Effects of delayed mating and male mating history on the reproductive potential of Leucoptera coffeella (Lepidoptera: Lyonetiidae)

Abstract 1 Despite the importance of Leucoptera coffeella (Guérin‐Mèneville) in coffee production worldwide, there is a lack of information on its reproduction. This knowledge will help in mass rearing, and support the development of behavioural control techniques for this insect. The present study determined the effects of delayed mating and previous matings of male L. coffeella on fecundity, egg viability and frequency of female remating. 2 The highest levels of fecundity and egg viability were obtained from matings of 1–3‐day‐old females. When females mated at 5 days of age, there were reductions of 40% in oviposition and of 43% in egg viability. 3 Females mated with 2‐day‐old virgin males were more fecund than those mated with older males; egg viability was also low (18%) from females mated with older males. 4 Virgin females that mated with virgin males laid a greater number of eggs than those mated with previously copulated males. Egg viability decreased with the increase in the number of previous male matings. 5 Five‐day‐old females remated in greater proportion than 2–3‐day‐old females. Females that copulated with males that had previously mated three times had higher rates of remating than those that copulated with virgin males. 6 The results obtained indicate that 1–3 days after emergence is the optimum age for mating. The implications of these findings for the control of L. coffeella by synthetic sex pheromone are discussed.     

Unusual sex roles in a highly promiscuous parrot: the Greater Vasa Parrot Caracopsis vasa

We describe the unusual mating system of the Greater Vasa Parrot Caracopsis vasa . The dull black plumage of males and females is similar but females are significantly larger than males. Females are promiscuous and copulated with at least five different males. Copulations were either short (1–3 s) or very long (mean 35.9 min), and long copulations involved a copulatory tie facilitated by the male's enlarged cloacal protrusion. Multilocus DNA fingerprinting of 17 broods showed that all were of mixed paternity, and that some broods had three fathers. Males never visited nests directly, but during the incubation and chick-rearing periods females came off the nest and were fed regurgitated fruit by multiple males. Four had band-sharing coefficients that suggested they were unrelated. Males copulated with and provided food for several widely separated females simultaneously. During the chick-rearing period females defended a territory around the nest from conspecific females, developed conspicuous orange skin on the head (through feather loss), and uttered loud, complex vocalizations that we refer to as 'song' from prominent perches near the nest. Males showed none of these traits. Females with high song rates attracted more males and as a result received more food than other females. Play-back experiments in which female song rates were either increased or decreased, attracted more or fewer males respectively. We propose that female song, conspicuous head colour and territoriality have all evolved as a result of competition between females for the food provided by males. The selective pressures favouring this highly unusual breeding system in the Greater Vasa Parrot are unclear but some sort of ecological constraint, such as food availability, may be important.     

Reproductive strategies of the larval parasitoid Microplitis croceipes

Mate choice may have important consequences for offspring sex ratio and fitness of haplodiploid insects. Mate preference of females of the solitary larval parasitoid Microplitis croceipes (Cresson) (Hymenoptera: Braconidae) for virgin and mated males, and vice versa, and the reproductive consequences (i.e., the sex ratio expressed as the proportion of male offspring) were examined in choice and non‐choice experiments. In addition, the effect of repeated rapid and daily copulation of an individual male on the sex ratio of offspring of the female mates was assessed. Males preferred virgins over mated females, whereas females copulated with a male irrespective of his mating status. In both the rapid and daily copulation assay, females copulating with a male that had copulated five times or more produced a higher sex ratio than females that had copulated with a virgin male. Females that copulated with virgin males once or twice produced a significantly and considerably lower sex ratio than females that first copulated with a sperm‐depleted male followed by a virgin male. This indicates that copulating with a sperm‐depleted male has costs and limits acquisition by the female of sperm from virgin males.     

Comparative effect of short term exposures of Callosobruchus subinnotatus to carbon dioxide, nitrogen, or low temperature on behavior and fecundity

The effect of anesthetizing with carbon dioxide, nitrogen, or low temperature on the reproductive behavior and fecundity of Callosobruchus subinnotatus was investigated. Males and females anesthetized with N2 as virgins had shorter recovery time, copulated earlier, and had shorter mating time than those subjected to CO2 or low temperature. Exposure of males and females to 2.0±0.5 °C for 10 min had the most profound effect on the recovery periods, copulation time, and duration of mating. Adults anesthetized with CO2 and low temperature laid fewer eggs than those anesthetized with N2. Females were more affected than males by the treatments, except when males were chilled for 10 min, and this prolonged the time for commencement of mating. The mode of anesthesia could affect egg production because vitellogenesis and choriogenesis take place in the imago stage of bruchids.     

