displacing a plug mated to ejaculation with the estrous female. In a mating system in which females typically mate with more than one male during estrous, often in succession, copulatory plug displacement may function to disrupt or preclude other males' successful insemination of estrous females. The effects of sperm plug displacement on paternity in Lemur catta are unknown, as no study had heretofore documented copulatory plug displacement in this species. The first-male mating advantage suggested for Lemur catta should be re-evaluated where mating order is known, and copulatory plug displacement during mating, or lack thereof, is identified. Because there is a tendency for first-mating males to mate-guard for longer periods of time in Lemur catta, the latency period between the first mate's ejaculation and that of subsequent mates may be an important determinant of male fertilization success.     

On Mertensiella luschani helverseni populations in Southwestern Anatolia

Abstract

The leech family Barbronidae is distinguished from the Salifidae and Erpobdellidae by the presence of two accessory copulatory pores, and by the ventral nerve cord with 19 free ganglia and 9 ganglia fused to form the posterior ganglionic mass. Some characteristics of Barbronia assiuti are revised, such as the presence of stylets and the annulation.     

Female walking during copulation reduces the likelihood of sperm transfer from males in the sweet potato weevil,Cylas formicarius

Behaviour during copulation can alter the fate of sperm of the mating males. This behaviour may exert selective pressure, resulting in the evolution of diverse reproductive behaviour, morphology, and physiology. This study examined the role of female copulatory behaviour on sperm fate in the sweet potato weevil, Cylas formicarius (Fabricius) (Coleoptera: Brentidae). In this species, males mount the female during copulation. The female frequently walks during copulation, carrying the male on her back. Here, we describe and quantify the copulatory behaviour of mating pairs and examine the sperm fate. Insemination success, as determined by the presence of sperm in the spermatheca, was lower when females walked for longer periods during copulation. This result emphasizes the value of studying variation in female copulatory behaviour in order to understand the factors that influence sperm fate. We discuss the implications of these results on sexual selection and utility in programs applying sterile insect techniques.     

EVIDENCE FOR WIDESPREAD COURTSHIP DURING COPULATION IN 131 SPECIES OF INSECTS AND SPIDERS,AND IMPLICATIONS FOR CRYPTIC FEMALE CHOICE

Male courtship behavior is generally thought to function prior to copulation, as an inducement to the female to allow the male to copulate with her; this study indicates however, that male courtship during and following copulation (“copulatory courtship”) is common in insects and spiders (81% of 131 species in 102 genera and 49 families, mostly Coleoptera, Hemiptera, Diptera, and Araneioidea). Copulatory courtship is apparently evolutionarily labile, as expected if it is under sexual selection; intrageneric variation occurred in all 17 genera in which more than one species was observed. In 81% of 94 species with copulatory courtship, the male abandoned the female soon after copulation ended; thus, copulatory courtship appears not to function generally to induce acceptance of further copulatory attempts. The most likely explanation for copulatory courtship is that it represents attempts by males to influence cryptic female choice. This suggests that an aspect of sexual selection by female choice not considered by Darwin may be more important than previously appreciated and that the common practice in evolutionary studies of measuring male reproductive success by counting numbers of copulations may sometimes be misleading because of cryptic female choice during and after copulation.     

Challenging the Aggressive Spillover Hypothesis: Is Pre‐Copulatory Sexual Cannibalism a Part of a Behavioural Syndrome?

