To change or not to change sex: A comparison between twoOphryotrocha species (Polychaeta)

Summary The polychaeteOphryotrocha puerilis is a male-to-female sex-changer, whereasO. labronica has separate sexes throughout life. In other respects the two species are remarkably similar: they occur in the same habitat, they look the same, they eat the same things, and in someO. labronica populations sex-changers can actually be found. The size-advantage hypothesis predicts that inO. puerilis males should not benefit reproductively from a size increase as much as males inO. labronica: henceO. puerilis but notO. labronica males should change to the female sex at a certain size. I experimentally compared reproductive success at different body sizes between the two species. In isolated male-female pairs reproductive rate increased significantly with female body size but not with male body size, and this pattern was the same in both species. Hence male fecundityper se cannot account for the difference in reproductive type between the two species. In other experiments I investigated if larger males gained access to more females because they were superior competitors for mates or were preferred by females, compared to small males. InO. puerilis the combined effect of these two factors conferred no size advantage to the males, whereas inO. labronica larger males acquired more females than did smaller males. Hence interactions among males and females, in accordance with the size-advantage hypothesis, can explain why sex change is maintained inO. puerilis, and why separate sexes are maintained inO. labronica.     

Sex change in either direction by growth-rate advantage in the monogamous coral goby, Paragobiodon echinocephalus

The size-advantage model predicts the evolution of sex changeif the relative reproductive success of the sexes changes withsize or age. In the goby (Paragobiodon echinocephalus) the largesttwo fish, a male and a female matched by size, breed monogamouslyin each host coral. Because the female fecundity and male abilityof egg care increase with body size in a similar way, no size-fecundityadvantage exists. However, we found both protogyny and infrequentprotandry in a natural population of this species in Okinawa.New pairs were often formed after movement between host coralsand also sex change or sex differentiation of one or both members.In most new pairs males were larger than females, and femalesgrew much faster than their mates until breeding (growth-rateadvantage). The smaller member of a new pair should be the femalethat grows faster, because the smaller limits the reproductivesuccess of the pair. To form such a pair, the goby changed sexaccording to the sex and relative size of a new mate, as a status-dependentconditional strategy. The growth-rate advantage predicts predominanceof protogyny, but movement between host corals provides opportunitiesalso for protandry.     

Mating system and protandrous sex change in the lizard flathead, <Emphasis Type="Italic">Inegocia japonica</Emphasis> (Platycephalidae)

Mating system and protandrous sex change of the lizard flathead Inegocia japonica were investigated off Nagashima, Kagoshima, Japan. The reproductive season of this species was estimated to be from June to September by a fluctuation of gonadsomatic index from monthly sampling data. The hermaphrodite fish, having testicular and ovarian parts, were collected from February to December, indicating that the sex change occurred in both reproductive and nonreproductive seasons. Females were larger than males and hermaphrodites. Moreover, five cases of the sex change from male to female were directly confirmed during the 1995–1997 field census in the behavioral observation area. Home ranges, where no aggressive interaction took place among any individuals, overlapped within and between sexes. Seven pair spawnings observed in the area were not of the same individual combination, and the size of pair fish was likely to be random. This species may thus have a promiscuous mating system, i.e., random pairing, which favors protandry as predicted in the size-advantage model.     

Size-dependent gender modification in a hermaphroditic perennial herb

The size-advantage model predicts that hermaphroditic organisms adjust sex allocation depending on their resource status. We investigated the relationship between size and sex allocation in the co-sexual perennial herbs Trillium erectum and Trillium grandiflorum at two sites in southern Ontario, Canada by measuring pollen and ovule production and biomass allocation at flowering and fruiting. In both species, there was a strong relationship between size and gender; larger plants allocated proportionately more biomass to female reproduction and produced fewer pollen grains relative to ovules than smaller plants. Variation in gender was better explained by size than age, although age and size were correlated. While the relationship between size and gender was similar between species, T. erectum allocated proportionately more to female reproduction than T. grandiflorum, independent of size. In the absence of pollen limitation, there was no evidence of secondary adjustment of gender at fruiting. The results are discussed in the context of models predicting size-dependent gender modification in animal-pollinated plants. Evidence about the pollination and seed dispersal biology of Trillium spp. suggests that the relative effects of local mate and resource competition may be important in driving size-dependent sex allocation in these species.     