Copulation type affects parturition in the guppy

In some animals, females are often compelled to mate with less desirable males due to the males' alternative mating tactics. Male guppies, Poecilia reticulata, exhibit courtship displays and cooperatively copulate with females. However, they also exhibit sneaking behaviors and coercively copulate with females. To examine the consequences of these two mating patterns, we investigated the influence of copulation type, i.e., cooperative or coercive, on parturition and brood size of females. A single female was allowed to freely contact and copulate with a single male only once. Males that cooperatively copulated with females had larger orange spot areas (an important criterion of female mate choice) than males that copulated coercively. Most females that were coerced into copulation did not give birth to offspring within 100 days after mating. The probability of parturition was high when females copulated cooperatively, and when their mates exhibited frequent postcopulatory jerking behavior. However, the results suggest that copulation type did not affect brood size. Brood size was positively influenced by both female body size and male orange spot area. These results suggest that parturient success is low when females are coerced into mating by less desirable males, whereas brood size is independent of copulation type.     

Function and genetics of long versus short copulations in the cactophilic fruit fly,drosophila mojavensis (diptera: drosophilidae)

Variation in copulation duration of Drosophila mojavensisstrains was influenced by both sexes. Males maintained predominant control, as copulation duration of pairs from different strains was more similar to that of the strain from which the male was derived, but female origin also contributed significantly to the duration of copulation. Variation among strains was controlled by genes acting additively in both sexes. The size of both males and females also affected copulation duration. Small males copulated longer on average than large males, while males paired with large females copulated longer than those paired with small females. The importance of copulation duration to fitness was tested by correlation analyses with male size, female size, female remating latency, and number of eggs laid prior to female remating. Longer copulations stimulated earlier oviposition, possibly by increasing accessory gland secretions that are passed by males during copulation.     

Female brown-headed cowbirds', Molothrus ater, organization and behaviour reflects male social dynamics

In four large aviaries, we studied social assortment and reproductive behaviour of female brown-headed cowbirds housed with males differing in age class and in corresponding levels of intrasexual interaction. Juvenile and adult females resided with either (1) adult males, (2) juvenile males, (3) adult and juvenile males, or (4) no males. We observed social behaviour of males and females from September 1999 through to the 2000 breeding season. Throughout the year, males in the different conditions showed different levels of social interaction, with adult males being most engaged in male-male interactions and juvenile males being least engaged in male-male interactions. Females in the four conditions differed in their associations with males and with other females. In conditions with adult males, females spent more time near males, approached males more often, and associated with females less. During the breeding season, females in conditions with more adult males copulated more and produced more fertile and viable eggs. In the condition where females were housed with juvenile males, they copulated less, laid fewer eggs and destroyed more of their eggs. Results indicate that throughout the year, females are sensitive to male age and behaviour in their social group, and that this sensitivity can have reproductive consequences. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Effects of morphological traits,age and copulation experience on mate choice in Ophraella communa

Ophraella communa is a biological control agent of invasive common ragweed, Ambrosia artemisiifolia. To understand the mate choice tactic of the beetle and improve mass-rearing of high-quality populations, the effects of morphological traits, age and copulation experience on mate choice in this beetle were studied in the laboratory. The results showed thatmate choice of male or female was related to certain adult morphological traits. Wing length, black streak width of frons and metaleg femur length of males were central to female mate choice. Females with longer wing length, black streak width of frons, pronotum width and foreleg tibia length were more attractive to males than those with the smaller ones. The chosen rate of older males was significantly higher than those of younger ones, and the chosen rate of females reached a maximum value on the 8th day after eclosion. The newly copulated females were less attractive to males than virgins, but females that have copulated in the distant past were more attractive than or were similar to virgins. The chosen rates of virgin males were significantly higher than those of newly copulated and 8 days past copulated ones, but no difference between virgin males and 3, 5 or 12 days past copulated ones in O. communa. The results of our study have added much needed empirical data regarding the significance of morphological traits, age and copulation experience as a source of variation in insect copulation signals. In addition, the results provide valuable information for mass-rearing of high-quality populations of O. communa.     

Gonadal hormones and sexual behavior in groups of adult talapoin monkeys (Miopithecus talapoin).