Pre‐copulatory cannibalism – females devouring males during courtship – may bring no benefit to either sex. The ‘aggressive spillover hypothesis’ (ASH) posits that pre‐copulatory cannibalism represents a spillover of female aggressiveness from the juvenile foraging context, when aggressiveness is advantageous, to the adult mating context, when aggressiveness may be non‐adaptive or maladaptive. The ASH suggests that individuals exhibit limited plasticity in aggressive behaviours because they are genetically canalised for indiscriminate aggressiveness towards prey and conspecifics, including males. Hence, a tendency to employ pre‐copulatory cannibalism is a part of the female aggression syndrome, an assertion generally accepted in the personality field. We here re‐evaluate the previous findings in the light of personality criteria, which we propose for ASH validation: between‐individual differences, repeatability and heritability in tendency for pre‐copulatory attacks (and pre‐copulatory cannibalism) and voracity towards prey, and their correlation. To re‐evaluate ASH and to allow for additional or alternative explanations, we ask whether pre‐copulatory cannibalism depends on female hunger, mating status, size and/or male quality. Finally, we ask whether cannibalistic females have a reduced reproductive success as predicted by the ASH. While repeatability and heritability in voracity towards prey and its correlation with the tendency to engage in pre‐copulatory cannibalism were found in certain systems, we lack any evidence for repeatability and heritability in pre‐copulatory cannibalistic attempts and for its maladaptiveness. Rather than only resorting to the ASH, foraging and mate choice hypotheses may also explain pre‐copulatory cannibalism. We suggest clarifying the use of the terms sexual cannibalism (effect) and female aggressiveness or tendency to attack and devour males (cause), and argue that male strategies to avoid cannibalism should be considered. We propose testing the ASH as the explanation for pre‐copulatory cannibalism in those cases where female tendency to devour males correlates with actual pre‐copulatory cannibalism and when all the above criteria are fulfilled. Finally, we propose future directions for studying the ASH.     

Serum 17β-estradiol and progesterone associated with mating behavior during early pregnancy in female rhesus monkeys

The phenomenon of postconception mating behavior was examined in a social group of rhesus monkeys living in an outdoor compound. Periodic blood samples and daily vaginal swabs were obtained from nine females beginning several weeks prior to conception and continuing through 6 weeks of pregnancy to permit an assessment of ovarian hormonal events associated with mating during early pregnancy. Each of the females showed a discrete period of copulatory activity during the periovulatory period which ceased within several days after the 17β-estradiol (E₂) ovulatory peak. In agreement with earlier reports, only a percentage of subjects (44%) exhibited a period of postconception mating, with copulatory activity beginning 19.8 (± 1.9) days following the E₂ peak and continuing for 9.5 (± 1.3 days). Implantation bleeding was detected in all of the subjects with the onset 19.5 (± 0.68) days after the E₂ peak. The interval between the E₂ peak and the onset of implantation bleeding was similar for all females. However, the duration of implantation bleeding was significantly shorter in females who exhibited postconception mating. The females who displayed postconception copulatory activity had significantly lower mean serum progesterone concentrations (2.33 ± 0.24 ng/ml vs. 3.64 ± 0.37 ng/ml) during the period associated with implantation bleeding and copulatory behavior. Although both groups had elevated concentrations of serum E₂ during this period, levels in the females who displayed postconception mating were significantly lower (173.8 ± 19.2 pg/ml vs 223.9 ± 28.8 pg/ml). These data demonstrate that the occurrence of postconception mating behavior in this environment is associated with a distinct pattern of ovarian hormonal events, and analysis suggests that differences in steroid concentrations probably account for the observed differences in implantation bleeding and copulatory behavior during pregnancy.     

Evaluating the post‐copulatory sexual selection hypothesis for genital evolution reveals evidence for pleiotropic harm exerted by the male genital spines of Drosophila ananassae

The contemporary explanation for the rapid evolutionary diversification of animal genitalia is that such traits evolve by post‐copulatory sexual selection. Here, we test the hypothesis that the male genital spines of Drosophila ananassae play an adaptive role in post‐copulatory sexual selection. Whereas previous work on two Drosophila species shows that these spines function in precopulatory sexual selection to initiate genital coupling and promote male competitive copulation success, further research is needed to evaluate the potential for Drosophila genital spines to have a post‐copulatory function. Using a precision micron‐scale laser surgery technique, we test the effect of spine length reduction on copulation duration, male competitive fertilization success, female fecundity and female remating behaviour. We find no evidence that male genital spines in this species have a post‐copulatory adaptive function. Instead, females mated to males with surgically reduced/blunted genital spines exhibited comparatively greater short‐term fecundity relative to those mated by control males, indicating that the natural (i.e. unaltered) form of the trait may be harmful to females. In the absence of an effect of genital spine reduction on measured components of post‐copulatory fitness, the harm seems to be a pleiotropic side effect rather than adaptive. Results are discussed in the context of sexual conflict mediating the evolution of male genital spines in this species and likely other Drosophila.     

