Male reproductive investment relative to age and flight behaviour in the monandrous butterfly Pararge aegeria

Male reproductive investment may signify a considerable cost to male insects that produce sperm packages or spermatophores. Male butterflies allocate much of their active time budget to mate location, and they may adopt different behavioural strategies to do so. In the speckled wood butterfly (Pararge aegeria L.), males adopt either a territorial wait-and-fight strategy (territorial perching) or a fly-and-search strategy in wider areas (patrolling). In this study, we analysed the impact of male age, male size and male behaviour (i.e. behavioural strategies and levels of activity) on spermatophore investment (i.e. spermatophore mass, number of eupyrene sperm bundles). As predicted, reproductive investment increased with male age and size. Nevertheless, the increase of spermatophore mass with age and the number of eupyrene sperm bundles (i.e. fertile sperm) was stronger in low-activity males compared to active flying males. This suggests that flight activity has a negative impact on male reproductive investment. However, males that were forced to fly in the laboratory produced more eupyrene sperm bundles than resting males. We discuss the potential effects of male–male competition and predation risk on current versus future male reproduction. Males adopting different mate-locating strategies (perching and patrolling) in outdoor cages did not differ in spermatophore traits as was predicted from their very different flight performances. Copulations of territorial perching males took somewhat longer than copulations with non-perching males. There was a significant family effect of spermatophore size and of the expression of male mate-locating strategies suggesting heritable variation. Female traits (i.e. age and size) did not strongly affect spermatophore production. We discuss the results relative to both ultimate and proximate explanations of the complex relationships between butterfly activity, behavioural strategies, age and spermatophore production.     

The cost of mating with a non-virgin male in a monandrous butterfly: experimental evidence from the speckled wood, Pararge aegeria

Contrary to vertebrates, sperm production in insects may bear considerable costs for males. This is especially true in species that donate spermatophores containing sperm and nutrient-rich accessory gland products like in butterflies. Hence, spermatophores at first and subsequent copulations can differ in a quantitative and qualitative way. Such effects have particularly been shown in polyandrous species providing large spermatophores. Here we experimentally tested the effect of male mating status (virgin male vs recently mated male) on copulation duration, spermatophore size and females’ fitness components in a monandrous butterfly Pararge aegeria that typically donates small spermatophores. Copulations with non-virgin males lasted on average five times longer than that with virgin males and resulted in a spermatophore which was on average three times smaller. Number of eggs laid and female life span were not affected by the mating status treatment, but there was a significant effect on the number of living caterpillars a female produced, as copulations with virgin males resulted in higher numbers of larval offspring. Interestingly, the difference in spermatophore mass at the first and the second copulation increased with male body size. This suggests differential spermatophore allocation decisions among males of different size. Consequences for females and potential mechanisms influencing female fitness components are discussed. Given the small absolute size of spermatophores in P. aegeria, components other than consumable nutrients (perhaps hormones) should cause the observed effects.     

Protein content of spermatophores in relation to monandry/polyandry in butterflies

The evolution of a mating system, and specifically mating frequency, is dependent on the costs and benefits to both sexes of mating once or several times. In butterflies, males transfer a spermatophore that contains both sperm and accessory gland products. Accessory gland substances contain nutrients which, in some species, females use to increase their reproductive output and longevity. Nutrients contained in these packaged ejaculates represent investment by males in reproduction. Consequently, the nutritional composition of spermatophores may vary depending on the mating system. There are two lines of arguments concerning the evolution of the nutrient content of ejaculates. One hypothesis argues that male nuptial gifts evolved in the context of certainty of paternity and ease of finding mates; thus spermatophores of polyandrous males (with lower certainty of paternity and greater ease of finding mates) should contain less protein than those of monandrous males, since more spermatophores are produced on average. The other hypothesis argues that polyandry evolved in the context of maximization of male transfer of nutrients to females, and hence spermatophores of polyandrous males should contain more protein than those of monandrous males. In an attempt to distinguish between these two hypotheses, we determined how protein content of ejaculates varied with the degree of polyandry in nine species of pierid and two species of satyrid butterflies. We found that both relative ejaculate mass and protein content increased with the degree of polyandry. Hence our results are consistent with the view that polyandry has evolved in the context of male transfer of nutrients to females, and provides another example of a male adaptation to multiple mating in butterflies.     

