Prior experience influences the fruit residence of male apple maggot flies,Rhagoletis pomonella

Male apple maggot flies spend considerable time residing on individual host fruit as territories on which they force-copulate arriving females in search of oviposition sites. Here, we present evidence from investigations in nature and the laboratory that shows the propensity of males to reside on a hawthorn or apple fruit as a territory is significantly modifiable through prior experience with fruit and, hence, involves learning. Previous studies revealed that after a female apple maggot fly, Rhagoletis pomonella, arrived on a host hawthorn or apple fruit, its propensity to accept or reject that fruit for egg-laying was similarly modifiable through prior fruit-exposure experience and also involved learning. We discuss how host fruit learning in males and females, in concert with genetic-based differences in host fruit residence and acceptance behavior between populations of flies originating from hawthorn and apple, could give rise to a reduction in gene flow between populations of flies on these two host types.     

Mating behavior and male mating success in wildAnastrepha Ludens (Diptera: Tephritidae) on a field-caged host tree

Mating behavior and factors affecting mating success of males were studied using wild Anastrepha ludens on a fieldcaged host tree. The most common courtship sequence had five components: (1) male calls from the underside of a leaf, (2) female arrives to the maleoccupied leaf, (3) male orients to female and stops calling, (4) one or both approach to a face-to-face position 1–3 cm apart, and (5) male mounts female after 1–2 s. Courtship behavior was almost identical to that of laboratoryculture flies observed previously under laboratory conditions. Most malefemale encounters occurred at a height of 1–2m, well inside the outer canopy of the tree. Differential mating success by males occurred. No male mated more than once per day, owing possibly to a very short sexual activity period. Factors favoring mating success of males were survival ability and tendency to join male aggregations and to fight other males. Thorax length and age (9–11 days difference) had no effects on male copulatory success. Overall win/loss percentage was not related to mating success because the males that were most successful at mating fought mostly among themselves, driving their win/loss percentage down. However, these successful males (at mating) won most of their fights against less successful males. Results confirmed a lek mating system: males aggregated, called, and defended territories; territories did not contain femalerequired resources; and females exercised mate choice, apparently through selection of sites within leks.     

Size Dependent Sexual Selection in Drosophila ananassae

Mate choice based on body size is widespread and can have numerous consequences. We present data, which show the effect of male and female body size on sexual selection in Drosophila ananassae. The relationships between wing size, locomotor activity, mating latency, courtship pattern, fertility and mating success were studied. Mating latency was negatively correlated with wing length and with locomotor activity, while wing length and locomotor activity was positively correlated in males as well as in females. In female- and male-choice, we found that mate choice influenced size-assortative mating by: (1) large and small males preferring to mate with large females, (2) large males successfully competing for large females, leaving small males to mate with small females. Males increased their reproductive success by mating with large and more fecund females. In addition, in pairs of long/short winged flies, long winged flies courted and mated more successfully than short winged flies and they also have longer duration of copulation and more progeny than short winged flies. We found sterile mating in pairs of small winged males and females.     

Effects of multiple matings on reproductive fitness of male and female Diaeretiella rapae

Mating frequency and the amount of sperm transferred during mating have important consequences on progeny sex ratio and fitness of haplodiploid insects. Production of female offspring may be limited by the availability of sperm for fertilizing eggs. This study examined multiple mating and its effect on fitness of the cabbage aphid parasitoid Diaeretiella rapae McIntosh (Hymenoptera: Aphidiidae). Female D. rapae mated once, whereas males mated with on average more than three females in a single day. The minimum time lag between two consecutive matings by a male was 3 min, and the maximum number of matings a male achieved in a day was eight. Sperm depletion occurred as a consequence of multiple mating in D. rapae. The number of daughters produced by females that mated with multiple‐mated males was negatively correlated with the number of matings achieved by these males. Similarly, the proportion of female progeny decreased in females that mated with males that had already mated three times. Although the proportion of female progeny resulting from multiple mating decreased, the decrease was quicker when the mating occurred on the same day than when the matings occurred once per day over several days. Mating success of males initially increased after the first mating, but then males became ‘exhausted’ in later matings; their mating success decreased with the number of prior matings. The fertility of females was affected by mating with multiple‐mated males. The study suggests that male mating history affects the fitness of male and female D. rapae.     

