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.     

The effect of male mating history on paternal investment, fecundity and female remating in the seed beetle Callosobruchus maculatus

1. In many organisms, males provide nutrients to females via ejaculates that can influence female fecundity, longevity and mating behaviour. The effect of male mating history on male ejaculate size, female fecundity, female longevity and female remating behaviour in the seed beetle Callosobruchus maculatus was determined.
2. The quantity of ejaculate passed to females declined dramatically with successive matings. Despite the decline, a male's ability to fertilize a female fully did not appear to decline substantially until his fourth mating.
3. When females multiply mated with males of a particular mated status, the pattern of egg production was cyclic, with egg production increasing after mating. Females multiply mated to virgins had higher fecundity than females mated to non-virgins, and females mated to twice-mated males had disproportionately increased egg production late in their life.
4. Females that mated to multiple virgins, and consequently laid more eggs, experienced greater mortality than females mated only once or mated to non-virgins, suggesting that egg production is costly, and rather than ameliorating these costs, male ejaculates may increase them by allowing or stimulating females to lay more eggs.
5. Females mating with non-virgin males remated more readily than did females mated to virgins. Females given food supplements were less likely to remate than females that were nutritionally stressed, suggesting that females remate in part to obtain additional nutrients.     

Inbreeding depresses sperm competitiveness,but not fertilization or mating success in male Tribolium castaneum

As populations decline to levels where reproduction among close genetic relatives becomes more probable, subsequent increases in homozygous recessive deleterious expression and/or loss of heterozygote advantage can lead to inbreeding depression. Here, we measure how inbreeding across replicate lines of the flour beetle Tribolium castaneum impacts on male reproductive fitness in the absence or presence of male–male competition. Effects on male evolution from mating pattern were removed by enforcing monogamous mating throughout. After inbreeding across eight generations, we found that male fertility in the absence of competition was unaffected. However, we found significant inbreeding depression of sperm competitiveness: non-inbred males won 57 per cent of fertilizations in competition, while inbred equivalents only sired 42 per cent. We also found that the P₂ ‘offence’ role in sperm competition was significantly more depressed under inbreeding than sperm ‘defence’ (P₁). Mating behaviour did not explain these differences, and there was no difference in the viability of offspring sired by inbred or non-inbred males. Sperm length variation was significantly greater in the ejaculates of inbred males. Our results show that male ability to achieve normal fertilization success was not depressed under strong inbreeding, but that inbreeding depression in these traits occurred when conditions of sperm competition were generated.     

Spermatophore displacement and male fertilization success in the ground beetle Carabus insulicola

Sperm displacement is an effective male mating tactic of spermcompetition to ensure paternity. However, the effectivenessof sperm displacement may depend on defensive mating tacticsemployed by rival males and also may be affected by females.To test this hypothesis, I conducted double-mating experimentswith microsatellite paternity analyses using the ground beetleCarabus insulicola. First, I examined how often spermatophoredisplacement by the second male occurs. Spermatophore depositionsuccess of the second male was high (85%) as was that of thefirst male (100%), in which 80% of the second males displacedthe first male's spermatophore, suggesting that spermatophoredisplacement is a highly effective offensive tactic of spermcompetition in C. insulicola. Second, I examined the effectsof postcopulatory guarding by the first male and remating intervaland sperm utilization pattern of the female on fertilizationsuccess of the second male. P₂ values were moderate on average(0.30–0.57) and negatively correlated with postcopulatoryguarding duration of the first male. Sperm use by the femalewas temporally nonrandom, such that sperm of the second maletended to be used first, although there was no significant effectof female-remating interval. These results suggest that theeffectiveness of spermatophore displacement depends on postcopulatoryguarding by the first male and the sperm utilization patternof the female. These findings provide empirical evidence forthe conflicting nature of selection for male offensive and defensivetactics in sperm competition and significant contribution offemales therein.     