Aspects of sexual biology of the anomuran crab emerita asiatica

Male Emerita asiatica Milne Edwards are smaller than females. The secondary sexual characters of males and females are described in detail. Applying the method of Wenner (1972), sex ratio is calculated. The overlap in size range between males and females is too wide to suggest sex reversal. An analysis of secondary sexual characters in post-larval juveniles during megalopa settlement corroborates the view that males and females develop separately from the megalopa. The males acquire their sexual maturity soon after metamorphosis from the megalopa stage, whereas the females attain their sexual maturity only after considerable body growth. the males continue to grow to attain a maximum carapace length of 11 mm; they do not show any tendency towards changing their sex. This is evidenced by the fact that the weight increase of the gonadal apparatus is directly related to the increase in the male carapace length (CL) and body weight. On the other hand, retention of larval characters such as short eye stalk, toothless chelae of the fifth leg (in males up to 5 mm CL), together with the very small size suggest that the males are neotenic. Only males up to 5 mm CL were found to take part in mating. During mating, as many as 5 tiny males deposit their spermatophores in the pleopodal region of one female. This type of mating assures effective transfer of spermatophores in the turbulent environment inhabited. Spermatophores are pedunculate and embedded in a gelatinous matrix to form a long ribbon. There are two types of spermatophores in the ribbon, and they differ in detail from those of other anomuran species. The spermatozoa are rod-like, with a club-shaped acrosomal head. Five spines originate from the junction between head and body. The mode of sperm release from the spermatophores is described; oviducal secretion may be responsible for opening the spermatophore.     

No cost of male mating experience on female reproductive success in the almond moth, <Emphasis Type="BoldItalic">Cadra cautella</Emphasis> (Lepidoptera; Pyralidae)

Male copulation experience may have a profound impact on female reproductive success if male reproductive investment declines over consecutive copulations and if females are unlikely to re-mate. Male reproductive investment is particularly interesting in lepidopterans because males produce dimorphic sperm: a fertilizing (eupyrene) and a non-fertilising (apyrene) sperm. In two experiments, we explored the lifetime reproductive investment of male almond moths, Cadra cautella (also known as Ephestia cautella) and examined its influence on female reproductive success. In the almond moth, females re-mate infrequently and males transfer sperm in a spermatophore. Attached to the spermatophore is a large chitinous process, the function of which is unknown. One group of males were permitted consecutive copulations with virgin females and the amount of sperm and size of the spermatophore transferred were compared for all females. We found that the number of both eupyrene and apyrene sperm per ejaculate decreased with his increased mating frequency, while the size of the spermatophore process decreased dramatically after the male’s first copulation. In a second experiment, we allowed males to mate with females throughout their lives and then compared female fecundity and fertilisation success. We found no obvious decrease in female fecundity and fertilisation success with increased male copulation experience, despite the likely reduction in male gametic investment. We discuss potential explanations for the development of this enlarged and elaborate first spermatophore of male almond moths given that it confers no clear fitness advantage to females.     

Some costs of reproduction for male bushcrickets,Requena verticalis (Orthoptera : Tettigoniidae) allocating resources to mate attraction and nuptial feeding

Summary The cost of reproductive effort is known to result in a trade-off between current and future reproduction. Similarly, trade-offs in energy allocation may occur between components of reproductive effort, mating and parental effort, within a single reproductive episode. We investigated the energy allocated to mating effort (calling to attract females) and parental effort (donation of spermatophore nutrients at mating) by male bushcrickets, Requena verticalis, under two dietary regimes. Males provided with a low quality diet reduced the daily energy allocated to calling activity while maintaining their investment in spermatophores. Males provided with a high quality diet did not allocate more resources per day to their spermatophores but stored excess energy for future reproduction. Thus, on a per day basis, males appear to hold constant their investment in the spermatophore at the cost of reduced mating effort when resources are limited. Males on both diets, however, increased the size of their spermatophore donations when the interval between female encounters was increased. One explanation for this pattern could be a frequency-dependent optimization of spermatophore size. Correspondence to: L.W. Simmons     