Male courtship success and female discrimination in the mediterranean fruit fly (Diptera: Tephritidae)

The mating success of individually marked male Mediterranean fruit flies was monitored over 6 consecutive days in the laboratory. Mating frequency was nonrandom, as the numbers of both males that failed to mate and males that mated many (more than four) times were much higher than expected by chance alone. Differential mating success resulted in part from intermale variation in activity level. Male copulatory success was positively correlated with the numbers of courtships performed, attempted copulations (mountings), and females courted. Male-male aggression, on the other hand, and a negligible effect on male mating success. Female choice also appeared to influence male mating frequency. Fewer than 10% of courtships resulted in mating, and in most cases females terminated courtship by simply moving away from the male. Females that did mate generally selected males having higher copulatory scores than previously rejected males.     

Starvation and the Mating Success of Wild Male Mediterranean Fruit Flies (Diptera: Tephritidae)

Recent evidence suggests that the nutritional state of male Mediterranean fruit flies, Ceratitis capitata (Wied.) (medfly), is an important influence on various components of their reproductive biology, including mating success. The objective of the present study was to examine experimentally the effect of temporary starvation on the mating success of wild male C. capitata. Males were maintained on protein–sugar or sugar-only diets, and for each diet we compared the mating success of continuously fed males versus males starved for 18 or 24 h immediately before testing. In trials conducted on field-caged, host trees, males starved for 24 h obtained only about half as many matings as fed males for both diets. However, when the starvation period was 18 h, starved males reared on the protein–sugar diet mated significantly less frequently than fed males, whereas starved males reared on sugar mated as often as fed males. Measurements of male pheromone calling and female attraction revealed that reduced mating success likely reflected the decreased signaling activity of starved males.     

Reproductive behaviour of the South American fruit fly Anastrepha fraterculus (Diptera: Tephritidae): laboratory and field studies

Abstract. The mate choice, courtship and oviposition behaviour of laboratory-reared and field-collected Anastrepha fraterculus (Wied.) were compared. In laboratory cultures in Southampton the duration of male calling activity in small leks increased gradually from 1-2h at 5 days old to up to 7 h at 10 days. This finding correlates with previous reports on the time at which male salivary glands, which are believed to produce sex pheromone, are fully developed. Wild flies which emerged from infested fruits in Brazil began to oviposit on the day they mated, whereas in laboratory flies oviposition began 1 day following the first mating. Both types of fly usually defended their position on a particular fruit throughout the day, and re-mated with either virgin or mated males. There was no significant difference in mating duration. Females did not copulate before the mean age (±SE) of 16.8±0.9 days. For both types of flies mating initiation occurred in the first 2h of photophase, with virgin females choosing mainly mated males. The average number of matings in the laboratory was three for females and four for males, and the interval between matings in females was significantly increased after the second mating. It is suggested that the tendency of virgin females to mate with mated males will lead to increased fitness, as males are on average 48 days old at their second mating. The potential life span of around 200 days for both sexes would allow adults to bridge the gap between seasonally available fruits in warm-temperate and sub-tropical South America.     

Effects of delayed mating on reproductive output of female oriental beetle Anomala orientalis (Coleoptera: Scarabaeidae)

1 Although management of the oriental beetle Anomala orientalis (Waterhouse) by mating disruption shows promise across a range of agricultural systems, relatively little is known about aspects of the reproductive biology of this species relevant to its management. We studied the effects of delayed mating on several aspects of the oviposition behaviour and biology of the oriental beetle using females mated in the laboratory at 4–13 days posteclosion. 2 Females exhibited a gradual decline in fecundity with increased age at mating that was largely a function of a decline in duration of the oviposition period, as well as lower female fertility: females mated at 11 days were less likely to lay any fertile eggs. However, egg fertility did not vary with female age at mating. 3 Because mating delay did not affect longevity, females mated at older ages experienced decreased oviposition periods; however, females laid more eggs per day with increased age at mating, which partially offset shorter oviposition periods. 4 A mating delay of ≥ 6 days relative to females mated within the first day of reaching sexual maturity resulted in an approximately 35–50% lower mean fecundity. 5 These results suggest that, for mating disruption to be a successful management tool for the oriental beetle, mating must be prevented rather than delayed. We compare the findings of the present study with the published research on delayed mating in moths and discuss the importance of these results in relation to management of the oriental beetle using female sex pheromones.     

Comparison of neonicotinoid insecticides for use with biodegradable and wooden spheres for control of key Rhagoletis species (Diptera: Tephritidae).