Patterns of male reproductive success in Crepidula fornicata provide new insight for sex allocation and optimal sex change

The size-advantage model and sex-allocation theory are frequently invoked to explain the evolution and maintenance of sequential hermaphroditism in many taxa. A test of current theory requires quantitative estimates of reproductive success and knowledge of the relationship between reproduction and size for each gender. Reproductive success can be difficult to measure. In species where polyandry occurs, it can be quantified only by determining paternity of offspring. We employed microsatellite loci to establish paternity for 12 families of Crepidula fornicata, where a family is defined as a single female, her brood, and the males stacked on top of her. Genetic data were analyzed and paternity was assigned to a single potential father for more than 83% of the offspring tested. Estimates of reproductive success revealed that one male within the family fathered the majority of offspring and that he was usually the largest male and the one closest to the brooding female. The dominant male's success also tended to decrease as the number of mature males within the family increased. Our results suggest that sperm competition could be a driving force in determining male reproductive success and the timing of sex change in C. fornicata.     

Lack of Sex-Biased Maternal Investment in spite of a Skewed Birth Sex Ratio in White-Headed Langurs (Trachypithecus leucocephalus)

Numerous hypotheses have been developed to explain sex allocation. In male-dispersing, female cooperatively breeding species, the local resource competition model predicts male-biased birth sex ratio, the local resource enhancement model predicts female-biased birth sex ratio, and the population adjustment model predicts that biased birth sex ratio should not be favored if the two sexes are equally costly to rear. The male quality model predicts that, in polygynous species, females in better physical condition will either produce more sons than daughters or invest more heavily in sons than in daughters. White-headed langurs are a female philopatry and female cooperatively breeding species. During a 11-yr study, a total of 133 births were recorded, among which birth sex ratio (M:F = 73:49) was significantly male-biased. This is consistent with the prediction of the local resource competition model. On the other hand, if mothers balanced their investment between the two sexes, according to Fisher's population adjustment model, males should be the less-costly-to-rear sex. However, we found no sex difference for infant mortality (12.3% in males and 12.2% in females), and sons induced slightly longer interbirth interval (son: 26.4 ± 1.1 mo, daughter: 24.1 ± 0.6 mo) and lactational period (son: 20.9 ± 1.0 mo, daughters: 19.6 ± 0.5 mo) for their mothers. Thus, the population adjustment model was not supported by this study. The local resource enhancement model was not supported because birth sex ratio did not bias to females who provided more reproductive assistance. On the individual level, probit regression showed no relation between birth sex ratio and group size. Because the group size was considered to be negatively related to female physical condition, our study did not support the male-quality model. We suggested several possibilities to explain these results.     

Local sex ratio affects the cost of reproduction

1.?Costs and benefits of reproduction are central to life-history theory, and the outcome of reproductive trade-offs may depend greatly on the ecological conditions in which they are estimated. In this study, we propose that costs and benefits of reproduction are modulated by social effects, and consequently that selection on reproductive rates depends on the social environment. 2.?We tested this hypothesis in a great tit Parus major population. Over 3 years, we altered parental reproductive effort via brood size manipulations (small, intermediate, large) and manipulated the local social environment via changes in the local fledgling density (decreased, increased) and the local sex ratio (female-biased, control, male-biased). 3.?We found that male-biased treatment consistently increased the subsequent local breeding densities over the 3-year study period. We also found that parents rearing small broods in these male-biased plots had increased survival rates compared with the other experimental groups. 4.?We conclude that reproductive costs are the product of an interaction between parental phenotypic quality after reproduction and the social environment: raising a small brood had long-lasting effects on some phenotypic traits of the parents and that this increased their survival chances in male-biased environment where habitat quality may have deteriorated (via increased disease/predation risk or intraspecific competition). 5.?Our results provide the first experimental evidence that local sex ratio can affect reproductive costs and thus optimal clutch size.     

Gregariousness and protandry promote reproductive insurance in the invasive gastropod Crepidula fornicata: evidence from assignment of larval paternity

