Lower Mating Success Research Articles

Alternative mating tactics in the Italian treefrog, Hyla intermedia

As in many lekking anurans, Italian treefrog males use two mating tactics: they can attract females by calling vigorously or be satellites, that is, they can remain silent in proximity of a calling male and try to intercept females attracted by their neighbour. We investigated the factors that affected the expression of this mating tactic. Consistent with the conditional mating tactic hypothesis, satellites were smaller than average and smaller than their parasitised calling males. They spent a larger-than-average number of nights at the breeding site, where most of them were also observed calling. Moreover, satellites showed lower call rates and lower mating success than those of males they parasitise but not lower than those of males they did not parasitise. Overall, these results, together with those derived from the analyses of the seasonal and spatial distribution of males, provide evidence for a non-random association between satellites and calling males and are consistent with the hypothesis that satellites have spectral and temporal acoustic preferences that parallel those of females. By adopting the less-successful satellite mating tactic, competitively inferior males can nevertheless maximise their potential reproductive fitness by sexually parasitising the most attractive chorusing males.

Two-Generation Reproductive Toxicity Study of Implanted Depleted Uranium (DU) in CD Rats

Depleted uranium (DU) munitions and armor plating have been used in several conflicts over the last 17 yr, including the Persian Gulf War and the Iraq War. Because of its effectiveness and availability, DU will continue to be used in military applications into the foreseeable future. There is much controversy over the use of DU in weapons and equipment because of its potential radiological and toxic hazards, and there is concern over the chronic adverse health effects of embedded DU shrapnel in war veterans and bystanders. This study evaluated the effects of long-term implantation of DU on the reproductive success of F0 generation adults and development and survival of subsequent F1 and F2 generations in a two-generation reproductive toxicity study. F0 generation Sprague-Dawley rats, 8 wk of age, were surgically implanted with 0, 4, 8, 12, or 20 DU pellets (1 × 2 mm). Inert implant control animals were implanted with 12 or 20 tantallum (Ta) pellets. The F0 generation was then mated at 120 d post DU implantation. In the F0 generation, when measured on postimplantation d 27 and 117, uranium was present in the urine of DU‐implanted animals in a dose-dependent manner. F0 reproductive success was similar across treatment groups and the maternal retrieval test revealed no changes in maternal behavior. DU implantation exerted no effect on the survival, health, or well-being of the F0 generation. Necropsy results of F0 animals were negative with the exception of a marked inflammatory response surrounding the implanted DU pellets. For the F1 generation, measures of F1 development through postnatal day (PND) 20 were unremarkable and no gross abnormalities were observed in F1 offspring. No uranium was detected in whole-body homogenates of PND 4 or PND 20 pups. Necropsy findings of F1 PND 20 pups were negative and no instances of ribcage malformation were observed in F1 PND 20 pups. Body weight and body weight gain of F1 rats through PND 120 were similar across treatment groups. Eight of 414 F1 animals observed from PND 20 to 120 died of unknown causes; 7 were from litters of DU-implanted F0 mating pairs. F1 mating success at 10 wk of age was an overall 70% compared with 91% for F0 mating pairs. Mating success was similar between F1 animals derived from DU-implanted F0 adults and those derived from F0 implant control adults suggesting that the comparatively low mating success was not due to F1 DU exposure. The gestational index of F1 animals derived from mid-dose F0 mating pairs was found to be lower compared with F1 controls. The average gestation duration of F1 animals derived from high-dose F0 mating pairs was found to be significantly longer than F1 controls. F1 sperm motility analyses did not differ among experimental groups and no gross abnormalities were identified at necropsy among surviving F1 animals at PND 120. Histopathology of kidneys, spleen, thymus, bone marrow, ovaries, and testes of F1 high-dose animals did not differ from F1 controls. F1 high-dose females had significantly higher mean relative liver and heart weights compared with F1 controls; the biological relevance of this finding could not be determined. For the F2 generation, measures of F2 development through PND 20 were unremarkable and no gross abnormalities were observed in F2 offspring. Necropsy findings of F2 PND 20 pups were negative and no instances of ribcage malformation were observed in F2 PND 20 pups. Body weight and body weight gain of F2 rats through PND 90 were similar across treatment groups. Mean relative heart weights of males derived from high-dose F0 parents were significantly lower compared with F2 controls. Sperm motility and concentration analysis of F2 males at PND 90 were similar across F2 groups. Overall, the consistent absence of positive findings in this study seems to suggest that DU is not a significant reproductive or developmental hazard, particularly when one considers that mid- and high-dose rats were implanted with the equivalent of 0.3 and 0.5 lb of DU in a 70-kg human, respectively. However, the findings that seven of eight F1 adults that died postweaning were from DU-implanted F0 mating pairs, and that mean relative heart weights were elevated in high-dose F1 and F2 pups, suggest conservatism is warranted in characterizing the reproductive and teratogenic hazards of embedded DU until further studies are completed.