Field-based studies and laboratory bioassays were conducted with apple maggot, Rhagoletis pomonella (Walsh), and blueberry maggot, Rhgoletis mendax Curran, flies to investigate the performance and duration of activity of insecticide-treated biodegradable and wooden spheres for control of Rhagoletis species. Four neonicotinoid insecticide treatments including imidacloprid, thiamethoxam, and thiocloprid at 2% (AI) were evaluated with biodegradable spheres. In 1999, significantly more apple maggot flies were found killed by imidacloprid-treated spheres compared with thiamethoxam-treated spheres during early and late season. In 2000, spheres treated with either of two formulations of imidacloprid killed significantly more apple maggot flies compared with thiamethoxam, thiocloprid, and untreated spheres. In blueberries, there were no significant differences between the numbers of blueberry maggot flies killed by both imidacloprid-treated or thiamethoxam-treated spheres in 1999. However, during the 2000 blueberry field season, both formulations of imidacloprid were significantly more effective in killing blueberry maggot flies compared with spheres treated with thiamethoxam, thiocloprid and untreated controls. Overall, spheres treated with thiocloprid were ineffective and did not kill significantly more apple maggot or blueberry maggot flies compared with the controls. Laboratory bioassays showed that the effectiveness of field-exposed spheres treated with imidacloprid at 4 and d 8% (AI) and thiamethoxam at 4% (AI) in killing apple maggot flies was not significantly reduced over a 12-wk aging period. Additionally, wooden spheres aged outdoors for 12 wk with and without mold maintained residual activity in laboratory tests, whereas biodegradable spheres of equal aging, with and without mold lost their effectiveness in killing apple maggot flies. In other studies, we confirmed that the addition of an external feeding stimulant (sucrose) significantly increases the effectiveness of both biodegradable and wooden spheres treated with imidacloprid at 2% (AI).     

Estimating a Key Parameter of Mammalian Mating Systems: The Chance of Siring Success for a Mated Male

Studies of multiple paternity in mammals and other animal species generally report proportion of multiple paternity among litters, mean litter sizes, and mean number of sires per litter. It is shown how these variables can be used to produce an estimate of the probability of reproductive success for a male that has mated with a female. This estimate of male success is more informative about the mating system that alternative measures, like the proportion of litters with multiple paternity or the mean number of sires per litter. The probability of success for a mated male can be measured both theoretically and empirically, and gives an estimate of the intensity of sperm competition and of a male's “confidence of paternity” upon mating. The probability of success for mated males for ten “exemplar” species of mammals is estimated and they are compared for insights into the functioning of their mating systems.     

The Starling Mating System as an Outcome of the Sexual Conflict

Many bird species demonstrate a variable mating system, with some males being monogamously mated and other males able to attract more than one mate. This variation in avian mating systems is often explained in terms of potential costs of sharing breeding partners and compensation for such costs. However, whenever there is a difference in the optimal mating system for males and females, a sexual conflict over the number of partners is expected. This paper contains a verbal model of how a conflict between male and female European starlings (Sturnus vulgaris),resulting from the fitness consequences of different mating systems for males and females differing over time, determines the mating system. We demonstrate that males and females have contrasting fitness interests regarding mating system, such that males gain from attracting additional mates whereas already mated females pay a cost in terms of reduced reproductive success if males are successful in attracting more mates. We demonstrate how this can be traced to the rules by which males allocate non-sharable care between different broods. Furthermore, we demonstrate that there exist male and female conflict behaviours with the potential to affect the mating system. For example, aggression from already mated females towards prospecting females can limit male mating success and males can circumvent this by spacing the nest-sites they defend. The realised mating system will emerge as a consequence of both the fitness value of the different mating systems for males and females, and the costs for males and females of intersexual competition. We discuss how this model can be developed and critically evaluated in the future.     

Female Age Determines Remating Behavior in Wild Mexican Fruit Flies

Male and female age are important factors that can influence mating and remating behavior. Females can discriminate against or prefer older males, but there have been relatively fewer studies on how female and male age influence female remating. Here we showed in wild flies of the Mexican fruit fly Anastrepha ludens (Loew), that when females were given a choice between males of different ages, younger females preferred to mate with younger males over older males, while older females were less selective. Also, when given a choice between males of different ages, older females had longer copulation durations than younger females. On the other hand, older males and females had lower mating success, compared with young and middle-aged flies under no choice conditions. However, middle-aged females mated faster compared to young females and young males mated faster compared to middle-aged males. Male age did not influence female remating, while female age strongly determined female remating, with no females remating when they were old. It is unclear if female receptivity mechanisms are switched off at older ages, or if females are reluctant to remate due to possible costs of mating. We discuss our results in terms of how male and female age can influence mating decisions.     