According to the size-advantage hypothesis, protandric sequential hermaphroditism is expected when the increase in reproductive success with age or size is small for males but large for females. Interestingly, some protandrous molluscs have developed gregarious strategies that might enhance male reproductive success but at the cost of intraspecific competition. The gastropod Crepidula fornicata, a European invading species, is ideal for investigating mating patterns in a sequential hermaphrodite in relation to grouping behaviour because individuals of different size (age) live in perennial stacks, fertilization is internal and embryos are brooded. Paternity analyses were undertaken in stacks sampled in three close and recently invaded sites in Brittany, France. Paternity assignment of 239 larvae, sampled from a set of 18 brooding females and carried out using five microsatellite loci, revealed that 92% of the crosses occurred between individuals located in the same stack. These stacks thus function as independent mating groups in which individuals may reproduce consecutively as male and female over a short time period, a pattern explained by sperm storage capacity. Gregariousness and sex reversal are promoting reproductive insurance in this species. In addition, females are usually fertilized by several males (78% of the broods were multiply sired) occupying any position within the stack, a result reinforcing the hypothesis of sperm competition. Our study pointed out that mating behaviours and patterns of gender allocation varied in concert across sites suggesting that multiple paternities might enhance sex reversal depending on sperm competition intensity.     

A dynamic model of size-dependent reproductive effort in a sequential hermaphrodite: a counterexample to Williams's conjecture

In 1966, G. C. Williams showed that for iteroparous organisms, the level of reproductive effort that maximizes fitness is that which balances the marginal gains through current reproduction against the marginal losses to expected future reproduction. When, over an organism's lifetime, the value of future reproduction declines relative to the value of current reproduction, the level of effort allocated to current reproduction should always increase with increasing age. Conversely, when the value of future reproduction increases relative to the value of current reproduction, the level of effort allocated to current reproduction should decrease or remain at zero. While this latter pattern occurs commonly in species that exhibit a delayed age at first reproduction, it may also occur following an initial period of reproduction in some sex-changing organisms that experience a dramatic increase in reproductive potential as they grow larger. Indeed, this schedule of reproductive effort is predicted by models of "early" sex change; however, these models may arrive at this result incidentally because they consider only two reproductive states: on and off. In order to examine the schedule of reproductive effort in greater detail in a system where the potential reproductive rate increases sharply, we adapt the logic and methods of time-dependent dynamic-programming models to develop a size-dependent model of reproductive effort for an example species that experiences a dramatic increase in reproductive potential at large sizes: the bluehead wrasse, Thalassoma bifasciatum. Our model shows that the optimal level of reproductive effort will decline with increasing size or age when increases to the residual reproductive value outpace the increases to current reproductive potential. This result confirms the logic of Williams's analysis of optimal life histories, while offering a realistic counterexample to his conjecture of ever-increasing allocation to current reproduction.     

Sexual development and reproductive seasonality of hogfish (Labridae: Lachnolaimus maximus), an hermaphroditic reef fish

The seasonality, size, age, colour phases and sexual dimorphism of 13 reproductive classes of hogfish Lachnolaimus maximus are described. Analysis of histological sections of gonads ( n = 1662) confirmed earlier conclusions that L. maximus is a monandric, protogynous hermaphrodite. Sex change was initiated at the end of the spawning season and over a broad range of sizes and ages. It occurred after a functional female phase (postmaturation) and proceeded more slowly (months) than previously believed. Eventually all individuals changed sex to a terminal male phase. Females were batch spawners, spawning as often as every day during winter and spring. There was no evidence of precocious sperm crypts in active females, sperm competition or other alternative male sexual strategies. Mating has been reported elsewhere to be haremic. The sexual development of L. maximus appears to be adaptive in terms of Ghiselin's size-advantage model, which links monandric protogyny and polygyny. The slow rate of sex change, however, poses problems when fishing pressure is high because harvest of a single male has the potential to reduce the reproductive output of an entire harem.     

No differential consequences of reproduction according to sex in Juniperus communis var. depressa (Cupressaceae)

In dioecious plant species, males and females are thought to have dissimilar allocation patterns. Females are believed to invest more in reproduction and less in growth and maintenance than males. This differential investment between sexes could result in distinct growth patterns and contrasting survival rates, thereby affecting the sex ratio of a population and the age and size distribution of males and females, possibly leading to habitat segregation according to sex. These effects might become more apparent under particularly limiting conditions, such as in nutrient-deficient soils or in climatically stressed environments. To verify these predictions, growth patterns, microsite characteristics, and age and size distribution of male and female individuals were compared, and population sex ratio was determined in three populations of the dioecious shrub Juniperus communis var. depressa (Cupressaceae, Pinophyta) along a short latitudinal gradient on the eastern coast of Hudson Bay (Northern Québec, Canada). We found that the northernmost population had a male-biased sex ratio, but that the southernmost one had a higher proportion of females. Our results failed to reveal any significant differences in radial growth patterns, mean sensitivity, annual elongation of the main axis, and size and age frequency distribution between males and females in any population. Furthermore, there was no evidence of microhabitat segregation according to sex as indicated by the lack of differences in the physicochemical characteristics of the substrate under males and females. Clearly, the expected ecological consequences of a presumed greater investment of females in reproduction were not apparent even under the very stressful conditions prevailing on subarctic dunes. Many factors could reduce differences in the cost of reproduction between males and females, such as the number and quality of reproductive structures produced annually by individuals of each sex, the possible photosynthetic activity of the immature female cones, and the complexity of the source/sink relationship within individuals. Alternatively, there may be no differences between sexes in their reproductive investment.     

