Male Age Affects Female Mate Preference and Reproductive Performance in the Cabbage Beetle, <Emphasis Type="Italic">Colaphellus bowringi</Emphasis>

The influence of male age on female mate preference and reproductive performance in the cabbage beetle, Colaphellus bowringi was examined, using male and female adults of varying ages (young, middle-aged and old) after a single mating. In a simultaneous choice test, females of all age class preferentially mated with middle-aged males. Mating duration was positively related to male age. Longevity of females was not significantly affected by male age. Young females paired to middle-aged males had significantly higher egg production than those paired to old males, and the eggs of females paired to middle-aged males exhibited significantly higher hatching success than the eggs of females mated to young or old males. These results suggest that middle-aged males are more advantageous for female fitness than young and old males.     

Age‐Related Male Reproductive Investment in Courtship Display and Nuptial Gifts in a Moth,Ostrinia scapulalis

Due to a trade‐off between current and future reproduction, costly reproductive investments should be increased towards the end of a lifespan when the probability of reproduction becomes low (terminal investment hypothesis). We investigated age‐related changes in male reproductive investment towards courtship display and the spermatophore in three age classes (young, middle‐aged and old) of a monandrous moth, Ostrinia scapulalis. As predicted, old males had higher mating success than young and middle‐aged males in no‐choice tests. Moreover, two‐choice tests revealed that middle‐aged males had a higher success rate than young males because of their higher courtship frequency rather than any female preference for them. It was found that old males produced a larger spermatophore than young and middle‐aged males, suggesting greater reproductive effort. The protein content of spermatophores also tended to increase with male age. Despite the age‐related variation in spermatophore size and protein content, age did not affect female fecundity or longevity. A decrease in the number of sperm in the older males might counteract the nutritional benefit of larger spermatophores. Alternatively, fitness components other than longevity and fecundity may be influenced by male age.     

The role of male age, sperm age and mating history on fecundity and fertilization success in the hide beetle

Models of age-related mate choice predict female preference for older males as they have proven survival ability. However, these models rarely address differences in sperm age and male mating history when evaluating the potential benefits to females from older partners. We used a novel experimental design to assess simultaneously the relative importance of these three parameters in the hide beetle, Dermestes maculatus. In a two-part experiment we first explored age-related male mating success and subsequently examined the consequences of male age, sperm age and male mating history on female fecundity and fertilization success. In a competitive mating environment, intermediate-age males gained significantly higher mating success than younger or older males. To test the consequences for females of aged-related male mating success, a second set of females were mated to males varying in age (young, intermediate-age and old), in numbers of matings and in timing of the most recent mating. We found that male age had a significant impact on female fecundity and fertilization success. Females mated to intermediate-age males laid more eggs and attained consistently higher levels of fertilization success than females with young and old mates. A male's previous mating history determined his current reproductive effort; virgin males spent longer in copula than males with prior mating opportunities. However, differences in copulation duration did not translate into increased fecundity or fertilization success. There was also little evidence to suggest that fertilization success was dependent on the age of a male's sperm. The experiment highlights the potential direct benefits accrued by females through mating with particular aged males. Such benefits are largely ignored by traditional viability models of age-related male mating success.     

Mating senescence and male reproductive organ size in the Mexican fruit fly

Ageing can reduce the probability that individuals reproduce. The present study investigates whether ageing influences the mating frequency of mass‐reared fertile and sterile Mexican fruit flies Anastrepha ludens (Loew). The ability of males of different ages to inhibit female remating is also determined, and the growth of male reproductive organs is measured as they age. Young males (6 days old) have a lower mating frequency than older males, and also have a lower capacity to inhibit female remating than older males. However, 7‐day‐old males are as likely to inhibit female remating as older males. Young males also have smaller reproductive organs than middle‐aged (21‐day‐old) or senescent males (57‐day‐old). These results have implications for the sterile insect technique because sterilized males of A. ludens are released in the field 6 days after emergence. The highest mating frequency, the lowest mating latency and the largest size of testes are observed at 21 days of age. Older males (57 days old) have more sperm in their seminal vesicles than young males (6 and 9 days old). Accessory glands take longer to grow to their complete size compared with testes, and mating frequency is more closely associated with accessory gland size than testes size. Furthermore, there are more sperm in the seminal vesicles during the afternoon period of peak sexual activity than during the morning when sexual activity is absent. These results indicate that, even at the onset of reproductive senescence, mass‐reared males of A. ludens are still capable of mating, as well as inhibiting remating in females.     