Sperm economy and limitation in spiny lobsters

Sperm limitation, when female fertilisation success is constrained by the supply of sperm, is generally perceived to be an uncommon feature of reproduction in species which directly transfer gametes during copulation. Male size, previous copulations, and the balance of expected reproductive return and future mating opportunity may, however, limit the amount of sperm males transfer to females. We used laboratory experiments where mate size could be manipulated and its consequences on spermatophore size and clutch size determined, to show that in two genera of spiny lobsters (Crustacea: Palinuridae) male reproductive output limits the size of clutches brooded by females. In Panulirus argus from the Florida Keys, we show that while male size affects spermatophore area, males also vary the amount of ejaculate positively with female size. Furthermore, the area of the spermatophore has a greater influence than female size on subsequent clutch weight. In Jasus edwardsii from New Zealand, female size, male size and mate order all affect clutch weight. In both species, clutches fertilised by small males in the laboratory are significantly smaller than clutches fertilised by large males. These results suggest that to ensure they receive sufficient sperm, females should either mate several times prior to oviposition, mate as early as possible in the reproductive season, or choose large, preferably unmated males as partners and thus compete with other females for preferred males. Sperm-limited female fecundity has the potential to limit the egg production of fished populations where large males are typically rare. Received: 18 May 1998 / Received in revised form: 20 November 1998 / Accepted: 30 November 1998     

Reversal and transfer of spermatophores by Octopus vulgaris and O. hummelincki

Copulatory behavior in the octopus consists of a patterned series of movements whose individual functions are not well understood. Observations and experiments on mating in Octopus vulgaris and O. hummelincki from Bimini, Bahamas and Haiti were made over a number of years in the laboratory. The study reveals how the male octopus reverses its spermatophore prior to transferring it to the female. The male’s terminal organ (penis) extrudes the sperm end of the spermatophore as it exits from Needham’s sac up to the cap or filament thread and then, holding onto the cap thread, inserts the ejaculatory end of the spermatophore into the groove of the hectocotylus, thus reversing it. It is hypothesized that the primary function of the cap or filament thread, one part of the spermatophore, is to be a handle by which the reversal and transfer is accomplished. The siphon is merely a conduit through which the terminal organ functions and plays no role whatsoever in either the reversal of the spermatophore or in its insertion into the groove. The stimulus to which the male orients to find the groove of the hectocotylus is the apex of the retracted interbrachial membrane between the third and fourth right arms, which bears a fixed spatial relationship to the origin of the groove. A number of deviations from the successful transfer process, which represent loss of sperm from the reproductive process, are illustrated. Despite these deviations, the delicately balanced integration of the various movements serves to transfer the spermatophore to the female to ensure reproductive success.     

Sex in the beach: spermatophores, dermal insemination and 3D sperm ultrastructure of the aphallic mesopsammic Pontohedyle milaschewitchii (Acochlidia, Opisthobranchia, Gastropoda)

Sperm transfer via spermatophores is common among organisms living in mesopsammic environments, and is generally considered to be an evolutionary adaptation to reproductive constraints in this habitat. However, conclusions about adaptations and trends in insemination across all interstitial taxa cannot be certain as differences in mode of insemination via spermatophores do exist, details of insemination are lacking for many species, and evolutionary relationships in many cases are poorly known. Opisthobranch gastropods typically transfer sperm via reciprocal copulation, but many mesopsammic Acochlidia are aphallic and transfer sperm via spermatophores, supposedly combined with dermal fertilisation. The present study investigates structural and functional aspects of sperm transfer in the Mediterranean microhedylacean acochlid Pontohedyle milaschewitchii. We show that spermatophore attachment is imprecise. We describe the histology and ultrastructure of the two-layered spermatophore and discuss possible functions. Using DAPI staining of the (sperm-) nuclei, we document true dermal insemination in situ under the fluorescence microscope. Ultrastructural investigation and computer-based 3D reconstruction from TEM sections visualise the entire spermatozoon including the exceptionally elongate, screw-like keeled sperm nucleus. An acrosomal complex was not detected. From their special structure and behaviour we conclude that sperm penetrate epithelia, tissues and cells mechanically by drilling rather than lysis. Among opisthobranchs, dermal insemination is limited to mesopsammic acochlidian species. In this spatially limited environment, a rapid though imprecise and potentially harmful dermal insemination is discussed as a key evolutionary innovation that could have enabled the species diversification of microhedylacean acochlidians.     