Copulation anatomy of Drosophila melanogaster (Diptera: Drosophilidae): wound-making organs and their possible roles

Males of several insect species inflict wounds on female genitalia during copulation. Such copulatory wounding also occurs in the fruit fly Drosophila melanogaster Meigen, 1830, one of the most important model organisms. Using a flash fixation technique with mating pairs of D. melanogaster, I examined the use and functions of the male phallic organ within the female reproductive tract. Paired components of the phallic organ (gonopods and two pairs of branches of the basal processes of the aedeagus) opened sequentially, from outer to inner components, during copulation. The dorsal branches of the aedeagal basal processes pierced the intima of the female reproductive tract at the lateral shallow folds. Consequently, mated females usually had a pair of melanized patches from repaired copulatory wounds. The sites that were stabbed by the dorsal branches were also clutched on the outside of the female oviscape (ovipositor) by the posterior process, which is a component of the periphallic organ. These structures likely function together as a mate-holding device. Male ejaculate labeled with rhodamine-B fluorescent dye entered the copulatory wounds in D. eugracilis Bock and Wheeler (Univ Texas Publ 7213:1–102, 1972), a related species, but not in D. melanogaster. Thus, copulatory wounds may function as an entrance for male seminal chemicals into the female circulatory system in D. eugracilis, but might not in D. melanogaster.     

Three new species of Philocorydoras Suriano, 1986 (Monogenoidea: Dactylogyridae) infecting the gills of callichthyids (Actinopterygii: Callichthyidae) from the Peruvian Amazonia

During a research on gill ectoparasites of callichthyids fishes from the Peruvian Amazonia, the following monogenoideans were found: Philocorydoras peruensis n. sp. from Corydoras splendens (Castelnau); Philocorydoras multiradiatus n. sp. and Philocorydoras jumboi n. sp. from Brochis multiradiatus (Orcés, V.). All new species described herein are mainly differentiated from their congeners based on the morphology of the copulatory complex. In P. peruensis n. sp. the cirrus is “J”-like shaped tube slightly tilted to one side, while in P. jumboi n. sp. is “J”-like shaped tube in a straight position and in P. multiradiatus n. sp. the cirrus is an arced tube with inflated base and distally narrow. Brochis (Orcés, V.) represents a new genus hosting species of Philocorydoras. All new species presented in this work represent the first species of Philocorydoras reported for Peru.

     

Mechanism of execution of sequential motor acts during copulation behavior in the male cricketGryllus bimaculatus DeGeer

Summary The roles of the mechanosensory afferents from the wings, cerci, tergites and genitalia in copulation behavior were examined by ablation and stimulation in the male cricketGryllus bimaculatus DeGeer.The sexually excited male cricket exhibited an intense posture (IP) upon contact stimulation of the elytra and the 4th to 9th abdominal tergites. This posture allowed the backward slipping (BWS) or hooking to take place subsequently.Backward slipping (BWS), which is the movement to get under the female, was elicited during IP by contact stimulation to the middle and distal regions of the dorsal surface of the cercus.Hooking, the coordinated movements for hanging the epiphallus onto the female's subgenital plate could be induced during IP by contact either on the dorsum, periproct or proximal 2 mm regions of the cercus. The latter two regions played a role in performing hooking accurately.Among four types of mechano-sensilla on the cercus the trichoid type was crucial for the initiation of BWS and hooking. Calculations revealed that about 60 trichoid hairs (4% of all the trichoid hairs in one cercus) were sufficient for the male to carry out hooking normally.The input from the bristle hairs on the epiphallus initiated the spermatophore extrusion (SPE) by swelling the endophallus.These results demonstrated that copulation behavior in the male cricket consisted of several motor acts and each act is triggered by specific input from the contact-sensitive sensilla on the elytra, tergites, cerci and genitalia. The sequential execution of each motor act is achieved because one motor act results in a positional change in contact with the female which in turn gives rise to another act. This type of motor control is a model of the so-called chain reaction in instinct behavior.Abbreviations BWS backward slipping - C cercus - EN endophallus - EP epiphallus - EPc epiphallic convexity - FW forewing - H hook - IP intense posture - HW hindwing - P pouch - PP periproct - SP spermatophore - SPE spermatophore extrusion - SEM scanning electron microscope     