Male age affects female mating preference but not fitness in the monandrous moth Dendrolimus punctatus Walker (Lepidoptera: Lasiocampidae)

It is widely accepted that male age can influence female mating preference and subsequent fitness consequences in many polyandrous species, yet this is seldom investigated in monandrous species. In the present study, we use the monandrous pine moth Dendrolimus punctatus to examine the effects of male age on female mating preference and future reproductive potential. In multiple male trials, when permitted free mating from an aggregation consisting of virgin males aged 0 (young), 2 (middle-aged) and 4 (old) days, virgin females preferentially mate with young and middle-aged males, although mating latency and mating duration are independent of male age. In single male trials, when virgin females are randomly assigned single virgin males of known age, a negative correlation is found between mating success and male age in this species. However, we find that male age also has no effect on mating latency and mating duration. Further fitness analysis reveals that females do not receive benefits in terms of oviposition period, total egg production, average daily egg production, percentage of egg hatching, longevity, expected reproduction and relative expected reproduction from mating with young and middle-aged males compared with mating with old males. The results of the present study are the first demonstrate that females mated preferentially with younger males but gain no apparent fitness benefits in a monandrous moth species.     

Pheromone Blend Does not Explain Old Male Mating Advantage in a Butterfly

In several insect species, male mating success is higher in older than in younger males, although condition diminishes dramatically with age. Two hypotheses are under debate to explain the counterintuitive pattern of old male mating advantage: first, an increased eagerness of older males to mate, driven by their low residual reproductive value, and second female preference for older males based on chemical cues such as sex pheromones (female choice hypothesis). In a series of experiments, we manipulated female olfaction, male pheromone blend and female age to test whether old male mating advantage prevails when the influence of male sex pheromones is controlled for, using the tropical butterfly Bicyclus anynana as model. We found that older males had a higher mating success than younger ones irrespective of female scent‐sensitivity and irrespective of male pheromone blend. Interestingly, older males were found to court more often and for longer time bouts than younger males. These results were independent of female age, although younger males courted younger females more often and for longer bouts than older females. Taken together, our results indicate that male courtship activity (1) is higher in older compared to younger males and (2) increases the mating success of older males. Olfaction and sensing pheromones, in contrast, were not a necessary prerequisite for old male mating advantage to occur and may use other cues than pheromones to assess male quality.     

The effect of mating frequency and mating pattern on female reproductive fitness in cabbage beetle,Colaphellus bowringi

It is generally thought that females can receive more of the material benefits from males by increasing mating frequency and polyandry can lead to greater reproductive success. The cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), is a highly promiscuous species, in which females or males can readily mate repeatedly with a given partner or multiple partners at a very high frequency. In the present study, the effect of mating frequency (number of matings) and mating pattern (polyandry vs. monogamy) on female reproductive fitness was investigated by measuring fecundity, fertility, and female longevity. The results indicated that increased female mating frequency with the same male did not result in variation in lifetime fecundity, but significantly increased fertility and decreased female longevity. Moreover, five copulations were sufficient to acquire maximal reproductive potential. Female lifetime fecundity also did not differ between polyandrous and monogamous treatments. However, monogamous females exhibited a significant increase in fertility and significant prolongation of longevity compared with polyandrous females, further demonstrating that monogamy is superior to polyandry in this beetle.     

Young Bighorn (Ovis canadensis) Males: can they Successfully Woo Females?