Sperm competition, alternative mating tactics and context-dependent fertilization success in the burying beetle, Nicrophorus vespilloides

Fertilization success in sperm competition is often determined by laboratory estimates of the proportion of offspring sired by the first (P1) or second (P2) male that mates. However, inferences from such data about how sexual selection acts on male traits in nature may be misleading if fertilization success depends on the biological context in which it is measured. We used the sterile male technique to examine the paternity of the same male in two mating contexts in the burying beetle, Nicrophorus vespilloides, a species where males have alternative mating strategies based on the presence or absence of resources. We found no congruence in the paternity achieved by a given male when mating under different social conditions. P2 estimates were extremely variable under both conditions. Body size was unrelated to success in sperm competition away from a carcass but, most probably through pre-copulatory male-male competition, influenced fertilization success on a carcass. The contribution of sperm competition is therefore dependent on the conditions under which it is measured. We discuss our findings in relation to sperm competition theory and highlight the need to consider biological context in order to link copulation and fertilization success for competing males.     

Influence of body and genital morphology on relative male fertilization success in oriental beetle

Although the frequently large variance in relative male fertilizationsuccess when females are mated by more than 1 male has beenappreciated for some time, the factors that influence relativepaternity are still poorly understood. Recently, experimentalevidence that morphology of male genitalia influences fertilizationsuccess has been documented in 2 water striders, a dung beetle,and a leaf beetle. We explored the role of male genital morphologyin postcopulatory sexual selection in the oriental beetle. Wemated females to 2 males in succession and assessed relativepaternity by the sterile male technique. Morphology of the malegenitalia was found to strongly influence relative paternitybut only for the first male to mate. Male body size influencedrelative fertilization success as well, but again, only forthe first male; surprisingly, smaller males achieved higherpaternity when mating first. We also found suggestive evidencethat copula duration of both the first and second male to mateinfluenced paternity. Other factors, including female size anddegree of asymmetry of hind tibiae length of males had no effecton relative fertilization success. Our results for the orientalbeetle are novel among sperm precedence studies for 2 reasons:1) traits of the first male appear to be more important in influencingpaternity than those of the second, and 2) smaller, not larger,males achieved greater relative paternity. Our results alsocontribute to the growing body of empirical evidence in supportof the hypothesis that male genitalia evolve by postcopulatorysexual selection.     

Effects of tapeworm infection on male reproductive success and mating vigor in the red flour beetle,Tribolium castaneum

Parasites may exert negative effects on host survivorship and reproductive success. The effects of parasites on female host fitness have been well documented; however, the effects of parasites on the reproductive success of male hosts and particularly the underlying mechanisms that alter male fitness are not well understood. Previous studies demonstrated that infection by rat tapeworm (Hymenolepis diminuta) reduced the fitness of male red flour beetles (Tribolium castaneum) in an environment of female mate choice and strong male-male competition. The present study determined the role of female mate choice and male insemination capacity on observed fitness reduction of male beetles by the tapeworm parasites. We found that infected males showed reduced mating vigor and consequently inseminated fewer females than did uninfected males. Specifically, tapeworm infection reduced the number of offspring sired by a male by 14-22% even when male-male competition and female mate choice were absent. Further, the insemination capacity of males diminished by 30% because of infection. Female beetles did not discriminate against infected males in precopulatory mate choice experiments. Copulatory courtship, a determinant of postcopulatory female choice, was not significantly different between infected and uninfected males. Hence, we concluded that female beetles did not show either pre- or postcopulatory choice against tapeworm-infected males. Therefore, tapeworm-induced reduction in the reproductive success of male beetles possibly results from altered reproductive biology, such as lower mating vigor and decreased sperm quantity or quality.     