Testing a new version of the size-advantage hypothesis for sex change: sperm competition and size-skew effects in the bucktooth parrotfish, Sparisoma radians

A variety of field studies suggest that sex change in animalsmay be more complicated than originally depicted by the size-advantagehypothesis. A modification of the size-advantage hypothesis,the expected reproductive success threshold model, proposesthat sperm competition and size-fecundity skew can stronglyaffect reproductive pay-offs. Size-fecundity skew occurs ifa large female's fecundity is markedly higher than the aggregateof the other members of her social group and, together withpaternity dilution from sperm competition, can produce situationsin which large females benefit by deferring sex change to smallerfemales. Deferral by large females can create sex-size distributionscharacterized by the presence of large females and small sex-changedmales, and it is precisely these distributions that the traditionalsize-advantage model cannot explain. We tested the predictionsof the new model with the bucktooth parrotfish, Sparisoma radians,on coral reefs in St. Croix, U.S. Virgin Islands. Collectionsand spawning observations determined that the local environmentalregime of S. radians is characterized by pervasive sperm competition(accompanying 30% of spawns) and factors that can produce substantialsize-fecundity skew in social groups. Dominant male removalexperiments demonstrate that the largest females in social groupsoften do not change sex when provided an opportunity. Instead,smaller, lower-ranking females change sex when a harem vacancyarises. This pattern of sex change is in contrast to virtuallyall previous studies of social control of sex change in fishes,but provides strong support for the general predictions of theexpected reproductive success threshold model.     

THE COSTS OF CHANGING SEX AND THE ONTOGENY OF MALES UNDER CONTEST COMPETITION FOR MATES

In its simplest form, the size-advantage hypothesis predicts that individuals should change sex in order to increase their reproductive success. In terms of lifetime expectations, this must be true for the hypothesis to hold. However, as we review here, some loss of reproductive success may occur immediately after sex change. Unavoidable costs (e.g., those resulting from a restructuring of the gonad) have not been adequately distinguished from adaptive allocations of resources which diminish current reproduction in favor of large increases in future mating success. This strategy can become particularly important for species in which a few males monopolize matings. To illustrate this idea, we describe the changes in mating frequency as mature females become sexually active males in three species of protogynous wrasses. In one species, a male defends a permanent, all-purpose territory composed of up to 12 females. When he is removed, a single female changes sex and successfully completes mating sequences with all females in the territory within an average of 5.6 days. This duration roughly corresponds to the time required for functional transformation of gonads; thus, individuals in this species suffer few reproductive losses as a result of changing sex. The largest males in two other species mate with an average of 25 to 50 females per day, but only by successfully defending reproductive territories. In one of those species, individuals that changed sex mated infrequently over a two-year period after sexual transformation and, by the end of the study, were still well below the average size of males that consistently obtained territories. Sex-changed individuals in the other species had very low reproductive success for up to 45% of the maximum lifespan as a male. It is improbable that the substantial cost of changing sex in the latter two species results from gonad restructuring or from mistakes due to imprecise cues for sex change. Instead, the cost appears to represent an investment in growth rather than current reproduction as a means of rapidly attaining a size advantage when individuals face intense competition for extraordinarily successful mating territories.     

Determinate growth in a paternal mouthbrooding fish whose reproductive success is limited by buccal capacity

The life history of the paternal mouthbrooding cardinal fish Apogon doederleini was investigated in the temperate waters of Japan, with particular reference to its growth and reproductive rate. Both males and females almost ceased to grow at age 3 years, although living to 7 years of age. Their growth pattern, represented by the relative size at sexual maturity to the asymptotic size and the von Bertalanffy growth coefficient, was among the most determinate in ectothermic vertebrates. Brood size just before hatching increased in proportion to the second power of the body size of the brooding male, and correlated more positively with the male's than the female's body size, suggesting that it was limited by the male's buccal capacity. The estimated total number of broods hatched in a breeding season showed a weak or no correlation with the body size or age in either sex. Using life-history parameters based on data of A. doederleini, a simulation model of energy allocation without considering sexual interaction revealed that the optimal growth pattern shows an indeterminate growth that differs greatly from the actual growth pattern of A. doederleini. This suggests that there are some brooding constraints to size-advantage of reproductive success in this species. The possible mechanism of such reproductive constraint is discussed.     