Male nest-building behaviour and mating success in the red bishop (Euplectes orix)

AbstractWe analysed the relation between male nest-building behaviour and mating success in the red bishop (Euplectes orix), a highly polygynous and colonially breeding weaverbird. Male mating success was mainly determined by the total number of nests built per male during a breeding season, and, to a lesser extent, by the proportion of unusable nests (i.e., nests that became squashed or deformed) per male. The number of nests built increased with longer territory tenures, larger proportions of unusable nests, shorter time delays between consecutive nest-building attempts, and shorter nest-building times. Nests were characterised by the density of fibres in the nest chamber and parameters describing size and deformation. Females preferred nests that were more densely woven and had a larger entrance roof overlap; the latter was also related to nest durability. However, this preference probably reflects only a general rejection of bad nests, since nest characteristics did not differ among males and had no detectable effect on male mating success. Young males had a lower number of nests built, and consequently lower mating success, compared to old males, while there were no differences with regard to other aspects of nest-building behaviour or nest characteristics between the two age groups.

Biased sex ratio and low population density increase male mating success in the bug Nysius huttoni (Heteroptera: Lygaeidae)

Demographic factors such as operational sex ratio (OSR) and local population density (LPD) are temporally and spatially dynamic in the natural environment but the influence of these variables on male mating success and the mechanisms behind it are still poorly understood and highly controversial. Here, we manipulated the OSR and LPD of a seed bug, Nysius huttoni, and carried out a series of mating trials to test how these variables affected male mating success. The two demographic factors had no significant interactions, suggesting that they affect male mating success independently in N. huttoni. In this species male mating success was significantly higher in both male- and female-biased OSR than in even OSR. It is suggested that, in male-biased OSR, the increased intensity of competition and interference does not result in lower male mating success; rather, males may make more effort in courting and females may have more chance to encounter better males, resulting in higher male mating success. In female-biased OSR, females may become less choosy and less likely to reject male mating attempt, leading to the higher male mating success. Lower male mating success in N. huttoni in high LPD may be due to increased interference between males and/or delayed female receptiveness for mating. OSR had a stronger effect on male mating success than LPD in N. huttoni, suggesting that OSR and LPD affect mating success in different ways and intensities.

Fitness Costs Associated with Cry1Ac-Resistant <I>Helicoverpa zea</I> (Lepidoptera: Noctuidae): A Factor Countering Selection for Resistance to Bt Cotton?

The heritability, stability, and fitness costs in a Cry1Ac-resistant Helicoverpa zea (Boddie) (Lepidoptera: Noctuidae) colony (AR) were measured in the laboratory. In response to selection, heritability values for AR increased in generations 4-7 and decreased in generations 11-19. AR had significantly increased pupal mortality, a male-biased sex ratio, and lower mating success compared with the unselected parental strain (SC). AR males had significantly more mating costs compared with females. AR reared on untreated diet had significantly increased fitness costs compared with rearing on Cry1Ac treated diet. AR had significantly higher larval mortality, lower larval weight, longer larval developmental period, lower pupal weight, longer pupal duration, and higher number of morphologically abnormal adults compared with SC. Due to fitness costs after 27 generations of selection as described above, AR was crossed with a new susceptible colony (SC1), resulting in AR1. After just two generations of selection, AR1 exhibited significant fitness costs in larval mortality, pupal weight and morphologically abnormal adults compared with SC1. Cry1Ac-resistance was not stable in AR in the absence of selection. This study demonstrates that fitness costs are strongly linked with selecting for Cry1Ac resistance in H. zea in the laboratory, and fitness costs remain, and in some cases, even increase after selection pressure is removed. These results support the lack of success of selecting, and maintaining Cry1Ac-resistant populations of H. zea in the laboratory, and may help explain why field-evolved resistance has yet to be observed in this major pest of Bacillus thuringiensis cotton, Gossypium hirsutum L.