Three heterosexual groups of six to eight monkeys were studied; all females were ovariectomized, whereas males were either intact or castrated. Aggressive hierarchies were evident in all groups, with females generally outranking males. When females were treated with estradiol, all males looked more frequently at the latters' sexual skin swellings, but only one male who was both dominant and intact copulated with them. Thus, either castration or low rank resulted in decreased levels of sexual behavior in male talapoins. The sexual behavior of dominant castrated males was restored by testosterone therapy, whereas subordinate castrates never copulated, even after large doses of testosterone, though penile erections and ejaculatory reflex (during masturbation) were restored. Following removal of a dominant male, the sexual behavior of the next male in rank was restored, provided he was not castrated and untreated. In contrast to males, female talapoins showed no consistent correlation between their rank and sexual activity. Estradiol therapy was without overall effect upon the frequency of female mounting behavior, though some females mounted and presented to one another more often. Estradiol treatment also caused females to present to males more frequently, but only to those that were sexually active (i.e., who mounted females).     

The adaptive significance of non-contact mate guarding by males of the dragonfly,Nannophya pygmaea Rambur (Odonata: Libellulidae)

InNannophya pygmaea, ovipositing females were frequently disturbed by conspecific males. Disturbed females often copulated with one of these males or flew away from the pool. Females which flew away from the pool due to male disturbance often returned later the same day and mated with different males. A territorial male would guard his ovipositing mate by hovering above her, presumably trying to prevent her from moving out of his territory. A non-territorial male would also guard his mate in a similar way, both at a vacant water area which was not occupied by any territorial males, or within the territory of a resident male. In addition, both territorial and non-territorial males chased intruding males in an attempt to prevent their mates from being stolen. Territorial males defended their mates better than non-territorial males. Both males and females often mated more than once in the course of a single day. Some territorial males copulated with a new female while another mate oviposited in their territories. This observation supported the “multiple mating hypothesis” proposed by Alcock (1979) and Uéda (1979) but other evidence suggested that this is an inadequate explanation for the non-contact guarding ofN. pygmaea.     

A male fitness advantage to wing reduction in the oriental chinch bug,Cavelerius saccharivorus okajima (heteroptera: Lygaeidae)

A male fitness advantage to wing reduction was investigated for the oriental chinch bug, Cavelerius saccharivorus, which is wing dimorphic. Field surveys for the frequency of matings between wing morphs showed that the mating probability was much higher in brachypterous males than in macropterous males in the early breeding season. Brachypterous males copulated with not only brachypterous females but also macropterous females in this season. This led to a considerable rate (30%) of insemination of macropterous females just before emigration even in the early breeding season. A rearing experiment for the pre-reproductive period of females revealed that females mated with brachypterous males copulated and oviposited earlier than females mated with macropterous males. These results indicate that there is a fitness advantage to wing reduction in the males of C. saccharivorus in terms of the earlier sexual maturation.     

Intrasexual copulation and mate discrimination in a population of Nodilittorina radiata (Gastropoda: Littorinidae)

Intrasexual copulation and mate discrimination by Nodilittorina radiata (Gastropoda: Littorinidae) were studied on a concrete breakwater at Hakodate Bay, southern Hokkaido, Japan. Intrasexual (male–male) copulations were observed in 4.7–21.1% of copulating pairs on the shore. As females were relatively larger than males and males copulated with females larger than themselves, we hypothesized that males choose potential mates larger than themselves. However, two male mates showed no significant size preference in intrasexual copulations, suggesting that males do not choose relatively larger individuals as mates. In a laboratory mate-choice experiment, male N. radiata preferred to mate with females, indicating precopulatory sex identification. They copulated with males, however, at the frequency of 37%, perhaps because of sex misidentification.     

Female sexual receptivity and male copula guarding during prolonged copulations in the damselflyIschnura senegalensis (Odonata:Coenagrionidae)

Laboratory experiments were conducted to clarify the relationship between female sexual receptivity and male copula guarding inI. senegalensis, a species that copulates for several hours. In insectaries, most copulations were initiated early in the morning, and terminated relatively synchronously between 11 00 and 13 00. Females refused males with wing-flutter display and oviposited alone in the afternoon regardless of copulation events of that morning. Females could sexually receive males only in the morning. Males copulated for several hours until 12 00 after which females could oviposit. To determine whether copulations that last for hours function as male copula guarding or only of sperm displacement, emerged males were kept at various densities and permitted to copulate with virgin and mated females in insectaries. Both with virgin and mated females, “social” (not solitary; 2–4 males / insectary) males initiated copulations early in the morning and always terminated at around 12 00. However, both with virgin and mated females, solitary (one male / insectary) males terminated copulations in the morning. In both cases, duration of copulations did not significantly differ for virgin females and mated females. Therefore, long (several hour) copulation is more likely to function as male copula guarding than as sperm displacement, and duration of copulations is predicted to be shortened when male density is very low.     