Microhabitat selection in the common lizard: implications of biotic interactions,age, sex,local processes,and model transferability among populations

Modeling species' habitat requirements are crucial to assess impacts of global change, for conservation efforts and to test mechanisms driving species presence. While the influence of abiotic factors has been widely examined, the importance of biotic factors and biotic interactions, and the potential implications of local processes are not well understood. Testing their importance requires additional knowledge and analyses at local habitat scale. Here, we recorded the locations of species presence at the microhabitat scale and measured abiotic and biotic parameters in three different common lizard (Zootoca vivipara) populations using a standardized sampling protocol. Thereafter, space use models and cross‐evaluations among populations were run to infer local processes and estimate the importance of biotic parameters, biotic interactions, sex, and age. Biotic parameters explained more variation than abiotic parameters, and intraspecific interactions significantly predicted the spatial distribution. Significant differences among populations in the relationship between abiotic parameters and lizard distribution, and the greater model transferability within populations than between populations are in line with effects predicted by local adaptation and/or phenotypic plasticity. These results underline the importance of including biotic parameters and biotic interactions in space use models at the population level. There were significant differences in space use between sexes, and between adults and yearlings, the latter showing no association with the measured parameters. Consequently, predictive habitat models at the population level taking into account different sexes and age classes are required to understand a specie's ecological requirements and to allow for precise conservation strategies. Our study therefore stresses that future predictive habitat models at the population level and their transferability should take these parameters into account.     

Age-dependent traits: a new statistical model to separate within- and between-individual effects

Evolutionary questions regarding aging address patterns of within-individual change in traits during a lifetime. However, most studies report associations between age and, for example, reproduction based on cross-sectional comparisons, which may be confounded with progressive changes in phenotypic population composition. Unbiased estimation of patterns of age-dependent reproduction (or other traits) requires disentanglement of within-individual change (improvement, senescence) and between-individual change (selective appearance and disappearance). We introduce a new statistical model that allows patterns of variance and covariance to differ between levels of aggregation. Our approach is simpler than alternative methods and can quantify the relative contributions of within- and between-individual changes in one framework. We illustrate our model using data on a long-lived bird species, the oystercatcher (Haematopus ostralegus). We show that for different reproductive traits (timing of breeding and egg size), either within-individual improvement or selective appearance can result in a positive association between age and reproductive traits at the population level. Potential applications of our methodology are manifold because within- and between-individual patterns are likely to differ in many biological situations.     

It takes two: Seasonal variation in sexually dimorphic weaponry results from divergent changes in males and females

Sexually dimorphic weaponry often results from intrasexual selection, and weapon size can vary seasonally when costs of bearing the weapon exceed the benefits outside of the reproductive season. Weapons can also be favored in competition over nonreproductive resources such as food or shelter, and if such nonreproductive competition occurs year‐round, weapons may be less likely to vary seasonally. In snapping shrimp (Alpheus angulosus), both sexes have an enlarged snapping claw (a potentially deadly weapon), and males of many species have larger claws than females, although females are more aggressive. This contrasting sexual dimorphism (larger weaponry in males, higher aggression in females) raises the question of whether weaponry and aggression are favored by the same mechanisms in males and females. We used field data to determine whether either sex shows seasonal variation in claw size such as described above. We found sexual dimorphism increased during the reproductive season due to opposing changes in both male and female claw size. Males had larger claws during the reproductive season than during the nonreproductive season, a pattern consistent with sexual selection. Females, however, had larger claws during the nonreproductive season than during the reproductive season—a previously unknown pattern of variation in weapon size. The observed changes in female weapon size suggest a trade‐off between claw growth and reproduction in the reproductive season, with investment in claw growth primarily in the nonreproductive season. Sexually dimorphic weaponry in snapping shrimp, then, varies seasonally due to sex differences in seasonal patterns of investment in claw growth, suggesting claws may be advantageous for both sexes but in different contexts. Thus, understanding sexual dimorphisms through the lens of one sex yields an incomplete understanding of the factors favoring their evolution.     