A role for female ornamentation in the facultatively polygynous mating system of collared flycatchers

In a polygynous mating system, females settling with already mated males often experience low mating success due to the reduced parental contribution of the male. However, there are numerous factors that may still make it advantageous for some females to choose this mating status. Facultative polygyny is believed to be dominated by male advertisement and female choice. Although quality differences and competition among females are increasingly recognized as important determinants of polygynous settlement patterns, the importance of signals of female quality in this mating system is largely unknown. Here we examined the relationship of the white wing patch size (WPS) of female collared flycatchers, a phenotypically plastic and age-dependent ornament, with social mating status, while controlling for settlement date and age. At the population level, monogamous, primary, and secondary females did not differ in WPS. However, the primary female of individual males was more ornamented than the secondary female, and this difference declined with increasing distance between primary and secondary nests. Secondary female ornamentation increased, whereas that of the primary female did not change with nest distance. These results suggest a subtle role for female ornamentation at polygynous mating. Future studies should therefore take into account mating status when assessing the costs and benefits of female signals. Moreover, patterns in quality-indicating female traits may contribute to the explanation of differences in reproductive success among females of different mating status. Key words: female sexual ornament, individual quality, polygyny, settlement date, territorial distance. [Behav Ecol]

Assortative Mating between European Corn Borer Pheromone Races: Beyond Assortative Meeting

BackgroundSex pheromone communication systems may be a major force driving moth speciation by causing behavioral reproductive isolation via assortative meeting of conspecific individuals. The ‘E’ and ‘Z’ pheromone races of the European corn borer (ECB) are a textbook example in this respect. ‘Z’ females produce and ‘Z’ males preferentially respond to a ‘Z’ pheromone blend, while the ‘E’ race communicates via an ‘E’ blend. Both races do not freely hybridize in nature and their populations are genetically differentiated. A straightforward explanation would be that their reproductive isolation is a mere consequence of “assortative meeting” resulting from their different pheromones specifically attracting males towards same-race females at long range. However, previous laboratory experiments and those performed here show that even when moths are paired in a small box – i.e., when the meeting between sexual partners is forced – inter-race couples still have a lower mating success than intra-race ones. Hence, either the difference in attractivity of E vs. Z pheromones for males of either race still holds at short distance or the reproductive isolation between E and Z moths may not only be favoured by assortative meeting, but must also result from an additional mechanism ensuring significant assortative mating at close range. Here, we test whether this close-range mechanism is linked to the E/Z female sex pheromone communication system.Methodology/Principal FindingsUsing crosses and backcrosses of E and Z strains, we found no difference in mating success between full-sisters emitting different sex pheromones. Conversely, the mating success of females with identical pheromone types but different coefficients of relatedness to the two parental strains was significantly different, and was higher when their genetic background was closer to that of their male partner's pheromone race.Conclusions/SignificanceWe conclude that the close-range mechanism ensuring assortative mating between the E and Z ECB pheromone races is unrelated to the difference in female sex pheromone. Although the nature of this mechanism remains elusive, our results show that it is expressed in females, acts at close range, segregates independently of the autosome carrying Pher and of both sex chromosomes, and is widely distributed since it occurs both in France and in the USA.