Mating by young males or low male‐to‐female ratios can decrease pregnancy rates and postpone birthdates in ungulates, thereby hindering population growth. Young (2.5–3.5 yr old) male bighorn (Ovis canadensis) behave differently than older males, and age, horn size, mating behavior, and social rank help determine reproductive success. We estimated birthdates in two populations of bighorn sheep in Utah, USA, to determine if mating by young males or low male‐to‐female ratios resulted in fewer young born per female, a shift in mean timing of births, or asynchronous births. When reintroduced, the Rock Canyon population consisted of four males (two each of 2.5 yr old and 1.5 yr old) and a 1 to 7.5 ratio of males (>2 yr old) to adult females (≥3.5 yr old); the Mount Nebo population consisted of four males ≤1.5 yr old and a 0 to 12 ratio of males to adult females. For both populations, the number of young born per female did not differ between the first parturition period after reintroduction (where females were impregnated by males from their source populations) and the second period of parturition (where females were impregnated by young, reintroduced males). Mean birthdates and synchrony (SD) of births did not differ for Rock Canyon (May 12, 2001 ± 4.5 d, May 14, 2002 ± 3.2 d) or Mount Nebo (May 23, 2005 ± 8.1 d, May 22, 2006 ± 10.2 d) between the first and second years following reintroduction. Mating by young males or low male‐to‐female ratios had no demonstrable effect on the number of young born per female or timing and synchrony of births in these populations.     

Age,Experience and Sex – Do Female Bulb Mites Prefer Young Mating Partners?

In species where advancing sire age is associated with decreased progeny fitness, female resistance to mating with old partners can be expected to evolve. In polyandrous species, such resistance may be contingent on female mating experience: virgins should be relatively indiscriminate to ensure egg fertility, whereas non‐virgins can be expected to base their re‐mating decisions on the age of their previous versus potential new partners, and ‘trade‐up’ if previously mated with old males. Here, we tested these predictions using a promiscuous and relatively long‐living bulb mite (Rhizoglyphus robini), in which old sire age is associated with decreased fecundity of daughters. In a fully factorial design, we applied two male treatments, young and old, and three female treatments, virgin, previously mated to an old male and previously mated to a young male. Consistent with earlier studies, we observed a reduced mating success of old males. However, we found no support for attributing this result to female discrimination, as female behavior in response to male mounting attempts was not affected by the age of the suitor, or by its interaction with the age of the female’s previous mate. Interestingly, females were passive during 93% of male mounting attempts observed, suggesting that once they are located by a male, they exert little control over copulation. Old males had lower mate‐searching activity and were less efficient in obtaining matings (lower success rate per mounting attempt), suggesting a decreased mate‐securing ability because of aging. Overall, our results suggest that in bulb mites, male ability to secure mates declines with age, whereas they do not support the prediction that females actively discriminate against old partners.     

Effect of age on reproductive attributes of an aphidophagous ladybird, Cheilomenes sexmaculata

The effect of both male and female age was investigated on certain reproductive attributes, viz. mating incidence, mating duration, fecundity, percent egg viability, ratio of reproductive and non‐reproductive periods and reproductive rate, of an aphidophagous ladybird, Cheilomenes sexmaculata (Fabricius). Females started mating at the age of 8 hours post‐emergence (PE) and males at the age of 2 days PE. Mating in the laboratory was a male‐dominated phenomenon. The mating duration and reproductive rate of 10‐day‐old females when mated with males of varying ages increased up to the male age of 60 days, and thereafter decreased, whereas, fecundity, egg viability and ratio of reproductive and non‐reproductive periods increased up to the male age of 50 days, and thereafter declined. However, when females of varying ages were mated with 10‐day‐old males, fecundity and reproductive rate increased up to 40 days of female age, respectively, then decreased. The ratio of reproductive and non‐reproductive periods increased with increasing age of females. Mating age for optimal reproductive output was 10J50‐day‐old males and NE to 40‐day‐old females. Reproductive cessation in males was recorded after 50 days PE, whereas in females at the age of 40 days PE. Higher mating durations lead to elevated reproductive rates. Delay in the reproductive phase was positively correlated with longevity. The results of this study may aid mass multiplication of this ladybird by identifying and promoting usage of adults of optimal age. Our results also enhance our understanding of the effect of age on reproductive attributes in ladybirds.     