RUNAWAY EVOLUTION TO SELF-EXTINCTION UNDER ASYMMETRICAL COMPETITION

We analyze a popular model of the evolution of traits related to performance in exploitative competition. This model has previously been used to explain a mechanism by which interspecific competition can cause taxon cycles. We show that purely intraspecific competition can cause evolution of extreme competitive abilities that ultimately result in extinction, without any influence from other species. The only change in the model required for this outcome is the assumption of a nonnormal distribution of resources of different sizes measured on a logarithmic scale. This suggests that taxon cycles, if they exist, may be driven by within- rather than between-species competition. Self-extinction does not occur when the advantage conferred by a large value of the competitive trait (e.g., size) is relatively small, or when the carrying capacity decreases at a comparatively rapid rate with increases in trait value. The evidence regarding these assumptions is discussed. The results suggest a need for more data on resource distributions and size-advantage in order to understand the evolution of competitive traits such as body size.     

Field studies of the behavioral ecology and agonistic behavior of Cichlasoma meeki (Pisces: Cichlidae)

Synopsis The behavior and ecology of the firemouth cichlid were investigated in southern Mexico. Observations were conducted primarily at Laguna Bacalar, Quintana Roo. The fish bred throughout the year, nesting in small rocky crevices. Territories were established either by wandering pairs or by lone males which subsequently attracted mates. Stationary territories were maintained for two to three weeks by both pair members until the young were free-swimming, after which the parents defended the young as they foraged. Brood care lasted approximately three months. The distances at which fish were attacked and the behavior performed varied with the age of the defended young and the sex of the parent, as well as the species and size of the intruder. The sexes differed in their reproductive and agonistic activities. Predation on young was rapid if the parents were diverted from their defense. Serious injuries to adults were not observed to result from intraspecific contests. Such interactions were very common, but were ritualized with physical contact between animals rare. Color patterning varied with breeding interval, sex and behavior.     

Delayed mating in tortricid leafroller species: simultaneously aging both sexes prior to mating is more detrimental to female reproductive potential than aging either sex alone

The effect of delayed mating on reproductive potential, longevity and oviposition period of female redbanded leafroller, Argyrotaenia velutinana (Walker) and Pandemis leafroller, Pandemis pyrusana Kearfott, was investigated in the laboratory. Virgin female or male moths of each species were held for 1, 2, 4, 6 or 10 days prior to pairing with one-day-old virgin conspecifics of the opposite sex. In addition, reproductive potential was assessed when both sexes of each species were aged for those periods prior to pairing. The expected reproduction of female A. velutinana was reduced by 34, 53, 71 and 81% for 2, 4, 6 and 10-day delays in female mating, respectively. For P. pyrusana, expected reproduction was reduced by 47, 74, 85 and 93% for 2, 4, 6 and 10-day delays in female mating, respectively. Increasing male age at mating in both species had a lesser effect on female reproductive output compared with increasing female age at mating. As male A. velutinana age at mating increased, the expected reproduction of female A. velutinana was reduced by 15, 45, 54 and 70% for 2, 4, 6 and 10-day delays, respectively. Comparing male P. pyrusana of various ages at mating, expected reproduction was reduced by 14, 42, 64 and 79% for 2, 4, 6 and 10-day delays in mating, respectively. The decrease in female reproduction when both sexes were aged prior to mating was higher than when either sex alone was aged prior to pairing with a one-day-old virgin of the opposite sex. The expected reproduction of female A. velutinana was reduced by 60, 83, 96 and 98% for 2, 4, 6 and 10-day delays in mating of both sexes, respectively. Only 7.5% of female eggs hatched when both sexes of A. velutinana were aged ten days prior to mating. When simultaneously aging both sexes of P. pyrusana prior to mating, expected reproduction was reduced by 71, 93, 96 and 99% for 2, 4, 6 and 10-day delays in mating, respectively. No P. pyrusana eggs hatched after a ten-day delay of mating for both sexes. For both species, female longevity increased and duration of oviposition period decreased with increasing female age at mating. Our results demonstrate that delayed mating in both females and males negatively affects female reproductive output in both species and that simultaneous aging of both sexes prior to mating has a greater effect than aging either sex alone. Our results suggest that laboratory studies that have paired aged females or aged males with conspecifics of optimal reproductive maturity have likely underestimated the effects of delayed mating on reproductive output.