Spermatophore characteristics in bushcrickets vary with parasitism and remating interval

Male bushcrickets provide females with a nuptial gift, a spermatophore, which is transferred to females at mating. The spermatophore consists of a gelatinous mass, the spermatophylax, and the sperm-containing ampulla. Male spermatophore size is positively correlated with insemination rate and female refractory period and therefore with male reproductive success. In this study, we examined spermatophylax weight, ampulla weight and sperm number in males of Poecilimon mariannae parasitized by the acoustically orienting fly Therobia leonidei. We show that in parasitized males, spermatophylax weight decreases with the level of parasitism. In line with the hypothesis that parasitism is a cost to reproduction, we found that spermatophylax weight was reduced at remating. In contrast, the replenishment of the spermatophylax in unparasitized males was complete after 2 days and was increased no further after 3 days. Both sperm number and ampulla weight showed an increase over time since last mating and sperm production was estimated at a constant rate of 500,000 per day in all individuals, regardless of parasitism. The allocation of investment in components of the spermatophore varies greatly with parasitism and remating. Both factors had rather independent effects on spermatophore constitution, revealing functional constraints acting on spermatophore characteristics in bushcrickets, which are important for understanding the selection pressures working on its components. Received: 13 September 1999 / Received in revised form: 4 February 2000 / Accepted: 13 March 2000     

Different ejaculate delivery strategies in first versus subsequent matings in the swallowtail butterfly Papilio machaon L.

Summary Evidence suggests that males incur costs in producing ejaculates. Therefore some discrimination should be expected with respect to their pattern of allocating ejaculates. In a laboratory study, males of the European swallowtail butterfly Papilio machaon were mated with conspecific virgin females up to six times. The weight of the spermatophore and of the accessory substances delivered by males, the duration of mating, and the number of sperm in ejaculates were measured. The weight of the first spermatophore was related to male weight but that of accessory substances was not. The weight of the spermatophore delivered by males upon remating was always less than that of the first spermatophore. Second spermatophores Produced by males were smallest when delivered on the day after the first mating and increased asymptotically to about half the initial size with time elapsed since the first mating. Copula duration was highest for males remating on the day after the first mating but decreased to the duration of the first mating for males that were remated on the third day after the first mating or later. The weight of accessory substances delivered by males and the number of sperm in ejaculates were also higher in first matings compared to all subsequent matings. Both of two males that were handpaired on three consecutive days after the first mating produced a cumulative ejaculate mass equalling that transferred at the first mating. Thus the fact that males delivered smaller ejaculates upon remating on the third day after the first mating or later requires explanation, especially in view of the fact that copulations were not prolonged. We argue that the male ejaculate delivery strategy in mildly polyandrous species should be to mate for the first time with virgin females, when a maximum size ejaculate should be delivered, and then to maximize the number of matings. The rationale behind the male shift towards smaller ejaculate sizes delivered at subsequent matings is that: (1) the size of the ejaculate that can be produced is dependent on the time elapsed since the last mating; (2) the abundance of virgin females decreases as the flight season proceeds; and (3) there is an inverse relationship between the size and number of ejaculates that a male can produce in his life-time.     

Male control of sperm transfer dynamics in a spermatophore-donating bushcricket

In many animal species, male and female interests often differ when it comes to decisions over mating and fertilization. However, it is intrinsically difficult to determine the degree to which males and females exert control over the various processes that determine the outcome of reproductive interactions, and thus to predict how such conflicts will be resolved. For example, in species where sperm are transferred to females via a spermatophore or other external sperm packaging device, it is unclear which sex determines subsequent sperm transfer dynamics to the female reproductive tract. To address this question, we used a reciprocal cross experimental design in a bushcricket species (Poecilimon veluchianus) comprising two subspecies differing in the dynamics of sperm transfer. The reciprocal crosses show that in these bushcrickets the timing of sperm transfer in inter-subspecies crosses closely resemble those typical of the subspecies of the male partner, indicating that it is the properties of the spermatophore rather than its handling by the female that determine sperm transfer dynamics. There was neither a significant female influence nor any indication of an interaction between males and females with regard to the number of sperm transferred after a set interval. Our study suggests that males rather than females appear to control the timing of the insemination process in this species.     

Courtship solicitation by females of the checkered white butterfly,Pieris protodice

Summary The willingness of some females of the checkered white butterfly to approach and chase conspecifies was studied in the field. Both virgin and mated females were observed approaching and chasing conspecifics. Such responses to males result in the solicitation or continuation of male courtship attempts, which in turn may lead to copulation.Mated females that approached conspecifics had significantly smaller spermatophores than females in a random sample from the same population.Females that approached tethered animals spent more time chasing tethered males than tethered females. Females with small spermatophores spent more time chasing tethered males than did females with larger spermatophores.These results confirm the hypothesis that mated females that approach and chase conspecifics are doing so in an effort to solicit male courtship attempts and to potentially receive a fresh spermatophore. The selective forces favoring courtship solicitation by females are discussed.     