Comparison of methods for deploying female gypsy moths to evaluate mating disruption treatments

Abstract1 Mating disruption is the primary tactic used to reduce rates of gypsy moth population spread in the United States Department of Agriculture’s Slow‐the‐Spread of the gypsy moth programme (STS). Because STS targets very low‐density gypsy moth populations within which it is extremely difficult to collect females or egg masses, mating success in native populations cannot be determined. Therefore, the evaluation of mating disruption treatments in field experiments such as those designed to test new formulations and application methods requires deploying and recovering laboratory‐reared female moths to determine mating success.2 Five methods of deploying females were evaluated for cost, rates of female and egg mass recovery, and female mating success. The deployment methods tested were: modified delta trap, square barrier, single and double trunk bands, and tethered females.3 Deployment of tethered females had the highest cost and mating success rate, but it did not yield the highest rates of female and egg mass recovery. Deployment of females in delta traps produced the lowest cost and mating success rate, but yielded the highest recovery rate. Neither of these deployment methods is recommended because of unacceptably high cost (tethered female) or low mating success (delta trap).4 There were no significant differences in cost or mating success among the other three deployment methods.5 The differences among the square barrier, single trunk band, and double trunk band methods in cost, female and egg mass recovery, and mating success are too small to recommend any one over the others.

SCARED TO DEATH? THE EFFECTS OF INTIMIDATION AND CONSUMPTION IN PREDATOR–PREY INTERACTIONS

Predation is a central feature of ecological communities. Most theoretical and empirical studies of predation focus on the consequences of predators consuming their prey. Predators reduce prey population densities through direct consumption (a density-mediated interaction, DMI), a process that may indirectly affect the prey's resources, competitors, and other predators. However, predators can also affect prey population density by stimulating costly defensive strategies. The costs of these defensive strategies can include reduced energy income, energetic investment in defensive structures, lower mating success, increased vulnerability to other predators, or emigration. Theoretical and empirical studies confirm the existence of these induced costs (trait-mediated interactions, TMIs); however, the relative importance of intimidation (TMI) and consumption (DMI) effects remains an open question. We conducted a meta-analysis assessing the magnitude of both TMIs and DMIs in predator–prey interactions. On average, the impact of intimidation on prey demographics was at least as strong as direct consumption (63% and 51% the size of the total predator effect, respectively). This contrast is even more pronounced when we consider the cascading effects of predators on their prey's resources: density effects attenuated through food chains, while TMIs remained strong, rising to 85% of the total predator effect. Predators can thus strongly influence resource density even if they consume few prey items. Finally, intimidation was more important in aquatic than terrestrial ecosystems. Our results suggest that the costs of intimidation, traditionally ignored in predator–prey ecology, may actually be the dominant facet of trophic interactions.

The sublethal effects of tebufenozide on the precopulatory and copulatory activities of Choristoneura fumiferana and C. rosaceana

AbstractThe sublethal effects of tebufenozide, an ecdysone agonist, on the reproductive biology of Choristoneura fumiferana (Clem) and of Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae), treated during the larval stage, were evaluated using two treatment methods: the force‐feeding method and the diet method. The percentage of mortality and the developmental time of survivors increased linearly with the concentration of tebufenozide used. This ecdysone analogue proved to be more toxic to C. fumiferana than to C. rosaceana. In C. rosaceana, the weight of males and females decreased proportionally with the dose ingested, but females were affected to a greater extent. This difference might be due to a greater consumption of the treated diet, or to a differential vulnerability to tebufenozide. Tebufenozide did not modify the pre‐copulatory activities associated with chemical communication in the females. However, the consumption of tebufenozide delayed ovarian maturation, causing a reduction in the fecundity of females. Treated males had smaller spermatophores and fewer eupyrene sperms in their bursa copulatrix and spermatheca, along with lower mating success. In C. fumiferana, tebufenozide delayed the females’ onset time of calling the first night after emergence, but did not affect the mean time spent calling or the production of the main component of the sex pheromone. The males showed significantly greater difficulty in executing oriented flight in a wind tunnel, although their mating success was not affected. We concluded that tebufenozide interferes with various aspects of the reproductive biology of males and females of C. fumiferana and C. rosaceana, including some pre‐copulatory behaviors associated with sex pheromone communication.