Effect of male and female age on the mating success of the obliquebanded leafrollerChoristoneura rosaceana (Lepidoptera: Tortricidae) under different ecological conditions

The present study was undertaken to test the hypothesis that male and female age, as well as temperature, can affect the mating success ofChoristoneura rosaceana, given the role that these two factors play in the pheromone biology of this species. In the laboratory, the mating success of females generally declined linearly with age, whereas in males, it increased during the first 3 days and then decreased. The decline in female mating success was more pronounced under warm than cool thermocycles, while the changes observed in males were greater under fluctuating than constant temperature regimes. The onset time of mating was unaffected by male age, however, older females always mated earlier than younger ones, with the advance being more pronounced at cool than warm temperatures. Similar results were also obtained under field conditions. When a single 3-day-old male was provided with 0-, 3-, and 5-day-old females simultaneously, older females obtained mates significantly more often than younger individuals at all temperatures, indicating that calling earlier may afford a reproductive advantage to older females. At both constant temperatures, the time spent mating was longer in older than in younger females, but not under fluctuating thermocycles. Very young and very old males generally spent more time in copula than middle-aged individuals both at constant temperatures and under the warm thermocycle. Under the cool thermocycle, the duration of mating was considerably prolonged at all ages, which could increase the risk of predation.     

Comparison of sexual compatibility in crosses between the southern and northern populations of the cabbage beetle Colaphellus bowringi

It is widely accepted that the genetic divergence and reproductive incompat- ibility between closely related species and/or populations is often viewed as an important step toward speciation. In this study, sexual compatibility in crosses between the southern XS population and the northern TA population of the polyandrous cabbage beetle Co- laphellus bowringi was investigated by testing their mating preferences, mating latency, copulation duration, and reproductive performances of post-mating. In choice mating ex- periments, the percentages ofmatings were significantly higher in intra-population crosses than in inter-population crosses. Both isolation index （/） and index of pair sexual isolation （/PSi） indicated partial mating incompatibility or assortative mating in crosses between the two different geographical populations. In single pair mating experiments, XS females in inter-population crosses mated significantly later and copulated significantly shorter than those in intra-population crosses. However, TA females in inter-population crosses mated significantly earlier and copulated longer than those in intra-population crosses, suggesting that larger XS males may enhance heterotypic mating. The lifetime fecundity was highest in XS homotypic matings, lowest in TA homotypic matings, and intermedi- ate in heterotypic rnatings between their parents. The inter-population crosses resulted in significantly lower egg hatching rate and shorter female longevity than intra-population crosses. These results demonstrated that there exist some incompatibilities in premating, postmating-prezygotic, and postzygotic stages between the southern XS population and northern TA population of the cabbage beetle Colaphellus bowringi.     

Methoprene application and diet protein supplementation to male melon fly,Bactrocera cucurbitae,modifies female remating behavior

Methoprene (an analogue of juvenile hormone) application and feeding on a protein diet is known to enhance male melon fly, Bactrocera cucurbitae Coquillett (Diptera: Tephritidae), mating success. In this study, we investigated the effect of these treatments on male B. cucurbitae's ability to inhibit female remating. While 14‐d‐old females were fed on protein diet, 6‐d‐old males were exposed to one of the following treatments: (i) topical application of methoprene and fed on a protein diet; (ii) no methoprene but fed on a protein diet; (iii) methoprene and sugar‐fed only; and (iv) sugar‐fed, 14‐d‐old males acted as controls. Treatments had no effect on a male's ability to depress the female remating receptivity in comparison to the control. Females mated with protein‐deprived males showed higher remating receptivity than females first mated with protein‐fed males. Methoprene and protein diet interaction had a positive effect on male mating success during the first and second mating of females. Significantly more females first mated with sugar‐fed males remated with protein‐fed males and females first mated with methoprene treated and protein‐fed males were more likely to remate with similarly treated males. Females mating latency (time to start mating) was significantly shorter with protein‐fed males, and mating duration was significantly longer with protein‐fed males compared with protein‐deprived males. These results are discussed in the context of methoprene and/or dietary protein as prerelease treatment of sterile males in area‐wide control of melon fly integrating the sterile insect technique (SIT).     