Role-reversal in katydids: Habitat influences reproductive behaviour (Orthoptera: Tettigoniidae,Metaballus sp.)

Summary Mating behaviour of the katydid Metaballus sp. varies. At sites QS and DB (in 1982) females competed for access to calling males and males chose mates by mating with heavier, more fecund females. At another site (BR) there was no evidence of role-reversal in reproductive behaviour, and males were observed to compete for mates. This species has a large spermatophore, a product of male reproductive glands, that is eaten by the female after mating. Males at the DB site had small reproductive glands. This suggests that some aspect of the QS and DB environments decreases spermatophore production; spermatophores become a limiting resource for females resulting in the reversal in reproductive roles observed at these sites. A field experiment that involved moving individuals from site BR to QS in 1983 determined that mating system was influerced by site (Table 1). At BR, males produced a continuous calling song, a third of the males observed attracted mates, and called for about 30 min before the female arrived; courtship duration was short. Males that were moved from BR to QS encountered a higher density of receptive females as all males attracted females after an average of just 3 min of calling. They changed their behaviour by producing short periodic bursts of song (zipping), and by courting females for long periods of time. The long courtship period may function as as a mate-assessment period for males. The reproductive behaviour of BR males moved to QS differed from that of native QS males only in the length of time spent in copula.     

The effects of gregarine parasites,body size,and time of day on spermatophore production and sexual selection in field crickets

Summary In laboratory experiments measuring the rate of spermatophore production in the field crickets Gryllus veletis and G. pennsylvanicus by confining single males with a conspecific female, 0–10 spermatophores were produced by each male within 24 h. The number of spermatophores produced was unrelated to a male's body size, but was significantly negatively correlated with the natural levels of gregarines, a protozoan gut parasite, in the males. Spermatophore production in the laboratory peaked between 0600 and 1000 h, as did the proportion of courtship songs given by male crickets in the field, suggesting that mating may occur more frequently in the morning. When single males were placed in jars with two conspecific females, 49% transferred spermatophores to both females, whereas 51% of males only gave spermatophores to one of the females. The results support the hypothesis that parasites are important in sexual selection, and are used to propose a new interpretation of post-copulatory guarding in crickets.     

The ultimate function of nuptial feeding in the bushcricket Poecilimon veluchianus (Orthoptera : Tettigoniidae : Phaneropterinae)

Summary During mating the males of the bushcricket Poecilimon veluchianus transfer a large spermatophore of about a quarter of their body weight to the female. Such nuptial feeding is often thought to function as paternal investment by increasing the fitness of the male's offspring. According to an alternative, though not mutually exclusive, hypothesis, the size of the spermatophore is maintained because of its function as a sperm protection device. In this case the cost to the male should be classified as mating effort. To discriminate between these two hypotheses we measured the duration of sperm transfer into the female spermatheca and the time taken for spermatophore consumption. A comparison of durations revealed that spermatophore consumption interferes with the process of sperm transfer (Fig. 4). There was no significant effect of spermatophore consumption on number of eggs laid, weight of eggs or absolute weight of hatched larvae. The relative dry weight of hatched larvae, however, was increased as a result of spermatophore consumption (Table 1). Thus spermatophylax size is adjusted in accordance with a sperm protection function and the spermatophylax therefore represents mating effort. The increase in relative dry weight indicates that there may also be a paternal investment effect of the spermatophylax, if the offspring that benefit from spermatophylax materials are fathered by the donating male. Correspondence to: K. Reinhold     

Fitness benefits of multiple mating versus female mate choice in the cellar spider (Pholcus phalangioides)