Copulatory wounds in the monandrous ant species <Emphasis Type="Italic">Formica japonica</Emphasis> (Hymenoptera,Formicidae)

Males of several insect species inflict wounds on female genitalia during copulation, but the significance of such copulatory wounds for males is not clear. I compared the genitalia of virgin and mated Formica japonica females and for the first time report the occurrence of copulatory wounds in this monandrous ant species. All inseminated females examined had two types of melanized patches, indicating wound repair, and the serrated penis valves and sharp-pointed volsellar digitus of male genitalia were the likely instruments of these wounds. Physically damaging mating in monandrous species supports the view that copulatory wounds do not necessarily contribute to postcopulatory fitness gains for males via advantages in sperm competition or cryptic female choice. Received 10 September 2007; revised 15 October 2007; accepted 16 October 2007.     

Female genital configuration in the classification of Lepidoptera

In Coleoptera, Neuroptera, and Megaloptera, and the panorpoid orders Diptera, Trichoptera, and Mecoptera the common oviduct is ventral to, or opens into, the vagina or genital chamber. In Lepidoptera, in the superfamilies Micropterigoidea, Eriocranioidea, Incurvarioidea, and Nepticuloidea, the common oviduct enters the copulatory chamber ventrally; in Mnesarchaeidae, Hepialoidea, and all Ditrysia auct. the common oviduct is dorsal to the copulatory chamber, and the vagina (that region posterior to the entry of the spermatheca) opens separately from the genital ostium. Lepidoptera are unique in the complex and various arrangements of the ectodermal elements in the female genitalia.

The arrangements of, and connections between rectal and genital structures in representatives of Megaloptera, Neuroptera, and the panorpoid orders are re‐examined for comparison with the systems found in Zeugloptera, Dacnonypha, Monotrysia, and Ditrysia auct.

The Zeugloptera are here included in Lepidoptera because they have a circumcloacal chamber. Other female zeuglopteran genital structures intergrade with those of Dacnonypha, here restricted to Eriocraniidae, Agathiphagidae, Lophocoronidae, and the divergent Acanthopteroctetes. Dacnonypha have a less specialised spermathecal and vaginal structure than do most of the Monotrysia, here restricted to Incurvarioidea and Nepticuloidea (including Tischeriidae); many Monotrysia lack a cloaca, whereas it is always present in Dacnonypha.

There is no basis for retaining either Mnesarchaeidae or Hepialoidea in Monotrysia auct., as the dorsal common oviduct and the two genital openings indicate that these groups are ditrysian. They are here regarded as exoporian Ditrysia, a group characterised by the lack of a free, tubular ductus seminalis. A fixed gutter or channel between ostium and ovipore characterises the Hepialoidea, and the absence of this channel (ostium and ovipore opposable within an external genital pouch) characterises Mnesarchaeidae.

The endoporian Ditrysia all have a free, tubular ductus seminalis; where a cloaca is present it is incomplete, i.e., combines ovipore and rectum but never copulatory structures, in contrast to the complete cloaca found in Zeugloptera, Dacnonypha, and many Monotrysia. The endoporian Ditrysia comprise all other superfamilies, i.e., about 97% of species of Lepidoptera.     