Effect of male age on sperm traits and sperm competition success in the guppy (Poecilia reticulata)

Deleterious mutations can accumulate in the germline with age, decreasing the genetic quality of sperm and imposing a cost on female fitness. If these mutations also affect sperm competition ability or sperm production, then females will benefit from polyandry as it incites sperm competition and, consequently, minimizes the mutational load in the offspring. We tested this hypothesis in the guppy (Poecilia reticulata), a species characterized by polyandry and intense sperm competition, by investigating whether age affects post‐copulatory male traits and sperm competition success. Females did not discriminate between old and young males in a mate choice experiment. While old males produced longer and slower sperm with larger reserves of strippable sperm, compared to young males, artificial insemination did not reveal any effect of age on sperm competition success. Altogether, these results do not support the hypothesis that polyandry evolved in response to costs associated with mating with old males in the guppy.     

Effects of age and repeated mating on male sperm supply and paternity in a parasitoid wasp

Post-copulatory paternity biases after female multiple mating are major constraints on both male and female reproductive systems. The outcome of paternity in certain situations is only controlled directly by male sperm stock. This was tested experimentally in the parasitoid wasp Anisopteromalus calandrae (Howard) (Hymenoptera: Pteromalidae), in which sperm stocks are small (several hundred) and the fertilizing efficiency of stored sperm is high (the ratio of sperm stored/fertilized eggs is about 0.75). Sperm in seminal vesicles and paternity of males of different status (virgin young, virgin old, or young previously mated) were measured after female single and double mating. The amount of sperm in the seminal vesicle differed according to male status (increasing from previously mated males to old males), but there was no difference in sperm stored by females after a single mating. In double mating experiments with two males of different status, paternity increased linearly with the relative amount of sperm in seminal vesicles. Paternity distribution conforms to 'a fair raffle' of sperm from both donors following complete mixing of sperm prior to fertilization. Thus, in a female multiple mating context, male fitness depends principally on their sperm stock, which in turn depends on life history parameters, such as age and previous mating.     

The effects of mating, age at mating, and plant stimuli, on the lifetime fecundity and fertility of the generalist herbivore Epiphyas postvittana

The effects of mating, age at mating, the presence or absence of a plant leaf, and the deprivation of a suitable ovipositional substrate during when the first ovipositional bout after mating would normally take place, on the lifetime fecundity and fertility (percentage of fertile eggs laid) of female Epiphyas postvittana were investigated. Mating had a significant effect on lifetime fecundity, with mated females laying 2.5 times more eggs than virgin females. Age at mating had a significant effect on both fecundity and fertility, both declining with increasing age when the female was mated. In the presence of a leaf of C. japonica, mated females had a greater lifetime fecundity than when no leaf was present; females in the presence of a C. japonica leaf consistently laid more eggs each day during the first 4–6 days after mating than females without a leaf. When females were deprived of a suitable ovipositional substrate, for the first 22 h after mating, they were significantly less fecund over their lifetime than were control females. Finally, in no-choice tests with three plants of different acceptability to females, the fecundity of females differed in the order C. japonica > Urtica ferox > Tibouchina multiflora. This different fecundity appeared to be inversely related to the pubescence of the leaves, suggesting that leaf texture may be a suitable antixenotic resistance factor for crops to be protected from this insect. These results suggest that strategies whereby mating is delayed or oviposition reduced within a critical period after mating, may result in significant reductions in pest populations.     

The effect of alternative mating tactics on the fertilization success of a hermaphroditic seabass

In the simultaneously hermaphroditic marine fish, Serranus subligarius, male role individuals are known to pair spawn, group spawn and streak spawn. While the effects of these common mating tactics on mating success in the male role have been well studied, their consequences for the reproductive success of the individuals taking the female role have received little attention. To investigate those consequences, I observed mating behaviors and quantified fertilization success in natural and experimental settings during the summers of 2005-2008 at three sites with different local population densities. I observed focal individuals in 15-min increments and recorded the total number of spawns, number of streak spawns, size of participating spawners, and fertilization rate. The occurrence of small-sized individuals in the local population is associated with higher frequencies of streaking behavior; these small fish are most often first-year individuals reaching sexual maturity late in the spawning season (August/September). Spawns that included one or more streak spawners had a significantly lower average fertilization rate (89?%) than pair spawns without a streak spawner (97?%). This pattern was confirmed with a field manipulation experiment in which spawning events that included streakers again showed lower fertilization rates (93?%) than spawning events that did not include streakers (98?%). These lower fertilization rates occurred despite the fact that spawns that included multiple males produced, on average, 20?% more sperm than produced in spawns with only a single male. These results indicate that females incur a significant fitness cost when streakers invade a spawning event, a cost not attributable to sperm limitation or any direct effects on the female.     