Relationship between morphometric and genetic variation in pure and hybrid populations of the smooth and Montandon's newt (Triturus vulgaris and T. montandoni)

AbstractThe variation in 10 morphometric characters and the degree of sexual dimorphism in size and shape was studied in pure and hybrid populations of smooth Triturus vulgaris and Montandon's T. montandoni newts. Multivariate analyses showed pronounced interspecific differences in both sexes. Females differed mainly in traits related to general size, whereas males differed mainly in body proportions. Triturus vulgaris males possessed a relatively longer trunk and tail. Sexual size dimorphism was much stronger in T. montandoni, the females being the larger sex. In hybrid zone populations a general correlation between morphometric traits and nuclear genotypes was found. This does not, however, fully explain the variation in size and shape. Whereas the majority of genetically identified hybrids fell into the range of morphologically pure species, many genetically pure individuals were classified as morphometric intermediates. The linear relationship between the morphometric and genetic variation suggested no strong differential selection on the molecular markers and loci responsible for morphometric differences between species. The differences in the level of sexual size and shape dimorphism, however, suggest that females of both species show dissimilar mate preferences, and thus hybrid males may experience reduced fitness resulting from lower mating success.

Depensation, probability of fertilization, and the mating system of Atlantic cod (Gadus morhua L.)

AbstractAtlantic cod (Gadus morhua L.) have been severely overexploited and are currently at historic population lows, having declined 90% in the North Sea and 99% off northeast Newfoundland in recent decades. Slow rates of recovery and continuing declines may be attributable to depensation, defined as a reduction in per capita growth rate concomitant with reduced population size. Several potential causes of depensation relate to low mating success and consequent reduced production of offspring. We explore the empirical basis of one of these in Atlantic cod using egg fertilization and male abundance data obtained from 21 experimental populations generated by three independent research programmes. We find support for the hypotheses that (a) fertilization rate declines with abundance and (b) variance in fertilization rate increases as population size declines. The former identifies one potential mechanism underlying depensation in Atlantic cod. The latter has negative genetic consequences for effective population size (Ne), resulting in a decline in the ratio of Ne to census population size (Ne/Nc) with declining abundance. Our results may have general implications for the conservation biology of broadcast-spawning marine fish, particularly those with mating systems similar to that of Atlantic cod.

Assessment of Ovarian Development and Its Relation to Mating in Wild and Pond‐Reared Litopenaeus vannamei Shrimp in a Commercial Hatchery

AbstractThe present study compares three scales for evaluation of ovary development stage in wild and pond‐reared shrimp. An external visual scale was assessed according to color and size of ovaries observed through the exoskeleton in live shrimp by staff of a commercial hatchery. Shrimp were then sacrificed to obtain gonadosomatic index, and gonads were scaled according to their color and size upon dissection. Finally, a portion of ovaries was processed for histological analysis, from which oocyte diameters were obtained, and females were evaluated with another scale based on cytological organization of gonads and on frequency of oocyte sizes. The certainties of the external and internal scales were analyzed, based on differences in gonadosomatic index and oocyte diameter and by comparison with the more accurate scaling obtained by histological analysis. The external scale was, in general, less accurate than the internal scale. Maturation stage in pond‐reared shrimp was underestimated or overestimated more frequently than in wild shrimp. The implications for hatchery operation and possible reasons are discussed. Pond‐reared shrimp were found to have a non‐linear relation between gonad weight and body weight, but this relation was linear for the larger wild shrimp. This reduced the validity of gonadosomatic index in pond‐reared shrimp, which were still growing. Thus, oocyte diameter, which was not affected by body weight in growing shrimp, represents a more accurate numeric measure of ovary development. Spermatophores were found attached to the thelycum in females only at the cortical stage of maturation, independent of origin. However, a significant proportion of fully mature, pond‐reared shrimp did not have an attached spermatophore, possibly indicating lower mating success or poorer spermatophore adherence in these shrimp.

Mating success and fidelity to territories in a fallow deer lek: a female removal experiment.

A lek is a clump of male territories that females visit only for mating; it is quite a rare mating system among mammals. As we wanted to investigate the role of past male mating success on their fidelity to lek territories, we displaced fallow deer females by a lek (Lek A), i.e. we removed the resources for rutting males. They moved to a new area followed by some of the territorial males ('mobile' males) where they established a new lek (Lek B). Other males ('faithful' males) stayed on Lek A. The former had lower mating success and dominance ranking in the early, undisturbed part of the rut. After disturbance, the MS of faithful males was much reduced, while that of mobile males was higher on Lek B than it had been on Lek A. We then disturbed Lek B. Females returned to Lek A, and the mating success of faithful males reverted to its original level. Past mating success was the main factor predicting whether a male was faithful or mobile. We illustrated for the first time the importance of past experience in promoting lek fidelity; this experiment gives an insight into the mechanisms leading to the traditional use of leks.