Reproductive interference by male Drosophila subobscura on female D. persimilis: A laboratory experiment

While females often reject courtship attempts by heterospecific males, reproductive interference by harassment from such males can nonetheless occur, potentially reducing female fitness. Such effects may be profound following a range expansion, when males from a new species may suddenly encounter (and perhaps even become abundant relative to) females of related native species. Drosophila subobscura recently invaded North America and may impact native species through reproductive interference and other processes. We test for the potential for reproductive interference by D. subobscura males on D. persimilis females in the laboratory. D. subobscura males aggressively copulated with D. persimilis females, including many females that exhibit rejection behaviors. Despite females attempting to dismount the males, the heterospecific copulations are on average longer than conspecific copulations, and females exhibit some reluctance to remate with conspecific males following this harassment. Females confined with both conspecific and heterospecific males produce fewer adult progeny than those with either conspecific males only or with conspecific males and distantly related D. simulans males that do not engage in female harassment. Overall, our results illustrate how reproductive interference by an invasive species can have negative effects on resident natural populations.     

Progesterone receptor in the forebrain of female gray short-tailed opossums: Effects of exposure to male stimuli

Progesterone receptor immunoreactivity (PRir) in brain areas involved in reproductive behavior in eutherian species was examined for the first time in a female marsupial, the gray short-tailed opossum (Monodelphis domestica, hereinafter, opossum). PRir in nuclei of neurons, measured as area covered by stained nuclei, was seen in the arcuate nucleus (Arc); anteroventral periventricular nucleus (AVPv); bed nucleus of the stria terminalis (BST); medial preoptic area (MPOA), and ventromedial hypothalamus (VMH), but not in control areas adjacent to the hypothalamus or cortex. Female opossums are induced into cytological, urogenital sinus (UGS), estrus by male pheromones and into behavioral estrus, i.e., receptivity, by pairing with a male, and both estradiol (E) and progesterone (P) are involved in induction of receptivity in intact and ovariectomized females. PRir in the AVPv, MPOA, and VMH was very low in females that had never been exposed to males or their scent marks, i.e., naïve anestrous (NVA) females, and either previous or current exposure to males or their scent marks was associated with elevated PRir. PRir was significantly higher in the AVPv and MPOA of anestrous females with previous but no current exposure to males and their scent marks, i.e., experienced anestrous (EXPA) females, than in NVA females, but PRir was significantly lower in the MPOA and VMH of EXPA females than in females that were behaviorally receptive and had recently copulated, i.e., behavioral receptive estrous (BRE) females. PRir was higher in the VMH of both UGS estrous (UGSE) and BRE females compared to that in EXPA animals, but PRir did not differ between UGSE and BRE females in any of the 3 brain areas examined, including the MPOA These results provide evidence that pheromonal induction of estrus and sexual receptivity in opossums is associated with elevation of PRir in the VMH and MPOA and that prior exposure to males or their pheromones, even in the absence of current male stimuli, is associated with persistent elevation of PRir in the AVPv and MPOA.     

Mating Frequency, Fecundity and Fertilization Success in the Hide Beetle, Dermestes Maculatus

While the immediate benefits accrued to females through multiple mating are well documented, the effect of sperm depletion for multiply mating males is rarely considered. We show that, in small mixed-sex laboratory aggregations, both male and female hide beetles, Dermestes maculatus (De Geer) mated multiply. There was considerable variation in the mating frequency of both sexes; however the skew in mating success was comparable for males and females. Several individuals that mated multiply also re-mated with a previous partner, but in a competitive environment no male copulated more than seven times. Mating success was unrelated to an individual's size, but males that had the most inter-sexual matings also engaged in the most intra-sexual mating attempts. In a second experiment, we show that, even in the absence of rivals, only a small number of males mated with all available virgin females. Moreover, even though males were mated twice to each female, males that copulated more than eight times failed to fertilize any eggs. We suggest that under natural conditions male hide beetles may refrain from mating either prior to, or at the point of, sperm depletion thereby reducing the selection pressure for females to discriminate against sperm depleted males. However, fecundity and fertilization success varied considerably across females and even those mating with sperm-replete males were unable to fertilize 100% of their egg batch. Thus, direct fertilization benefits accrued by females through mating more than once with the same male may play a key role in the maintenance of polyandry in this species.     