Recent studies have demonstrated that mating with multiple males can be beneficial for females and her offspring even if males contribute nothing but sperm. This was mainly established for species in which sperm from several males mix in the reproductive tract of the female, thus allowing sperm competition and/or female sperm choice. However, in species with last male sperm precedence, female re-mating decides against the previous male by strongly limiting his reproductive success. We tested the effect of female re-mating behaviour using the cellar spider Pholcus phalangioides, which shows strong last males sperm precedence and moderate levels of polyandry under natural situations. We predicted that females prevented from remating even though they are receptive would show reduced reproductive success compared to females that accept two copulations and females that reject a second male, since the latter two treatments were allowed to behave according to their decisions. However, if the number of matings per se had an effect on oviposition or on offspring performance, double-mated females should perform better compared to both treatments of once-mated females. We measured female fecundity and fertility over a period of 140 days, comparable to the species' natural reproductive peak season. Two thousand one hundred and fifty-two offspring from 67 first egg sacs were reared under two feeding levels. We registered development time and survival, and measured offspring adult size and mass. We found a positive effect of double mating, as in this treatment, oviposition probability was higher compared to the other treatments. Interestingly, adult female offspring of the DM treatment that were raised under low food level had a higher condition index compared to those from FS and RM, but development time, size and mass at adulthood were not affected by mating treatment. Female choice only seemed to affect hatching latency of the offspring. Overall, the main predictor of female reproductive output and success was female body size.     

Ventilation rate and copulation in Octopus vulgaris

Typically three events occur when Octopus vulgaris Cuvier mates. The male inserts its hectocotylus into the mantle cavity of the female; the hectocotylus and the bodies of the male and female become quiescent; and the female increases its ventilation rate about 2.5 times, on the average, above its precopulatory base rate. It is hypothesized that the increased ventilation rate of the female provides the stimulus to the male to transfer its spermatophore to the oviduct. Since the insertion of the hectocotylus and the female's increased ventilation rate may be dissociated, it is suggested that copulation be defined as the attachment of the hectocotylus to the oviduct. A conditioned ventilation-rate increase in the female was observed with repeated testing. Observation of arching and pumping movements of the male as well as measurements of its ventilation rate before and during copulation indicate that only a few spermatophores are transferred per copulation.     

Sperm precedence, mating interval, and a novel mechanism of paternity bias in a beetle (Tenebrio molitor L.)

When females mate with more than one male, the ensuing sperm competition leads to the evolution of male mechanisms that skew paternity. Males of the yellow mealworm beetle (Tenebrio molitor) transfer a spermatophore to females during copulation, but sperm release and storage occur later. We investigated how the interval between two matings with different males affects sperm precedence by varying the interval between the copulations so that the second mating was either: (1) before sperm release from the first spermatophore (<5 min); (2) after sperm release but before spermatophore ejection (15–20 min); (3) after spermatophore ejection but before sperm storage (4 h), or (4) after complete sperm storage (24 h). We collected offspring over a period of 2 weeks and determined paternity by protein electrophoresis. There was second-male sperm precedence in all treatments, but when the interval was <5 min, the second male usually (86% of cases) had complete sperm precedence (i.e., P ₂=1). Investigations into the mechanism of second-male sperm precedence during <5-min mating intervals indicate that sperm release from the first spermatophore is inhibited, a phenomenon which has not been previously documented. Received: 31 January 2000 / Revised: 9 June 2000 / Accepted: 26 August 2000     

Competition,fluctuating asymmetry and sperm transfer in male gryllid crickets (<Emphasis Type="Italic">Gryllus bimaculatus</Emphasis> and <Emphasis Type="Italic">Gryllodes sigillatus</Emphasis>)

Sperm competition theory predicts that ejaculate expenditure by males should increase with the risk of competition from rivals. The tendency to modulate expenditure, however, may reflect various constraints acting on different individuals. We test this idea here by looking at the effects of competition and fluctuating asymmetry (FA) in two species of gryllid cricket. Male Gryllus bimaculatus and Gryllodes sigillatus modulated the sperm content of their spermatophores in relation to both apparent competition and FA in limb size. Males of both species increased the amount of sperm transferred when an apparent male competitor was present, sperm number increasing regardless of the species of apparent competitor in G. bimaculatus, but significantly more with a conspecific competitor in G. sigillatus. In addition, the relative size of the spermatophylax in G. sigillatus increased in the presence of apparent competition, while spermatophore transfer time decreased in G. bimaculatus. Sperm number showed opposite relationships with limb asymmetry in the two species, decreasing with increasing asymmetry in males in G. sigillatus, but increasing with asymmetry in both males and females in G. bimaculatus. G. bimaculatus, but not G. sigillatus, also transferred more sperm when paired with a larger female. Relationships between reproductive measures and limb asymmetry were significant only when competitors were present in G. bimaculatus, but were significant regardless of competition in G. sigillatus. The differences may reflect the additional burden of producing a spermatophylax in G. sigillatus.