Female reproductive decisions and parasite burden in a calopterygid damselfly (Insecta: Odonata)

There is currently a gap in sexual selection theory about how much the environment drives female mating decisions. We present field data that suggest that female sexual behaviour in the damselfly Calopteryx haemorrhoidalis is influenced by parasite burden. Male wing pigmentation in Calopteryx is a sexually selected trait that signals a male's ability to cope with eugregarine parasites (an intestinal parasite that feeds on the adult's ingested food). Because adult C.haemorrhoidalis females also show wing pigmentation, we examined whether this trait is similarly influenced by parasite burden and whether it may signal the female's reproductive value. MaleC.haemorrhoidalis defend riverine substrates that females use for oviposition. After copulation and during oviposition, females are guarded by the copulating male against intruder males. Alternatively, females may avoid mating and ‘steal’ an oviposition site within a male's territory. In the present study, we found that the amount of female wing pigmentation was negatively correlated with the number of eugregarines present. Females with more parasites produced fewer eggs, survived fewer days, spent less time during courtship, ‘inspected’ fewer males before mating, had a lower mating success, were guarded for less time during oviposition and engaged in fewer ‘stealing’ events during oviposition. The reduced egg production and survival of heavily infected females may result from eugregarine depletion of the females' consumed food reserves. Thus, to offset reduced longevity, heavily infected females may accept a mating more rapidly and mate with fewer males. ‘Stealing’ behaviour may be related to the female's differential use of sperm from some males, particularly high-quality males. Interestingly, males that mated with low-pigmented females showed greater variance in wing pigmentation than did males that mated with high-pigmented females. Possibly, female wing pigmentation may signal a female's reproductive value, which provides females with longer mate-guarding episodes and reduced interference from intruder males. This study points out one possible constraint, intestine parasites, that females may face during mating decisions. Because females in bad condition mate with males in both good and bad condition, this constraint may be pervasive enough to weaken the intensity of selection for a male sexually selected trait, wing pigmentation, and help to maintain its variation in phenotypic expression. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.

Nonlocal male mountain white-crowned sparrows have lower paternity and higher parasite loads than males singing local dialect

Bird song often varies geographically, and when this geographic variation has distinct boundaries, the shared song types are referred to as song dialects. We investigated the role of song dialect in male mating success in a wild breeding population of mountain white-crowned sparrows (Zonotrichia leucophrys oriantha). In 2 of 3 years, males singing unusual songs (“nonlocal” males) had lower total fertilization success (measured by microsatellite paternity analysis) than did males singing the local dialect (“local” males). Similarly, females produced disproportionately more young with local than with nonlocal males. However, dialect was not a significant predictor of male mating success when controlling for other factors that might affect paternity. Instead, the low mating success of nonlocal males was apparently due to an interaction between song dialect and parasite load. Nonlocal males were more severely infected by bloodborne Haemoproteus than were local males, although they did not differ in any other measured aspect of quality. Immigrant birds may be immunologically disadvantaged, possibly due to a lack of previous experience with the local parasite fauna, resulting in low mating success.

Modified pheromone communication associated with insecticidal resistance in the obliquebanded leafroller, Choristoneura rosaceana (Lepidoptera: Tortricidae)

This study reports on the impact of insecticidal resistance on the diel periodicity of the calling behaviour and pheromone production of different-aged virgin females of the obliquebanded leafroller (OBL), Choristoneura rosaceana. While both resistant (R) and susceptible (S) females initiated calling on the first night following emergence, the periodicity of the calling behaviour, as determined by the mean onset time of calling (MOTC) and the mean time spent calling (MTSC) over the first six nights of calling, differed between the two strains. R females started calling significantly later in the night. However, as the MOTC of R females advanced with age but did not do so in S individuals, the difference between strains was more pronounced in younger than older females. Furthermore, R females spent less time calling than S individuals. However, the MTSC increased as a function of age in both R and S females, so the difference between strains remained fairly constant for each night of calling. The major component of OBL sex pheromone, the Z11-14: Ac, determined at peak calling activity, significantly declined with female age. Overall, pheromone production was lower in R females than in S females, with the difference being more pronounced in younger than in older individuals. Thus, resistant females may have a lower mating success. The mating success of both R and S strain males did not vary with the number of previous matings acquired. With regard to males, although there was a significant decline in spermatophore size with successive matings, there was no significant difference between strains. However, R males are smaller and may be disadvantaged through female choice and/or may respond differently to pheromone source compared with S individuals. If the reproductive success of both sexes is affected, this may have a profound influence on the dynamics of insecticidal resistance in the presence or absence of selection in OBL populations.