The effects of the antiestrogen CI-628 on sexual behavior activated by androgen or estrogen in quail

This series of experiments sought to determine whether conversion of androgen to estrogen is important in the activation of male sexual behavior in quail by seeing if an antiestrogen will block androgen stimulated copulation in this species. Experiment I compared the ability of two antiestrogens, MER-25 (5 mg/day) and CI-628 (2 mg/day), to block estrogen stimulated characteristics in female quail. Both treatments greatly reduced oviduct growth in “photically castrated” females given estradiol benzoate (EB, 50 μg/day), but only CI-628 reduced receptivity in these birds. In Experiment II surgically castrated males given 50 μg/day EB together with 2 mg/day CI-628 were much less receptive than castrated males given EB alone, and in addition copulated in fewer tests. In Experiments III, IV, and V, castrated males given testosterone propionate (TP) together with CI-628 were compared with males given TP alone. The ability of CI-628 to suppress TP-stimulated copulation increased with increasing CI/TP dosage ratio, and at the highest ratio (4:1), CI-628 effectively blocked copulation in five out of seven birds. Those birds that did copulate did so in fewer tests and performed fewer cloacal contact movements. CI-628 had no antiandrogenic effects in these experiments. These results suggest that estrogens may be important active metabolites of testosterone with respect to quail copulation.     

Quantity of the male ejaculate influenced by female unreceptivity in the fly, Culicoides melleus

The number of sperm placed in the spermatophore by virgin Culicoides melleus males mated with virgin females was compared with the number deposited in matings with previously once or twice copulated females. When compared to an all-virgin mating, there was a significant reduction (P<0·001) in sperm complement (of 15 per cent) when the female had already been mated once as compared to a virgin, and a greater reduction (of 21 per cent) when the female had already been mated twice. The mean numbers (± S.E.) of sperm ejaculated with virgin, once mated, and twice mated females were 857 ± 24, 732 ± 23, and 673 ± 25, respectively.When a major activity in the unreceptive behaviour of females (kicking at the male in the early stages of copulation) was prevented by amputation of the hind legs, the mean number of sperm ejaculated (788 ± 29) was significantly larger (P<0·01) compared with twice mated females having both legs intact, and also larger, though not significantly so, than with intact once mated females. In addition to the kicking activity, there was evidence that other forms of unreceptive behaviour contributed to reducing the quantity of sperm ejaculated.     

Multiple Copulations and Post-Copulatory Guarding in a Free-Living Population of Sika Deer (Cervus nippon)

Multiple copulations by females and post‐copulatory guarding by males, were studied in a free‐living sika deer population, on Nozaki, an island of the Goto Group, off the coast of Kyushu, Japan. In 1990, 1991, and 1993, observations were carried out mainly on open grassland in the central section of the island. It was found, that 10 of 22 females in estrus copulated more than once and five of them copulated with several males (multi‐male copulations), while the remainder copulated with a single male (repeated copulation). Almost all copulations were followed by guarding behavior. Dominant males (DMs) guarded significantly longer and more effectively than subordinate males (SMs). Guarding was interrupted in four of the SMs, when the guarding male was forcibly driven away from a female by higher ranked males (‘take‐over’) and in three cases when the female left the guarding male spontaneously (‘escape’). The only interruption of guarding of a DM was caused when a lower ranked male sneaked towards the guarded female and mated with her briefly (sneak). Interruptions of guarding initiated by females (escapes) occurred more when they were guarded by SMs than by DMs. Our results suggest that female sika deer indulge in multiple copulations to seek better genetic quality, rather than insuring fertility, enhancing genetic diversity, or avoiding harassment. The post‐copulatory guarding by DMs appears to be more effective in the prevention of additional copulations by females with other males than guarding by SMs. Moreover, SMs decrease the duration of the pre‐copulatory phase to achieve copulation before having to give way to DMs.     

The effects of body size on mate choice in a grapsid crab,Gaetice depressus (Crustacea,Decapoda)

Male and female mate choices were investigated in the grapsid crab,Gaetice depressus (Crustacea, Decapoda) in a laboratory experiment. Males mated indiscriminately with regard to the body size of the females, and frequently copulated with the first females they encountered. In contrast, females showed mate discrimination with regard to the body size of males. The females tended to sample potential mates prior to copulation, and showed both a preference for the larger males and a tendency toward the rejection of males with body sizes smaller than their own. However, they did not discriminate between two males that were either larger or smaller than they themselves. Mate choice by the females of this species is though to be based upon threshold-criterion tactics, in which the body size of the female itself is used as a threshold value.