Old‐male paternity advantage is a function of accumulating sperm and last‐male precedence in a butterfly

Old‐male mating advantage has been convincingly demonstrated in Bicyclus anynana butterflies. This intriguing pattern may be explained by two alternative hypotheses: (i) an increased aggressiveness and persistence of older males during courtship, being caused by the older males' low residual reproductive value; and (ii) an active preference of females towards older males what reflects a good genes hypothesis. Against this background, we here investigate postcopulatory sexual selection by double‐mating Bicyclus anynana females to older and younger males, thus allowing for sperm competition and cryptic mate choice, and by genotyping the resulting offspring. Virgin females were mated with a younger virgin (2–3 days old) and afterwards an older virgin male (12–13 days old) or vice versa. Older males had a higher paternity success than younger ones, but only when being the second (=last) mating partner, while paternity success was equal among older and younger males when older males were the first mating partner. Older males produced larger spermatophores with much higher numbers of fertile sperm than younger males. Thus, we found no evidence for cryptic female mate choice. Rather, the findings reported here seem to result from a combination of last‐male precedence and the number of sperm transferred upon mating, both increasing paternity success.     

Chemosensory discrimination of male age by female Psammodromus algirus lizards based on femoral secretions and feces

Chemical communication plays an essential role in several social and reproductive behaviors of many animals. In lizards, the main sources of semiochemicals are femoral or pre‐anal gland secretions and feces. In male lizards Psammodromus algirus, there are age‐related differences in the chemical composition of femoral gland secretions and in the reproductive strategies, with older males defending territories and females, while younger males adopting sneak‐mating strategies. Females flee more often from mating advances of young males than from those of old males, which are more successful in obtaining matings. This suggests that age discrimination of males may be important for females. We tested here whether females showed differential chemosensory responses to chemical cues (femoral gland secretion and feces) of males of two age classes, and whether females use information from substrate scent marks of males of different ages to select where to stay. We found that females elicited more tongue‐flicks to the secretion and feces of old males than to control or secretion and feces of young males. Also, the time spent by females on a scented paper depended on the treatment, suggesting that females tended to spend more time on scent marks made with femoral secretions of old males. Adult females seemed capable to discriminate between young and old males based on chemical cues alone and showed more interest in scents of old males. However, substrate scent marks did not seem to entirely determine site selection by females, suggesting that females might need additional cues to perform the choice. These results can be explained by the different age‐dependent reproductive strategies of males, which can affect differentially to females.     

Male age and strain affect ejaculate quality in the Mexican fruit fly

Aging in all organisms is inevitable. Male age can have profound effects on mating success and female reproduction, yet relatively little is known on the effects of male age on different components of the ejaculate. Furthermore, in mass‐reared insects used for the Sterile Insect Technique, there are often behavioral differences between mass‐reared and wild males, while differences in the ejaculate have been less studied. The ejaculate in insects is composed mainly of sperm and accessory gland proteins. Here, we studied how male age and strain affected (i) protein quantity of testes and accessory glands, (ii) the biological activity of accessory gland products injected into females, (iii) sperm viability, and (iv) sperm quantity stored by females in wild and mass‐reared Anastrepha ludens (Diptera: Tephritidae). We found lower protein content in testes of old wild males and lower sperm viability in females mated with old wild males. Females stored more sperm when mated to young wild males than with young mass‐reared males. Accessory gland injections of old or young males did not inhibit female remating. Knowledge of how male age affects different ejaculate components will aid our understanding on investment of the ejaculate and possible postcopulatory consequences on female behavior.     