Reproductive behavior of the honeydew moth,Cryptoblabes gnidiella

Summary

The reproductive behavior of the honeydew moth, Cryptoblabes gnidiella (Millière) (Lepidoptera: Pyralidae), was studied in the laboratory. The sex ratio was 1.1:1, males to females, in both laboratory and field stocks. Most of the females that mated did so during the first night after emergence; males began mating on the following night. Mating occurred 1–2 h before dawn and averaged 100 min. Both sexes mated only once in one night. Most females mated only once in their lifetime, a few mated 2–4 times, whereas males mated up to six times per lifetime. Insects that lived longer also mated more times. When the sex ratio was altered from 3:1 to 1:3, males to females, the percentage of females that mated in one night dropped from 90 to 65, whereas the number of matings per male rose from 0.32 to 2.25. When fresh one-day-old females were provided daily at a ratio of three per male, the males averaged 1.4 matings per lifetime vs. 2.6 with 2- to 3-day-old females. A delay in mating did not affect the percentages of males and females that mated; highest percentages were obtained with 2- to 4-day-old males and females, but a delay in mating resulted in egg fertility dropping from 91 % to 73 %. The preoviposition period lasted a full day after mating, and then most of the eggs were laid during the first night. Average fecundity was 105 eggs per female (maximum: 230).     

The likely fate of hybrids of Bactrocera tryoni and Bactrocera neohumeralis

Bactrocera tryoni (Froggatt) and B. neohumeralis (Hardy) (Diptera: Tephritidae) are sympatric species which hybridise readily in the laboratory yet remain distinct in the field. B. tryoni mates only at dusk and B. neohumeralis mates only during the day, but hybrids can mate at both times. We investigated the inheritance of mating time in successively backcrossed hybrid stocks to establish whether mating with either species is more likely. The progeny of all backcrosses to B. tryoni mated only at dusk. The majority of the progeny of the first and a minority of the progeny of the second backcross to B. neohumeralis also mated at dusk, but the third successive B. neohumeralis backcross produced flies that mated only during the day. This trend towards dominance of the B. tryoni trait was also reflected in a diagnostic morphological character. We discuss the possible genetic background for these phenomena and propose that unidirectional gene flow might explain how the two species remain distinct in the face of natural hybridisation.     

Effects of autodisseminated Metarhizium guizhouense PSUM02 on mating propensity and mating competitiveness of Bactrocera cucurbitae (Diptera: Tephritidae)

We investigated the effects of inoculation by Metarhizium guizhouense PSUM02 on mating propensity and mating competitiveness of Bactrocera cucurbitae, with a view on pest management. On day 4 postinoculation, the M. guizhouense-treated male flies had significantly lowered mating propensity and mating competitiveness, while the treated female flies had reduced mating propensity on day 4 and reduced mating competitiveness on day 5. The mating propensity and competitiveness of treated male and female flies then further declined until death. Kaplan–Meier survival analysis of treated male and female flies gave average survival times (AST) of 6.2 ± 0.2 and 5.4 ± 0.3 days in the mating propensity assay, and about 5.0 ± 0.1 and 4.4 ± 0.2 days in the mating competitiveness assay. The AST of untreated flies ranged from 12.8 ± 0.1 to 14.7 ± 0.2 days for comparison (observation up to 15 days). Untreated flies had decreased AST and mating characteristics when exposed to contact with treated male flies, indicating transmission of the fungal infection by such contact also to untreated male flies. Surprisingly, contact with treated female flies did not affect the AST of untreated males or females in the same cage. These results corroborate the potential for pest control by autodissemination with treated male flies, which transmit the fungus to a healthy population better than the treated female flies.     

Paternity costs from polyandry compensated by increased fecundity in the hide beetle

Polyandry-induced sperm competition is assumed to impose costson males through reduced per capita paternity success. In contrast,studies focusing on the consequences of polyandry for femalesreport increased oviposition rates and fertility. For thesespecies, there is potential for the increased female fecundityassociated with polyandry to offset the costs to males of sharedpaternity. We tested this hypothesis by comparing the proportionand number of offspring sired by males mated with monandrousand polyandrous females in the hide beetle, Dermestes maculates,both for males mating with different females and for males rematingwith the same female. In 4 mating treatments, monandrous femalesmated either once or twice with the same male and polyandrousfemales mated either twice with 2 different males or thricewith 2 males (where 1 male mated twice). Polyandrous and twice-matingmonandrous females displayed greater fecundity and fertilitythan singly mating monandrous females. Moreover, males rematedto the same female had greater paternity regardless of whetherthat female mated with another male. In both polyandrous treatments,male mating order did not affect paternity success. Finally,although the proportion of eggs sired decreased if a male matedwith a polyandrous female, multiply mating females or femalesthat remated with a previous mate laid significantly more eggsand thus the actual number of eggs sired was comparable. Thus,males do not necessarily accrue a net fitness loss when matingwith polyandrous females. This may explain the absence of anyobvious defensive paternity-protection traits in hide beetlesand other species.     