Paternal effort and its relation to mating success in the European ground squirrel

The necessity for parental care is a driving force for determining mating systems and social organization. The European ground squirrel, Spermophilus citellus, is a polygynous, gregarious species in which male parental behaviour would not be expected. We had observed males digging in litter burrows that were later occupied by females and their offspring. Males never stayed overnight within these burrows. To determine whether this was some kind of paternal effort we tested the following hypotheses: (1) that male burrowing behaviour was directed towards the male's own offspring or towards the pregnant or lactating mother of the male's offspring; (2) that this behaviour had costs in terms of condition, decreased survival or fecundity; and (3) that it benefited offspring condition or survival. All three assumptions were met. Males worked on the litter burrow of their copulatory partners. Thus, this behaviour was directed towards the male's potential offspring. Male burrowing costs were seen in decreased foraging time and increased body mass loss. Offspring benefits were evident in increased mass at natal emergence. We conclude that male digging at litter burrows can be considered as paternal effort. Lastly, we considered the effects of polygyny on this male parental effort by comparing mating effort, mating success and paternal effort. High mating success was associated with high mating effort and low paternal effort. Moderate to low mating success was associated with lower mating effort and higher paternal effort, indicating a trade-off between the two.

Lifetime mating success, sexual selection and life history of fallow bucks ( Dama dama )

We used data from a long-term study (15 years) of fallow deer to report for the first time the lifetime mating success, overall variance in lifetime mating success, and age-specific mortality levels of males. Fallow bucks that gain matings have higher social dominance rank, higher rates of fighting, and invest more in vocal display during the breeding season than unsuccessful males. Therefore, we examined if mating was associated with trade-offs in terms of survival, lifespan, and mating potential. We found that the variance in lifetime mating success was very high: 34 (10.7%) males mated, and of those, the 10 most successful males gained 73% of all matings (n=934). Mortality rates were generally high and only 22.3% (71/318) of males reached social maturity, i.e., 4 years. The oldest male was 13 years old. We found that fallow bucks that mated were not more likely to die during the following year, did not suffer from a reduction in lifespan, and did not incur lower mating success later in life as a result of mating during the early years of social maturity. Our results show that mating males at age 5 years (and possibly 9 years) may be more likely to survive than non-mating males. Additionally, the number of matings gained by males during the first years of social maturity was positively correlated with lifespan. We suggest that mating males are of higher quality than non-mating males because they are not more likely to incur trade-offs as a result of their increased reproductive efforts.

Male signalling and lek attractiveness in the Mediterranean fruit fly

I tested the null hypothesis that females of the Mediterranean fruit fly, Ceratitis capitata, were equally likely to visit leks composed of males with high versus low mating success. In laboratory trials, the observed distribution of matings among males differed significantly from that expected by chance, owing primarily to the higher than expected numbers of individuals with low (mated 0–1 days over 6 consecutive observation days) or high (mated 4 or more days) mating scores. I termed these two groups as ‘low’ and ‘high’ maters, respectively. In the field, greater numbers of female sightings were made at artificial leks of high maters than low maters. This result apparently reflected a greater calling propensity among high maters. Slopes of female sightings versus calling level did not differ significantly between leks of low and high maters, suggesting that the observed relationship between calling activity and female sightings was independent of male mating status. Following the same protocol, I conducted a second experiment to examine whether males used the signals of conspecific males to locate lek sites and, if so, whether signal attractiveness varied with male mating ability. Attraction of males to calling conspecifics was far weaker than that observed for females, and over five different trials a total of only seven male sightings were made at any of the established leks.