Mating Group Size and Stability in Reindeer Rangifer tarandus: The Effects of Male Characteristics,Sex Ratio and Male Age Structure

In polygynous mating systems, males compete intensely for mates and may mate several females during a single reproductive season. Accordingly, factors influencing the ability of males to control a larger number of females during the breeding season can provide information on the processes underlying sexual selection. In ungulates, age, body mass and social rank are considered good predictors of the reproductive success of males, but how male age structure and sex ratio in the population influence mating group (MG) dynamics has received little empirical testing. Between 1996 and 2005, we manipulated male age‐ and sex structure and monitored MG dynamics in a reindeer (Rangifer tarandus) population. We investigated the influence of male characteristics, percentage of males and male age structure on MG size and stability. We found that males with higher social rank (that were also older and heavier) controlled larger MGs (therefore had greater mating opportunities) and had more stable MGs (corresponding to a higher ability to maintain and control females) than males of lower social rank. Moreover, MG size and MG stability decreased as the percentage of males in the population increased, most likely resulting from greater male–male competition and increased female movements. Male age structure did not influence MG stability. Given the positive relationship between mating success and MG size (and likely MG stability), frequent female movements and intense competition among males to control females seem to be the principal components of reindeer MGs dynamic.     

Adaptive female choice for middle-aged mates in a lekking sandfly

Most theoretical models of age-related mate choice predict that females should prefer older males because they have proven survival ability. An alternative view is that older males represent inferior mates because of negative genetic correlations between early and late fitness components, or because older males have traded off longevity against other fitness components, have accumulated deleterious germ-line mutations, or are less well adapted to current conditions than more recently born individuals. While numerous studies have reported female choice for older males, few have explicitly examined the fitness consequences of such a preference. We present evidence from a lekking sandfly, Lutzomyia longipalpis, showing that choosy females discriminate against older males and gain a fitness benefit from their choice. When permitted free choice from an aggregation consisting of males aged zero to two days (young), four to six days (middle-aged) and eight to ten days (old), females preferentially mated with middle-aged males, but all measures of female reproductive success were independent of male age. In contrast, when a second set of females was randomly assigned single virgin males of known age, the eggs of those paired to old mates exhibited lower hatching success than the eggs of females mated to young or middle-aged males. These results suggest that females avoid mating with older males because they represent poorer quality mates. Age-related differences in male quality may have a genetic basis, but could equally well arise through a phenotypic decline in sperm quality or sperm transfer ability with male age. The lack of evidence of female discrimination against older males from other studies may be because these did not explore the reproductive success of the full age range of males.     

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.     

Age-related sperm transfer and sperm competitive ability in the male hide beetle

The influence of male age on reproductive success after a singlemating has been explored widely; however, few studies have investigatedwhether quantitative or qualitative differences in male spermare responsible for the observed patterns. Moreover, the roleof male age on sperm competitive ability has been largely ignored.We examined the importance of male age on the probability andamount of sperm transferred during a single mating and exploredwhether sperm competitive ability varies with male age in thehide beetle Dermestes maculatus, a species where sperm viabilitydoes not vary with male age. We also investigated whether spermtransfer rates varied with female age. We found that the probabilityof sperm transfer and the amount of sperm transferred variedwith male, but not female, age. All males performed behaviorallysuccessful copulations, but intermediate-age and old males weremore likely to transfer sperm successfully and also transferreda greater quantity of sperm than young males. Old males wereless likely to transfer sperm than intermediate-age males, butif they did transfer sperm successfully, they transferred comparableamounts. Sperm competitive ability varied with male age andreflected the quantity of sperm transferred. On average, intermediate-agemales achieved greater fertilization success when competingagainst young or old males than when competing against otherintermediate-age males. Old males were poor competitors againstintermediate-age males, but they achieved significantly higherrates of fertilization when competing against young males. Ourfindings suggest that quantitative differences in the amountof sperm transferred determine male success in sperm competitionin the hide beetle.