A novel mating system in a solitary carnivore: the fossa

The mating strategies of male mammals have long been treated as broadly predictable on the basis of just two factors: the dispersion of females and the benefit of paternal care to male reproductive success. Female strategies and finer scale variations in mating systems remain poorly understood. In the fossa Cryptoprocta ferox , we had the rare opportunity to examine the mating system of a wild solitary carnivore directly, and identified features not classified or predicted by mating system theory. Males competed for mating opportunities at a traditional site monopolized by a female, high in a tree. The female mated with multiple males, repeatedly mated with some individuals and appeared to express mate choice. We observed three females thus, one replacing another on the site after each was seen to mate with four to five males over a period of 1–6 days. Copulations were prolonged (up to 3 h 8 min), involving a weak copulatory tie, and males appeared to guard females briefly after mating. Fossas are at low population density and do not use a den regularly; we suggest that both these factors impede individuals from locating a mate. We hypothesize that the observed mating system reduces this problem for both sexes, and increases the number of mates available to a female while ensuring a low risk of sexual harassment.     

Effects of pyriproxyfen on ovarian development and fecundity ofRhagoletis pomonella flies

Three-day old female apple maggot flies,Rhagoletis pomonella (Walsh), were topically exposed to different doses (0.1, 1, 10, and 100 μg per fly) of a juvenile hormone mimic, pyriproxyfen, in the laboratory. Pyriproxyfen had little lethal effect on females except at the extremely high dose of 100 μg. It also had no significant effect on egg viability of treated females. A non-lethal dose of 1 μg per fly did, however, enhance significantly the fecundity (egg production) as well as the ovarian development (number of eggs in ovaries and length of egg folicles) of treated flies. We conclude that pyriproxyfen could be a useful aid in exploring endocrine regulation of feeding and reproductive physiology behavior inR. pomonella, about which current knowledge is scant.     

Courtship song, male agonistic encounters, and female mate choice in the house cricket,Acheta domesticus (Orthoptera: Gryllidae)

Whether female crickets choose among males based on characteristics of the courtship song is uncertain, but in many species, males not producing courtship song do not mate. In the house cricket,Acheta domesticus, we examined whether a female chose or rejected a male based on his size, latency to chirp, latency to produce courtship song, or rate of the high-frequency pulse of courtship song (“court rate”). We confirmed that females mated only with males that produced courtship song, but we found no evidence that the other factors we measured affected a female’s decision to mate. In addition, we investigated whether the outcome of male agonistic encounters affected the subsequent production of courtship song. In one experiment, we observed courtship and mating behavior when a single female was placed with a pair of males following a 10-min interaction period between the two males. Winners of male agonistic encounters had higher mating success. However, winners and losers of agonistic encounters were not different in their likelihood or latency to produce courtship song or in the number of times they were disrupted by the other male in the pair. In a second experiment, we allowed two males to interact for a 10-min period, but following this interaction period, we placed a female with each male separately and observed courtship and mating behavior. The mating success of winners and losers was not different under these circumstances, and we found no differences between winners and losers in any subsequent courtship or mating behavior examined. We conclude that winning agonistic encounters influences a male’s mating success in ways other than his production of courtship song and this effect is lost when winning and losing males are separated and each is given an opportunity to mate.     

Extreme Promiscuity in a Mating System Dominated by Sexual Conflict

Coelopids live in wrack beds consisting of seaweed washed up on beaches. Their mating system is characterized by sexual conflict and convenience polyandry, with females resisting male mating attempts. We estimated the level of harassment by males and the success rate of rejection by females collected from a high density wild population. Males mounted a female every 8.41 min. Of these mounts 35% resulted in copulation. This suggests that females could be mated up to 5 times every 2 h. Females typically live for 3 weeks, and thus, could mate with hundreds of males during their lifetime. We found a 50:50 sex ratio throughout the wrack bed revealing that females do not avoid male harassment by leaving the wrack bed when not ovipositing.