Impact of male mating history on the temporal sperm dynamics of Choristoneura rosaceana and C. fumiferana females

In the oblique-banded leafroller, Choristoneura rosaceana, and the spruce budworm, C. fumiferana, male reproductive performance decreases with consecutive matings. While the onset time of mating did not vary, the time spent mating was longer in mated than in virgin males. Furthermore, a decline observed in the spermatophore mass with successive matings was associated with a concomitant decline in its apyrene and eupyrene spermatozoa content. In the hours following mating, spermatozoa migrate from the spermatophore, located in the bursa copulatrix, to the spermatheca. Regardless of the male's previous mating history, the number of apyrene sperm dropped rapidly in the days following mating whereas the number of eupyrene spermatozoa declined gradually. As the temporal pattern of sperm movement was similar in all treatments, females mated with previously-mated males would suffer from sperm shortage sooner than those mated with virgins. Large C. rosaceana females stored more apyrene spermatozoa in their spermatheca than small ones, irrespective of the time after mating or male mating history, while only large females mated with once-mated males received more apyrene sperm and accessory gland secretions than small ones mated with virgin or twice-mated males. The results obtained in this study are discussed in relation with their potential impact on the reproductive success of both sexes.     

Effects of male fecundity, interindividual distance and anisotropic pollen dispersal on mating success in a Scots pine (Pinus sylvestris) seed orchard

Quantifying the effect of pollen dispersal and flowering traits on mating success is essential for understanding evolutionary responses to changing environments and establishing strategies for forest tree breeding. This study examined, quantitatively, the effects of male fecundity, interindividual distance and anisotropic pollen dispersal on the mating success of Scots pine (Pinus sylvestris), utilizing a well-mapped Scots pine seed orchard. Paternity analysis of 1021 seeds sampled from 87 trees representing 28 clones showed that 53% of the seeds had at least one potential pollen parent within the orchard. Pronounced variation in paternal contribution was observed among clones. Variations in pollen production explained up to 78% of the variation in mating success, which was 11.2 times greater for clones producing the largest amount of pollen than for clones producing the least pollen. Mating success also varied with intertree distance and direction, which explained up to 28% of the variance. Fertilization between neighboring trees 2.3 m apart was 2.4 times more frequent than between trees 4.6 m apart, and up to 12.4 times higher for trees downwind of the presumed prevailing wind direction than for upwind trees. The effective number of pollen donors recorded in the seed orchard (12.2) was smaller than the theoretical expectation (19.7). Based on the empirical observations, a mating model that best describes the gene dispersal pattern in clonal seed orchards was constructed.     

Mating behaviour and male mating success in the green treefrog

Green treefrogs, Hyla cinerea, at a study site in eastern Georgia behaved as prolonged breeders: there were many more males present each night than females, the arrival of which was unpredictable. Individual males were usually consistent in their behaviour during a single night, and were most often observed calling. There was a weak, negative correlation between male size (snout-vent length, unless otherwise stated) and the frequency of satellite behaviour, defined as a non-calling male's being situated near a calling male. In 2 of 3 years, the mean size of males observed as satellites was less than that of the calling males with which they were associated on a given night; however, many satellites were the same size or larger than calling males. The distributions of matings per male during three seasons were not significantly different than Poisson distributions with the same means. Individual mating success was best predicted by attendance at the breeding site; mating success was negatively related to the frequency of satellite behaviour in some analyses, but very little of the variance was explained. An exhaustive series of analyses failed to show any pattern of size-dependent mating, including weight-assortative mating, even when males that were frequently satellites were excluded. The relative sizes of males and females in amplexus had little effect on fertilization success. On a seasonal basis, males were clumped in their spatial distribution, and matings tended to occur where male density was highest. Individual males were highly variable in their patterns of movement, which were not correlated with size or mating success. Mating success was also unrelated to the frequency (Hz) of the low-frequency spectral peak in the advertisement call of the male; male size and weight were negatively correlated with the frequency of the low-frequency peak.     