Dedifferentiation and redifferentiation of the penis of the freshwater planarian Bdellocephala brunnea

Following decapitation of Bdellocephala brunnea Ijima et Kaburaki and subsequent repeated removal of regenerating head-blastemas, the copulatory apparatus degenerated within 40–60 days. The copulatory apparatus disintegrated into cell clusters that further divided into separate neoblast-like cells that, in turn, eventually transformed into ordinary parenchymal cells by 60 days after decapitation. When the head of worms with a degenerate copulatory apparatus was allowed to regenerate by discontinuing excision of the head blastema, a new copulatory apparatus differentiated, ostensibly from the dedifferentiated cells. The neoblast-like cells appear to participate in the redifferentiation of the penis and of other parts of the copulatory apparatus.     

Female behaviour and the interaction of male and female genital traits mediate sperm transfer during mating

Natural selection and post‐copulatory sexual selection, including sexual conflict, contribute to genital diversification. Fundamental first steps in understanding how these processes shape the evolution of specific genital traits are to determine their function experimentally and to understand the interactions between female and male genitalia during copulation. Our experimental manipulations of male and female genitalia in red‐sided garter snakes (Thamnophis sirtalis parietalis) reveal that copulation duration and copulatory plug deposition, as well as total and oviductal/vaginal sperm counts, are influenced by the interaction between male and female genital traits and female behaviour during copulation. By mating females with anesthetized cloacae to males with spine‐ablated hemipenes using a fully factorial design, we identified significant female–male copulatory trait interactions and found that females prevent sperm from entering their oviducts by contracting their vaginal pouch. Furthermore, these muscular contractions limit copulatory plug size, whereas the basal spine of the male hemipene aids in sperm and plug transfer. Our results are consistent with a role of sexual conflict in mating interactions and highlight the evolutionary importance of female resistance to reproductive outcomes.     

Copulatory Courtship in Drosophila birchii and D. serrata,Species Recognition and Sexual Selection

Two endemic Australian Drosophila species, D. birchii and D. serrata, have a copulatory courtship, i.e., the males court the female mainly during copulation. In the present study we found the males of both species to mount their prospective mating partners selectively, exhibiting both sex and species recognition. The males began to sing after mounting the female, and they often exhibited also postcopulatory displays typical to copulatory courtship. D. birchii and D. serrata females discriminated against males which did not sing during mounting/copulation, which suggests that the females utilize cryptic female choice. Our findings raise the question of how widespread a phenomenon cryptic female choice is in Drosophila species.     

Paternity analyses in wild‐caught and laboratory‐reared Caribbean cricket females reveal the influence of mating environment on post‐copulatory sexual selection

Polyandry is ubiquitous in insects and provides the conditions necessary for male‐ and female‐driven forms of post‐copulatory sexual selection to arise. Populations of Amphiacusta sanctaecrucis exhibit significant divergence in portions of the male genitalia that are inserted directly into the female reproductive tract, suggesting that males may exercise some post‐copulatory control over fertilization success. We examine the potential for male–male and male–female post‐copulatory interactions to influence paternity in wild‐caught females of A. sanctaecrucis and contrast our findings with those obtained from females reared in a high‐density laboratory environment. We find that female A. sanctaecrucis exercise control by mating multiple times (females mount males), but that male–male post‐copulatory interactions may influence paternity success. Moreover, post‐copulatory interactions that affect reproductive success of males are not independent of mating environment: clutches of wild‐caught females exhibit higher sire diversity and lower paternity skew than clutches of laboratory‐reared females. There was no strong evidence for last male precedence in either case. Most attempts at disentangling the contributions of male–male and male–female interactions towards post‐copulatory sexual selection have been undertaken in a laboratory setting and may not capture the full context in which they take place – such as the relationship between premating and post‐mating interactions. Our results reinforce the importance of designing studies that can capture the multifaceted nature of sexual selection for elucidating the role of post‐copulatory sexual selection in driving the evolution of male and female reproductive traits, especially when different components (e.g. precopulatory and post‐copulatory interactions) do not exert independent effects on reproductive outcomes.