Male age, dominance, and mating success among rhesus macaques

Some previous primate studies have found a positive correlation between male dominance and mating success when data from subadult males were included in the analyses. The information in this paper suggests that an unconscious bias may have been introduced when data on subadult males were included because of the lower dominance rank of these animals. Data from a study of rhesus monkeys on Cayo Santiago showed that subadult males mated significantly less than adults. Because these monkeys are not fully mature, data on them should not be used in any test for correlation between adult male dominance and mating success. The only significant correlation found for adult male mating success was an inverse one with relation to age. Based on behavioral data young, fully adult males have the best chance of fathering offspring regardless of their dominance rank.     

Genital morphology and fertilization success in the dung beetle Onthophagus taurus: an example of sexually selected male genitalia

In animals with internal fertilization and promiscuous mating, male genitalia show rapid and divergent evolution. Three hypotheses have been suggested to explain the evolutionary processes responsible for genital evolution: the lock-and-key hypothesis, the pleiotropy hypothesis and the sexual-selection hypothesis. Here, we determine whether variation in male genital morphology influences fertilization success in the dung beetle Onthophagus taurus, as predicted by the sexual-selection hypothesis. Variation in four out of five genital sclerites of the endophallus influenced a male's fertilization success, supporting the general hypothesis that male genitalia can evolve under sexual selection. Furthermore, different genital sclerites were found to enhance first versus second male paternity, indicating that different sclerites serve offensive and defensive roles. Genital-trait variability was comparable to that in other species but was less variable than a non-genital sexually selected trait (head horns). We suggest that directional selection for genital elaboration may be countered by natural selection, which should favour genitalia of a size and shape necessary for efficient coupling and sperm transfer.     

Size-assortative mating, male choice and female choice in the curculionid beetle Diaprepes abbreviatus

In the beetle Diaprepes abbreviatus (L.) females are larger on average than males, as indicated by elytra length. Size-assortative matings were observed in wild populations in Florida and in laboratory mating experiments. We tested three mechanisms for this size-assortative mating: (1) mate availability; (2) mating constraints; and (3) mate 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; and (3) females accepting large males as mates more readily than small males. Males increased their reproductive success by mating with larger, more fecund females. They transferred protein to females during mating. Copyright 1999 The Association for the Study of Animal Behaviour.     

The mating system and male mating success of the Virginia opossum (Didelphis virginiana) in Florida

The mating system of the Virginia opossum, a solitary, nocturnal marsupial, was studied over a period of two years in north–central Florida. There were two mating seasons each year, both lasting about four weeks and starting in mid-January and late April, respectively. Radio-tracking males and females revealed that, independent of body size, males doubled their home ranges during these periods to include an average of 5.9 females within their new ranges. Females were visited regularly and, on the night of oestrus, males directly competed for access to the female. Dominance and hence mating success was strongly related to body mass but not age. Dominant males weighed on average 3.18 kg (N = 20) compared to 2.65 kg of unsuccessful males. The largest of the competing males was successful in 20 of 23 cases involving at least two competitors. Individual male mating success varied from zero to three matings during a breeding season, but the range was probably underestimated. Mate searching activities were costly, as most males lost weight during both mating seasons. Weight changes were a function of body mass, with males over 2.2 kg losing up to 23% in weight and smaller males gaining weight. There was also evidence that male breeding activities entailed a survival cost.